Quantifying animal movements is necessary for answering a wide array of research questions in ecology and conservation biology. Consequently, ecologists have made considerable efforts to identify the best way to estimate an animal’s home range, and many methods of estimating home ranges have arisen over the past half century. Most of these methods fall into two distinct categories of estimators that have only recently been described in statistical detail: those that measure range distributions (methods such as Kernel Density Estimation that quantify the long-run behavior of a movement process that features restricted space use) and those that measure occurrence distributions (methods such as Brownian Bridge Movement Models and the Correlated Random Walk Library that quantify uncertainty in an animal movement path during a specific period of observation). In this paper, we use theory, simulations, and empirical analysis to demonstrate the importance of applying these two classes of space use estimators appropriately and distinctly. Conflating range and occurrence distributions can have serious consequences for ecological inference and conservation practice. For example, in most situations, home-range estimates quantified using occurrence estimators are too small, and this problem is exacerbated by ongoing improvements in tracking technology that enable more frequent and more accurate data on animal movements. We encourage researchers to use range estimators to estimate the area of home ranges and occurrence estimators to answer other questions in movement ecology, such as when and where an animal crosses a linear feature, visits a location of interest, or interacts with other animals. Open Research Statement Tracking data on Aepyceros melampus, Beatragus hunteri, Bycanistes bucinator, Cerdocyon thous, Eulemur rufifrons, Glyptemys insculpta, Gyps coprotheres, Madoqua guentheri, Ovis canadensis, Propithecus verreauxi, Sus scrofa, and Ursus arctos are publicly archived in the Dryad repository (Noonan et al. 2018; https://doi.org/10.5061/dryad.v5051j2), as are data from Procapra gutturosa (Fleming et al. 2014a; https://doi.org/10.5061/dryad.45157). Data on Panthera onca were taken from (Morato et al. 2018). Additional data are publicly archived in the Movebank repository under the following identifiers: Canis latrans, 8159699; Canis lupus, 8159399; Chrysocyon brachyurus, 18156143; Felis silvestris, 40386102; Gyps africanus, 2919708; Lepus europaeus, 25727477; Martes pennanti, 2964494; Panthera leo, 220229; Papio cynocephalus, 222027; Syncerus caffer, 1764627; Tapirus terrestris, 443607536; Torgos tracheliotus, 2919708; and Ursus americanus, 8170674.
Raptor populations in Africa are rapidly declining, yet causes of mortality are insufficiently known for most resident species understudied beyond poisoning. This study aimed to investigate the causes of raptor mortality on the African continent. We compiled 1012 mortality records from 43 raptor species tracked with GPS transmitters in Africa between 2003 and 2026 to identify the main causes of mortality, estimate the relative contribution of anthropogenic vs natural causes of mortality, and the factors affecting mortality. Human-induced mortality accounted for 58.8% of known mortality in migratory raptors (n = 233) and 74.3% in non-migratory raptors (n = 241). The main causes of known mortality were direct killing (24.4% of 474 records), poisoning (23.6%), and electrocution (10.1%). Predatory raptors were at higher risk of direct killing than vultures, had higher risk in urbanised than rural landscapes, for non-migrants compared to migrants, and during the non-breeding season compared to the breeding and migration seasons. Vultures had a higher risk of poisoning among adults compared to immatures and in more rural/protected areas relative to urbanised areas. Among poisoned vultures (n = 103), 40.2% were poisoned intentionally. Electrocution occurred at similar rates in vultures and predatory raptors, and more frequently with more urbanisation. The proportion of tracked birds for which cause of mortality could be identified increased over time but remained lower for non-migrants compared to migrants. Our results show the predominantly anthropogenic origin of raptor mortality in Africa, stressing the need for stronger conservation actions directed against direct killing, poisoning, and electrocution.
Physiological parameters have the potential to serve as valuable early warning indicators for the conservation of animal populations. However, measuring physiological adaptations in wildlife is often challenging, due to intrinsic differences causing natural variations of physiological measures between individuals across species. This study is aimed at addressing this by investigating the influence of intrinsic factors, including sex, age, body condition, and the incubation of eggs on the H/L ratio of a forest bird community. As physiological measure, we used the heterophil to lymphocyte (H/L) ratio of individuals belonging to different species in the forest bird community, which was assessed using a novel deep learning approach based on convolutional neural networks (CNNs) applied to whole blood smear scans. Using phylogenetically controlled Bayesian analyses across the bird species, we found higher H/L ratios in adult birds than in juveniles and observed slightly higher H/L ratios in females than in males. While body condition had no effect on the H/L ratio, incubating birds tended to have higher H/L ratios than non‐reproductive birds, regardless of their sex. Furthermore, we found a robust phylogenetic signal of the H/L ratio in the studied bird community. Our results reveal significant general patterns of the effect of intrinsic factors on the H/L ratio across a bird community.
1. Comprehensive estimations of relative abundances and changes therein are one of the most important measures of conservation monitoring. The use of passive acoustic monitoring (PAM) has the potential to greatly enhance information availability, yet has mainly focused on species diversity or coverage, yielding results comparable to those from expert observers. Comparisons with conventional monitoring data have been made mainly for point counts, even though area-wide territory mapping is a widely used standard in many monitoring programs.2. We compare data derived from the combination of PAM and automated identification via machine learning with an area-wide conventional breeding bird survey conducted by expert observers across the Marburg Open Forest in Hesse, Germany. By varying the number of survey intervals, recording duration and locations, we then determined the sampling effort needed to adequately reflect both species coverage and community composition of the conventional survey.3. Only small subsamples of PAM data were required to reach maximum species richness of the conventional survey; minimum requirements were as low as 1) one survey interval or 2) 30 min duration or 3) three recording locations if other factors were allowed to vary accordingly. Revealing the community composition of the conventional survey, however, required sampling over three to six survey intervals with recording durations between 10 and 100 h. While communities were similar between methods in terms of species activity and relative abundance, PAM also partially reflected the composition of territories in the conventional survey.4. Our study demonstrates the relative importance of greater sampling effort especially for monitoring community composition, requiring more recording locations, duration and intervals in comparison to species richness. We provide insight into the current applicability of PAM in monitoring practice and present best-use scenarios on how to make the most of high spatio-temporal resolution acoustic monitoring data.
Long-distance seed dispersal plays a crucial role in shaping plant distributions. The annual migration of millions of frugivorous birds holds substantial potential for the long-distance dispersal of fleshy-fruited plants, which are a major component of forests and woodlands globally. However, quantitative estimations of seed-dispersal distances mediated by migrant frugivorous birds are still lacking. Here, we uniquely combine GPS-tracked movement data of thrushes migrating across Europe (n = 38 individuals) with empirical data on fruit consumption, flight departure time and flight speed, and seed retention times in the birds' gut to mechanistically simulate dispersal events. Our results reveal that, during migratory flights, frugivorous birds can disperse seeds over tens to hundreds of kilometres, depending on the plant species (range across species: mean = 9-45 km; percentile P99.9 = 42-221 km; max. = 58-521 km). In contrast, most dispersal distances (P99.9) by non-migratory movements were within 1 km. Smaller-seeded species disperse farther owing to their longer seed retention times. Our study fills an important knowledge gap on the large-scale dispersal of fleshy-fruited plants by migratory birds and uncovers interspecific, trait-based differences in their long-distance dispersal capacity.
Ecosystem functions and services are severely threatened by unprecedented global loss in biodiversity. To counteract these trends, it is essential to develop systems to monitor changes in biodiversity for planning, evaluating, and implementing conservation and mitigation actions. However, the implementation of monitoring systems suffers from a trade-off between grain (i.e., the level of detail), extent (i.e., the number of study sites), and temporal repetition. Here, we present an applied and realized networked sensor system for integrated biodiversity monitoring in the Nature 4.0 project as a solution to these challenges, which considers plants and animals not only as targets of investigation, but also as parts of the modular sensor network by carrying sensors. Our networked sensor system consists of three main closely interlinked components with a modular structure: sensors, data transmission, and data storage, which are integrated into pipelines for automated biodiversity monitoring. We present our own real-world examples of applications, share our experiences in operating them, and provide our collected open data. Our flexible, low-cost, and open-source solutions can be applied for monitoring individual and multiple terrestrial plants and animals as well as their interactions. Ultimately, our system can also be applied to area-wide ecosystem mapping tasks, thereby providing an exemplary cost-efficient and powerful solution for biodiversity monitoring. Building upon our experiences in the Nature 4.0 project, we identified ten key challenges that need to be addressed to better understand and counteract the ongoing loss of biodiversity using networked sensor systems. To tackle these challenges, interdisciplinary collaboration, additional research, and practical solutions are necessary to enhance the capability and applicability of networked sensor systems for researchers and practitioners, ultimately further helping to ensure the sustainable management of ecosystems and the provision of ecosystem services.
Avian blood analysis is a fundamental method for investigating a wide range of topics concerning individual birds and populations of birds. Determining precise blood cell counts helps researchers gain insights into the health condition of birds. For example, the ratio of heterophils to lymphocytes (H/L ratio) is a well-established index for comparing relative stress load. However, such measurements are currently often obtained manually by human experts. In this article, we present a novel approach to automatically quantify avian red and white blood cells in whole slide images. Our approach is based on two deep neural network models. The first model determines image regions that are suitable for counting blood cells, and the second model is an instance segmentation model that detects the cells in the determined image regions. The region selection model achieves up to 97.3% in terms of F1 score (i.e., the harmonic mean of precision and recall), and the instance segmentation model achieves up to 90.7% in terms of mean average precision. Our approach helps ornithologists acquire hematological data from avian blood smears more precisely and efficiently.
Human-induced direct mortality affects huge numbers of birds each year, threatening hundreds of species worldwide. Tracking technologies can be an important tool to investigate temporal and spatial patterns of bird mortality as well as their drivers. We compiled 1704 mortality records from tracking studies across the African-Eurasian flyway for 45 species, including raptors, storks, and cranes, covering the period from 2003 to 2021. Our results show a higher frequency of human-induced causes of mortality than natural causes across taxonomic groups, geographical areas, and age classes. Moreover, we found that the frequency of human-induced mortality remained stable over the study period. From the human-induced mortality events with a known cause (n = 637), three main causes were identified: electrocution (40.5 %), illegal killing (21.7 %), and poisoning (16.3 %). Additionally, combined energy infrastructure-related mortality (i.e., electrocution, power line collision, and wind-farm collision) represented 49 % of all human-induced mortality events. Using a random forest model, the main predictors of human-induced mortality were found to be taxonomic group, geographic location (latitude and longitude), and human footprint index value at the location of mortality. Despite conservation efforts, human drivers of bird mortality in the African-Eurasian flyway do not appear to have declined over the last 15 years for the studied group of species. Results suggest that stronger conservation actions to address these threats across the flyway can reduce their impacts on species. In particular, projected future development of energy infrastructure is a representative example where application of planning, operation, and mitigation measures can enhance bird conservation.
Seed dispersal by frugivores is a fundamental function for plant community dynamics in fragmented landscapes, where forest remnants are typically embedded in a matrix of anthropogenic habitats. Frugivores can mediate both connectivity among forest remnants and plant colonization of the matrix. However, it remains poorly understood how frugivore communities change from forest to matrix due to the loss or replacement of species with traits that are less advantageous in open habitats and whether such changes ultimately influence the composition and traits of dispersed plants via species interactions. Here, we close this gap by using a unique dataset of seed-dispersal networks that were sampled in forest patches and adjacent matrix habitats of seven fragmented landscapes across Europe. We found a similar diversity of frugivores, plants, and interactions contributing to seed dispersal in forest and matrix, but a high turnover (replacement) in all these components. The turnover of dispersed seeds was smaller than that of frugivore communities because different frugivore species provided complementary seed dispersal in forest and matrix. Importantly, the turnover involved functional changes toward larger and more mobile frugivores in the matrix, which dispersed taller, larger-seeded plants with later fruiting periods. Our study provides a trait-based understanding of frugivore-mediated seed dispersal through fragmented landscapes, uncovering nonrandom shifts that can have cascading consequences for the composition of regenerating plant communities. Our findings also highlight the importance of forest remnants and frugivore faunas for ecosystem resilience, demonstrating a high potential for passive forest restoration of unmanaged lands in the matrix.
IntroductionParasites play important roles in ecosystems. Through their interactions with host and vector species, they are capable of changing the behavior and population dynamics of their host species, and the shape of entire communities. Over the past years, many studies have acknowledged the role of parasitism for host populations and communities and discovered their important regulatory functions for many vertebrate populations. Although birds are a well-studied group of vertebrates, the infection patterns of endoparasites at the community level are not fully understood. Some bird species and families are known to have a higher susceptibility to certain endoparasites than others, which may be driven by their abundance in the community.MethodsOver the course of four consecutive breeding seasons (2019–2022), we monitored the patterns of endoparasite infections in a bird community of a temperate forest ecosystem. We sampled 483 birds belonging to 29 Palearctic species and investigated the prevalence of blood parasites (haemosporidian parasites) andTrichomonasspp. using molecular methods.ResultsWe found an overall prevalence of 48.1% of haemosporidians belonging to 53 genetic lineages of the three generaHaemoproteus,Leucocytozoon, andPlasmodiumspp. While the bird families Turdidae (94%) and Paridae (76%) showed a high prevalence of haemosporidians, Certhiidae, and Picidae were not infected (0%). Host–parasite network analysis detected high variability in interactions. Infections withTrichomonasspp. were not observed in the forest bird community.DiscussionWe found that the prevalence and lineage diversity of haemosporidian parasites differed between avian families and that the parasite prevalence of a family could not serve as a predictor of lineage diversity. To further assess the consequences of these host–parasite interactions for bird communities, future research should aim to disentangle the infection pathways in different ecosystems while also considering the vector community and environmental factors.
The rapid development of wind energy in southern Africa represents an additional threat to the already fragile populations of African vultures. The distribution of the vulnerable Cape Vulture Gyps coprotheres overlaps considerably with wind energy development areas in South Africa, creating conflicts that can hinder both vulture conservation and sustainable energy development. To help address this conflict and aid in the safe placement of wind energy facilities, we map the utilization distribution (UD) of this species across its distributional range. Using tracking data from 68 Cape Vultures collected over the last 20 years, we develop a spatially explicit habitat use model to estimate the expected UDs around known colonies. Scaling the UDs by the number of vultures expected to use each of the colonies, we estimate the Cape Vulture population utilization distribution (PUD) and determine its exposure to wind farm impacts. To complement our results, we model the probability of a vulture flying within the rotor sweep area of a wind turbine throughout the species range and use this to identify areas that are particularly prone to collisions. Overall, our estimated PUD correlates well with reporting rates of the species from the Southern African Bird Atlas Project, currently used to assess potential overlap between Cape Vultures and wind energy developments, but it adds important benefits, such as providing a spatial gradient of activity estimates over the entire species range. We illustrate the application of our maps by analyzing the exposure of Cape Vultures in the Renewable Energy Development Zones (REDZs) in South Africa. This application is a scalable procedure that can be applied at different planning phases, from strategic, nationwide planning to project-level assessments.
The most basic behavioural states of animals can be described as active or passive. However, while high-resolution observations of activity patterns can provide insights into the ecology of animal species, few methods are able to measure the activity of individuals of small taxa in their natural environment. We present a novel approach in which the automated VHF radio-tracking of small vertebrates fitted with lightweight transmitters (< 0.2 g) is used to distinguish between active and passive behavioural states. A dataset containing > 3 million VHF signals was used to train and test a random forest model in the assignment of either active or passive behaviour to individuals from two forest-dwelling bat species (Myotis bechsteinii (n = 50) and Nyctalus leisleri (n = 20)). The applicability of the model to other taxonomic groups was demonstrated by recording and classifying the behaviour of a tagged bird and by simulating the effect of different types of vertebrate activity with the help of humans carrying transmitters. The random forest model successfully classified the activity states of bats as well as those of birds and humans, although the latter were not included in model training (F-score 0.96–0.98). The utility of the model in tackling ecologically relevant questions was demonstrated in a study of the differences in the daily activity patterns of the two bat species. The analysis showed a pronounced bimodal activity distribution of N. leisleri over the course of the night while the night-time activity of M. bechsteinii was relatively constant. These results show that significant differences in the timing of species activity according to ecological preferences or seasonality can be distinguished using our method. Our approach enables the assignment of VHF signal patterns to fundamental behavioural states with high precision and is applicable to different terrestrial and flying vertebrates. To encourage the broader use of our radio-tracking method, we provide the trained random forest models together with an R-package that includes all necessary data-processing functionalities. In combination with state-of-the-art open-source automated radio-tracking, this toolset can be used by the scientific community to investigate the activity patterns of small vertebrates with high temporal resolution, even in dense vegetation.
Context The assessment of suitable habitat for a species is often evaluated based merely on where the species is most abundant. However, a variety of environmental factors such as temperature, resources and human disturbances might force animals to avoid otherwise high-quality habitats. Additionally, individuals’ capacities to cope with such factors – as reflected in physiological stress responses – may also contribute to avoidance. Combining measures of stress response of individual animals with habitat suitability models may contribute essential insights to the development of species conservation plans. Aims We used a landscape-level approach to assess whether habitat quality, abiotic factors and human disturbance affect the physiological stress response of a forest specialist, the capercaillie (Tetrao urogallus), and to what extent these relationships are determined by variation between individuals. Methods We applied a non-invasive approach by collecting 1096 droppings across the entire species range in two adjacent national parks spanning the border between the Czech Republic and Germany. We measured faecal corticosterone metabolites (FCMs) to evaluate the physiological stress response. To test the influences of temperature, habitat quality, season and sex, as well as forest management and tourism intensity on FCM levels, we used generalised additive mixed effect models (GAMMs). The same models were applied for a reduced subset of randomly selected samples that were attributed to individuals via DNA profiling using 10 microsatellite loci. This approach allowed us to control for possible individual differences (through repeated measures within individual genotypes). Key results Both models revealed decreasing FCM levels with increasing habitat quality and showed no relationship between increasing forestry intensity and FCMs. When accounting for individual identity, we found a significant positive effect from increasing tourism intensity, although this relationship was not significant in the full dataset. Conclusions Our results highlight three conclusions for the conservation of capercaillie: (1) high-quality habitats reduce stress levels; (2) tourism-free refuge areas are important, and (3) individuality appears to be a strong determinant of response to environmental factors. Implications Generalising management recommendations from observations for single individuals to an entire population may be inappropriate.
Calls for urgent action to conserve biodiversity under global change are increasing, and conservation of migratory species in this context poses special challenges. In the last two decades the Convention on the Conservation of Migratory Species of Wild Animals (CMS) has provided a framework for several subsidiary instruments including action plans for migratory bird species, but the effectiveness and transferability of these plans remain unclear. Such laws and policies have been credited with positive outcomes for the conservation of migratory species, but the lack of international coordination and on-ground implementation pose major challenges. While research on migratory populations has received growing attention, considerably less emphasis has been given to integrating ecological information throughout the annual cycle for examining strategies to conserve migratory species at multiple scales in the face of global change. We fill this gap through a case study examining the ecological status and conservation of a migratory raptor and facultative scavenger, the red kite (Milvus milvus), whose current breeding range is limited to Europe and is associated with agricultural landscapes and restricted to the temperate zone. Based on our review, conservation actions have been successful at recovering red kite populations within certain regions. Populations however remain depleted along the southern-most edge of the geographic range where many migratory red kites from northern strongholds overwinter. This led us to a forward-looking and integrated strategy that emphasizes international coordination involving researchers and conservation practitioners to enhance the science-policy-action interface. We identify and explore key issues for conserving the red kite under global change, including enhancing conservation actions within and outside protected areas, recovering depleted populations, accounting for climate change, and transboundary coordination in adaptive conservation and management actions. The integrated conservation strategy is sufficiently general such that it can be adapted to inform conservation of other highly mobile species subject to global change.
We present Bird@Edge, an Edge AI system for recognizing bird species in audio recordings to support real-time biodiversity monitoring. Bird@Edge is based on embedded edge devices operating in a distributed system to enable efficient, continuous evaluation of sound-scapes recorded in forests. Multiple ESP32-based microphones (called Bird@Edge Mics) stream audio to a local Bird@Edge Station, on which bird species recognition is performed. The results of several Bird@Edge Stations are transmitted to a backend cloud for further analysis, e.g., by biodiversity researchers. To recognize bird species in soundscapes, a deep neural network based on the EfficientNet-B3 architecture is trained and optimized for execution on embedded edge devices and deployed on a NVIDIA Jetson Nano board using the DeepStream SDK. Our experiments show that our deep neural network outperforms the state-of-the-art BirdNET neural network on several data sets and achieves a recognition quality of up to 95.2% mean average precision on soundscape recordings in the Marburg Open Forest, a research and teaching forest of the University of Marburg, Germany. Measurements of the power consumption of the Bird@Edge components highlight the real-world applicability of the approach. All software and firmware components of Bird@Edge are available under open source licenses.
Migration is used by many species as a strategy to deal with a seasonally changing environment. For some species, migration patterns can vary across different or even within the same breeding area. The Common Woodpigeon Columba palumbus, an abundant and widespread Palearctic species, exhibits three migratory strategies (strictly migratory, partially migratory and resident) across its European breeding grounds. Based on ring recoveries and satellite tracking data, we investigated the migration and foraging behaviour of Woodpigeons breeding in Southwestern Europe (Portugal) and Central Europe (Germany). We found that individuals could be classified as residents (Portugal) or partial migrants (Germany), with migrating individuals following the European sector of the East Atlantic flyway, and mainly wintering in France. In addition to general data on migration phenology, we provide evidence for different migration strategies (migration of varying distances or resident behaviour), low wintering site fidelity and the use of multiple wintering sites. Furthermore, tracking data provided information on migratory behaviour in consecutive years, clearly showing that individuals may switch migratory strategies (resident vs. migrant) between years, i.e. are facultative partial migrants. While individuals from Portugal mainly stayed within a large park ('green urban area') year-round, Woodpigeons from the city of Giessen (Germany) regularly left the urban area to forage on surrounding farmland (with an average distance covered of 5.7 km), particularly from July to September. Overall, our results highlight the behavioural plasticity in Woodpigeons in terms of foraging and migration strategies within and amongst individuals as well as populations.
Protected areas are intended as tools in reducing threats to wildlife and preserving habitat for their long-term population persistence. Studies on ranging behavior provide insight into the utility of protected areas. Vultures are one of the fastest declining groups of birds globally and are popular subjects for telemetry studies, but continent-wide studies are lacking. To address how vultures use space and identify the areas and location of possible vulture safe zones, we assess home range size and their overlap with protected areas by species, age, breeding status, season, and region using a large continent-wide telemetry datasets that includes 163 individuals of three species of threatened Gyps vulture. Immature vultures of all three species had larger home ranges and used a greater area outside of protected areas than breeding and non-breeding adults. Cape vultures had the smallest home range sizes and the lowest level of overlap with protected areas. Ruppell's vultures had larger home range sizes in the wet season, when poisoning may increase due to human-carnivore conflict. Overall, our study suggests challenges for the creation of Vulture Safe Zones to protect African vultures. At a minimum, areas of 24,000 km2 would be needed to protect the entire range of an adult African White-backed vulture and areas of more than 75,000 km2 for wider-ranging Ruppell's vultures. Vulture Safe Zones in Africa would generally need to be larger than existing protected areas, which would require widespread conservation activities outside of protected areas to be successful.
Intrinsic and extrinsic drivers shape the space use of wide‐ranging raptors. A large proportion of raptors are migrants that shift their activity ranges between summer and winter habitats, where they encounter different environmental conditions. Analysing the effects of intrinsic and extrinsic drivers on the space use in summer and winter habitats provides crucial insights into the ecology of migratory raptors. Here, we investigated the seasonal space use by 43 red kites Milvus milvus tracked by GPS‐transmitters across central and south‐western Europe over seven consecutive years. We compared space use patterns, i.e. activity range sizes and mean daily distances of the birds between summer and winter, and analysed the influence of extrinsic (landscape diversity, primary productivity) and intrinsic factors (sex). Within summer, we investigated the influence of breeding success and sex on activity range sizes. We further analysed differences in habitat availability and habitat selection between seasons. We found that space use was smaller in summers than in winters. When compared to those of males, activity ranges of female red kites were larger in summers and smaller in winters, with shorter mean daily distances in both seasons. Within summer, successfully breeding red kites had smaller activity range sizes in both sexes, but this effect was stronger in females than in males. Regardless of the season, landscape diversity was positively correlated with space use, whereas primary productivity was negatively correlated with it. The habitat use differed between seasons, with agricultural landscapes being less proportionally used in summers than in winters. Overall, we showed that both intrinsic and extrinsic drivers shaped space use in both seasons, leading to differences in space use patterns and habitat use in migratory raptors between their summer and winter habitats. Our findings underline the importance of consideration of the entire annual cycle of migratory species for conservation management.