Deer Behavior Affects Density Estimates With Camera Traps, but Is Outwighted by Spatial Variability

Density is a key trait of populations and an essential parameter in ecological research, wildlife conservation and management. Several models have been developed to estimate population density based on camera trapping data, including the random encounter model (REM) and camera trap distance sampling (CTDS). Both models need to account for variation in animal behavior that depends, for example, on the species and sex of the animals along with temporally varying environmental factors. We examined whether the density estimates of REM and CTDS can be improved for Europe’s most numerous deer species, by adjusting the behavior-related model parameters per species and accounting for differences in movement speeds between sexes, seasons, and years. Our results showed that bias through inadequate consideration of animal behavior was exceeded by the uncertainty of the density estimates, which was mainly influenced by variation in the number of independent observations between camera trap locations. The neglection of seasonal and annual differences in movement speed estimates for REM overestimated densities of red deer in autumn and spring by ca. 14%. This GPS telemetry-derived parameter was found to be most problematic for roe deer females in summer and spring when movement behavior was characterized by small-scale displacements relative to the intervals of the GPS fixes. In CTDS, density estimates of red deer improved foremost through the consideration of behavioral reactions to the camera traps (avoiding bias of max. 19%), while species-specific delays between photos had a larger effect for roe deer. In general, the applicability of both REM and CTDS would profit profoundly from improvements in their precision along with the reduction in bias achieved by exploiting the available information on animal behavior in the camera trap data.Deer Behavior Affects Density Estimates With Camera Traps, but Is Outwighted by Spatial VariabilitypublishedVersio

Spatial variation in red deer density in a transboundary forest ecosystem

Forests in Europe are exposed to increasingly frequent and severe disturbances. The resulting changes in the structure and composition of forests can have profound consequences for the wildlife inhabiting them. Moreover, wildlife populations in Europe are often subjected to differential management regimes as they regularly extend across multiple national and administrative borders. The red deer Cervus elaphus population in the Bohemian Forest Ecosystem, straddling the Czech-German border, has experienced forest disturbances, primarily caused by windfalls and bark beetle Ips typographus outbreaks during the past decades. To adapt local management strategies to the changing environmental conditions and to coordinate them across the international border, reliable estimates of red deer density and abundance are highly sought-after by policymakers, wildlife managers, and stakeholders. Covering a 1081-km2 study area, we conducted a transnational non-invasive DNA sampling study in 2018 that yielded 1578 genotyped DNA samples from 1120 individual red deer. Using spatial capture-recapture models, we estimated total and jurisdiction-specific abundance of red deer throughout the ecosystem and quantified the role of forest disturbance and differential management strategies in shaping spatial heterogeneity in red deer density. We hypothesised that (a) forest disturbances provide favourable habitat conditions (e.g., forage and cover), and (b) contrasting red deer management regimes in different jurisdictions create a differential risk landscape, ultimately shaping density distributions. Overall, we estimated that 2851 red deer (95% Credible Interval = 2609–3119) resided in the study area during the sampling period, with a relatively even overall sex ratio (1406 females, 95% CI = 1229–1612 and 1445 males, 95% CI = 1288–1626). The average red deer density was higher in Czechia (3.5 km−2, 95% CI = 1.2–12.3) compared to Germany (2 km−2, 95% CI = 0.2–11). The effect of forest disturbances on red deer density was context-dependent. Forest disturbances had a positive effect on red deer density at higher elevations and a negative effect at lower elevations, which could be explained by partial migration and its drivers in this population. Density of red deer was generally higher in management units where hunting is prohibited. In addition, we found that sex ratios differed between administrative units and were more balanced in the non-intervention zones. Our results show that the effect of forest disturbances on wild ungulates is modulated by additional factors, such as elevation and ungulate management practices. Overall density patterns and sex ratios suggested strong gradients in density between administrative units. With climate change increasing the severity and frequency of forest disturbances, population-level monitoring and management are becoming increasingly important, especially for wide-ranging species as both wildlife and global change transcend administrative boundaries.publishedVersio

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Data from: Impact of winter enclosures on the gut bacterial microbiota of red deer in the Bavarian Forest National Park

High numbers of red deer (Cervus elaphus) pose a challenge for natural forests because of their high browsing intensities, especially during winter months. To mitigate this human–wildlife conflict, conservation management in Central Europe involves luring red deer into fenced winter-feeding sites. The supplementary diet provided in these so-called winter enclosures strongly differs from the natural diet of red deer. Dietary shifts, however, can lead to an imbalance of the gut microbiota, which could promote bacterial pathogens. Moreover, increased inter-individual contact in winter enclosures enhances the exchange of symbiotic but also pathogenic bacteria. In this study, we used high-throughput sequencing of the 16S rRNA gene in fecal samples of red deer inhabiting the Bavarian Forest National Park to investigate differences in the gut bacterial microbiota between individuals in winter enclosures and individuals that ranged freely in the forests in winter. We also investigated the occurrence of potential zoonotic bacterial pathogens in both study groups. Our results revealed that proportions of bacterial taxa, alpha- and beta-diversities, and relative abundances of amplicon sequence variants in the gut bacterial microbiota of the two groups differed. These differences were attributed to the enrichment of bacterial taxa involved in the digestion of the supplementary food and to different natural diets consumed before entering the winter enclosures. We detected sequences with high similarities to known red deer pathogens in both study groups, but their relative abundances were low, which suggests that the population of red deer of the National Park Bavarian Forest is healthy

Reducing bias in density estimates for unmarked populations that exhibit reactive behaviour towards camera traps

Abstract Density estimates guide wildlife management, and camera traps are commonly used to estimate sizes of unmarked populations. Unfortunately, animals often alter their natural behaviour in the presence of camera traps, which may bias subsequent density estimates. We simulated 100 populations of known density to test several new and existing methods that aimed to reduce bias in density estimates from camera trap distance sampling (CTDS) and the random encounter model (REM). Within our simulated populations, we introduced different behavioural reactions including attraction towards cameras, freezing when near cameras and fleeing from cameras. CTDS and REM provided density estimates with decent coverage of confidence intervals (CTDS = 94%, REM = 87%), mean coefficient of variation (CTDS = 0.121, REM = 0.071) and minimal bias (root‐mean squared error: CTDS = 1.336, REM = 0.913) for simulated populations with no reactive behaviour. However, failure to implement a method to account for reactive behaviour resulted in low coverage, large bias and potentially imprecise density estimates when 30% of the simulated population potentially reacted by attraction to or fleeing from camera traps. We identified a corrective strategy that enhanced confidence interval coverage, increased precision and reduced bias for every behavioural reaction except when individuals potentially fled from cameras. Synthesis and applications. We provide empirically tested methods for reducing bias of density estimates. Wildlife managers requiring population estimates of animals that exhibit reactive behaviour can use our methods to reduce inaccuracy. We encourage future studies to quantify behavioural responses to camera traps and to implement, test and possibly extend our methods to reduce bias through simulation

Estimating effective survey duration in camera trap distance sampling surveys

This study was funded by the German Centre for Integrative Biodiversity Research (iDiv) Halle‐Jena‐Leipzig and the Bavarian State Ministry of the Environment and Consumer Protection (project ID 77262). The field work was financed by the program Ziel ETZ FreeState of Bavaria—Czech Republic 2014–2020 (INTERREG V, project number 184). We thank our interns for their support with the field work and data processing. Open Access funding enabled and organized by Projekt DEAL.Among other approaches, camera trap distance sampling (CTDS) is used to estimate animal abundance from unmarked populations. It was formulated for videos and observation distances are measured at predetermined ‘snapshot moments’. Surveys recording still images with passive infrared motion sensors suffer from frequent periods where animals are not photographed, either because of technical delays before the camera can be triggered again (i.e. ‘camera recovery time’) or because they remain stationary and do not immediately retrigger the camera following camera recovery time (i.e. ‘retrigger delays’). These effects need to be considered when calculating temporal survey effort to avoid downwardly biased abundance estimates. Here, we extend the CTDS model for passive infrared motion sensor recording of single images or short photo series. We propose estimating ‘mean time intervals between triggers’ as combined mean camera recovery time and mean retrigger delays from the time interval distribution of pairs of consecutive pictures, using a Gamma and Exponential function, respectively. We apply the approach to survey data on red deer, roe deer and wild boar. Mean time intervals between triggers were very similar when estimated empirically and when derived from the model-based approach. Depending on truncation times (i.e. the time interval between consecutive pictures beyond which data are discarded) and species, we estimated mean time intervals between retriggers between 8.28 and 15.05 s. Using a predefined snapshot interval, not accounting for these intervals, would lead to underestimated density by up to 96% due to overestimated temporal survey effort. The proposed approach is applicable to any taxa surveyed with camera traps. As programming of cameras to record still images is often preferred over video recording due to reduced consumption of energy and memory, we expect this approach to find broad application, also for other camera trap methods than CTDS.Publisher PDFPeer reviewe

The influence of camera trap flash type on the behavioural reactions and trapping rates of red deer and roe deer

Camera traps have become an important tool in wildlife monitoring. However, an issue in interpreting their data in statistical analyses of population densities, demography or behaviour is that the probability of detecting the target animals and their behaviours may vary depending on environmental and methodological factors. A specific problem is the type of flash used in the camera trap, as animals may react differently to different flash types and change their avoidance or habituation behaviour accordingly over time. Here, we provide the first systematic test of the impact of flash type in studies of red deer (Cervus elaphus) and roe deer (Capreolus capreolus), based on an analysis of behavioural responses to white, standard infrared and black flashes during 900 camera trap deployments in the Bavarian Forest National Park and the Northern Black Forest. The results revealed that both deer species were more likely to react to standard infrared than to black flash, but trigger delays prevented comparisons to white flash. Red deer reacted more frequently to camera traps than did roe deer, and responses were more common in the Northern Black Forest than in the Bavarian Forest National Park, where hunting is severely restricted. Contrary to our expectations, camera trapping rates of free-ranging deer did not significantly decline over time for any flash type or species. Despite the lack of evidence for avoidance behaviour, we recommend the use of black flash for behavioural studies of deer populations to minimize the risk of introducing a source of disturbance whereas infrared and white flash are equally suitable for determinations of demographic parameters

Deer Behavior Affects Density Estimates With Camera Traps, but Is Outwighted by Spatial Variability

Density is a key trait of populations and an essential parameter in ecological research, wildlife conservation and management. Several models have been developed to estimate population density based on camera trapping data, including the random encounter model (REM) and camera trap distance sampling (CTDS). Both models need to account for variation in animal behavior that depends, for example, on the species and sex of the animals along with temporally varying environmental factors. We examined whether the density estimates of REM and CTDS can be improved for Europe’s most numerous deer species, by adjusting the behavior-related model parameters per species and accounting for differences in movement speeds between sexes, seasons, and years. Our results showed that bias through inadequate consideration of animal behavior was exceeded by the uncertainty of the density estimates, which was mainly influenced by variation in the number of independent observations between camera trap locations. The neglection of seasonal and annual differences in movement speed estimates for REM overestimated densities of red deer in autumn and spring by ca. 14%. This GPS telemetry-derived parameter was found to be most problematic for roe deer females in summer and spring when movement behavior was characterized by small-scale displacements relative to the intervals of the GPS fixes. In CTDS, density estimates of red deer improved foremost through the consideration of behavioral reactions to the camera traps (avoiding bias of max. 19%), while species-specific delays between photos had a larger effect for roe deer. In general, the applicability of both REM and CTDS would profit profoundly from improvements in their precision along with the reduction in bias achieved by exploiting the available information on animal behavior in the camera trap data