Back and forth: day–night alternation between cover types reveals complementary use of habitats in a large herbivore

Context The Complementary Habitat Hypothesis posits that animals access resources for different needs by moving between complementary habitats that can be seen as ‘resource composites’. These movements can occur over a range of temporal scales, from diurnal to seasonal, in response to multiple drivers such as access to food, weather constraints, risk avoidance and human disturbance. Within this framework, we hypothesised that large herbivores cope with human-altered landscapes through the alternate use of complementary habitats at both daily and seasonal scales. Objectives We tested the Complementary Habitat Hypothesis in European roe deer (Capreolus capreolus) by classifying 3900 habitat-annotated movement trajectories of 154 GPS-monitored individuals across contrasting landscapes. Methods We considered day-night alternation between open food-rich and closed refuge habitats as a measure of complementary habitat use. We first identified day–night alternation using the Individual Movement - Sequence Analysis Method, then we modelled the proportion of day–night alternation over the year in relation to population and individual characteristics. Results We found that day-night alternation is a widespread behaviour in roe deer, even across markedly different landscapes. Day–night alternation followed seasonal trends in all populations, partly linked to vegetation phenology. Within populations, seasonal patterns of open/closed habitat alternation differed between male and female adults, but not in juveniles. Conclusion Our results support the Complementary Habitat Hypothesis by showing that roe deer adjust their access to the varied resources available in complex landscapes by including different habitats within their home range, and sequentially alternating between them in response to seasonal changes and individual life history.publishedVersio

Back and forth: day–night alternation between cover types reveals complementary use of habitats in a large herbivore

Context The Complementary Habitat Hypothesis posits that animals access resources for different needs by moving between complementary habitats that can be seen as ‘resource composites’. These movements can occur over a range of temporal scales, from diurnal to seasonal, in response to multiple drivers such as access to food, weather constraints, risk avoidance and human disturbance. Within this framework, we hypothesised that large herbivores cope with human-altered landscapes through the alternate use of complementary habitats at both daily and seasonal scales. Objectives We tested the Complementary Habitat Hypothesis in European roe deer (Capreolus capreolus) by classifying 3900 habitat-annotated movement trajectories of 154 GPS-monitored individuals across contrasting landscapes. Methods We considered day-night alternation between open food-rich and closed refuge habitats as a measure of complementary habitat use. We first identified day–night alternation using the Individual Movement - Sequence Analysis Method, then we modelled the proportion of day–night alternation over the year in relation to population and individual characteristics. Results We found that day-night alternation is a widespread behaviour in roe deer, even across markedly different landscapes. Day–night alternation followed seasonal trends in all populations, partly linked to vegetation phenology. Within populations, seasonal patterns of open/closed habitat alternation differed between male and female adults, but not in juveniles. Conclusion Our results support the Complementary Habitat Hypothesis by showing that roe deer adjust their access to the varied resources available in complex landscapes by including different habitats within their home range, and sequentially alternating between them in response to seasonal changes and individual life history

Settle Down! Ranging Behaviour Responses of Roe Deer to Different Capture and Release Methods

Simple Summary The study of animal movement in wild, free ranging species is fundamental for advancing knowledge on ecosystem relationships and for conservation. The deployment of bio-logging devices to this purpose (often GPS-collars in large mammals) requires relatively invasive procedures, such as capture, handling and release. Capture and manipulation cause behavioural modifications that are largely understudied in wild species and may affect both the welfare of animals and the output of the studies. We evaluated post capture and release ranging behaviour responses of a small deer species (roe deer Capreolus capreolus) for five different capture methods across 14 study sites within the EURODEER collaborative project. Roe deer showed modifications in their movement behaviour, independently of the capture method. However, individuals recovered rapidly, converging towards the average behaviour within a relatively short interval of time (between 10 days and one month), demonstrating a general resilience to such stressful events. We encourage researchers to continually adapt capture and handling methods so as to minimize stress and prioritize animal welfare. The fitting of tracking devices to wild animals requires capture and handling which causes stress and can potentially cause injury, behavioural modifications that can affect animal welfare and the output of research. We evaluated post capture and release ranging behaviour responses of roe deer (Capreolus capreolus) for five different capture methods. We analysed the distance from the centre of gravity and between successive locations, using data from 14 different study sites within the EURODEER collaborative project. Independently of the capture method, we observed a shorter distance between successive locations and contextual shift away from the home range centre of gravity after the capture and release event. However, individuals converged towards the average behaviour within a relatively short space of time (between 10 days and one month). If researchers investigate questions based on the distance between successive locations of the home range, we recommend (1) initial investigation to establish when the animals start to behave normally again or (2) not using the first two to three weeks of data for their analysis. We also encourage researchers to continually adapt methods to minimize stress and prioritize animal welfare wherever possible, according to the Refinement of the Three R's

Settle down! Ranging behaviour responses of roe deer to different capture and release methods

16openInternationalInternational coauthor/editorThe fitting of tracking devices to wild animals requires capture and handling which causes stress and can potentially cause injury, behavioural modifications that can affect animal welfare and the output of research. We evaluated post capture and release ranging behaviour responses of roe deer (Capreolus capreolus) for five different capture methods. We analysed the distance from the centre of gravity and between successive locations, using data from 14 different study sites within the EURODEER collaborative project. Independently of the capture method, we observed a shorter distance between successive locations and contextual shift away from the home range centre of gravity after the capture and release event. However, individuals converged towards the average behaviour within a relatively short space of time (between 10 days and one month). If researchers investigate questions based on the distance between successive locations of the home range, we recommend (1) initial investigation to establish when the animals start to behave normally again or (2) not using the first two to three weeks of data for their analysis. We also encourage researchers to continually adapt methods to minimize stress and prioritize animal welfare wherever possible, according to the Refinement of the Three R’sopenBergvall, Ulrika A; Morellet, Nicolas; Kjellander, Petter; Rauset, Geir R; Groeve, Johannes De; Borowik, Tomasz; Brieger, Falko; Gehr, Benedikt; Heurich, Marco; Hewison, A J Mark; Kröschel, Max; Pellerin, Maryline; Saïd, Sonia; Soennichsen, Leif; Sunde, Peter; Cagnacci, FrancescaBergvall, U.A.; Morellet, N.; Kjellander, P.; Rauset, G.R.; Groeve, J.D.; Borowik, T.; Brieger, F.; Gehr, B.; Heurich, M.; Hewison, A.J.M.; Kröschel, M.; Pellerin, M.; Saïd, S.; Soennichsen, L.; Sunde, P.; Cagnacci, F

Large herbivore migration plasticity along environmental gradients in Europe: life-history traits modulate forage effects

The most common framework under which ungulate migration is studied predicts that it is driven by spatio–temporal variation in plant phenology, yet other hypotheses may explain differences within and between species. To disentangle more complex patterns than those based on single species/ single populations, we quantified migration vari-ability using two sympatric ungulate species differing in their foraging strategy, mating system and physiological constraints due to body size. We related observed variation to a set of hypotheses. We used GPS-collar data from 537 individuals in 10 roe Capreolus capreolus and 12 red deer Cervus elaphus populations spanning environmental gra-dients across Europe to assess variation in migration propensity, distance and tim-ing. Using time-to-event models, we explored how the probability of migration varied in relation to sex, landscape (e.g. topography, forest cover) and temporally-varying environmental factors (e.g. plant green-up, snow cover). Migration propensity varied across study areas. Red deer were, on average, three times more migratory than roe deer (56% versus 18%). This relationship was mainly driven by red deer males which were twice as migratory as females (82% versus 38%). The probability of roe deer migration was similar between sexes. Roe deer (both sexes) migrated earliest in spring. While territorial male roe deer migrated last in autumn, male and female red deer migrated around the same time in autumn, likely due to their polygynous mating system. Plant productivity determined the onset of spring migration in both species, but if plant productivity on winter ranges was sufficiently high, roe deer were less likely to leave. In autumn, migration coincided with reduced plant productivity for both species. This relationship was stronger for red deer. Our results confirm that ungulate migration is influenced by plant phenology, but in a novel way, that these effects appear to be modulated by species-specific traits, especially mating strategies.publishedVersio

Evaluating expert-based habitat suitability information of terrestrial mammals with GPS-tracking data

Aim Macroecological studies that require habitat suitability data for many species often derive this information from expert opinion. However, expert-based information is inherently subjective and thus prone to errors. The increasing availability of GPS tracking data offers opportunities to evaluate and supplement expert-based information with detailed empirical evidence. Here, we compared expert-based habitat suitability information from the International Union for Conservation of Nature (IUCN) with habitat suitability information derived from GPS-tracking data of 1,498 individuals from 49 mammal species. Location Worldwide. Time period 1998-2021. Major taxa studied Forty-nine terrestrial mammal species. Methods Using GPS data, we estimated two measures of habitat suitability for each individual animal: proportional habitat use (proportion of GPS locations within a habitat type), and selection ratio (habitat use relative to its availability). For each individual we then evaluated whether the GPS-based habitat suitability measures were in agreement with the IUCN data. To that end, we calculated the probability that the ranking of empirical habitat suitability measures was in agreement with IUCN's classification into suitable, marginal and unsuitable habitat types. Results IUCN habitat suitability data were in accordance with the GPS data (> 95% probability of agreement) for 33 out of 49 species based on proportional habitat use estimates and for 25 out of 49 species based on selection ratios. In addition, 37 and 34 species had a > 50% probability of agreement based on proportional habitat use and selection ratios, respectively. Main conclusions We show how GPS-tracking data can be used to evaluate IUCN habitat suitability data. Our findings indicate that for the majority of species included in this study, it is appropriate to use IUCN habitat suitability data in macroecological studies. Furthermore, we show that GPS-tracking data can be used to identify and prioritize species and habitat types for re-evaluation of IUCN habitat suitability data

Moving in the anthropocene: global reductions in terrestrial mammalian movements

Animal movement is fundamental for ecosystem functioning and species survival, yet the effects of the anthropogenic footprint on animal movements have not been estimated across species. Using a unique GPS-tracking database of 803 individuals across 57 species, we found that movements of mammals in areas with a comparatively high human footprint were on average one-half to one-third the extent of their movements in areas with a low human footprint. We attribute this reduction to behavioral changes of individual animals and to the exclusion of species with long-range movements from areas with higher human impact. Global loss of vagility alters a key ecological trait of animals that affects not only population persistence but also ecosystem processes such as predator-prey interactions, nutrient cycling, and disease transmission

Behavioral responses of terrestrial mammals to COVID-19 lockdowns

DATA AND MATERIALS AVAILABILITY : The full dataset used in the final analyses (33) and associated code (34) are available at Dryad. A subset of the spatial coordinate datasets is available at Zenodo (35). Certain datasets of spatial coordinates will be available only through requests made to the authors due to conservation and Indigenous sovereignty concerns (see table S1 for more information on data use restrictions and contact information for data requests). These sensitive data will be made available upon request to qualified researchers for research purposes, provided that the data use will not threaten the study populations, such as by distribution or publication of the coordinates or detailed maps. Some datasets, such as those overseen by government agencies, have additional legal restrictions on data sharing, and researchers may need to formally apply for data access. Collaborations with data holders are generally encouraged, and in cases where data are held by Indigenous groups or institutions from regions that are under-represented in the global science community, collaboration may be required to ensure inclusion.COVID-19 lockdowns in early 2020 reduced human mobility, providing an opportunity to disentangle its effects on animals from those of landscape modifications. Using GPS data, we compared movements and road avoidance of 2300 terrestrial mammals (43 species) during the lockdowns to the same period in 2019. Individual responses were variable with no change in average movements or road avoidance behavior, likely due to variable lockdown conditions. However, under strict lockdowns 10-day 95th percentile displacements increased by 73%, suggesting increased landscape permeability. Animals’ 1-hour 95th percentile displacements declined by 12% and animals were 36% closer to roads in areas of high human footprint, indicating reduced avoidance during lockdowns. Overall, lockdowns rapidly altered some spatial behaviors, highlighting variable but substantial impacts of human mobility on wildlife worldwide.The Radboud Excellence Initiative, the German Federal Ministry of Education and Research, the National Science Foundation, Serbian Ministry of Education, Science and Technological Development, Dutch Research Council NWO program “Advanced Instrumentation for Wildlife Protection”, Fondation Segré, RZSS, IPE, Greensboro Science Center, Houston Zoo, Jacksonville Zoo and Gardens, Nashville Zoo, Naples Zoo, Reid Park Zoo, Miller Park, WWF, ZCOG, Zoo Miami, Zoo Miami Foundation, Beauval Nature, Greenville Zoo, Riverbanks zoo and garden, SAC Zoo, La Passarelle Conservation, Parc Animalier d’Auvergne, Disney Conservation Fund, Fresno Chaffee zoo, Play for nature, North Florida Wildlife Center, Abilene Zoo, a Liber Ero Fellowship, the Fish and Wildlife Compensation Program, Habitat Conservation Trust Foundation, Teck Coal, and the Grand Teton Association. The collection of Norwegian moose data was funded by the Norwegian Environment Agency, the German Ministry of Education and Research via the SPACES II project ORYCS, the Wyoming Game and Fish Department, Wyoming Game and Fish Commission, Bureau of Land Management, Muley Fanatic Foundation (including Southwest, Kemmerer, Upper Green, and Blue Ridge Chapters), Boone and Crockett Club, Wyoming Wildlife and Natural Resources Trust, Knobloch Family Foundation, Wyoming Animal Damage Management Board, Wyoming Governor’s Big Game License Coalition, Bowhunters of Wyoming, Wyoming Outfitters and Guides Association, Pope and Young Club, US Forest Service, US Fish and Wildlife Service, the Rocky Mountain Elk Foundation, Wyoming Wild Sheep Foundation, Wild Sheep Foundation, Wyoming Wildlife/Livestock Disease Research Partnership, the US National Science Foundation [IOS-1656642 and IOS-1656527, the Spanish Ministry of Economy, Industry and Competitiveness, and by a GRUPIN research grant from the Regional Government of Asturias, Sigrid Rausing Trust, Batubay Özkan, Barbara Watkins, NSERC Discovery Grant, the Federal Aid in Wildlife Restoration act under Pittman-Robertson project, the State University of New York, College of Environmental Science and Forestry, the Ministry of Education, Youth and Sport of the Czech Republic, the Ministry of Agriculture of the Czech Republic, Rufford Foundation, an American Society of Mammalogists African Graduate Student Research Fund, the German Science Foundation, the Israeli Science Foundation, the BSF-NSF, the Ministry of Agriculture, Forestry and Food and Slovenian Research Agency (CRP V1-1626), the Aage V. Jensen Naturfond (project: Kronvildt - viden, værdier og værktøjer), the Deutsche Forschungsgemeinschaft (DFG, German Research Foundation) under Germany’s Excellence Strategy, National Centre for Research and Development in Poland, the Slovenian Research Agency, the David Shepherd Wildlife Foundation, Disney Conservation Fund, Whitley Fund for Nature, Acton Family Giving, Zoo Basel, Columbus, Bioparc de Doué-la-Fontaine, Zoo Dresden, Zoo Idaho, Kolmården Zoo, Korkeasaari Zoo, La Passarelle, Zoo New England, Tierpark Berlin, Tulsa Zoo, the Ministry of Environment and Tourism, Government of Mongolia, the Mongolian Academy of Sciences, the Federal Aid in Wildlife Restoration act and the Illinois Department of Natural Resources, the National Science Foundation, Parks Canada, Natural Sciences and Engineering Research Council, Alberta Environment and Parks, Rocky Mountain Elk Foundation, Safari Club International and Alberta Conservation Association, the Consejo Nacional de Ciencias y Tecnología (CONACYT) of Paraguay, the Norwegian Environment Agency and the Swedish Environmental Protection Agency, EU funded Interreg SI-HR 410 Carnivora Dinarica project, Paklenica and Plitvice Lakes National Parks, UK Wolf Conservation Trust, EURONATUR and Bernd Thies Foundation, the Messerli Foundation in Switzerland and WWF Germany, the European Union’s Horizon 2020 research and innovation program under the Marie Skłodowska-Curie Actions, NASA Ecological Forecasting Program, the Ecotone Telemetry company, the French National Research Agency, LANDTHIRST, grant REPOS awarded by the i-Site MUSE thanks to the “Investissements d’avenir” program, the ANR Mov-It project, the USDA Hatch Act Formula Funding, the Fondation Segre and North American and European Zoos listed at http://www.giantanteater.org/, the Utah Division of Wildlife Resources, the Yellowstone Forever and the National Park Service, Missouri Department of Conservation, Federal Aid in Wildlife Restoration Grant, and State University of New York, various donors to the Botswana Predator Conservation Program, data from collared caribou in the Northwest Territories were made available through funds from the Department of Environment and Natural Resources, Government of the Northwest Territories. The European Research Council Horizon2020, the British Ecological Society, the Paul Jones Family Trust, and the Lord Kelvin Adam Smith fund, the Tanzania Wildlife Research Institute and Tanzania National Parks. The Eastern Shoshone and Northern Arapahoe Fish and Game Department and the Wyoming State Veterinary Laboratory, the Alaska Department of Fish and Game, Kodiak Brown Bear Trust, Rocky Mountain Elk Foundation, Koniag Native Corporation, Old Harbor Native Corporation, Afognak Native Corporation, Ouzinkie Native Corporation, Natives of Kodiak Native Corporation and the State University of New York, College of Environmental Science and Forestry, and the Slovenia Hunters Association and Slovenia Forest Service. F.C. was partly supported by the Resident Visiting Researcher Fellowship, IMéRA/Aix-Marseille Université, Marseille. This work was partially funded by the Center of Advanced Systems Understanding (CASUS), which is financed by Germany’s Federal Ministry of Education and Research (BMBF) and by the Saxon Ministry for Science, Culture and Tourism (SMWK) with tax funds on the basis of the budget approved by the Saxon State Parliament. This article is a contribution of the COVID-19 Bio-Logging Initiative, which is funded in part by the Gordon and Betty Moore Foundation (GBMF9881) and the National Geographic Society.https://www.science.org/journal/sciencehj2023Mammal Research InstituteZoology and Entomolog

Predator-prey Interactions in a Human-dominated Landscape

Understanding animal space us is a fundamental concern in Ecology. Predator-prey interactions are a behavioral response game where predators try to find prey, and prey are trying to avoid the predators, thereby shaping animal space use and habitat selection. In the process both predator and prey tradeoff between risk avoidance and food acquisition in order to maximize their fitness. As such predator-prey dynamics have fascinated many generations of ecologists. At the same time the impact of human activities on ecosystem processes have become increasingly evident over the years and almost all habitats on earth today have been altered to some degree by humans. As a consequence humans can be considered as ecological players of ecosystem processes that shape the behavior and demography of their commensal species. In the presence of humans large top predators may be downgraded in the food chain or compete with humans for their prey. Furthermore prey species themselves have to tradeoff between risk avoidance towards humans or their natural predators. It was the goal of this thesis to look at various aspects involved in shaping the space use patterns of the Eurasian lynx (Lynx lynx) and its main prey, the European roe deer (Capreolus capreolus) and to characterize the spatiotemporal drivers of habitat selection and risk avoidance of both players in a human dominated environment. By focusing on both predators and prey I identify different key components of the behavioral response game between Eurasian lynx and European roe deer. In the first chapter I investigated how tradeoffs between risk avoidance and resource abundance in human-altered environments affect habitat selection of Eurasian lynx. In particular, I studied how spatial scale interacts with the intrinsic behavioral state of Eurasian lynx in shaping the response to temporal and spatial fluctuations in human activity and prey availability. I show that lynx tradeoff between anthropogenic risk and prey density by using areas of high prey density during times of low human disturbance. This tradeoff differs among behavioral states and is dependent on the spatial scale considered. In the second chapter I focused on the prey and tested how lynx and humans interact in shaping space use patterns of roe deer during the hunting season in fall. Multiple predators of a single prey may positively or negatively interact with each other via their prey. The same may be true for human hunters that compete with natural predators for the same prey. I explored how roe deer tradeoff between hunting and lynx predation risk and whether hunting increases the deer’s susceptibility to lynx predation. I found that roe deer avoid areas of high hunting risk during the hunting season at the expense of higher exposure to lynx predation risk. Furthermore there was strong evidence that lynx increased their hunting effort and hunting success during this time period. The results provided strong evidence that hunters facilitated lynx predation which resulted in superadditive mortality for deer during the hunting season. In environments with limited resources and time antipredator behavior may come at costs that ultimately affect prey demography. The strength of such non-consumptive effects (NCE) largely depends on the hunting mode of the predator. Ambush predators exert stronger NCE than active predators as they provide more persistent cues about predation risk than their widely roaming counterparts. While evidence comes from different taxonomic groups, evidence from large mammalian predator-prey systems is rare and equivocal. In the third chapter I looked at the occurrence and strength of NCE of the ambush predator, the Eurasian lynx on its main prey the European roe deer. According to ecological prediction I found that lynx evoked strong NCE that increased non-lynx-related mortality of roe deer in the study area. In summary, I focused in this thesis on identifying key drivers involved in shaping space use patterns of a large ambush predator and one of its main prey. Furthermore I characterized the spatiotemporal drivers of food acquisition and risk avoidance of both players in a human dominated environment. The presented results illustrate the tradeoffs between risk avoidance and food acquisition that affect decision making of all animals and show the potential costs of these tradeoffs in the form of reduced survival or reproduction. Finally the findings highlight the role of humans as ecological drivers of Ecosystem processes and may help developing efficient wildlife management plans that are needed for the conservation of large predators and their prey alike in human altered environements

Evidence for nonconsumptive effects from a large predator in an ungulate prey?

Predators can indirectly affect prey survival and reproduction by evoking costly antipredator responses. Such nonconsumptive effects may be as strong or stronger than consumptive predator effects. However, evidence for this in large terrestrial vertebrate systems is equivocal and few studies quantify the actual fitness costs of nonconsumptive effects. Here, we investigated whether nonconsumptive effects elicited by Eurasian lynx (Lynx lynx), a large terrestrial predator, reduced survival in an ungulate prey, the European roe deer (Capreolus capreolus). To reveal the behavioral processes underlying nonconsumptive effects, we distinguished between proactive risk avoidance of areas with high lynx encounter probability, and reactive risk avoidance in response to actual lynx encounters and analyzed these responses using step selection functions. We also quantified the consequences of these behaviors for deer survival. Deer reacted differently at day and at night, but avoided high-risk areas proactively during the day and at night in the summer. During a predator encounter, deer increased avoidance of high-risk areas at night but not during the day. Thus, roe deer exhibited a behavioral response race that involved temporally and spatially varying tradeoffs with environmental constraints. We found evidence that nonconsumptive effects of lynx predation risk reduced deer survival and that survival was more sensitive to variation in nonconsumptive effects of lynx than to variation in human proximity. Our findings highlight that nonconsumptive effects may depend on the spatiotemporal distribution of risks and the environmental context, and we discuss how human factors contribute to predator–prey dynamics in human dominated landscapes