1.Density estimation is of fundamental importance in wildlife management. The use of camera traps to estimate animal density has so far been restricted to capture-recapture analysis of species with individually identifiable markings. This study developed a method that eliminates the requirement for individual recognition of animals by modelling the underlying process of contact between animals and cameras. 2. The model provides a factor that linearly scales trapping rate with density, depending on two key biological variables (average animal group size and day range) and two characteristics of the camera sensor (distance and angle within which it detects animals). 3. We tested the approach in an enclosed animal park with known abundances of four species, obtaining accurate estimates in three out of four cases. Inaccuracy in the fourth species was because of biased placement of cameras with respect to the distribution of this species. 4. Synthesis and applications. Subject to unbiased camera placement and accurate measurement of model parameters, this method opens the possibility of reduced labour costs for estimating wildlife density and may make estimation possible where it has not been previously. We provide guidelines on the trapping effort required to obtain reasonably precise estimates.
Summary1. Activity level (the proportion of time that animals spend active) is a behavioural and ecological metric that can provide an indicator of energetics, foraging effort and exposure to risk. However, activity level is poorly known for free-living animals because it is difficult to quantify activity in the field in a consistent, cost-effective and non-invasive way. 2. This article presents a new method to estimate activity level with time-of-detection data from camera traps (or more generally any remote sensors), fitting a flexible circular distribution to these data to describe the underlying activity schedule, and calculating overall proportion of time active from this. 3. Using simulations and a case study for a range of small-to medium-sized mammal species, we find that activity level can reliably be estimated using the new method. 4. The method depends on the key assumption that all individuals in the sampled population are active at the peak of the daily activity cycle. We provide theoretical and empirical evidence suggesting that this assumption is likely to be met for many species, but may be less likely met in large predators, or in high-latitude winters. Further research is needed to establish stronger evidence on the validity of this assumption in specific cases; however, the approach has the potential to provide an effective, non-invasive alternative to existing methods for quantifying population activity levels.
Summary1. Abundance estimation is a pervasive goal in ecology. The rate of detection by motion-sensitive camera traps can, in principle, provide information on the abundance of many species of terrestrial vertebrates that are otherwise difficult to survey. The random encounter model (REM, Rowcliffe et al. 2008) provides a means estimating abundance from camera trap rate but requires camera sensitivity to be quantified. 2. Here, we develop a method to estimate the area effectively monitored by cameras, which is one of the most important codeterminants of detection rate. Our method borrows from distance sampling theory, applying detection function models to data on the position (distance and angle relative to the camera) where the animals are first detected. Testing the reliability of this approach through simulation, we find that bias depends on the effective detection angle assumed but was generally low at less than 5% for realistic angles typical of camera traps. 3. We adapted standard detection functions to allow for the possibility of smaller animals passing beneath the field of view close to the camera, resulting in reduced detection probability within that zone. Using a further simulation to test this approach, we find that detection distance can be estimated with little or no bias if detection probability is certain for at least some distance from the camera. 4. Applying this method to a 1-year camera trapping data set from Barro Colorado Island, Panama, we show that effective detection distance is related strongly positively to species body mass and weakly negatively to species average speed of movement. There was also a strong seasonal effect, with shorter detection distance during the wet season. Effective detection angle is related more weakly to species body mass, and again strongly to season, with a wider angle in the wet season. 5. This method represents an important step towards practical application of the REM, including abundance estimation for relatively small (<1 kg) species.
Animals that forage socially often stand to gain from coordination of their behaviour. Yet it is not known how group members reach a consensus on the timing of foraging bouts. Here we demonstrate a simple process by which this may occur. We develop a state-dependent, dynamic game model of foraging by a pair of animals, in which each individual chooses between resting or foraging during a series of consecutive periods, so as to maximize its own individual chances of survival. We find that, if there is an advantage to foraging together, the equilibrium behaviour of both individuals becomes highly synchronized. As a result of this synchronization, differences in the energetic reserves of the two players spontaneously develop, leading them to adopt different behavioural roles. The individual with lower reserves emerges as the 'pace-maker' who determines when the pair should forage, providing a straightforward resolution to the problem of group coordination. Moreover, the strategy that gives rise to this behaviour can be implemented by a simple 'rule of thumb' that requires no detailed knowledge of the state of other individuals.
Mammalian carnivores fall into two broad dietary groups: smaller carnivores (<20 kg) that feed on very small prey (invertebrates and small vertebrates) and larger carnivores (>20 kg) that specialize in feeding on large vertebrates. We develop a model that predicts the mass-related energy budgets and limits of carnivore size within these groups. We show that the transition from small to large prey can be predicted by the maximization of net energy gain; larger carnivores achieve a higher net gain rate by concentrating on large prey. However, because it requires more energy to pursue and subdue large prey, this leads to a 2-fold step increase in energy expenditure, as well as increased intake. Across all species, energy expenditure and intake both follow a three-fourths scaling with body mass. However, when each dietary group is considered individually they both display a shallower scaling. This suggests that carnivores at the upper limits of each group are constrained by intake and adopt energy conserving strategies to counter this. Given predictions of expenditure and estimates of intake, we predict a maximum carnivore mass of approximately a ton, consistent with the largest extinct species. Our approach provides a framework for understanding carnivore energetics, size, and extinction dynamics.
Countries committed to implementing the Convention on Biological Diversity's 2011–2020 strategic plan need effective tools to monitor global trends in biodiversity. Remote cameras are a rapidly growing technology that has great potential to transform global monitoring for terrestrial biodiversity and can be an important contributor to the call for measuring Essential Biodiversity Variables. Recent advances in camera technology and methods enable researchers to estimate changes in abundance and distribution for entire communities of animals and to identify global drivers of biodiversity trends. We suggest that interconnected networks of remote cameras will soon monitor biodiversity at a global scale, help answer pressing ecological questions, and guide conservation policy. This global network will require greater collaboration among remote‐camera studies and citizen scientists, including standardized metadata, shared protocols, and security measures to protect records about sensitive species. With modest investment in infrastructure, and continued innovation, synthesis, and collaboration, we envision a global network of remote cameras that not only provides real‐time biodiversity data but also serves to connect people with nature.
Fatal amphibian chytridiomycosis has typically been associated with the direct costs of infection. However the relationship between exposure to the pathogen, infection and mortality may not be so straightforward. Using results from both field work and experiments we report how exposure of common toads to Batrachochytrium dendrobatidis influences development and survival and how developmental stage influences host responses. Our results show that costs are accrued in a dose dependent manner during the larval stage and are expressed at or soon after metamorphosis. Exposure to B. dendrobatidis always incurs a growth cost for tadpoles and can lead to larval mortality before or soon after metamorphosis even when individuals do not exhibit infection at time of death. In contrast, exposure after metamorphosis almost always results in infection, but body size dictates survival to a greater extent than does dose. These data show that amphibian survival in the face of challenge by an infectious agent is dependent on host condition as well as life history stage. Under current models of climate change, many species of amphibia are predicted to increasingly occur outside their environmental optima. In this case, condition‐dependent traits such as we have demonstrated may weigh heavily on species survival.
Summary 1.The distance travelled by animals is an important ecological variable that links behaviour, energetics and demography. It is usually measured by summing straight-line distances between intermittently sampled locations along continuous animal movement paths. The extent to which this approach underestimates travel distance remains a rarely addressed and unsolved problem, largely because true movement paths are rarely, if ever, available for comparison. Here, we use simulated movement paths parameterized with empirical movement data to study how estimates of distance travelled are affected by sampling frequency. 2. We used a novel method to obtain fine-scale characteristics of animal movement from camera trap videos for a set of tropical forest mammals and used these characteristics to generate detailed movement paths. We then sampled these paths at different frequencies, simulating telemetry studies, and quantified the accuracy of sampled travel distance estimation. 3. For our focal species, typical telemetry studies would underestimate distances travelled by 67-93%, and extremely high sampling frequencies (several fixes per minute) would be required to get tolerably accurate estimates. The form of the relationship between tortuosity, sample frequency, and distance travelled was such that absolute distance cannot accurately be estimated by the infrequent samples used in typical tracking studies. 4. We conclude that the underestimation of distance travelled is a serious but underappreciated problem. Currently, there is no reliable, widely applicable method to obtain approximately unbiased estimates of distance travelled by animals. Further research on this problem is needed.
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