The Ideal Free Distribution with travel costs.

This article studies the effect of travel costs on population distribution in a patchy environment. The Ideal Free Distribution with travel costs is defined in the article as the distribution under which it is not profitable for individuals to move, i.e., the movement between patches ceases. It is shown that depending on the travel costs between patches, the Ideal Free Distribution may be unique, there may be infinitely many possible IFDs, or no Ideal Free Distribution exists. In the latter case, animal distribution can converge to an equilibrium of distributional dynamics at which individuals do disperse, but the net movement between patches ceases. Such distributional equilibrium corresponds to balanced dispersal.

Fish microbiota repel ovipositing mosquitoes.

The mere presence of predators causes prey organisms to display predation-avoidance strategies. Predator presence is often communicated through predator-released chemical signals. Ovipositing female mosquitoes of several species are repelled by unknown signals released from larvivorous fish. It was previously suggested that in many cases, a predator's microbiota plays an important role in the release of these signals; however, this mechanism is still poorly understood. In this study, we looked into the effects of the microbiota originating from the larvivorous Gambusia affinis (Baird and Girard) on the oviposition behaviour of gravid female mosquitoes. We used fish with altered microbiota and bacterial isolates in a set of outdoor mesocosm experiments to address this aim. We show that interference with the fish microbiota significantly reduces fish's repellent effect. We further show that the bacterium Pantoea pleuroti, isolated from the skin of the fish, repels oviposition of Culex laticinctus Edwards and Culiseta longiareolata Macquart mosquitoes similarly to the way in which live fish repel them. Our results highlight the importance of bacteria in the interspecies interactions of their hosts. Furthermore, this finding may lead to the development of an ecologically friendly mosquito repellent, that may reduce the use of larvivorous fish for mosquito control.

Intraguild processes drive space-use patterns in a large-bodied marine predator community.

Interspecific interactions, including predator-prey, intraguild predation (IGP) and competition, may drive distribution and habitat use of predator communities. However, elucidating the relative importance of these interactions in shaping predator distributions is challenging, especially in marine communities comprising highly mobile species. We used individual-based models (IBMs) to predict the habitat distributions of apex predators, intraguild (IG) prey and prey. We then used passive acoustic telemetry to test these predictions in a subtropical marine predator community consisting of eight elasmobranch (i.e. shark and ray) species in Bimini, The Bahamas. IBMs predicted that prey and IG prey will preferentially select habitats based on safety over resources (food), with stronger selection for safe habitat by smaller prey. Elasmobranch space-use patterns matched these predictions. Species with predator-prey and asymmetrical IGP (between apex and small mesopredators) interactions showed the clearest spatial separation, followed by asymmetrical IGP among apex and large mesopredators. Competitors showed greater spatial overlap although with finer-scale differences in microhabitat use. Our study suggests space-use patterns in elasmobranchs are at least partially driven by interspecific interactions, with stronger spatial separation occurring where interactions include predator-prey relationships or IGP.

Estimating encounter-habitat relationships with scale-integrated resource selection functions.

Encounters between animals occur when animals are close in space and time. Encounters are important in many ecological processes including sociality, predation and disease transmission. Despite this, there is little theory regarding the spatial distribution of encounters and no formal framework to relate environmental characteristics to encounters. The probability of encounter could be estimated with resource selection functions (RSFs) by comparing locations where encounters occurred to available locations where they may have occurred, but this estimate is complicated by the hierarchical nature of habitat selection. We developed a method to relate resources to the relative probability of encounter based on a scale-integrated habitat selection framework. This framework integrates habitat selection at multiple scales to obtain an appropriate estimate of availability for encounters. Using this approach, we related encounter probabilities to landscape resources. The RSFs describe habitat associations at four scales, home ranges within the study area, areas of overlap within home ranges, locations within areas of overlap, and encounters compared to other locations, which can be combined into a single scale-integrated RSF. We apply this method to intraspecific encounter data from two species: white-tailed deer (Odocoileus virginianus) and elk (Cervus elaphus) and interspecific encounter data from a two-species system of caribou (Rangifer tarandus) and coyote (Canis latrans). Our method produced scale-integrated RSFs that represented the relative probability of encounter. The predicted spatial distribution of encounters obtained based on this scale-integrated approach produced distributions that more accurately predicted novel encounters than a naïve approach or any individual scale alone. Our results highlight the importance of accounting for the conditional nature of habitat selection in estimating the habitat associations of animal encounters as opposed to 'naïve' comparisons of encounter locations with general availability. This method has direct relevance for testing hypotheses about the relationship between habitat and social or predator-prey behaviour and generating spatial predictions of encounters. Such spatial predictions may be vital for understanding the distribution of encounters driving disease transmission, predation rates and other population and community-level processes.

Counter-strategies to infanticide: The importance of cubs in determining lion habitat selection and social interactions.

Animal social and spatial behaviours are inextricably linked. Animal movements are driven by environmental factors and social interactions. Habitat structure and changing patterns of animal space use can also shape social interactions. Animals adjust their social and spatial behaviours to reduce the risk of offspring mortality. In territorial infanticidal species, two strategies are possible for males: they can stay close to offspring to protect them against rivals (infant-defence hypothesis) or patrol the territory more intensively to prevent rival intrusions (territorial-defence hypothesis). Here, we tested these hypotheses in African lions (Panthera leo) by investigating how males and females adjust their social and spatial behaviours in the presence of offspring. We combined datasets on the demography and movement of lions, collected between 2002 and 2016 in Hwange National Park (Zimbabwe), to document the presence of cubs (field observations) and the simultaneous movements of groupmates and competitors (GPS tracking). We showed a spatial response of lions to the presence of offspring, with females with cubs less likely to select areas close to waterholes or in the periphery of the territory than females without cubs. In contrast, these areas were more selected by males when there were cubs in the pride. We also found social responses. Males spent more time with females as habitat openness increased but the presence of cubs in the pride did not influence the average likelihood of observing males with females. Furthermore, rival males relocated further after an encounter with pride males when cubs were present in the prides, suggesting that the presence of cubs leads to a more vigorous repulsion of competitors. Males with cubs in their pride were more likely to interact with male competitors on the edge of the pride's home range and far from the waterholes, suggesting that they are particularly assiduous in detecting and repelling rival males during these periods. In general, the strategies to avoid infanticide exhibited by male lions supported the territorial-defence hypothesis. Our study contributes to answer the recent call for a behavioural ecology at the spatial-social interface.

Insect-mediated apparent competition between mammals in a boreal food web.

While the important role of animal-mediated interactions in the top-down restructuring of plant communities is well documented, less is known of their ensuing repercussions at higher trophic levels. We demonstrate how typically decoupled ecological interactions may become intertwined such that the impact of an insect pest on forest structure and composition alters predator-prey interactions among large mammals. Specifically, we show how irruptions in a common, cyclic insect pest of the boreal forest, the spruce budworm (Choristoneura fumiferana), modulated an indirect trophic interaction by initiating a flush in deciduous vegetation that benefited moose (Alces alces), in turn strengthening apparent competition between moose and threatened boreal caribou (Rangifer tarandus caribou) via wolf (Canis lupus) predation. Critically, predation on caribou postoutbreak was exacerbated by human activity (salvage logging). We believe our observations of significant, large-scale reverberating consumer-producer-consumer interactions are likely to be common in nature.

A shift in habitat use patterns of brown trout (Salmo trutta): A behavioural response to macrophyte removal.

Mass development of macrophytes is an increasing problem worldwide and they are frequently removed where they are in conflict with local waterway users. Yet, macrophytes can provide important refuge and nursery habitats for fish. Little is known about the consequences of macrophyte removal for fish behavioural space use and habitat selection. We hypothesised that macrophyte removal would affect brown trout (Salmo trutta) movement during the partial removal of the aquatic plant Juncus bulbosus (L.) in an oligotrophic impounded Norwegian river.We tagged 94 brown trout and tracked them using passive acoustic telemetry for 10 months and mapped the cover of J. bulbosus. Trout behavioural patterns were quantified as space use (utilisation areas 50% and 95%) which was linked to habitat use and selection for J. bulbosus. Removal of J. bulbosus influenced space use of brown trout by reducing the core utilisation area by 22%. Habitat use and selection were likewise influenced by removal with increased use and selection of areas with low J. bulbosus cover (<25%) with corresponding reduction in high J. bulbosus cover (>25-75%). Finally, diurnal differences in space use and habitat use were found, with 19% larger utilisation areas at night and higher use of areas with low J. bulbosus during daytime. Yet, all effect sizes were relatively small compared to the size of the study area. This research provides a detailed case study on the effects of macrophyte removal on fish behavioural patterns in a section of a large Norwegian river with macrophyte mass development. We found no large effects of removal on trout behaviour but noted an increased use of areas with low macrophyte cover. This research is relevant for water managers and policy makers of freshwater conservation and provides a template for using acoustic telemetry to study the effects of macrophyte removal on fish.

Density-dependent habitat selection alters drivers of population distribution in northern Yellowstone elk.

Although it is well established that density dependence drives changes in organismal abundance over time, relatively little is known about how density dependence affects variation in abundance over space. We tested the hypothesis that spatial trade-offs between food and safety can change the drivers of population distribution, caused by opposing patterns of density-dependent habitat selection (DDHS) that are predicted by the multidimensional ideal free distribution. We addressed this using winter aerial survey data of northern Yellowstone elk (Cervus canadensis) spanning four decades. Supporting our hypothesis, we found positive DDHS for food (herbaceous biomass) and negative DDHS for safety (openness and roughness), such that the primary driver of habitat selection switched from food to safety as elk density decreased from 9.3 to 2.0 elk/km2 . Our results demonstrate how population density can drive landscape-level shifts in population distribution, confounding habitat selection inference and prediction and potentially affecting community-level interactions.

Resource selection by New York City deer reveals the effective interface between wildlife, zoonotic hazards and humans.

Although the role of host movement in shaping infectious disease dynamics is widely acknowledged, methodological separation between animal movement and disease ecology has prevented researchers from leveraging empirical insights from movement data to advance landscape scale understanding of infectious disease risk. To address this knowledge gap, we examine how movement behaviour and resource utilization by white-tailed deer (Odocoileus virginianus) determines blacklegged tick (Ixodes scapularis) distribution, which depend on deer for dispersal in a highly fragmented New York City borough. Multi-scale hierarchical resource selection analysis and movement modelling provide insight into how deer's movements contribute to the risk landscape for human exposure to the Lyme disease vector-I. scapularis. We find deer select highly vegetated and accessible residential properties which support blacklegged tick survival. We conclude the distribution of tick-borne disease risk results from the individual resource selection by deer across spatial scales in response to habitat fragmentation and anthropogenic disturbances.

Coevolution of Patch Selection in Stochastic Environments.

AbstractSpecies interact in landscapes where environmental conditions vary in time and space. This variability impacts how species select habitat patches. Under equilibrium conditions, evolution of this patch selection can result in ideal free distributions where per capita growth rates are zero in occupied patches and negative in unoccupied patches. These ideal free distributions, however, do not explain why species occupy sink patches, why competitors have overlapping spatial ranges, or why predators avoid highly productive patches. To understand these patterns, we solve for coevolutionarily stable strategies (coESSs) of patch selection for multispecies stochastic Lotka-Volterra models accounting for spatial and temporal heterogeneity. In occupied patches at the coESS, we show that the differences between the local contributions to the mean and the variance of the long-term population growth rate are equalized. Applying this characterization to models of antagonistic interactions reveals that environmental stochasticity can partially exorcize the ghost of competition past, select for new forms of enemy-free and victimless space, and generate hydra effects over evolutionary timescales. Viewing our results through the economic lens of modern portfolio theory highlights why the coESS for patch selection is often a bet-hedging strategy coupling stochastic sink populations. Our results highlight how environmental stochasticity can reverse or amplify evolutionary outcomes as a result of species interactions or spatial heterogeneity.

Changes in phenology can alter patterns of natural selection: the joint evolution of germination time and postgermination traits.

The timing of a developmental transition (phenology) can influence the environment experienced by subsequent life stages. When phenology causes an organism to occupy a particular habitat as a consequence of the developmental cues used, it can act as a form of habitat tracking. Evolutionary theory predicts that habitat tracking can alter the strength, direction, and mode of natural selection on subsequently expressed traits. To test whether germination phenology altered natural selection on postgermination traits, we manipulated germination time by planting seedlings in seven germination cohorts spanning 2 yr. We measured selection on postgermination traits relating to drought, freezing, and heat tolerance using a diverse combination of Arabidopsis thaliana mutants and naturally occurring ecotypes. Germination cohorts experienced variable selection: when dry, cold, and hot environments were experienced by seedlings, selection was intensified for drought, freezing, and heat tolerance, respectively. Reciprocally, postgermination traits modified the optimal germination time; genotypes had maximum fitness after germinating in environments that matched their physiological tolerances. Our results support the theoretical predictions of feedbacks between habitat tracking and traits expressed after habitat selection. In natural populations, whether phenological shifts alter selection on subsequently expressed traits will depend on the effectiveness of habitat tracking through phenology.

Habitat fragmentation reduces survival and drives source-sink dynamics for a large carnivore.

Rigorous understanding of how environmental conditions impact population dynamics is essential for species conservation, especially in mixed-use landscapes where source-sink dynamics may be at play. Conservation of large carnivore populations in fragmented, human-dominated landscapes is critical for their long-term persistence. However, living in human-dominated landscapes comes with myriad costs, including direct anthropogenic mortality and sublethal energetic costs. How these costs impact individual fitness and population dynamics are not fully understood, partly due to the difficulty in collecting long-term demographic data for these species. Here, we analyzed an 11-year dataset on puma (Puma concolor) space use, mortality, and reproduction in the Santa Cruz Mountains, California, USA, to quantify how living in a fragmented landscape impacts individual survival and population dynamics. Long-term exposure to housing density drove mortality risk for female pumas, resulting in an 18-percentage-point reduction in annual survival for females in exurban versus remote areas. While the overall population growth rate appeared stable, reduced female survival in more developed areas resulted in source-sink dynamics across the study area, with 42.1% of the Santa Cruz Mountains exhibiting estimated population growth rates <1. Since habitat selection is often used as a proxy for habitat quality, we also assessed whether puma habitat selection predicted source and sink areas. Patterns of daytime puma habitat selection predicted source areas, while time-of-day-independent habitat selection performed less well as a proxy. These results illuminate the individual- and population-level consequences of habitat fragmentation for large carnivores, illustrating that habitat fragmentation can produce source-sink dynamics that may not be apparent from other metrics of habitat quality. Locally, conserving high-quality source habitat within the Santa Cruz Mountains is necessary to support long-term puma population persistence. More broadly, source-sink dynamics may at play for other carnivore populations in similar fragmented systems, and linking landscape conditions to population dynamics is essential for effective conservation. Caution should be used in inferring habitat quality from habitat selection alone, but these results shed light on metrics of selection that may be better or worse proxies to identify source areas for large carnivores.

Effects of vehicle traffic on space use and road crossings of caribou in the Arctic.

Assessing the effects of industrial development on wildlife is a key objective of managers and conservation practitioners. However, wildlife responses are often only investigated with respect to the footprint of infrastructure, even though human activity can strongly mediate development impacts. In Arctic Alaska, there is substantial interest in expanding energy development, raising concerns about the potential effects on barren-ground caribou (Rangifer tarandus granti). While caribou generally avoid industrial infrastructure, little is known about the role of human activity in moderating their responses, and whether managing activity levels could minimize development effects. To address this uncertainty, we examined the influence of traffic volume on caribou summer space use and road crossings in the Central Arctic Herd within the Kuparuk and Milne Point oil fields on the North Slope of Alaska. We first modeled spatiotemporal variation in hourly traffic volumes across the road system from traffic counter data using gradient-boosted regression trees. We then used generalized additive models to estimate nonlinear step selection functions and road-crossing probabilities from collared female caribou during the post-calving and insect harassment seasons, when they primarily interact with roads. Step selection analyses revealed that caribou selected areas further from roads (~1-3 km) during the post-calving and mosquito seasons and selected areas with lower traffic volumes during all seasons, with selection probabilities peaking when traffic was <5 vehicles/h. Using road-crossing models, we found that caribou were less likely to cross roads during the insect seasons as traffic increased, but that response dissipated as insect harassment became more severe. Past studies suggested that caribou exhibit behavioral responses when traffic exceeds 15 vehicles/h, but our results demonstrate behavioral responses at much lower traffic levels. Our results illustrate that vehicle activity mediates caribou responses to road infrastructure, information that can be used in future land-use planning to minimize the behavioral responses of caribou to industrial development in sensitive Arctic landscapes.

Habitat detection, habitat choice copying or mating benefits: What drives conspecific attraction in a nomadic songbird?

Conspecific attraction during habitat selection is common among animals, but the ultimate (i.e. fitness-related) reasons for this behaviour often remain enigmatic. We aimed to evaluate the following three hypotheses for conspecific attraction during the breeding season in male Wood Warblers (Phylloscopus sibilatrix): the habitat detection hypothesis, the habitat choice copying hypothesis and the female preference hypothesis. These hypotheses make different predictions with respect to the relative importance of social and nonsocial information during habitat assessment, and whether benefits accrue as a consequence of aggregation. We tested the above hypotheses using a combination of a 2-year playback experiment, spatial statistics and mate choice models. The habitat detection hypothesis was the most likely explanation for conspecific attraction and aggregation in male Wood Warblers, based on the following results: (1) males were attracted to conspecific song playbacks, but fine-scale habitat heterogeneity was the better predictor of spatial patterns in the density of settling males; (2) male pairing success did not increase, but instead slightly decreased, as connectivity with other males (i.e. the number and proximity of neighbouring males) increased. Our study highlights how consideration of the process by which animals detect and assess habitat, together with the potential fitness consequences of resulting aggregations, are important for understanding conspecific attraction and spatially clustered distributions.

LiDAR reveals a preference for intermediate visibility by a forest-dwelling ungulate species.

Visibility (viewshed) plays a significant and diverse role in animals' behaviour and fitness. Understanding how visibility influences animal behaviour requires the measurement of habitat visibility at spatial scales commensurate to individual animal choices. However, measuring habitat visibility at a fine spatial scale over a landscape is a challenge, particularly in highly heterogeneous landscapes (e.g. forests). As a result, our ability to model the influence of fine-scale visibility on animal behaviour has been impeded or limited. In this study, we demonstrate the application of the concept of three-dimensional (3D) cumulative viewshed in the study of animal spatial behaviour at a landscape level. Specifically, we employed a newly described approach that combines terrestrial and airborne light detection and ranging (LiDAR) to measure fine-scale habitat visibility (3D cumulative viewshed) on a continuous scale in forested landscapes. We applied the LiDAR-derived visibility to investigate how visibility in forests affects the summer habitat selection and the movement of 20 GPS-collared female red deer Cervus elaphus in a temperate forest in Germany. We used integrated step selection analysis to determine whether red deer show any preference for fine-scale habitat visibility and whether visibility is related to the rate of movement of red deer. We found that red deer selected intermediate habitat visibility. Their preferred level of visibility during the day was substantially lower than that of night and twilight, whereas the preference was not significantly different between night and twilight. In addition, red deer moved faster in high-visibility areas, possibly mainly to avoid predation and anthropogenic risk. Furthermore, red deer moved most rapidly between locations in the twilight. For the first time, the preference for intermediate habitat visibility and the adaption of movement rate to fine-scale visibility by a forest-dwelling ungulate species at a landscape scale was revealed. The LiDAR technique used in this study offers fine-scale habitat visibility at the landscape level in forest ecosystems, which would be of broader interest in the fields of animal ecology and behaviour.

A life-history spectrum of population responses to simultaneous change in climate and land use.

Climate and land use change are two of the primary threats to global biodiversity; however, each species within a community may respond differently to these facets of global change. Although it is typically assumed that species use the habitat that is advantageous for survival and reproduction, anthropogenic changes to the environment can create ecological traps, making it critical to assess both habitat selection (e.g. where species congregate on the landscape) and the influence of selected habitats on the demographic processes that govern population dynamics. We used a long-term (1958-2011), large-scale, multi-species dataset for waterfowl that spans the United States and Canada to estimate species-specific responses to climate and land use variables in a landscape that has undergone significant environmental change across space and time. We first estimated the effects of change in climate and land use variables on habitat selection and population dynamics for nine species. We then hypothesized that species-specific responses to environmental change would scale with life-history traits, specifically: longevity, nesting phenology and female breeding site fidelity. We observed species-level heterogeneity in the demographic and habitat selection responses to climate and land use change, which would complicate community-level habitat management. Our work highlights the importance of multi-species monitoring and community-level analysis, even among closely related species. We detected several relationships between life-history traits, particularly nesting phenology, and species' responses to environmental change. One species, the early-nesting northern pintail (Anas acuta), was consistently at the extreme end of responses to land use and climate predictors and has been a species of conservation concern since their population began to decline in the 1980s. They, and the blue-winged teal, also demonstrated a positive habitat selection response to the proportion of cropland on the landscape that simultaneously reduced abundance the following year, indicative of susceptibility to ecological traps. By distilling the diversity of species' responses to environmental change within a community, our methodological approach and findings will help improve predictions of community responses to global change and can inform multi-species management and conservation plans in dynamic landscapes that are based on simple tenets of life-history theory.

Accounting for behaviour in fine-scale habitat selection: A case study highlighting methodological intricacies.

Animal habitat selection-central in both theoretical and applied ecology-may depend on behavioural motivations such as foraging, predator avoidance, and thermoregulation. Step-selection functions (SSFs) enable assessment of fine-scale habitat selection as a function of an animal's movement capacities and spatiotemporal variation in extrinsic conditions. If animal location data can be associated with behaviour, SSFs are an intuitive approach to quantify behaviour-specific habitat selection. Fitting SSFs separately for distinct behavioural states helped to uncover state-specific selection patterns. However, while the definition of the availability domain has been highlighted as the most critical aspect of SSFs, the influence of accounting for behaviour in the use-availability design has not been quantified yet. Using a predator-free population of high-arctic muskoxen Ovibos moschatus as a case study, we aimed to evaluate how (1) defining behaviour-specific availability domains, and/or (2) fitting separate behaviour-specific models impacts (a) model structure, (b) estimated selection coefficients and (c) model predictive performance as opposed to behaviour-unspecific approaches. To do so, we first applied hidden Markov models to infer different behavioural modes (resting, foraging, relocating) from hourly GPS positions (19 individuals, 153-1062 observation days/animal). Using SSFs, we then compared behaviour-specific versus behaviour-unspecific habitat selection in relation to terrain features, vegetation and snow conditions. Our results show that incorporating behaviour into the definition of the availability domain primarily impacts model structure (i.e. variable selection), whereas fitting separate behaviour-specific models mainly influences selection strength. Behaviour-specific availability domains improved predictive performance for foraging and relocating models (i.e. behaviours with medium to large spatial displacement), but decreased performance for resting models. Thus, even for a predator-free population subject to only negligible interspecific competition and human disturbance we found that accounting for behaviour in SSFs impacted model structure, selection coefficients and predictive performance. Our results indicate that for robust inference, both a behaviour-specific availability domain and behaviour-specific model fitting should be explored, especially for populations where strong spatiotemporal selection trade-offs are expected. This is particularly critical if wildlife habitat preferences are estimated to inform management and conservation initiatives.

Carrying eggs uphill: are costs of reproduction stronger on steeper slopes?

Locomotor impairment during pregnancy is a well-documented cost of reproduction, but most empirical studies have not incorporated ecological complexity, such as locomotion on sloping inclines rather than horizontal surfaces. Biomechanical factors suggest that carrying a heavy burden-including shifts in the body's centre of mass-may impair locomotor ability even more when an animal is running uphill. If so, then measuring costs of reproduction on horizontal racetracks may underestimate these costs in nature for arboreal species. To evaluate this prediction, we measured the pregnancy-induced reduction in speed for jacky dragons (Amphibolurus muricatus) at inclines ranging from 0 to 45°. Both pregnancy and steeper slopes reduced lizard performance, but pregnancy did not exacerbate the locomotor decrement on steeper racetracks. An ability to maintain mobility on steep slopes during pregnancy may be a target of selection in arboreal taxa. To understand the evolutionary context of locomotion-based costs of reproduction, we also need studies on the relationship between organismal performance and ecologically relevant measures such as predation risk.

Using high-resolution LiDAR-derived canopy structure and topography to characterise hibernaculum locations of the hazel dormouse.

The hazel dormouse is predominantly an arboreal species that moves down to the ground to hibernate in the autumn in temperate parts of its distributional ranges at locations not yet well understood. The main objective of this study is to test whether environmental characteristics surrounding hazel dormouse hibernacula can be identified using high-resolution remote sensing and data collected in situ. To achieve this, remotely sensed variables, including canopy height and cover, topographic slope, sky view, solar radiation and cold air drainage, were modelled around 83 dormouse hibernacula in England (n = 62) and the Netherlands (n = 21), and environmental characteristics that may be favoured by pre-hibernating dormice were identified. Data on leaf litter depth, temperature, canopy cover and distance to the nearest tree were collected in situ and analysed at hibernaculum locations in England. The findings indicated that remotely sensed data were effective in identifying attributes surrounding the locations of dormouse hibernacula and when compared to in situ information, provided more conclusive results. This study suggests that remotely sensed topographic slope, canopy height and sky view have an influence on hazel dormice choosing suitable locations to hibernate; whilst in situ data suggested that average daily mean temperature at the hibernaculum may also have an effect. Remote sensing proved capable of identifying localised environmental characteristics in the wider landscape that may be important for hibernating dormice. This study proposes that this method can provide a novel progression from habitat modelling to conservation management for the hazel dormouse, as well as other species using habitats where topography and vegetation structure influence fine-resolution favourability.

Hawk owl irruptions: spatial and temporal variation in rodent abundance drive push and pull dynamics.

Bird irruptions are thought to be triggered by low food availability in breeding areas, thereby causing emigration (push factor). However, few studies have tested whether emigrating individuals are drawn towards areas of high food availability (pull factor). The Northern hawk owl (Surnia ulula), a rodent specialist, occurs irruptively to southern parts of Fennoscandia. We analysed whether irruption size during 1980-2020 in southeastern Norway was related to rodent abundance at four sites 450-990 km to the north-northeast (potential source areas) and at two sites in southeastern Norway to test push and pull dynamics of irruptions. Irruptions occurred when rodent abundance in potential source areas were low, supporting the push hypothesis. High rodent abundance in potential source areas 1-2 years before irruptions suggested that irruptions were preceded by high reproduction. Upon arrival to southeastern Norway, hawk owls did not encounter high rodent abundance in their main habitat (boreal forest). However, hawk owls stayed in boreal forest in hills in years with higher microtine rodent abundance, but occurred in farmland areas in the lowlands when microtine rodents were less abundant. Use of lowlands coincided with higher than median numbers of wood mouse (Apodemus sylvaticus) for 87% of the hawk owls settling in the lowlands, thus suggesting support for the pull hypothesis. In conclusion, hawk owl irruptions to southern Fennoscandia were triggered by low food availability in northern areas (push factor), and appeared to be drawn by high food availability in southeastern Norway to some degree (pull factor).