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Artificial intelligence (AI) and machine learning (ML) present revolutionary opportunities to enhance our understanding of animal behaviour and conservation strategies. Using elephants, a crucial species in Africa and Asia's protected areas, as our focal point, we delve into the role of AI and ML in their conservation. Given the increasing amounts of data gathered from a variety of sensors like cameras, microphones, geophones, drones and satellites, the challenge lies in managing and interpreting this vast data. New AI and ML techniques offer solutions to streamline this process, helping us extract vital information that might otherwise be overlooked. This paper focuses on the different AI-driven monitoring methods and their potential for improving elephant conservation. Collaborative efforts between AI experts and ecological researchers are essential in leveraging these innovative technologies for enhanced wildlife conservation, setting a precedent for numerous other species.

There are powerful tools for modelling swarms that have strong spatial structures like flocks of birds, schools of fish and formations of drones, but relatively little work on developing formalisms for other swarm structures like hub-based colonies doing foraging, maintaining a nest or selecting a new nest site. We present a method for finding low-dimensional representations of swarm state for simulated homogeneous hub-based colonies solving the best-of-N problem. The embeddings are obtained from latent representations of convolution-based graph neural network architectures and have the property that swarm states which have similar performance have very similar embeddings. Such embeddings are used to classify swarm state into binned estimates of success probability and time to completion. We demonstrate how embeddings can be obtained in a sequence of experiments that progressively require less information, which suggests that the methods can be extended to larger swarms in more complicated environments. This article is part of the theme issue 'The road forward with swarm systems'. [ABSTRACT FROM AUTHOR]

Ageing affects almost all aspects of life and therefore is an important process across societies, human and non-human animal alike. This article introduces new research exploring the complex interplay between individual-level ageing and demography, and the consequences this interplay holds for the structure and functioning of societies across various natural populations. We discuss how this Special Issue provides a foundation for integrating perspectives from evolutionary biology, behavioural ecology and demography to provide new insights into how ageing shapes individuals' social behaviour and social associations, and how this in turn impacts social networks, social processes (such as disease or information transfer) and fitness. Through examining these topics across taxa, from invertebrates to birds and mammals, we outline how contemporary studies are using natural populations to advance our understanding of the relationship between age and society in innovative ways. We highlight key emerging research themes from this Special Issue, such as how sociality affects lifespan and health, the genetic and ecological underpinnings of social ageing and the adaptive strategies employed by different species. We conclude that this Special Issue underscores the importance of studying social ageing using diverse systems and interdisciplinary approaches for advancing evolutionary and ecological insights into both ageing and sociality more generally. This article is part of the discussion meeting issue 'Understanding age and society using natural populations '. [ABSTRACT FROM AUTHOR]

The age of individuals has consequences not only for their fitness and behaviour but also for the functioning of the groups they form. Because social behaviour often changes with age, population age structure is expected to shape the social organization, the social environments individuals experience and the operation of social processes within populations. Although research has explored changes in individual social behaviour with age, particularly in controlled settings, there is limited understanding of how age structure governs sociality in wild populations. Here, we synthesize previous research into age-related effects on social processes in natural populations, and discuss the links between age structure, sociality and ecology, specifically focusing on how population age structure might influence social structure and functioning. We highlight the potential for using empirical data from natural populations in combination with social network approaches to uncover pathways linking individual social ageing, population age structure and societal functioning. We discuss the broader implications of these insights for understanding the social impacts of anthropogenic effects on animal population demography and for building a deeper understanding of societal ageing in general. This article is part of the discussion meeting issue 'Understanding age and society using natural populations'. [ABSTRACT FROM AUTHOR]

In long-lived organisms, experience can accumulate with age, such that older individuals may act as repositories of ecological and social knowledge. Such knowledge is often beneficial and can spread via social transmission, leading to the expectation that ageing individuals will remain socially well-integrated. However, social ageing involves multiple processes that modulate the relationship between age and social connectivity in complex ways. We developed a generative model to explore how social ageing may drive changes in social network position and shape older individuals' capacity to transmit knowledge to others. We further employ novel hypernetwork analyses that capture higher-order interactions (i.e. involving ≥ 3 participants) to reveal potential relationships between age and sociality that conventional dyadic networks may overlook. We find that older individuals in our simulations effectively facilitate transmission across a range of scenarios, especially when transmission resembles a complex contagion or when social selectivity (i.e. prioritization of key relationships) rapidly emerges with age. These patterns result from the formation of tight-knit sets of older associates that co-occur in multiple groups, thereby reinforcing one another's capacity to transmit knowledge. Our findings suggest key avenues for future empirical work and illustrate the use of hypernetworks in advancing the study of social behaviour. This article is part of the discussion meeting issue 'Understanding age and society using natural populations'. [ABSTRACT FROM AUTHOR]

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'. [ABSTRACT FROM AUTHOR]

To benefit from group living, individuals need to maintain cohesion and coordinate their activities. Effective communication thus becomes critical, facilitating rapid coordination of behaviours and reducing consensus costs when group members have differing needs and information. In many bird and mammal species, collective decisions rely on acoustic signals in some contexts but on movement cues in others. Yet, to date, there is no clear conceptual framework that predicts when decisions should evolve to be based on acoustic signals versus movement cues. Here, we first review how acoustic signals and movement cues are used for coordinating activities. We then outline how information masking, discrimination ability (Weber's Law) and encoding limitations, as well as trade-offs between these, can identify which types of collective behaviours likely rely on acoustic signals or movement cues. Specifically, our framework proposes that behaviours involving the timing of events or expression of specific actions should rely more on acoustic signals, whereas decisions involving complex choices with multiple options (e.g. direction and destination) should generally use movement cues because sounds are more vulnerable to information masking and Weber's Law effects. We then discuss potential future avenues of enquiry, including multimodal communication and collective decision-making by mixed-species animal groups. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamic'. [ABSTRACT FROM AUTHOR]

Cooperation may emerge from intrinsic factors such as social structure and extrinsic factors such as environmental conditions. Although these factors might reinforce or counteract each other, their interaction remains unexplored in animal populations. Studies on multilevel societies suggest a link between social structure, environmental conditions and individual investment in cooperative behaviours. These societies exhibit flexible social configurations, with stable groups that overlap and associate hierarchically. Structure can be seasonal, with upper-level units appearing only during specific seasons, and lower-level units persisting year-round. This offers an opportunity to investigate how cooperation relates to social structure and environmental conditions. Here, we study the seasonal multilevel society of superb fairy-wrens (Malurus cyaneus), observing individual responses to experimental playback of conspecific distress calls. Individuals engaged more in helping behaviour and less in aggressive/territorial song during the harsher non-breeding season compared to the breeding season. The increase in cooperation was greater for breeding group members than for members of the same community, the upper social unit, comprised of distinct breeding groups in association. Results suggest that the interaction between social structure and environmental conditions drives the seasonal switch in cooperation, supporting the hypothesis that multilevel societies can emerge to increase cooperation during harsh environmental conditions. [ABSTRACT FROM AUTHOR]

Repeated use of the same areas may benefit animals as they exploit familiar sites, leading to consistent home ranges over time that can span generations. Changing risk landscapes may reduce benefits associated with home range fidelity, however, and philopatric animals may alter movement in response to new pressures. Despite the importance of range changes to ecological and evolutionary processes, little tracking data have been collected over the long-term nor has range change been recorded in response to human pressures across generations. Here, we investigate the relationships between ecological, demographic and human variables and elephant ranging behaviour across generations using 16 years of tracking data from nine distinct female social groups in a population of elephants in northern Kenya that was heavily affected by ivory poaching during the latter half of the study. Nearly all groups--including those that did not experience loss of mature adults--exhibited a shift north over time, apparently in response to increased poaching in the southern extent of the study area. However, loss of mature adults appeared to be the primary indicator of range shifts and expansions, as generational turnover was a significant predictor of range size increases and range centroid shifts. Range expansions and northward shifts were associated with higher primary productivity and lower poached carcass densities, while westward shifts exhibited a trend to areas with higher values of primary productivity and higher poached carcass densities relative to former ranges. Together these results suggest a trade-off between resource access, mobility and safety. We discuss the relevance of these results to elephant conservation efforts and directions meriting further exploration in this disrupted society of a keystone species. [ABSTRACT FROM AUTHOR]

Social instability frequently arises in group-living species, but the potential costs have rarely been investigated in free-living cooperative breeders, especially across different timeframes. Using natural observations, body mass measurements and life-history data from dwarf mongooses (Helogale parvula), we determined the short- and long-term consequences of a change in one of the dominant breeding pairs. We found that a new breeder led to alterations in both collective and individual behaviours (i.e. increases in communal scent-marking, engagement in intergroup interactions, sentinel activity and within-group grooming), as well as reduced body mass gain, further demographic changes and decreased reproductive success (i.e. fewer pups surviving to adulthood). The effects were particularly apparent when it was the female breeder who changed; new female breeders were younger than more experienced counterparts. Our findings support the idea that stability and cooperation are strongly linked and provide potential reasons for previously documented health and fitness benefits of social stability. [ABSTRACT FROM AUTHOR]

Africa is experiencing extensive biodiversity loss due to rapid changes in the environment, where natural resources constitute the main instrument for socioeconomic development and a mainstay source of livelihoods for an increasing population. Lack of data and information deficiency on biodiversity, but also budget constraints and insufficient financial and technical capacity, impede sound policy design and effective implementation of conservation and management measures. The problem is further exacerbated by the lack of harmonized indicators and databases to assess conservation needs and monitor biodiversity losses. We review challenges with biodiversity data (availability, quality, usability and database access) as a key limiting factor that impacts funding and governance. We also evaluate the drivers of both ecosystems change and biodiversity loss as a central piece of knowledge to develop and implement effective policies. While the continent focuses more on the latter, we argue that the two are complementary in shaping restoration and management solutions. We thus underscore the importance of establishing monitoring programmes focusing on biodiversity–ecosystem linkages in order to inform evidence-based decisions in ecosystem conservation and restoration in Africa. This article is part of the theme issue 'Detecting and attributing the causes of biodiversity change: needs, gaps and solutions'. [ABSTRACT FROM AUTHOR]

Despite decades of evidence revealing a multitude of ways in which animals are adapted to minimize the energy cost of locomotion, little is known about how energy expenditure shapes adaptive gait over complex terrain. Here, we show that the principle of energy optimality in human locomotion can be generalized to complex task-level locomotor behaviours requiring advance decision-making and anticipatory control. Participants completed a forced-choice locomotor task requiring them to choose between discrete multi-step obstacle negotiation strategies to cross a 'hole' in the ground. By modelling and analysing mechanical energy cost of transport for preferred and non-preferred manoeuvres over a wide range of obstacle dimensions, we showed that strategy selection was predicted by relative energy cost integrated across the complete multi-step task. Vision-based remote sensing was sufficient to select the strategy associated with the lowest prospective energy cost in advance of obstacle encounter, demonstrating the capacity for energetic optimization of locomotor behaviour in the absence of online proprioceptive or chemosensory feedback mechanisms. We highlight the integrative hierarchic optimizations that are required to facilitate energetically efficient locomotion over complex terrain and propose a new behavioural level linking mechanics, remote sensing and cognition that can be leveraged to explore locomotor control and decision-making. [ABSTRACT FROM AUTHOR]

Planning for the future is a complex skill that is often considered uniquely human. This cognitive ability has never been investigated in wild gibbons (Hylobatidae). Here we evaluated the movement patterns from sleeping trees to out-of-sight breakfast trees in two groups of endangered skywalker gibbons (Hoolock tianxing). These Asian apes inhabit a cold seasonal montane forest in southwestern China. After controlling for possible confounding variables including group size, sleeping pattern (sleep alone or huddle together), rainfall and temperature, we found that food type (fruits or leaves) of the breakfast tree was the most important factor affecting gibbon movement patterns. Fruit breakfast trees were more distant from sleeping trees compared with leaf trees. Gibbons left sleeping trees and arrived at breakfast trees earlier when they fed on fruits compared with leaves. They travelled fast when breakfast trees were located further away from the sleeping trees. Our study suggests that gibbons had foraging goals in mind and plan their departure times accordingly. This ability may reflect a capacity for route-planning, which would enable them to effectively exploit highly dispersed fruit resources in high-altitude montane forests. [ABSTRACT FROM AUTHOR]

Ivory poaching continues to threaten African elephants. We (1) used criminology theory and literature evidence to generate hypotheses about factors that may drive, facilitate or motivate poaching, (2) identified datasets representing these factors, and (3) tested those factors with strong hypotheses and sufficient data quality for empirical associations with poaching. We advance on previous analyses of correlates of elephant poaching by using additional poaching data and leveraging new datasets for previously untested explanatory variables. Using data on 10 286 illegally killed elephants detected at 64 sites in 30 African countries (2002–2020), we found strong evidence to support the hypotheses that the illegal killing of elephants is associated with poor national governance, low law enforcement capacity, low household wealth and health, and global elephant ivory prices. Forest elephant populations suffered higher rates of illegal killing than savannah elephants. We found only weak evidence that armed conflicts may increase the illegal killing of elephants, and no evidence for effects of site accessibility, vegetation density, elephant population density, precipitation or site area. Results suggest that addressing wider systemic challenges of human development, corruption and consumer demand would help reduce poaching, corroborating broader work highlighting these more ultimate drivers of the global illegal wildlife trade. [ABSTRACT FROM AUTHOR]

The science guiding design and evaluation of restoration interventions in tropical landscapes is dominated by ecological processes and outcomes and lacks indicators and methods that integrate human wellbeing into the restoration process. We apply a new systems approach framework for tree restoration in forest-agricultural landscapes to showhowthis shortcoming can be addressed. Demonstrating 'proof of concept', we tested statistical models underlying the framework pathways with data collected from a case study in Tanzania. Local community perceptions of nature's values were not affected by levels of self-reported wildlife-induced crop damage. But mapped predictions from the systems approach under a tree restoration scenario suggested differential outcomes for biodiversity indicators and altered spatial patterns of crop damage risk, expected to jeopardize human wellbeing. The predictions map anticipated trade-offs in costs and benefits of restoration scenarios, which we have started to explore with stakeholders to identify restoration opportunities that consider local knowledge, value systems and human wellbeing. We suggest that the framework be applied to other landscapes to identify commonalities and differences in forest landscape restoration outcomes under varying governance and land use systems. This should form a foundation for evidence-based implementation of the global drive for forest landscape restoration, at local scales. [ABSTRACT FROM AUTHOR]

The Cretaceous–Palaeogene (K-Pg) extinction of the non-avian dinosaurs (66 Ma) led to a 25 million year gap of megaherbivores (>1000 kg) before the evolution of megaherbivorous mammals in the Late Eocene (40 Ma). The botanical consequences of this 'Palaeocene megaherbivore gap' (PMHG) remain poorly explored. We hypothesize that the absence of megaherbivores should result in changes in the diversification and trait evolution of associated plant lineages. We used phylogenetic time- and trait-dependent diversification models with palms (Arecaceae) and show that the PMHG was characterized by speciation slowdowns, decreased evolution of armature and increased evolution of megafaunal (≥4 cm) fruits. This suggests that the absence of browsing by megaherbivores during the PMHG may have led to a loss of defence traits, but the absence of megaherbivorous seed dispersers did not lead to a loss of megafaunal fruits. Instead, increases in PMHG fruit sizes may be explained by simultaneously rising temperatures, rainforest expansion, and the subsequent radiation of seed-dispersing birds and mammals. We show that the profound impact of the PMHG on plant diversification can be detected even with the overwriting of adaptations by the subsequent Late Eocene opening up of megaherbivore-associated ecological opportunities. Our study provides a quantitative, comparative framework to assess diversification and adaptation during one of the most enigmatic periods in angiosperm history. [ABSTRACT FROM AUTHOR]

Human encroachment into natural habitats is typically followed by conflicts derived from wildlife damage to agriculture and livestock. Spatial risk modelling is a useful tool to gain the understanding of wildlife damage and mitigate conflicts. Although resource selection is a hierarchical process operating at multiple scales, risk models usually fail to address more than one scale, which can result in the misidentification of the underlying processes. Here, we addressed the multi-scale nature of wildlife damage occurrence by considering ecological and management correlates interacting from household to landscape scales. We studied brown bear (Ursus arctos) damage to apiaries in the North-eastern Carpathians as our model system. Using generalized additive models, we found that brown bear tendency to avoid humans and the habitat preferences of bears and beekeepers determine the risk of bear damage at multiple scales. Damage risk at fine scales increased when the broad landscape context also favoured damage. Furthermore, integrated-scale risk maps resulted in more accurate predictions than single-scale models. Our results suggest that principles of resource selection by animals can be used to understand the occurrence of damage and help mitigate conflicts in a proactive and preventive manner. [ABSTRACT FROM AUTHOR]

African elephants (Loxodonta africana) use many sensory modes to gather information about their environment, including the detection of seismic, or ground-based, vibrations. Seismic information is known to include elephant-generated signals, but also potentially encompasses biotic cues that are commonly referred to as 'noise'. To investigate seismic information transfer in elephants beyond communication, here we tested the hypothesis that wild elephants detect and discriminate between seismic vibrations that differ in their noise types, whether elephant- or human-generated. We played three types of seismic vibrations to elephants: seismic recordings of elephants (elephant-generated), white noise (human-generated) and a combined track (elephant- and human-generated). We found evidence of both detection of seismic noise and discrimination between the two treatments containing human-generated noise. In particular, we found evidence of retreat behaviour, where seismic tracks with human-generated noise caused elephants to move further away from the trial location. We conclude that seismic noise are cues that contain biologically relevant information for elephants that they can associate with risk. This expands our understanding of how elephants use seismic information, with implications for elephant sensory ecology and conservation management. [ABSTRACT FROM AUTHOR]

Social structure is a fundamental aspect of animal populations. In order to understand the function and evolution of animal societies, it is important to quantify how individual attributes, such as age and sex, shape social relationships. Detecting these influences in wild populations under natural conditions can be challenging, especially when social interactions are difficult to observe and broad-scale measures of association are used as a proxy. In this study, we use unoccupied aerial systems to observe association, synchronous surfacing, and physical contact within a pod of southern resident killer whales (Orcinus orca). We show that interactions do not occur randomly between associated individuals, and that interaction types are not interchangeable. While age and sex did not detectably influence association network structure, both interaction networks showed significant social homophily by age and sex, and centrality within the contact network was higher among females and young individuals. These results suggest killer whales exhibit interesting parallels in social bond formation and social life histories with primates and other terrestrial social mammals, and demonstrate how important patterns can be missed when using associations as a proxy for interactions in animal social network studies. [ABSTRACT FROM AUTHOR]

Protecting wild places is conservation's most pressing task given rapid contemporary declines in biodiversity and massive land use changes. We suggest that behavioural ecology has a valuable, albeit limited, role to play in this agenda. Behaviourally based empiricism and modelling, especially of animal movements and habitat preferences have enjoyed wide applicability in delineating reserve boundaries. In protected areas that sanction exploitation, it may also be important to understand individuals' behavioural and life-history responses to management decisions. We also argue, however, that the in-depth studies of behavioural ecologists may have an important role in conservation by elevating species' status from mundane to charismatic and often sparking public empathy, and their mere presence in field generates local (or broader) intrigue. More generally behavioural ecologists will only be listened to, and their contributions considered of conservation importance, if they become more involved in decision-making processes as witnessed by several prominent examples that have supported the establishment of protected areas. This article is part of the theme issue 'Linking behaviour to dynamics of populations and communities: application of novel approaches in behavioural ecology to conservation'. [ABSTRACT FROM AUTHOR]

Wildlife tracking is one of the most frequently employed approaches to monitor and study wildlife populations. To date, the application of tracking data to applied objectives has focused largely on the intensity of use by an animal in a location or the type of habitat. While this has provided valuable insights and advanced spatial wildlife management, such interpretation of tracking data does not capture the complexity of spatio-temporal processes inherent to animal behaviour and represented in the movement path. Here, we discuss current and emerging approaches to estimate the behavioural value of spatial locations using movement data, focusing on the nexus of conservation behaviour and movement ecology that can amplify the application of animal tracking research to contemporary conservation challenges. We highlight the importance of applying behavioural ecological approaches to the analysis of tracking data and discuss the utility of comparative approaches, optimization theory and economic valuation to gain understanding of movement strategies and gauge population-level processes. First, we discuss innovations in the most fundamental movement-based valuation of landscapes, the intensity of use of a location, namely dissecting temporal dynamics in and means by which to weight the intensity of use. We then expand our discussion to three less common currencies for behavioural valuation of landscapes, namely the assessment of the functional (i.e. what an individual is doing at a location), structural (i.e. how a location relates to use of the broader landscape) and fitness (i.e. the return from using a location) value of a location. Strengthening the behavioural theoretical underpinnings of movement ecology research promises to provide a deeper, mechanistic understanding of animal movement that can lead to unprecedented insights into the interaction between landscapes and animal behaviour and advance the application of movement research to conservation challenges. This article is part of the theme issue 'Linking behaviour to dynamics of populations and communities: application of novel approaches in behavioural ecology to conservation'. [ABSTRACT FROM AUTHOR]

The rarity of female-biased kinship organization in human societies raises questions about ancestral hominin family structures. Such questions require grounding in the form and function of kin relationships in our close phylogenetic relatives, the non-human primates. Common features of primate societies, such as low paternity certainty and lack of material wealth, are consistent with features that promote matriliny in humans. In this review, I examine the role of kinship in three primate study systems (socially monogamous species, female-bonded cercopithecines and great apes) that, each for different reasons, offer insights into the evolutionary roots of matriliny. Using these and other examples, I address potential analogues to features of female-biased kinship organization, including residence, descent and inheritance. Social relationships are biased towards matrilineal kin across primates, even where female dispersal limits access to them. In contrast to the strongly intergenerational nature of human kinship, most primate kin relationships function laterally as the basis for cooperative networks and require active reinforcement. There is little evidence that matrilineal kin relationships in primates are functionally equivalent to descent or true inheritance, but further research is needed to understand whether human cultural constructs of kinship produce fundamentally different biological outcomes from their antecedents in primates. This article is part of the theme issue 'The evolution of female-biased kinship in humans and other mammals'. [ABSTRACT FROM AUTHOR]

Poaching fuelled by international trade in horn caused the deaths of over 1000 African rhinoceros (Ceratotherium simum and Diceros bicornis) per year between 2013 and 2017. Deterrents, which act to establish avoidance behaviours in animals, have the potential to aid anti-poaching efforts by moving at-risk rhinos away from areas of danger (e.g. near perimeter fences). To evaluate the efficacy of deterrents, we exposed a population of southern white rhinos (C. simum simum) to acoustic- (honeybee, siren, turtle dove), olfactory- (chilli, sunflower), and drone-based stimuli on a game reserve in South Africa. We exposed rhinos to each stimulus up to four times. Stimuli were considered effective deterrents if they repeatedly elicited avoidance behaviour (locomotion away from the deterrent). Rhinos travelled significantly further in response to the siren than to the honeybee or turtle dove stimulus, and to low-altitude drone flights than to higher altitude flights. We found the drone to be superior at manipulating rhino movement than the siren owing to its longer transmission range and capability of pursuit. By contrast, the scent stimuli were ineffective at inciting avoidance behaviour. Our findings indicate that deterrents are a prospective low-cost and in situ method to manage rhino movement in game reserves. [ABSTRACT FROM AUTHOR]

Many animal groups exhibit signatures of persistent internal modular structure, whereby individuals consistently interact with certain groupmates more than others. In such groups, information relevant to a collective decision may spread unevenly through the group, but how this impacts the quality of the resulting decision is not well understood. Here, we explicitly model modularity within animal groups and examine how it affects the amount of information represented in collective decisions, as well as the accuracy of those decisions. We find that modular structure necessarily causes a loss of information, effectively silencing the input from a fraction of the group. However, the effect of this information loss on collective accuracy depends on the informational environment in which the decision is made. In simple environments, the information loss is detrimental to collective accuracy. By contrast, in complex environments, modularity tends to improve accuracy. This is because small group sizes typically maximize collective accuracy in such environments, and modular structure allows a large group to behave like a smaller group (in terms of its decision-making). These results suggest that in naturalistic environments containing correlated information, large animal groups may be able to exploit modular structure to improve decision accuracy while retaining other benefits of large group size. This article is part of the theme issue 'Liquid brains, solid brains: How distributed cognitive architectures process information'. [ABSTRACT FROM AUTHOR]

In this paper, a mechanism called proximity inheritance is introduced in the birth–death process of a networked population involving the Prisoner's Dilemma game. Different from the traditional birth–death process, in the proposed model, players are distributed in a spatial space and offspring is distributed in the neighbourhood of its parents. That is, offspring inherits not only the strategy but also the proximity of its parents. In this coevolutionary game model, a cooperative neighbourhood gives more neighbouring cooperative offspring and a defective neighbourhood gives more neighbouring defective offspring, leading to positive feedback among cooperative interactions. It is shown that with the help of proximity inheritance, natural selection will favour cooperation over defection under various conditions, even in the presence of mutation. Furthermore, the coevolutionary dynamics could lead to self-organized substantial network clustering, which promotes an assortment of cooperative interactions. This study provides a new insight into the evolutionary mechanism of cooperation in the absence of social attributions such as reputation and punishment. [ABSTRACT FROM AUTHOR]

Ecologists regularly use animal contact networks to describe interactions underlying pathogen transmission, gene flow, and information transfer. However, empirical descriptions of contact often overlook some features of individual movement, and decisions about what kind of network to use in a particular setting are commonly ad hoc. Here, we relate individual movement trajectories to contact networks through a tripartite network model of individual, space, and time nodes. Most networks used in animal contact studies (e.g. individual association networks, home range overlap networks, and spatial networks) are simplifications of this tripartite model. The tripartite structure can incorporate a broad suite of alternative ecological metrics like home range sizes and patch occupancy patterns into inferences about contact network metrics such as modularity and degree distribution. We demonstrate the model's utility with two simulation studies using alternative forms of ecological data to constrain the tripartite network's structure and inform expectations about the harder-to-measure metrics related to contact. [ABSTRACT FROM AUTHOR]

Evolutionary thanatology benefits from broad taxonomic comparisons of non-human animals' responses to death. Furthermore, exploring the sensory and cognitive bases of these responses promises to allow classification of the underlying mechanisms on a spectrum from phylogenetically ancient to more derived traits.We draw on studies of perception and cognition in invertebrate and vertebrate taxa (with a focus on arthropods, corvids, proboscids, cetaceans and primates) to explore the cues that these animals use to detect life and death in others, and discuss proximate and ultimate drivers behind their capacities to do so. Parallels in thanatological behaviour exhibited by the last four taxa suggest similar sensory-cognitive processing rules for dealing with corpses, the evolution of which may have been driven by complex social environments. Uniting these responses is a phenomenon we term 'animacy detection malfunction', whereupon the corpse, having both animate and inanimate attributes, creates states of fear/curiosity manifested as approach/ avoidance behaviours in observers.We suggest that integrating diverse lines of evidence (including the 'uncanny valley' effect originating from the field of robotics) provides a promising way to advance the field, and conclude by proposing avenues for future research. [ABSTRACT FROM AUTHOR]

Observations of some mammals and birds touching their dead provoke questions about the motivation and adaptive value of this potentially risky behaviour. Here, we use controlled experiments to determine if tactile interactions are characteristic of wild American crow responses to dead crows, and what the prevalence and nature of tactile interactions suggests about their motivations. In Experiment 1, we test if food or information acquisition motivates contact by presenting crows with taxidermy-prepared dead crows, and two species crows are known to scavenge: dead pigeons and dead squirrels. In Experiment 2, we test if territoriality motivates tactile interactions by presenting crows with taxidermy crows prepared to look either dead or upright and life-like. In Experiment 1, we find that crows are significantly less likely to make contact but more likely to alarm call and recruit other birds in response to dead crows than to dead pigeons and squirrels. In addition, we find that aggressive and sexual encounters with dead crows are seasonally biased. These findings are inconsistent with feeding or information acquisition-based motivation. In Experiment 2, we find that crows rarely dive-bomb and more often alarm call and recruit other crows to dead than to life-like crows, behaviours inconsistent with responses given to live intruders. Consistent with a danger response hypothesis, our results show that alarm calling and neighbour recruitment occur more frequently in response to dead crows than other stimuli, and that touching dead crows is atypical. Occasional contacts, which take a variety of aggressive and sexual forms, may result from an inability to mediate conflicting stimuli. [ABSTRACT FROM AUTHOR]

Some aquaticmammals appear to care for their dead, whereas others abandon their live offspring when conditions are unfavourable. This incredible variety in behaviours suggests the importance of comparing and contrasting mechanisms driving death-related behaviours among these species. We reviewed 106 cases of aquatic mammals (81 cetaceans and 25 non-cetaceans) reacting to a death event, and extrapolated 'participant' (age class, sex, relationship and decomposition) and 'social' characteristics (escorting, calf dependence, alloparental care, herding and dispersal patterns) from published and unpublished literature. A multiple correspondence analysis (MCA) was performed to explore the relationships between these characteristics and death-related behaviours, with species clustered based on MCA scores. Results showed that both cetaceans and non-cetaceans react to death but in different ways. Non-cetaceans, characterized by a short maternal investment, were observed to protect the dead (defending it from external attacks), while cetaceans spent much longer with their offspring and display carrying (hauling, spinning, mouthing with the carcass and diving with it) and breathing-related (lifting and sinking the carcass) activities with the dead generally in association with other conspecifics. Our work emphasizes the need of increased documentation of death-related cases around theworld to improve our understanding of aquatic mammals and their responses to death. [ABSTRACT FROM AUTHOR]

Many animals socialize in two or more major ecological contexts. In nature, these contexts often involve one situation in which space is more constrained (e.g. shared refuges, sleeping cliffs, nests, dens or burrows) and another situation in which animal movements are relatively free (e.g. in open spaces lacking architectural constraints). Although it is widely recognized that an individual's characteristics may shape its social life, the extent to which architecture constrains social decisions within and between habitats remains poorly understood. Here we developed a novel, automated-monitoring system to study the effects of personality, life-history stage and sex on the social network structure of a facultatively social mammal, the California ground squirrel (Otospermophilus beecheyi) in two distinct contexts: aboveground where space is relatively open and belowground where it is relatively constrained by burrow architecture. Aboveground networks reflected affiliative social interactions whereas belowground networks reflected burrow associations. Network structure in one context (belowground), along with preferential juvenile-adult associations, predicted structure in a second context (aboveground). Network positions of individuals were generally consistent across years (within contexts) and between ecological contexts (within years), suggesting that individual personalities and behavioural syndromes, respectively, contribute to the social network structure of these free-living mammals. Direct ties (strength) tended to be stronger in belowground networks whereas more indirect paths (betweenness centrality) flowed through individuals in aboveground networks. Belowground, females fostered significantly more indirect paths than did males. Our findings have important potential implications for disease and information transmission, offering new insights into the multiple factors contributing to social structures across ecological contexts. [ABSTRACT FROM AUTHOR]

Social networks can be adaptive for members and a recent model (Ilany and Akçay 2016 Nat. Comm. 7, 12084 (doi:10.1038/ncomms12084)) has demonstrated that network structure can be maintained by a simple process of social inheritance. Here, we ask how juvenile vervet monkeys integrate into their adult grooming networks, using the model to test whether observed grooming patterns replicate network structure. Female juveniles, who are philopatric, increased their grooming effort towards adults more than males, although this was not reciprocated by the adults themselves. While more consistent maternal grooming networks, together with maternal network strength, predicted increasing similarity in the patterning of mother-daughter grooming allocations, daughters' grooming networks generally did not match closely those of their mothers. However, maternal networks themselves were not very consistent across time, thus presenting youngsters with a moving target that may be difficult to match. Observed patterns of juvenile female grooming did not replicate the adult network, for which increased association with adults not groomed by their mothers would be necessary. These results suggest that network flexibility, not stability, characterizes our groups and that juveniles are exposed to, and must learn to cope with, temporal shifts in network structure. We hypothesize that this may lead to individual variation in behavioural flexibility, which in turn may help explain why and how variation in sociability influences fitness. [ABSTRACT FROM AUTHOR]

How individuals move through their environment dictates which other individuals they encounter, determining their social and reproductive interactions and the extent to which they experience sexual selection. Specifically, females rarely have the option of mating with all males in a population--they can only choose among the males they encounter. Further, quantifying phenotypic differences between the males that females encounter and those that sire females' offspring lends insight into how social and reproductive interactions shape male phenotypes. We used an explicitly spatio-temporal Markov chain model to estimate the number of potential mates of Anolis sagrei lizards from their movement behaviour, and used genetic paternity assignments to quantify sexual selection on males. Females frequently encountered and mated with multiple males, offering ample opportunity for female mate choice. Sexual selection favoured males that were bigger and moved over larger areas, though the effect of body size cannot be disentangled from last-male precedence. Our approach corroborates some patterns of sexual selection previously hypothesized in anoles based on describing them as territorial, whereas other results, including female multiple mating itself, are at odds with territorial polygyny, offering insight into discrepancies in other taxa between behavioural and genetic descriptions of mating systems. [ABSTRACT FROM AUTHOR]

Though morphologically very similar, equids across the extant species occupy ecological niches that are surprisingly non-overlapping. Occupancy of these distinct niches appears related to subtle physiological and behavioural adaptations which, in turn, correspond to significant differences in the social behaviours and emergent social systems characterizing the different species. Although instances of intraspecific behavioural variation in equids demonstrate that the same body plan can support a range of social structures, each of these morphologically similar species generally shows robust fidelity to its evolved social system. The pattern suggests a subtle relationship between physiological phenotypes and behavioural flexibility. While environmental conditions can vary widely within relatively short temporal or spatial scales, physiological changes and changes to the behaviours that regulate physiological processes, are constrained to longer cycles of adaptation. Physiology is then the limiting variable in the interaction between ecological variation and behavioural and socio-structural flexibility. Behavioural and socio-structural flexibility, in turn, will generate important feedbacks that will govern physiological function, thus creating a coupled web of interactions that can lead to changes in individual and collective behaviour. Longitudinal studies of equid and other large-bodied ungulate populations under environmental stress, such as those discussed here, may offer the best opportunities for researchers to examine, in real time, the interplay between individual behavioural plasticity, socio-structural flexibility, and the physiological and genetic changes that together produce adaptive change. [ABSTRACT FROM AUTHOR]

One of the key questions regarding the underlying mechanisms ofmammalian land migrations is how animals select where to go. Most studies assume perception of resources as the navigational mechanism. The possible role of memory that would allow forecasting conditions at distant locations and times based on information about environmental conditions from previous years has been little studied. We study migrating zebra in Botswana using an individual-based simulation model, where perceptually guided individuals use currently sensed resources at different perceptual ranges, while memoryguided individuals use long-term averages of past resources to forecast future conditions. We compare simulated individuals guided by perception or memory on resource landscapes of remotely sensed vegetation data to trajectories of GPS-tagged zebras. Our results show that memory provides a clear signal that best directs migrants to their destination compared to perception at even the largest perceptual ranges. Zebras modelled with memory arrived two to four times, or up to 100 km, closer to the migration destination than those using perception. We suggest that memory in addition to perception is important for directing ungulate migration. Furthermore, our findings are important for the conservation of migratory mammals, as memory informing direction suggests migration routes could be relatively inflexible. [ABSTRACT FROM AUTHOR]

Collective foraging confers benefits in terms of reduced predation risk and access to social information, but it heightens local competition when resources are limited. In social insects, resource limitation has been suggested as a possible cause for the typical decrease in per capita productivity observed with increasing colony size, a phenomenon known as Michener's paradox. Polydomy (distribution of a colony's brood and workers across multiple nests) is believed to help circumvent this paradox through its positive effect on foraging efficiency, but there is still little supporting evidence for this hypothesis. Here, we showexperimentally that polydomy enhances the foraging performance of food-deprived Temnothorax nylanderi ant colonies via several mechanisms. First, polydomy influences task allocation within colonies, resulting in faster retrieval of protein resources. Second, communication between sister nests reduces search times for far away resources. Third, colonies move queens, brood and workers across available nest sites in response to spatial heterogeneities in protein and carbohydrate resources. This suggests that polydomy represents a flexible mechanism for space occupancy, helping ant colonies adjust to the environment. [ABSTRACT FROM AUTHOR]

Temporal variability in primary productivity can change habitat quality for consumer species by affecting the energy levels available as food resources. However, it remains unclear how habitat-quality fluctuations may determine the dynamics of spatially structured populations, where the effects of habitat size, quality and isolation have been customarily assessed assuming static habitats. We present the first empirical evaluation on the effects of stochastic fluctuations in primary productivity--a major outcome of ecosystem functions--on the meta population dynamics of a primary consumer. A unique 13-year dataset from an herbivore rodent was used to test the hypothesis that inter-annual variations in primary productivity determine spatiotemporal habitat occupancy patterns and colonization and extinction processes. Inter-annual variability in productivity and in the growing season phenology significantly influenced habitat colonization patterns and occupancy dynamics. These effects lead to changes in connectivity to other potentially occupied habitat patches, which then feed back into occupancy dynamics. According to the results, the dynamics of primary productivity accounted for more than 50% of the variation in occupancy probability, depending on patch size and landscape configuration. Evidence connecting primary productivity dynamics and spatiotemporal population processes has broad implications for metapopulation persistence in fluctuating and changing environments. [ABSTRACT FROM AUTHOR]

The use of social and contact networks to answer basic and applied questions about infectious disease transmission in wildlife and livestock is receiving increased attention. Through social network analysis, we understand that wild animal and livestock populations, including farmed fish and poultry, often have a heterogeneous contact structure owing to social structure or trade networks. Network modelling is a flexible tool used to capture the heterogeneous contacts of a population in order to test hypotheses about the mechanisms of disease transmission, simulate and predict disease spread, and test disease control strategies. This review highlights how to use animal contact data, including social networks, for network modelling, and emphasizes that researchers should have a pathogen of interest in mind before collecting or using contact data. This paper describes the rising popularity of network approaches for understanding transmission dynamics in wild animal and livestock populations; discusses the common mismatch between contact networks as measured in animal behaviour and relevant parasites to match those networks; and highlights knowledge gaps in how to collect and analyse contact data. Opportunities for the future include increased attention to experiments, pathogen genetic markers and novel computational tools. [ABSTRACT FROM AUTHOR]

Large herbivore populations respond strongly to remotely sensed measures of primary productivity. Whereas most studies in seasonal environments have focused on the effects of spring plant phenology on juvenile survival, recent studies demonstrated that autumn nutrition also plays a crucial role. We tested for both direct and indirect (through body mass) effects of spring and autumn phenology on winter survival of 2315 mule deer fawns across a wide range of environmental conditions in Idaho, USA. We first performed a functional analysis that identified spring and autumn as the key periods for structuring the among-population and among-year variation of primary production (approximated from 1 km Advanced Very High Resolution Radiometer Normalized Difference Vegetation Index (NDVI)) along the growing season. A path analysis showed that early winter precipitation and direct and indirect effects of spring and autumn NDVI functional components accounted for 45% of observed variation in overwinter survival. The effect size of autumn phenology on body mass was about twice that of spring phenology, while direct effects of phenology on survival were similar between spring and autumn. We demonstrate that the effects of plant phenology vary across ecosystems, and that in semi-arid systems, autumn may be more important than spring for overwinter survival. [ABSTRACT FROM AUTHOR]

Partial migration of some, but not all, members of a population is a common form of migration. We evaluated how migration costs influence which members migrate in 10 populations of two salmonid species. The migratory patterns of both species were evaluated based on the size at maturity for resident males, which is the threshold trait that determines the migratory tactics used within a population. In both species, this sizewas smaller in males located further from the sea, where migration costs are presumably higher. Moreover, the threshold sizes at maturity in males were correlated between both species. Our results suggest that migration costs are a significant convergent selective force on migratory tactics and life-history traits in nature. [ABSTRACT FROM AUTHOR]

Culture is increasingly being understood as a driver of mammalian phenotypes. Defined as group-specific behaviour transmitted by social learning, culture is shaped by social structure. However, culture can itself affect social structure if individuals preferentially interact with others whose behaviour is similar, or cultural symbols are used to mark groups. Using network formalism, this interplay can be depicted by the coevolution of nodes and edges together with the coevolution of network topology and transmission patterns. We review attempts to model the links between the spread, persistence and diversity of culture and the network topology of non-human societies. We illustrate these processes using cetaceans. The spread of socially learned begging behaviour within a population of bottlenose dolphins followed the topology of the social network, as did the evolution of the song of the humpback whale between breeding areas. In three bottlenose dolphin populations, individuals preferentially associated with animals using the same socially learned foraging behaviour. Homogeneous behaviour within the tight, nearly permanent social structures of the large matrilineal whales seems to result from transmission bias, with cultural symbols marking social structures. We recommend the integration of studies of culture and society in species for which social learning is an important determinant of behaviour. [ABSTRACT FROM AUTHOR]

We examine the relationship between mesowear variables and carbon and nitrogen isotopes in 16 species of African antelope (Mammalia: Bovidae). We show significant differences in carbon and nitrogen isotope values between individuals exhibiting sharp versus round cusps, and high versus low occlusal relief. We show significant correlations between mesowear variables and both carbon and nitrogen isotopes. We find significant correlations between mesowear score and nitrogen, but not carbon isotopes. Finally, we find no significant correlations between hypsodonty index and either isotope examined. Our results provide strong support for the use of mesowear variables in palaeodietary reconstructions of antelopes. Our results further suggest that for the antelopes examined here, mesowear signals are a direct result of diet, while hyposodonty may be the result of phylogenetic legacy. [ABSTRACT FROM AUTHOR]

Elephants, dolphins, as well as some carnivores and primates maintain social links despite their frequent splitting and merging in groups of variable composition, a phenomenon known as fission-fusion. Information on the dynamics of social links and interactions among individuals is of high importance to the understanding of the evolution of animal sociality, including that of humans. However, detailed longterm data on such dynamics in wild mammals with fully known demography and kin structures are scarce. Applying a weighted network analysis on 20 500 individual roosting observations over 5 years, we show that in two wild Bechstein's bat colonies with high fission-fusion dynamics, individuals of different age, size, reproductive status and relatedness maintain long-term social relationships. In the larger colony, we detected two stable subunits, each comprising bats from several family lineages. Links between these subunits were mainly maintained by older bats and persisted over all years. Moreover, we show that the full details of the social structure become apparent only when large datasets are used. The stable multi-level social structures in Bechstein's bat colonies resemble that of elephants, dolphins and some primates. Our findings thus may shed new light on the link between social complexity and social cognition in mammals. [ABSTRACT FROM AUTHOR]

Terrestrial plants are powerful climate sentinels because their annual cycles of growth, reproduction and senescence are finely tuned to the annual climate cycle having a period of one year. Consistency in the seasonal phasing of terrestrial plant activity provides a relatively low-noise background from which phenological shifts can be detected and attributed to climate change. Here, we ask whether phytoplankton biomass also fluctuates over a consistent annual cycle in lake, estuarine-coastal and ocean ecosystems and whether there is a characteristic phenology of phytoplankton as a consistent phase and amplitude of variability. We compiled 125 time series of phytoplankton biomass (chlorophyll a concentration) from temperate and subtropical zones and used wavelet analysis to extract their dominant periods of variability and the recurrence strength at those periods. Fewer than half (48%) of the series had a dominant 12-month period of variability, commonly expressed as the canonical spring-bloom pattern. About 20 per cent had a dominant six-month period of variability, commonly expressed as the spring and autumn or winter and summer blooms of temperate lakes and oceans. These annual patterns varied in recurrence strength across sites, and did not persist over the full series duration at some sites. About a third of the series had no component of variability at either the six- or 12-month period, reflecting a series of irregular pulses of biomass. These findings show that there is high variability of annual phytoplankton cycles across ecosystems, and that climate-driven annual cycles can be obscured by other drivers of population variability, including human disturbance, aperiodic weather events and strong trophic coupling between phytoplankton and their consumers. Regulation of phytoplankton biomass by multiple processes operating at multiple time scales adds complexity to the challenge of detecting climate-driven trends in aquatic ecosystems where the noise to signal ratio is high. [ABSTRACT FROM AUTHOR]

Autocorrelation has been viewed as a problem in telemetry studies because sequential observations are not independent in time or space, therefore violating assumptions for statistical inference. Yet nearly all ecological and behavioural data are autocorrelated in both space and time. We argue that there is much to learn about the structure of ecological and behavioural data from patterns of autocorrelation. Such patterns include periodicity in movement and patchiness in spatial data, which can be characterized by an autocorrelogram, semivariogram or spectrum. We illustrate the utility of temporal autocorrelation functions (ACFs) for analysing step-length data from GPS telemetry of wolves (Canis lupus), cougars (Puma concolor), grizzly bears (Ursus arctos) and elk (Cervus elaphus) in western Alberta. ACFs often differ by season, reflecting differences in foraging behaviour. In wilderness landscapes, step-length ACFs for predators decay slowly to apparently random patterns, but sometimes display strong daily rhythms in areas of human disturbance. In contrast, step lengths of elk are consistently periodic, reflecting crepuscular activity. [ABSTRACT FROM AUTHOR]

With the advent of new technologies, animal locations are being collected at ever finer spatiotemporal scales. We review analytical methods for dealing with correlated data in the context of resource selection, including post hoc variance inflation techniques, 'two-stage' approaches based on models fit to each individual, generalized estimating equations and hierarchical mixed-effects models. These methods are applicable to a wide range of correlated data problems, but can be difficult to apply and remain especially challenging for use-availability sampling designs because the correlation structure for combinations of used and available points are not likely to follow common parametric forms. We also review emerging approaches to studying habitat selection that use finescale temporal data to arrive at biologically based definitions of available habitat, while naturally accounting for autocorrelation by modelling animal movement between telemetry locations. Sophisticated analyses that explicitly model correlation rather than consider it a nuisance, like mixed effects and state-space models, offer potentially novel insights into the process of resource selection, but additional work is needed to make them more generally applicable to large datasets based on the use-availability designs. Until then, variance inflation techniques and two-stage approaches should offer pragmatic and flexible approaches to modelling correlated data. [ABSTRACT FROM AUTHOR]

Global positioning system (GPS) telemetry technology allows us to monitor and to map the details of animal movement, securing vast quantities of such data even for highly cryptic organisms. We envision an exciting synergy between animal ecology and GPS-based radiotelemetry, as for other examples of new technologies stimulating rapid conceptual advances, where research opportunities have been paralleled by technical and analytical challenges. Animal positions provide the elemental unit of movement paths and show where individuals interact with the ecosystems around them. We discuss how knowing where animals go can help scientists in their search for a mechanistic understanding of key concepts of animal ecology, including resource use, home range and dispersal, and population dynamics. It is probable that in the not-so-distant future, intense sampling of movements coupled with detailed information on habitat features at a variety of scales will allow us to represent an animal's cognitive map of its environment, and the intimate relationship between behaviour and fitness. An extended use of these data over long periods of time and over large spatial scales can provide robust inferences for complex, multi-factorial phenomena, such as meta-analyses of the effects of climate change on animal behaviour and distribution. [ABSTRACT FROM AUTHOR]

In the past decade, ecologists have witnessed vast improvements in our ability to collect animal movement data through animal-borne technology, such as through GPS or ARGOS systems. However, more data does not necessarily yield greater knowledge in understanding animal ecology and conservation. In this paper, we provide a review of the major benefits, problems and potential misuses of GPS/Argos technology to animal ecology and conservation. Benefits are obvious, and include the ability to collect fine-scale spatio-temporal location data on many previously impossible to study animals, such as ocean-going fish, migratory songbirds and long-distance migratory mammals. These benefits come with significant problems, however, imposed by frequent collar failures and high cost, which often results in weaker study design, reduced sample sizes and poorer statistical inference. In addition, we see the divorcing of biologists from a field-based understanding of animal ecology to be a growing problem. Despite these difficulties, GPS devices have provided significant benefits, particularly in the conservation and ecology of wide-ranging species.We conclude by offering suggestions for ecologists on which kinds of ecological questions would currently benefit the most from GPS/Argos technology, and where the technology has been potentially misused. Signifi- cant conceptual challenges remain, however, including the links between movement and behaviour, and movement and population dynamics. [ABSTRACT FROM AUTHOR]