Predicting kill sites of an apex predator from GPS data in different multiprey systems.

Kill rates are a central parameter to assess the impact of predation on prey species. An accurate estimation of kill rates requires a correct identification of kill sites, often achieved by field-checking GPS location clusters (GLCs). However, there are potential sources of error included in kill-site identification, such as failing to detect GLCs that are kill sites, and misclassifying the generated GLCs (e.g., kill for nonkill) that were not field checked. Here, we address these two sources of error using a large GPS dataset of collared Eurasian lynx (Lynx lynx), an apex predator of conservation concern in Europe, in three multiprey systems, with different combinations of wild, semidomestic, and domestic prey. We first used a subsampling approach to investigate how different GPS-fix schedules affected the detection of GLC-indicated kill sites. Then, we evaluated the potential of the random forest algorithm to classify GLCs as nonkills, small prey kills, and ungulate kills. We show that the number of fixes can be reduced from seven to three fixes per night without missing more than 5% of the ungulate kills, in a system composed of wild prey. Reducing the number of fixes per 24 h decreased the probability of detecting GLCs connected with kill sites, particularly those of semidomestic or domestic prey, and small prey. Random forest successfully predicted between 73%-90% of ungulate kills, but failed to classify most small prey in all systems, with sensitivity (true positive rate) lower than 65%. Additionally, removing domestic prey improved the algorithm's overall accuracy. We provide a set of recommendations for studies focusing on kill-site detection that can be considered for other large carnivore species in addition to the Eurasian lynx. We recommend caution when working in systems including domestic prey, as the odds of underestimating kill rates are higher.

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.

Body size and digestive system shape resource selection by ungulates: A cross-taxa test of the forage maturation hypothesis.

The forage maturation hypothesis (FMH) states that energy intake for ungulates is maximised when forage biomass is at intermediate levels. Nevertheless, metabolic allometry and different digestive systems suggest that resource selection should vary across ungulate species. By combining GPS relocations with remotely sensed data on forage characteristics and surface water, we quantified the effect of body size and digestive system in determining movements of 30 populations of hindgut fermenters (equids) and ruminants across biomes. Selection for intermediate forage biomass was negatively related to body size, regardless of digestive system. Selection for proximity to surface water was stronger for equids relative to ruminants, regardless of body size. To be more generalisable, we suggest that the FMH explicitly incorporate contingencies in body size and digestive system, with small-bodied ruminants selecting more strongly for potential energy intake, and hindgut fermenters selecting more strongly for surface water.

Harvest models of small populations of a large carnivore using Bayesian forecasting.

Harvesting large carnivores can be a management tool for meeting politically set goals for their desired abundance. However, harvesting carnivores creates its own set of conflicts in both society and among conservation professionals, where one consequence is a need to demonstrate that management is sustainable, evidence-based, and guided by science. Furthermore, because large carnivores often also have high degrees of legal protection, harvest quotas have to be carefully justified and constantly adjusted to avoid damaging their conservation status. We developed a Bayesian state-space model to support adaptive management of Eurasian lynx harvesting in Scandinavia. The model uses data from the annual monitoring of lynx abundance and results from long-term field research on lynx biology, which has provided detailed estimates of key demographic parameters. We used the model to predict the probability that the forecasted population size will be below or above the management objectives when subjected to different harvest quotas. The model presented here informs decision makers about the policy risks of alternative harvest levels. Earlier versions of the model have been available for wildlife managers in both Sweden and Norway to guide lynx harvest quotas and the model predictions showed good agreement with observations. We combined monitoring data with data on vital rates and were able to estimate unobserved additional mortality rates, which are most probably due to poaching. In both countries, the past quota setting strategy suggests that there has been a de facto threshold strategy with increasing proportion, which means that there is no harvest below a certain population size, but above this threshold there is an increasing proportion of the population harvested as the population size increases. The annual assessment of the monitoring results, the use of forecasting models, and a threshold harvest approach to quota setting will all reduce the risk of lynx population sizes moving outside the desired goals. The approach we illustrate could be adapted to other populations of mammals worldwide.

Ungulate management in European national parks: Why a more integrated European policy is needed.

1. Primary objectives of national parks usually include both, the protection of natural processes and species conservation. When these objectives conflict, as occurs because of the cascading effects of large mammals (i.e., ungulates and large carnivores) on lower trophic levels, park managers have to decide upon the appropriate management while considering various local circumstances. 2. To analyse if ungulate management strategies are in accordance with the objectives defined for protected areas, we assessed the current status of ungulate management across European national parks using the naturalness concept and identified the variables that influence the management. 3. We collected data on ungulate management from 209 European national parks in 29 countries by means of a large-scale questionnaire survey. Ungulate management in the parks was compared by creating two naturalness scores. The first score reflects ungulate and large carnivore species compositions, and the second evaluates human intervention on ungulate populations. We then tested whether the two naturalness score categories are influenced by the management objectives, park size, years since establishment, percentage of government-owned land, and human impact on the environment (human influence index) using two generalized additive mixed models. 4. In 67.9% of the national parks, wildlife is regulated by culling (40.2%) or hunting (10.5%) or both (17.2%). Artificial feeding occurred in 81.3% of the national parks and only 28.5% of the national parks had a non-intervention zone covering at least 75% of the area. Furthermore, ungulate management differed greatly among the different countries, likely because of differences in hunting traditions and cultural and political backgrounds. Ungulate management was also influenced by park size, human impact on the landscape, and national park objectives, but after removing these variables from the full model the reduced models only showed a small change in the deviance explained. In areas with higher anthropogenic pressure, wildlife diversity tended to be lower and a higher number of domesticated species tended to be present. Human intervention (culling and artificial feeding) was lower in smaller national parks and when park objectives followed those set by the International Union for the Conservation of Nature (IUCN). 5. Our study shows that many European national parks do not fulfil the aims of protected area management as set by IUCN guidelines. In contrast to the USA and Canada, Europe currently has no common ungulate management policy within national parks. This lack of a common policy together with differences in species composition, hunting traditions, and cultural or political context has led to differences in ungulate management among European countries. To fulfil the aims and objectives of national parks and to develop ungulate management strategies further, we highlight the importance of creating a more integrated European ungulate management policy to meet the aims of national parks.

Border Security Fencing and Wildlife: The End of the Transboundary Paradigm in Eurasia?

The ongoing refugee crisis in Europe has seen many countries rush to construct border security fencing to divert or control the flow of people. This follows a trend of border fence construction across Eurasia during the post-9/11 era. This development has gone largely unnoticed by conservation biologists during an era in which, ironically, transboundary cooperation has emerged as a conservation paradigm. These fences represent a major threat to wildlife because they can cause mortality, obstruct access to seasonally important resources, and reduce effective population size. We summarise the extent of the issue and propose concrete mitigation measures.

Mainstreaming human and large carnivore coexistence through institutional collaboration.

Achieving coexistence between large carnivores and humans in human-dominated landscapes (HDLs) is a key challenge for societies globally. This challenge cannot be adequately met with the current sectoral approaches to HDL governance and an academic community largely dominated by disciplinary sectors. Academia (universities and other research institutions and organizations) should take a more active role in embracing societal challenges around conservation of large carnivores in HDLs by facilitating cross-sectoral cooperation to mainstream coexistence of humans and large carnivores. Drawing on lessons from populated regions of Europe, Asia, and South America with substantial densities of large carnivores, we suggest academia should better embrace the principles and methods of sustainability sciences and create institutional spaces for the implementation of transdisciplinary curricula and projects; reflect on research approaches (i.e., disciplinary, interdisciplinary, or transdisciplinary) they apply and how their outcomes could aid leveraging institutional transformations for mainstreaming; and engage with various institutions and stakeholder groups to create novel institutional structures that can respond to multiple challenges of HDL management and human-large carnivore coexistence. Success in mainstreaming this coexistence in HDL will rest on the ability to think and act cooperatively. Such a conservation achievement, if realized, stands to have far-reaching benefits for people and biodiversity.

Incorporación de la Coexistencia entre Humanos y Carnívoros Mayores por Medio de la Colaboración Institucional Resumen Un reto importante para las sociedades mundiales es lograr la coexistencia entre los carnívoros mayores y los humanos en los paisajes dominados por el hombre (HDL, en inglés). Este reto no puede enfrentarse adecuadamente con las actuales estrategias sectoriales que se usan en la gobernanza de los HDL y con una comunidad académica dominada principalmente por sectores disciplinarios. La academia (las universidades y demás instituciones y organizaciones de investigación) debería realizar un papel más activo en la aceptación de los retos sociales que rodean a la conservación de los carnívoros mayores en los HDL al facilitar la cooperación intersectorial para incorporar la coexistencia entre humanos y dichos carnívoros. A partir de las lecciones aprendidas en las regiones pobladas por densidades abundantes de carnívoros mayores en América del Sur, Asia y Europa, sugerimos que la academia debería aceptar de mejor manera los principios y métodos de las ciencia de la sustentabilidad y crear espacios institucionales para la implementación de currículos y proyectos; reflexionar sobre las estrategias de investigación (es decir, disciplinaria, interdisciplinaria o transdisciplinaria) que aplican y cómo sus resultados podrían ayudar en fomentar las transformaciones institucionales para la incorporación; y participar junto a varias instituciones y grupos de accionistas para crear estructuras institucionales novedosas que puedan responder a los múltiples retos del manejo de los HDL y de la coexistencia entre humanos y carnívoros mayores. El éxito en la incorporación de esta coexistencia en los HDL dependerá de la habilidad para pensar y actuar cooperativamente. Tal logro de conservación, si se alcanza, promete tener beneficios de largo alcance para las personas y para la biodiversidad.

Truly sedentary? The multi-range tactic as a response to resource heterogeneity and unpredictability in a large herbivore.

Much research on large herbivore movement has focused on the annual scale to distinguish between resident and migratory tactics, commonly assuming that individuals are sedentary at the within-season scale. However, apparently sedentary animals may occupy a number of sub-seasonal functional home ranges (sfHR), particularly when the environment is spatially heterogeneous and/or temporally unpredictable. The roe deer (Capreolus capreolus) experiences sharply contrasting environmental conditions due to its widespread distribution, but appears markedly sedentary over much of its range. Using GPS monitoring from 15 populations across Europe, we evaluated the propensity of this large herbivore to be truly sedentary at the seasonal scale in relation to variation in environmental conditions. We studied movement using net square displacement to identify the possible use of sfHR. We expected that roe deer should be less sedentary within seasons in heterogeneous and unpredictable environments, while migratory individuals should be seasonally more sedentary than residents. Our analyses revealed that, across the 15 populations, all individuals adopted a multi-range tactic, occupying between two and nine sfHR during a given season. In addition, we showed that (i) the number of sfHR was only marginally influenced by variation in resource distribution, but decreased with increasing sfHR size; and (ii) the distance between sfHR increased with increasing heterogeneity and predictability in resource distribution, as well as with increasing sfHR size. We suggest that the multi-range tactic is likely widespread among large herbivores, allowing animals to track spatio-temporal variation in resource distribution and, thereby, to cope with changes in their local environment.

Framing the relationship between people and nature in the context of European conservation.

A key controversy in conservation is the framing of the relationship between people and nature. The extent to which the realms of nature and human culture are viewed as separate (dualistic view) or integrated is often discussed in the social sciences. To explore how this relationship is represented in the practice of conservation in Europe, we considered examples of cultural landscapes, wildlife (red deer, reindeer, horses), and protected area management. We found little support, for a dualistic worldview, where people and nature are regarded as separate in the traditional practice of conservation in Europe. The borders between nature and culture, wild and domestic, public land and private land, and between protected areas and the wider landscape were blurred and dynamic. The institutionalized (in practice and legislation) view is of an interactive mutualistic system in which humans and nature share the whole landscape. However, more dualistic ideals, such as wilderness and rewilding that are challenging established practices are expanding. In the context of modern day Europe, wilderness conservation and rewilding are not valid for the whole landscape, although it is possible to integrate some areas of low-intervention management into a wider matrix. A precondition for success is to recognize and plan for a plurality of values concerning the most valid approaches to conservation and to plan for this plurality at the landscape scale.

Enmarcando la Relación entre las Personas y la Naturaleza en el Contexto de la Conservación en Europa Resumen Una controversia clave en la conservación es el marco de la relación entre las personas y la naturaleza. En las ciencias sociales se discute con frecuencia el alcance al cual los reinos de la naturaleza y la cultura humana son vistos como separados (visión dualista) o íntegros. Para explorar cómo se representa esta relación en la práctica de la conservación en Europa consideramos ejemplos de paisajes culturales, vida silvestre (ciervos rojos, renos, caballos) y manejo de áreas protegidas. Encontramos poco apoyo, para una visión dual global, en donde las personas y la naturaleza son vistos como separados en la práctica tradicional de la conservación en Europa. Las fronteras entre la naturaleza y la cultura, lo silvestre y lo doméstico, el suelo público y el suelo privado, y entre las áreas protegidas y el terreno en conjunto, fueron dinámicas y borrosas. La visión institucionalizada (en la práctica y en la legislación) es de un sistema mutualista interactivo en el cual los humanos y la naturaleza comparten todo el terreno. Sin embargo, se están expandiendo más ideales dualistas, como la naturaleza y la reintroducción, los cuales están obstaculizando a las prácticas establecidas. En el contexto de la Europa contemporánea, la conservación de la naturaleza y la reintroducción no son válidas para todo el terreno, aunque es posible integrar algunas áreas de manejo de baja intervención a una matriz más amplia. Un prerrequisito para el éxito es reconocer y planear una pluralidad de valores con respecto a las estrategias de conservación en la escala de paisaje.

One size fits all: Eurasian lynx females share a common optimal litter size.

Lack proposed that the average clutch size of altricial species should be determined by the average maximum number of young the parents can raise such that all females in a given population should share a common optimal clutch size. Support for this model remains equivocal and recent studies have suggested that intra-population variation in clutch size is adaptive because each female has its own optimal clutch size associated with its intrinsic ability to raise offspring. Although Lack litter size and condition-dependent litter size are presented as two competing models, both are based on the concept of individual optimization. We propose a unified optimal litter size model (called 'adaptive litter size') and identify a set of conditions under which a common vs. a state-dependent optimal litter size should be observed. We test whether females of Eurasian lynx (Lynx lynx) have a common optimal litter size, or whether they adjust their litter size according to their state. We used a detailed individual-based data set collected from contrasting populations of Eurasian lynx in Scandinavia. Observed reproductive patterns in female lynx provide strong support for the existence of a common optimal litter size. Litter size did not vary according to female body mass or reproductive category, or among contrasted populations and years. A litter size of 2 was associated with a higher fitness than both smaller and larger litters, and thus corresponded to the 'adaptive litter size' for female lynx. We suggest that the reproductive pattern of female lynx might correspond to a risk avoidance tactic common to all individuals, which has evolved in response to strong environmental constraints generated by a highly unpredictable food supply during lactation.

A one night stand? Reproductive excursions of female roe deer as a breeding dispersal tactic.

Breeding dispersal, defined as the net movement between successive breeding sites, remains a poorly understood and seldom reported phenomenon in mammals, despite its importance for population dynamics and genetics. In large herbivores, females may be more mobile during the breeding season, undertaking short-term trips (excursions) outside their normal home range. If fertilisation occurs, leading to gene flow of the male genome, this behaviour could be considered a form of breeding dispersal from a genetic point of view. Here, we investigated ranging behaviour of 235 adult roe deer using intensive GPS monitoring in six populations across Europe within the EURODEER initiative. We show that excursions are common from June to August among females, with 41.8% (vs. 18.1% of males) making at least one excursion. Most individuals performed only one excursion per season and departure dates for females were concentrated in time, centred on the rutting period, suggesting a link with reproduction. The distance females travelled during excursions was significantly greater than the site-specific average diameter of a male home range, while travel speed decreased once they progressed beyond this diameter, indicating search behaviour or interaction with other male(s) outside the resident male's territory. Because adults are normally highly sedentary, the potential for mating with relatives is substantial; hence, we conclude that rut excursions could be an alternative tactic enabling females to avoid mating with a closely related male. To understand better the ultimate drivers at play, it will be crucial to explore the genetic causes and consequences of this behaviour.

Loss of Sunda clouded leopards and forest integrity drive potential impacts of mesopredator release on vulnerable avifauna.

Amongst the unintended consequences of anthropogenic landscape conversion is declining apex predator abundance linked to loss of forest integrity, which can potentially re-order trophic networks. One such re-ordering, known as mesopredator release, occurs when medium-sized predators, also called mesopredators, rapidly increase in abundance following the decline in apex predator abundance, consequently reducing the abundance of mesopredator prey, notably including terrestrial avifauna. We examine the cascading impacts of declining Sunda clouded leopard abundance, itself consequent upon a reduction in forest integrity, on the mesopredator community of Sabah, Malaysia, to determine whether the phenomenon of mesopredator release is manifest and specifically whether it impacts the terrestrial avifauna community of pheasants and pittas. To explore this trophic interaction, we used a piecewise structural equation model to compare changes in the relative abundance of organisms. Our results suggest that loss of forest integrity may have broad impacts on the community and trigger mesopredator release, the two acting additively in their impact on already vulnerable species of terrestrial avifauna: a result not previously documented in tropical systems and rarely detected even on a global scale. The limiting effect that the Sunda clouded leopard has on the Sunda leopard cat could illuminate the mechanism whereby mesopredator release impacts this system. Both Bulwer's pheasant and pittas appear to be significantly impacted by the increase in Sunda leopard cats, while the great argus pheasant shows similar compelling, although not statistically significant, declines as Sunda leopard cats increase. The inverse relationship between Sunda clouded leopards and Sunda leopard cats suggests that if a mesopredator release exists it could have downstream consequences for some terrestrial avifauna. These results suggest the under-studied interface between mammalian carnivores and avifauna, or more broadly species interactions in general, could offer important conservation tool for holistic ecosystem conservation efforts.

Decomposing risk: landscape structure and wolf behavior generate different predation patterns in two sympatric ungulates.

Recolonizing carnivores can have a large impact on the status of wild ungulates, which have often modified their behavior in the absence of predation. Therefore, understanding the dynamics of reestablished predator-prey systems is crucial to predict their potential ecosystem effects. We decomposed the spatial structure of predation by recolonizing wolves (Canis lupus) on two sympatric ungulates, moose (Alces alces) and roe deer (Capreolus capreolus), in Scandinavia during a 10-year study. We monitored 18 wolves with GPS collars, distributed over 12 territories, and collected records from predation events. By using conditional logistic regression, we assessed the contributions of three main factors, the utilization patterns of each wolf territory, the spatial distribution of both prey species, and fine-scale landscape structure, in determining the spatial structure of moose and roe deer predation risk. The reestablished predator-prey system showed a remarkable spatial variation in kill occurrence at the intra-territorial level, with kill probabilities varying by several orders of magnitude inside the same territory. Variation in predation risk was evident also when a spatially homogeneous probability for a wolf to encounter a prey was simulated. Even inside the same territory, with the same landscape structure, and when exposed to predation by the same wolves, the two prey species experienced an opposite spatial distribution of predation risk. In particular, increased predation risk for moose was associated with open areas, especially clearcuts and young forest stands, whereas risk was lowered for roe deer in the same habitat types. Thus, fine-scale landscape structure can generate contrasting predation risk patterns in sympatric ungulates, so that they can experience large differences in the spatial distribution of risk and refuge areas when exposed to predation by a recolonizing predator. Territories with an earlier recolonization were not associated with a lower hunting success for wolves. Such constant efficiency in wolf predation during the recolonization process is in line with previous findings about the naive nature of Scandinavian moose to wolf predation. This, together with the human-dominated nature of the Scandinavian ecosystem, seems to limit the possibility for wolves to have large ecosystem effects and to establish a behaviorally mediated trophic cascade in Scandinavia.

Seasonality, weather and climate affect home range size in roe deer across a wide latitudinal gradient within Europe.

1. Because many large mammal species have wide geographical ranges, spatially distant populations may be confronted with different sets of environmental conditions. Investigating how home range (HR) size varies across environmental gradients should yield a better understanding of the factors affecting large mammal ecology. 2. We evaluated how HR size of a large herbivore, the roe deer (Capreolus capreolus), varies in relation to seasonality, latitude (climate), weather, plant productivity and landscape features across its geographical range in Western Europe. As roe deer are income breeders, expected to adjust HR size continuously to temporal variation in food resources and energetic requirements, our baseline prediction was for HR size to decrease with proxies of resource availability. 3. We used GPS locations of roe deer collected from seven study sites (EURODEER collaborative project) to estimate fixed-kernel HR size at weekly and monthly temporal scales. We performed an unusually comprehensive analysis of variation in HR size among and within populations over time across the geographical range of a single species using generalized additive mixed models and linear mixed models, respectively. 4. Among populations, HR size decreased with increasing values for proxies of forage abundance, but increased with increases in seasonality, stochastic variation of temperature, latitude and snow cover. Within populations, roe deer HR size varied over time in relation to seasonality and proxies of forage abundance in a consistent way across the seven populations. Thus, our findings were broadly consistent across the distributional range of this species, demonstrating a strong and ubiquitous link between the amplitude and timing of environmental seasonality and HR size at the continental scale. 5. Overall, the variability in average HR size of roe deer across Europe reflects the interaction among local weather, climate and seasonality, providing valuable insight into the limiting factors affecting this large herbivore under contrasting conditions. The complexity of the relationships suggests that predicting ranging behaviour of large herbivores in relation to current and future climate change will require detailed knowledge not only about predicted increases in temperature, but also how this interacts with factors such as day length and climate predictability.

Home range size variation in a recovering wolf population: evaluating the effect of environmental, demographic, and social factors.

Home range size in mammals is a key ecological trait and an important parameter in conservation planning, and has been shown to be influenced by ecological, demographic and social factors in animal populations. Information on space requirements is especially important for carnivore species which range over very large areas and often come into direct conflict with human interest. We used long-term telemetry-location data from a recovering wolf population in Scandinavia to investigate variation in home range size in relation to environmental and social characteristics of the different packs. Wolves showed considerable variation in home range size, which ranged from 259 to 1,676 km(2). Although wolf density increased fourfold during the study period, we found no evidence that intraspecific competition influenced range size. Local variation in moose density, which was the main prey for most packs, did not influence wolf home range size. Home ranges increased with latitude and elevation and decreased with increased roe deer density. Although prey biomass alone did not influence range size, our data suggest that there is a correlation between habitat characteristics, choice of prey species and possible hunting success, which currently combine to shape home range size in Scandinavian wolves.

Building a resilient coexistence with wildlife in a more crowded world.

There is an urgent need to sustainably coexist with wildlife. However, realizing this goal is hampered by scant understanding of the processes that facilitate and maintain coexistence. Here, we synthesize human-wildlife interactions into eight archetypal outcomes, from eradication to sustained co-benefits, which collectively serve as a heuristic for forms of coexistence across a wide range of species and systems worldwide. We utilize resilience theory to elucidate how and why human-wildlife systems shift between these archetypes, yielding insights on research and policy priorities. We underscore the importance of governance structures that actively enhance the resilience of coexistence.

Altitude, latitude and climate zone as determinants of mountain hare (Lepus timidus) coat colour change.

Local adaptation to annually changing environments has evolved in numerous species. Seasonal coat colour change is an adaptation that has evolved in multiple mammal and bird species occupying areas that experience seasonal snow cover. It has a critical impact on fitness as predation risk may increase when an individual is mismatched against its habitat's background colour. In this paper, we investigate the correlation between landscape covariates and moult timing in a native winter-adapted herbivore, the mountain hare (Lepus timidus), throughout Norway. Data was collected between 2011 and 2019 at 678 camera trap locations deployed across an environmental gradient. Based on this data, we created a Bayesian multinomial logistic regression model that quantified the correlations between landscape covariates and coat colour phenology and analysed among season and year moult timing variation. Our results demonstrate that mountain hare moult timing is strongly correlated with altitude and latitude with hares that live at higher latitudes and altitudes keeping their winter white coats for longer than their conspecifics that inhabit lower latitudes and altitudes. Moult timing was also weakly correlated with climate zone with hares that live in coastal climates keeping their winter white coats for longer than hares that live in continental climates. We found evidence of some among year moult timing variation in spring, but not in autumn. We conclude that mountain hare moult timing has adapted to local environmental conditions throughout Norway.

Why humans kill animals and why we cannot avoid it.

Killing animals has been a ubiquitous human behaviour throughout history, yet it is becoming increasingly controversial and criticised in some parts of contemporary human society. Here we review 10 primary reasons why humans kill animals, discuss the necessity (or not) of these forms of killing, and describe the global ecological context for human killing of animals. Humans historically and currently kill animals either directly or indirectly for the following reasons: (1) wild harvest or food acquisition, (2) human health and safety, (3) agriculture and aquaculture, (4) urbanisation and industrialisation, (5) invasive, overabundant or nuisance wildlife control, (6) threatened species conservation, (7) recreation, sport or entertainment, (8) mercy or compassion, (9) cultural and religious practice, and (10) research, education and testing. While the necessity of some forms of animal killing is debatable and further depends on individual values, we emphasise that several of these forms of animal killing are a necessary component of our inescapable involvement in a single, functioning, finite, global food web. We conclude that humans (and all other animals) cannot live in a way that does not require animal killing either directly or indirectly, but humans can modify some of these killing behaviours in ways that improve the welfare of animals while they are alive, or to reduce animal suffering whenever they must be killed. We encourage a constructive dialogue that (1) accepts and permits human participation in one enormous global food web dependent on animal killing and (2) focuses on animal welfare and environmental sustainability. Doing so will improve the lives of both wild and domestic animals to a greater extent than efforts to avoid, prohibit or vilify human animal-killing behaviour.