Using global remote camera data of a solitary species complex to evaluate the drivers of group formation.

The social system of animals involves a complex interplay between physiology, natural history, and the environment. Long relied upon discrete categorizations of "social" and "solitary" inhibit our capacity to understand species and their interactions with the world around them. Here, we use a globally distributed camera trapping dataset to test the drivers of aggregating into groups in a species complex (martens and relatives, family Mustelidae, Order Carnivora) assumed to be obligately solitary. We use a simple quantification, the probability of being detected in a group, that was applied across our globally derived camera trap dataset. Using a series of binomial generalized mixed-effects models applied to a dataset of 16,483 independent detections across 17 countries on four continents we test explicit hypotheses about potential drivers of group formation. We observe a wide range of probabilities of being detected in groups within the solitary model system, with the probability of aggregating in groups varying by more than an order of magnitude. We demonstrate that a species' context-dependent proclivity toward aggregating in groups is underpinned by a range of resource-related factors, primarily the distribution of resources, with increasing patchiness of resources facilitating group formation, as well as interactions between environmental conditions (resource constancy/winter severity) and physiology (energy storage capabilities). The wide variation in propensities to aggregate with conspecifics observed here highlights how continued failure to recognize complexities in the social behaviors of apparently solitary species limits our understanding not only of the individual species but also the causes and consequences of group formation.

Restoring vertebrate predator populations can provide landscape-scale biological control of established invasive vertebrates: Insights from pine marten recovery in Europe.

Invasive species pose one of the greatest global threats to biodiversity. There has been a long history of importing coevolved natural enemies to act as biological control agents to try to suppress densities of invasive species, with historically limited success and frequent adverse impacts on native biodiversity. Our understanding of the processes and drivers of successful biological control has been focussed on invertebrates and is evidently limited and potentially ill-suited with respect to biological control of vertebrate populations. The restoration of native vertebrate predator populations provides a promising nature-based solution for slowing, halting, or even reversing the spread of some invasive vertebrates over spatial scales relevant to the management of wildlife populations. Here, we first review the growing literature and data from the pine marten-red and grey squirrel system in Europe. We synthesise a multi-decadal dataset to show that the recovery of a native predator has resulted in rapid, landscape-scale declines of an established invasive species. We then use the model system, predator-prey interaction theory, and examples from the literature to develop ecological theory relating to natural biological control in vertebrates and evolutionary processes in native-invasive predator-prey interactions. We find support for the hypotheses that evolutionary naivety of invasive species to native predators and lack of local refuges results in higher predation of naive compared to coevolved prey. We apply lessons learnt from the marten-squirrel model system to examine the plausibility of specific native predator solutions to some of the Earth's most devastating invasive vertebrates. Given the evidence, we conclude that depletion of vertebrate predator populations has increased ecosystem vulnerability to invasions and thus facilitated the spread of invasive species. Therefore, restoration of vertebrate predator populations is an underappreciated, fundamental, nature-based solution to the crisis of invasive species and should be a priority for vertebrate invasive species management globally.

Habitat mediates coevolved but not novel species interactions.

Ongoing recovery of native predators has the potential to alter species interactions, with community and ecosystem wide implications. We estimated the co-occurrence of three species of conservation and management interest from a multi-species citizen science camera trap survey. We demonstrate fundamental differences in novel and coevolved predator-prey interactions that are mediated by habitat. Specifically, we demonstrate that anthropogenic habitat modification had no influence on the expansion of the recovering native pine marten in Ireland, nor does it affect the predator's suppressive influence on an invasive prey species, the grey squirrel. By contrast, the direction of the interaction between the pine marten and a native prey species, the red squirrel, is dependent on habitat. Pine martens had a positive influence on red squirrel occurrence at a landscape scale, especially in native broadleaf woodlands. However, in areas dominated by non-native conifer plantations, the pine marten reduced red squirrel occurrence. These findings suggest that following the recovery of a native predator, the benefits of competitive release are spatially structured and habitat-specific. The potential for past and future landscape modification to alter established interactions between predators and prey has global implications in the context of the ongoing recovery of predator populations in human-modified landscapes.

Animal lifestyle affects acceptable mass limits for attached tags.

Animal-attached devices have transformed our understanding of vertebrate ecology. To minimize any associated harm, researchers have long advocated that tag masses should not exceed 3% of carrier body mass. However, this ignores tag forces resulting from animal movement. Using data from collar-attached accelerometers on 10 diverse free-ranging terrestrial species from koalas to cheetahs, we detail a tag-based acceleration method to clarify acceptable tag mass limits. We quantify animal athleticism in terms of fractions of animal movement time devoted to different collar-recorded accelerations and convert those accelerations to forces (acceleration × tag mass) to allow derivation of any defined force limits for specified fractions of any animal's active time. Specifying that tags should exert forces that are less than 3% of the gravitational force exerted on the animal's body for 95% of the time led to corrected tag masses that should constitute between 1.6% and 2.98% of carrier mass, depending on athleticism. Strikingly, in four carnivore species encompassing two orders of magnitude in mass (ca 2-200 kg), forces exerted by '3%' tags were equivalent to 4-19% of carrier body mass during moving, with a maximum of 54% in a hunting cheetah. This fundamentally changes how acceptable tag mass limits should be determined by ethics bodies, irrespective of the force and time limits specified.

Limitations of using surrogates for behaviour classification of accelerometer data: refining methods using random forest models in Caprids.

BACKGROUND: Animal-attached devices can be used on cryptic species to measure their movement and behaviour, enabling unprecedented insights into fundamental aspects of animal ecology and behaviour. However, direct observations of subjects are often still necessary to translate biologging data accurately into meaningful behaviours. As many elusive species cannot easily be observed in the wild, captive or domestic surrogates are typically used to calibrate data from devices. However, the utility of this approach remains equivocal. METHODS: Here, we assess the validity of using captive conspecifics, and phylogenetically-similar domesticated counterparts (surrogate species) for calibrating behaviour classification. Tri-axial accelerometers and tri-axial magnetometers were used with behavioural observations to build random forest models to predict the behaviours. We applied these methods using captive Alpine ibex (Capra ibex) and a domestic counterpart, pygmy goats (Capra aegagrus hircus), to predict the behaviour including terrain slope for locomotion behaviours of captive Alpine ibex. RESULTS: Behavioural classification of captive Alpine ibex and domestic pygmy goats was highly accurate (> 98%). Model performance was reduced when using data split per individual, i.e., classifying behaviour of individuals not used to train models (mean ± sd = 56.1 ± 11%). Behavioural classifications using domestic counterparts, i.e., pygmy goat observations to predict ibex behaviour, however, were not sufficient to predict all behaviours of a phylogenetically similar species accurately (> 55%). CONCLUSIONS: We demonstrate methods to refine the use of random forest models to classify behaviours of both captive and free-living animal species. We suggest there are two main reasons for reduced accuracy when using a domestic counterpart to predict the behaviour of a wild species in captivity; domestication leading to morphological differences and the terrain of the environment in which the animals were observed. We also identify limitations when behaviour is predicted in individuals that are not used to train models. Our results demonstrate that biologging device calibration needs to be conducted using: (i) with similar conspecifics, and (ii) in an area where they can perform behaviours on terrain that reflects that of species in the wild.

Native and invasive squirrels show different behavioural responses to scent of a shared native predator.

Invasive species pose a serious threat to native species. In Europe, invasive grey squirrels (Sciurus carolinensis) have replaced native red squirrels (Sciurus vulgaris) in locations across Britain, Ireland and Italy. The European pine marten (Martes martes) can reverse the replacement of red squirrels by grey squirrels, but the underlying mechanism of how pine martens suppress grey squirrels is little understood. Research suggests the reversal process is driven by direct predation, but why the native red squirrel may be less susceptible than the invasive grey squirrel to predation by a commonly shared native predator, is unknown. A behavioural difference may exist with the native sciurid being more effective at avoiding predation by the pine marten with which they have a shared evolutionary history. In mammals, olfactory cues are used by prey species to avoid predators. To test whether anti-predator responses differ between the native red squirrel and the invasive grey squirrel, we exposed both species to scent cues of a shared native predator and quantified the responses of the two squirrel species. Red squirrels responded to pine marten scent by avoiding the feeder, increasing their vigilance and decreasing their feeding activity. By contrast, grey squirrels did not show any anti-predator behaviours in response to the scent of pine marten. Thus, differences in behavioural responses to a shared native predator may assist in explaining differing outcomes of species interactions between native and invasive prey species depending on the presence, abundance and exposure to native predators.

Increasing land-use intensity reverses the relative occupancy of two quadrupedal scavengers.

Human land use is continuously altering the natural environment, yet the greater ecological implications of this change for many groups that are key to healthy ecosystem functioning remains uncharacterised in the tropics. Terrestrial scavenging vertebrates are one such group, providing integral ecosystem services through the removal of carrion which is a crucial component of both nutrient cycling and disease dynamics. To explore how anthropogenic processes may affect forest scavengers, we investigated the changes in the relative occupancy of two important terrestrial scavengers along a gradient of land use intensity, ranging from protected forest to oil palm plantation in Borneo. We found the Malay civet (Viverra tangalunga) had highest, albeit variable, occupancy in areas of low land use intensity and the Southeast Asian water monitor (Varanus salvator macromaculatus) had highest occupancy in areas of high land use intensity. Land use had no effect on the combined occupancy of the two species. In high land use intensity sites, individual water monitors were larger and had better body condition, but at population level had a highly biased sex ratio with more males than females and increased signs of intraspecific conflict. We did not assess scavenging rate or efficiency as a process, but the high occupancy rates and apparent health of the scavengers in high land use intensity landscapes suggests this ecological process is robust to land use change.