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.

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.

Environment and physiology shape Arctic ungulate population dynamics.

Species conservation in a rapidly changing world requires an improved understanding of how individuals and populations respond to changes in their environment across temporal scales. Increased warming in the Arctic puts this region at particular risk for rapid environmental change, with potentially devastating impacts on resident populations. Here, we make use of a parameterized full life cycle, individual-based energy budget model for wild muskoxen, coupling year-round environmental data with detailed ontogenic metabolic physiology. We show how winter food accessibility, summer food availability, and density dependence drive seasonal dynamics of energy storage and thus life history and population dynamics. Winter forage accessibility defined by snow depth, more than summer forage availability, was the primary determinant of muskox population dynamics through impacts on calf recruitment and longer term carryover effects of maternal investment. Simulations of various seasonal snow depth and plant biomass and quality profiles revealed that timing of and improved/limited winter forage accessibility had marked influence on calf recruitment (±10-80%). Impacts on recruitment were the cumulative result of condition-driven reproductive performance at multiple time points across the reproductive period (ovulation to calf weaning) as a trade-off between survival and reproduction. Seasonal and generational condition effects of snow-rich winters interacted with age structure and density to cause pronounced long-term consequences on population growth and structure, with predicted population recovery times from even moderate disturbances of 10 years or more. Our results show how alteration in winter forage accessibility, mediated by snow depth, impacts the dynamics of northern herbivore populations. Further, we present here a mechanistic and state-based model framework to assess future scenarios of environmental change, such as increased or decreased snowfall or plant biomass and quality to impact winter and summer forage availability across the Arctic.

Long-term patterns in winter habitat selection, breeding and predation in a density-fluctuating, high Arctic lemming population.

Habitat selection is expected to balance benefits and costs that maximizes fitness. Using a rare data set on collared lemming (Dicrostonyx groenlandicus) winter nest location spanning more than two decades, we show that lemmings actively select for Salix snow beds, likely due to its favorable micro-climate, and that lemming habitat selection was density-dependent. Lemmings nevertheless exhibited some flexibility in their habitat selection, which appeared to be influenced by the year-to-year variation in snow conditions. The likelihood of both lemming breeding and nest predation by stoats (Mustela erminea) was not directly linked to habitat despite a delicate interplay between habitat, nest size, breeding, and predation. Hence, the larger lemming nests were found in Salix snow beds, and these were more often used for breeding, but both larger nests and nests used for breeding were also predated more often than other nests. Our study provides a clear example of how density-dependent habitat selection acts to balance fitness in the various habitats utilized by collared lemmings.

Habitat suitability analysis reveals high ecological flexibility in a "strict" forest primate.

BACKGROUND: Research of many mammal species tends to focus on single habitats, reducing knowledge of ecological flexibility. The Javan lutung (Trachypithecus auratus) is considered a strict forest primate, and little is known about populations living in savannah. In 2017-2018, we investigated the density and distribution of Javan lutung in Baluran National Park, Indonesia. We conducted ad libitum follows and line transect distance sampling with habitat suitability analysis of Javan lutung. RESULTS: Estimated density was 14.91 individuals km- 2 (95% CI 7.91-28.08), and estimated population size was 3727 individuals (95% CI 1979 - 7019). Long-tailed macaque (Macaca fascicularis) habitat suitability was the main driver of lutung habitat suitability as the probability of lutung occurrence increased greatly with macaque habitat suitability. Distance to roads, and distance to secondary forest had a negative relationship with lutung occurrence. Lutung habitat suitability decreased with increasing elevation, however, Mt Baluran and the primary forest on Mt Baluran was under-sampled due to treacherous conditions. Follows of six focus groups revealed considerable use of savannah, with terrestrial travel. The follows also revealed polyspecific associations with long-tailed macaques through shared sleeping sites and inter-specific vocalisations. CONCLUSIONS: Our study provides new knowledge on the general ecology of Javan lutung, such as use of savannah habitats, underlining our need to branch out in our study sites to understand the flexibility and adaptability of our study species. Another undocumented behaviour is the polyspecific association with long-tailed macaques. We encourage more research on this subject.

Bioaccumulation of mining derived metals in blood, liver, muscle and otoliths of two Arctic predatory fish species (Gadus ogac and Myoxocephalus scorpius).

Mining activities can cause adverse and long-lasting environmental impacts and detailed monitoring is therefore essential to assess the pollution status of mining impacted areas. Here we evaluated the efficacy of two predatory fish species (Gadus ogac i.e. Greenland cod and Myoxocephalus scorpius i.e. shorthorn sculpin) as biomonitors of mining derived metals (Pb, Zn, Cd and Hg) by measuring concentrations in blood, liver, muscle and otoliths along a distance gradient near the former Black Angel Pb-Zn mine (West Greenland). We detected metals in all tissues (except Cd and Hg in otoliths) and sculpin generally displayed higher concentrations than cod. For both species, concentrations were generally highest closest to the dominant pollution source(s) and gradually decreased away from the mine. The clearest gradient was observed for Pb in blood and liver (both species), and for Pb in otoliths (sculpin only). Similar to dissolved concentrations in seawater (but in contrast to bottom sediment), no significant decrease was found for Zn, Cd and Hg in any of the tissues. This demonstrates that by including tissues of blood (representing recent accumulation) and otolith (representing more long-term exposure signals) in the sampling collection, the temporal information on contaminant exposure and accumulation can be extended. We therefore conclude that both fish species are suitable as biomonitors near Arctic mine sites and, moreover, that blood and otoliths can serve as important supplementary monitoring tissues (in addition to liver and muscle traditionally sampled) as they provide extended temporal information on recent to long-term contaminant exposure.

Decision-Support Tools Used in the Baltic Sea Area: Performance and End-User Preferences.

Decision-support tools (DSTs) synthesize complex information to assist environmental managers in the decision-making process. Here, we review DSTs applied in the Baltic Sea area, to investigate how well the ecosystem approach is reflected in them, how different environmental problems are covered, and how well the tools meet the needs of the end users. The DSTs were evaluated based on (i) a set of performance criteria, (ii) information on end user preferences, (iii) how end users had been involved in tool development, and (iv) what experiences developers/hosts had on the use of the tools. We found that DSTs frequently addressed management needs related to eutrophication, biodiversity loss, or contaminant pollution. The majority of the DSTs addressed human activities, their pressures, or environmental status changes, but they seldom provided solutions for a complete ecosystem approach. In general, the DSTs were scientifically documented and transparent, but confidence in the outputs was poorly communicated. End user preferences were, apart from the shortcomings in communicating uncertainty, well accounted for in the DSTs. Although end users were commonly consulted during the DST development phase, they were not usually part of the development team. Answers from developers/hosts indicate that DSTs are not applied to their full potential. Deeper involvement of end users in the development phase could potentially increase the value and impact of DSTs. As a way forward, we propose streamlining the outputs of specific DSTs, so that they can be combined to a holistic insight of the consequences of management actions and serve the ecosystem approach in a better manner.

Age and seasonal variation in testis and baculum morphology in East Greenland polar bears (Ursus maritimus) in relation to high concentrations of persistent organic pollutants.

Persistent organic pollutants (POPs) are found in high concentrations in the Artic. Polar bears (Ursus maritimus) are one of the most exposed mammals in the Arctic and are thereby vulnerable to reproductive disruption. The aim of this study was to investigate male polar bear reproduction based on a detailed evaluation of testis histology and to assess possible effects of environmental chemicals on male polar bear reproduction. Reproductive groups that were identified based on histology were as follows: actively reproductive (REP), non-reproductive either with degenerated testes (DEG), undeveloped seminiferous tubules (UND), or morphology in-transition (INT). Categorization into these groups was supported by significant differences in testis and baculum measurements among REP, DEG, and UND, as well as differences in the area and diameter of seminiferous tubules among REP, DEG, and UND. These results show that it is possible to identify the reproductive stage in polar bears even if capture date and or age is lacking. Based on testis morphology we suggest that adult male polar bears from East Greenland have active spermatogenesis in February to June, and inactive degenerated testes in August to January. January to February was the main period of reproductive transition, characterised by a shift between inactive and active spermatogenesis. Baculum and testis size measurements decreased significantly with increasing concentrations of the chlordane metabolite oxychlordane, suggesting a potential impact on male reproductive success. Half of the investigated polar bears in REP group displayed signs of disorganization of the spermatogenesis which might be a sign of disrupted reproduction. However, no correlations with levels of the investigated POPs were detected. Reproductive organ measurements in polar bears differed significantly between REP and DEG groups, which cannot be explained by age, and therefore should be considered when investigating the effect of POPs on male reproduction.

Increasing density leads to generalization in both coarse-grained habitat selection and fine-grained resource selection in a large mammal.

Density is a fundamental driver of many ecological processes including habitat selection. Theory on density-dependent habitat selection predicts that animals should be distributed relative to profitability of habitat, resulting in reduced specialization in selection (i.e. generalization) as density increases and competition intensifies. Despite mounting empirical support for density-dependent habitat selection using isodars to describe coarse-grained (interhabitat) animal movements, we know little of how density affects fine-grained resource selection of animals within habitats [e.g. using resource selection functions (RSFs)]. Using isodars and RSFs, we tested whether density simultaneously modified habitat selection and within-habitat resource selection in a rapidly growing population of feral horses (Equus ferus caballus Linnaeus; Sable Island, Nova Scotia, Canada; 42% increase in population size from 2008 to 2012). Among three heterogeneous habitat zones on Sable Island describing population clusters distributed along a west-east resource gradient (west-central-east), isodars revealed that horses used available habitat in a density-dependent manner. Intercepts and slopes of isodars demonstrated a pattern of habitat selection that first favoured the west, which generalized to include central and east habitats with increasing population size consistent with our understanding of habitat quality on Sable Island. Resource selection functions revealed that horses selected for vegetation associations similarly at two scales of extent (total island and within-habitat zone). When densities were locally low, horses were able to select for sites of the most productive forage (grasslands) relative to those of poorer quality. However, as local carrying capacity was approached, selection for the best of available forage types weakened while selection for lower-quality vegetation increased (and eventually exceeded that of grasslands). Isodars can effectively describe coarse-grained habitat selection in large mammals. Our study also shows that the main predictions of density-dependent habitat selection are highly relevant to our interpretation of RSFs in space and time. At low but not necessarily high population size, density will be a leading indicator of habitat quality. Fitness maximization from specialist vs. generalist strategies of habitat and resource selection may well be apparent at multiple spatial extents and grains of resolution.

Density-dependent habitat selection and partitioning between two sympatric ungulates.

Theory on density-dependent habitat selection predicts that as population density of a species increases, use of higher quality (primary) habitat by individuals declines while use of lower quality (secondary) habitat rises. Habitat partitioning is often considered the primary mechanism for coexistence between similar species, but how this process evolves with changes in population density remains to be empirically tested for free-ranging ungulates. We used resource-selection functions to quantify density effects on landscape-scale habitat selection of two sympatric species of ungulates [moose (Alces alces) and elk (Cervus canadensis manitobensis)] in Riding Mountain National Park, Manitoba, Canada (2000-2011). The density of elk was actively reduced from 1.2 to 0.4 elk km(-2) through increased hunting effort during the period of study, while moose density decreased without additional human influence from 1.6-0.7 moose km(-2). Patterns of habitat selection during winter by both species changed in accordance to expectations from density-dependent habitat-selection theory. At low intraspecific density, moose and elk did not partition habitat, as both species selected strongly for mixed forest (primary habitat providing both food and cover), but did so in different areas segregated across an elevational gradient. As intraspecific density increased, selection for primary habitat by both species decreased, while selection for secondary, lower quality habitat such as agricultural fields (for elk) and built-up areas (for moose) increased. We show that habitat-selection strategies during winter for moose and elk, and subsequent effects on habitat partitioning, depend heavily on the position in state space (density) of both species.

Reproductive success and failure: the role of winter body mass in reproductive allocation in Norwegian moose.

A life history strategy that favours somatic growth over reproduction is well known for long-lived iteroparous species, especially in unpredictable environments. Risk-sensitive female reproductive allocation can be achieved by a reduced reproductive effort at conception, or the subsequent adjustment of investment during gestation or lactation in response to unexpected environmental conditions or resource availability. We investigated the relative importance of reduced investment at conception compared with later in the reproductive cycle (i.e. prenatal, perinatal or neonatal mortality) in explaining reproductive failure in two high-density moose (Alces alces) populations in southern Norway. We followed 65 multiparous, global positioning system (GPS)-collared females throughout the reproductive cycle and focused on the role of maternal nutrition during gestation in determining reproductive success using a quasi-experimental approach to manipulate winter forage availability. Pregnancy rates in early winter were normal (≥0.8) in all years while spring calving rates ranged from 0.4 to 0.83, with prenatal mortality accounting for most of the difference. Further losses over summer reduced autumn recruitment rates to 0.23-0.69, despite negligible predation. Over-winter mass loss explained variation in both spring calving and autumn recruitment success better than absolute body mass in early or late winter. Although pregnancy was related to body mass in early winter, overall reproductive success was unrelated to pre-winter body condition. We therefore concluded that reproductive success was limited by winter nutritional conditions. However, we could not determine whether the observed reproductive allocation adjustment was a bet-hedging strategy to maximise reproduction without compromising survival or whether females were simply unable to invest more resources in their offspring.

Emergent interactions in the management of multiple threats to the conservation of harbour porpoises.

Human activities at sea are intensifying and diversifying. This is leading to more complex interactions of anthropogenic impacts requiring adaptable management interventions to mitigate their cumulative effects on biodiversity conservation and restoration objectives. Bycatch remains the dominant conservation threat for coastal cetaceans. Additionally, the indirect impact of repeated exposure to disturbances, particularly acoustic disturbances, can affect cetacean population growth and therefore conservation objectives. Pingers are used to ensonify nets to provide an effective mitigation of bycatch risk. As those become more prevalent across fisheries at risk to catch for example harbour porpoises, pingers become contributors to the anthropogenic noise landscape which may affect the vital rates of this species as well. Currently, we do not know how to best balance pinger prevalence to minimise both bycatch rate and the population consequences of acoustic disturbance (PCoD). Here we use an agent-based model to determine how pinger prevalence in nets can be adjusted to minimise bycatch rate and noise disturbance propagating to affect population growth for harbour porpoises. We show that counter-intuitively bycatch rate can increase at lower pinger prevalence. When ecological conditions are such that PCOD can emerge, higher prevalence of pingers can lead to indirect effects on population growth. This would result from condition-mediated decreased reproductive potential. Displacing fishing effort, via time-area closure, can be an effective mitigation strategy in these circumstances. These findings have important implications for current management plans which, for practical consideration, may lead to lower overall pinger prevalence at sea. This study also shows that estimating the reproductive potential of the species should be incorporated in bycatch monitoring programmes. We now need to better understand how physiological condition affect reproductive decisions and behavioural responses to noise in cetaceans to better appraise and estimate the cumulative impacts of bycatch and its mitigations.

Direct and Indirect Linkages Between Trace Element Status and Health Indicators - a Multi-tissue Case-Study of Two Deer Species in Denmark.

Measuring trace element concentrations in tissue can be a valuable approach to monitor animal health status. Temporal variation in the absorption, transport, and storage of elements between different tissues can, however, complicate the assessment of element-health relationships. Here, we measured concentrations of selected essential (copper (Cu), zinc (Zn), selenium (Se)) and non-essential (arsenic (As), cadmium (Cd), lead (Pb)) trace elements within blood, liver, kidney, and hair of fallow deer (Dama dama; N=20) and red deer (Cervus elaphus; N=21). Using multivariate regression and structural equation models, we estimated direct and indirect linkages between tissue-specific trace element profiles and long- (body condition) and short-term (serum protein biomarkers for acute inflammation, infection, and malnutrition) health indicators. Trace element concentrations varied markedly and were weakly correlated among tissues, with the exception of Se. After accounting for sex- and site-differences in trace element concentrations, body condition of red deer was directly, and positively, associated to trace element status in liver and hair, but not in kidney. For both deer species, trace element status in blood was directly linked to serum protein status with an indirect positive association to deer body condition. For fallow deer, no direct association between trace element status and body condition was detected in any of the tissues, possibly because of elemental homeostasis, and because all individuals were in good clinical health. This study shows that hair can serve as an effective, non-invasive, biomarker in deer health assessments, yet, to fully uncover trace element-health relationships a variety of sample matrices is preferred.

Estimating parasite-condition relationships and potential health effects for fallow deer (Dama dama) and red deer (Cervus elaphus) in Denmark.

Parasites can exert a substantial influence on the ecology of wildlife populations by altering host condition. Our objectives were to estimate single and multiparasite-condition relationships for fallow deer (Dama dama) and red deer (Cervus elaphus) in Denmark and to assess potential health effects along the parasite burden gradient. Fallow deer hosted on average two endoparasite taxa per individual (min = 0, max = 5) while red deer carried on average five parasite taxa per individual (min = 2, max = 9). Body condition of both deer species was negatively related to presence of Trichuris ssp. eggs while body condition of red deer was positively related to antibodies of the protozoan Toxoplasma gondii. For the remaining parasite taxa (n = 12), we either found weak or no apparent association between infection and deer body condition or low prevalence levels restricted formal testing. Importantly, we detected a strong negative relationship between body condition and the sum of endoparasite taxa carried by individual hosts, a pattern that was evident in both deer species. We did not detect systemic inflammatory reactions, yet serology revealed reduced total protein and iron concentrations with increased parasite load in both deer species, likely due to maldigestion of forage or malabsorption of nutrients. Despite moderate sample sizes, our study highlights the importance of considering multiparasitism when assessing body condition impacts in deer populations. Moreover, we show how serum chemistry assays are a valuable diagnostic tool to detect subtle and sub-clinical health impacts of parasitism, even at low-level infestation.

Geochemical landscapes as drivers of wildlife reproductive success: Insights from a high-Arctic ecosystem.

The bioavailability of essential and non-essential elements in vegetation is expected to influence the performance of free-ranging terrestrial herbivores. However, attempts to relate the use of geochemical landscapes by animal populations directly to reproductive output are currently lacking. Here we measured concentrations of 14 essential and non-essential elements in soil and vegetation samples collected in the Zackenberg valley, northeast Greenland, and linked these to environmental conditions to spatially predict and map geochemical landscapes. We then used long-term (1996-2021) survey data of muskoxen (Ovibos moschatus) to quantify annual variation in the relative use of essential and non-essential elements in vegetated sites and their relationship to calf recruitment the following year. Results showed that the relative use of the geochemical landscape by muskoxen varied substantially between years and differed among elements. Selection for vegetated sites with higher levels of the essential elements N, Cu, Se, and Mo was positively linked to annual calf recruitment. In contrast, selection for vegetated sites with higher concentrations of the non-essential elements As and Pb was negatively correlated to annual calf recruitment. Based on the concentrations measured in our study, we found no apparent associations between annual calf recruitment and levels of C, Mn, Co, Zn, Cd, Ba, Hg, and C:N ratio in the vegetation. We conclude that the spatial distribution and access to essential and non-essential elements are important drivers of reproductive output in muskoxen, which may also apply to other wildlife populations. The value of geochemical landscapes to assess habitat-performance relationships is likely to increase under future environmental change.

Forecasting shifts in habitat suitability of three marine predators suggests a rapid decline in inter-specific overlap under future climate change.

Understanding how environmental and climate change can alter habitat overlap of marine predators has great value for the management and conservation of marine ecosystems. Here, we estimated spatiotemporal changes in habitat suitability and inter-specific overlap among three marine predators: Baltic gray seals (Halichoerus grypus), harbor seals (Phoca vitulina), and harbor porpoises (Phocoena phocoena) under contemporary and future conditions. Location data (>200 tagged individuals) were collected in the southwestern region of the Baltic Sea; one of the fastest-warming semi-enclosed seas in the world. We used the maximum entropy (MaxEnt) algorithm to estimate changes in total area size and overlap of species-specific habitat suitability between 1997-2020 and 2091-2100. Predictor variables included environmental and climate-sensitive oceanographic conditions in the area. Sea-level rise, sea surface temperature, and salinity data were taken from representative concentration pathways [RCPs] scenarios 6.0 and 8.5 to forecast potential climate change effects. Model output suggested that habitat suitability of Baltic gray seals will decline over space and time, driven by changes in sea surface salinity and a loss of currently available haulout sites following sea-level rise in the future. A similar, although weaker, effect was observed for harbor seals, while suitability of habitat for harbor porpoises was predicted to increase slightly over space and time. Inter-specific overlap in highly suitable habitats was also predicted to increase slightly under RCP scenario 6.0 when compared to contemporary conditions, but to disappear under RCP scenario 8.5. Our study suggests that marine predators in the southwestern Baltic Sea may respond differently to future climatic conditions, leading to divergent shifts in habitat suitability that are likely to decrease inter-specific overlap over time and space. We conclude that climate change can lead to a marked redistribution of area use by marine predators in the region, which may influence local food-web dynamics and ecosystem functioning.

What determines variation in home range size across spatiotemporal scales in a large browsing herbivore?

1. Most studies of intraspecific variation in home range size have investigated only a single or a few factors and often at one specific scale. However, considering multiple spatial and temporal scales when defining a home range is important as mechanisms that affect variation in home range size may differ depending on the scale under investigation. 2. We aim to quantify the relative effect of various individual, forage and climatic determinants of variation in home range size across multiple spatiotemporal scales in a large browsing herbivore, the moose (Alces alces), living at the southern limit of its distribution in Norway. 3. Total home range size and core home range areas were estimated for daily to monthly scales in summer and winter using both local convex hull (LoCoH) and fixed kernel home range methods. Variance in home range size was analysed using linear mixed-effects models for repeated measurements. 4. Reproductive status was the most influential individual-level factor explaining variance in moose home range size, with females accompanied by a calf having smaller summer ranges across all scales. Variation in home range size was strongly correlated with spatiotemporal changes in quantity and quality of natural food resources. Home range size decreased with increasing browse density at daily scales, but the relationship changed to positive at longer temporal scales. In contrast, browse quality was consistently negatively correlated with home range size except at the monthly scale during winter when depletion of high-quality forage occurs. Local climate affected total home range size more than core areas. Temperature, precipitation and snow depth influenced home range size directly at short temporal scales. 5. The relative effects of intrinsic and extrinsic determinants of variation in home range size differed with spatiotemporal scale, providing clear evidence that home range size is scale dependent in this large browser. Insight into the behavioural responses of populations to climatic stochasticity and forage variability is essential in view of current and future climate change, especially for populations with thermoregulatory restrictions living at the southern limit of their distribution.

On the uncertainty and confidence in decision support tools (DSTs) with insights from the Baltic Sea ecosystem.

Ecosystems around the world are increasingly exposed to multiple, often interacting human activities, leading to pressures and possibly environmental state changes. Decision support tools (DSTs) can assist environmental managers and policy makers to evaluate the current status of ecosystems (i.e. assessment tools) and the consequences of alternative policies or management scenarios (i.e. planning tools) to make the best possible decision based on prevailing knowledge and uncertainties. However, to be confident in DST outcomes it is imperative that known sources of uncertainty such as sampling and measurement error, model structure, and parameter use are quantified, documented, and addressed throughout the DST set-up, calibration, and validation processes. Here we provide a brief overview of the main sources of uncertainty and methods currently available to quantify uncertainty in DST input and output. We then review 42 existing DSTs that were designed to manage anthropogenic pressures in the Baltic Sea to summarise how and what sources of uncertainties were addressed within planning and assessment tools. Based on our findings, we recommend future DST development to adhere to good modelling practise principles, and to better document and communicate uncertainty among stakeholders.

Quantifying energetic and fitness consequences of seasonal heterothermy in an Arctic ungulate.

Animals have adapted behavioral and physiological strategies to conserve energy during periods of adverse conditions. Heterothermy is one such adaptation used by endotherms. While heterothermy-fluctuations in body temperature and metabolic rate-has been shown in large vertebrates, little is known of the costs and benefits of this strategy, both in terms of energy and in terms of fitness. Hence, our objective was to model the energetics of seasonal heterothermy in the largest Arctic ungulate, the muskox (Ovibos moschatus), using an individual-based energy budget model of metabolic physiology. We found that the empirically based drop in body temperature (winter max ~-0.8°C) overwinter in adult females resulted in substantial fitness benefits in terms of reduced daily energy expenditure and body mass loss. Body mass and energy reserves were 8.98% and 14.46% greater in modeled heterotherms compared to normotherms by end of winter. Based on environmental simulations, we show that seasonal heterothermy can, to some extent, buffer the negative consequences of poor prewinter body condition or reduced winter food accessibility, leading to greater winter survival (+20%-30%) and spring energy reserves (+10%-30%), and thus increased probability of future reproductive success. These results indicate substantial adaptive short-term benefits of seasonal heterothermy at the individual level, with potential implications for long-term population dynamics in highly seasonal environments.

Forage quantity, quality and depletion as scale-dependent mechanisms driving habitat selection of a large browsing herbivore.

1. Mechanisms that affect the spatial distribution of animals are typically scale-dependent and may involve forage distribution. Forage quality and quantity are often inversely correlated and a much discussed trade-off is whether or not to select for high-quality forage at the expense of forage abundance. This discussion has rarely involved scale-dependence or been applied to Northern browsing herbivores. At small spatial scales, browsers are assumed to select for the best quality forage. But, as high-quality forage resources are often scarce and may become depleted, coarse-scale habitat selection is assumed to be driven by forage availability. 2. To evaluate if moose selection for forage quantity and quality is scale-dependent we modelled summer and winter habitat selection of 32 GPS-marked female moose (Alces alces) at two spatial scales (landscape-scale vs. within-home range-scale). We used mixed-effects resource selection functions (RSFs) and landscape-scale forage availability models of six tree species of varying quality for moose. We considered silver birch (Betula pendula), downy birch (Betula pubescens.), Scots pine (Pinus sylvestris) as low quality browse species and rowan (Sorbus aucuparia), aspen (Populus tremula), willow (Salix spp.) as high-quality species. 3. As expected, the overall selection patterns for available browse biomass and quality varied across spatiotemporal scales. At the landscape-scale, moose selected for habitat with high available browse biomass of low quality species while at the within-home range-scale moose selected for sites with the highest quality browse species available. Furthermore, selection patterns during summer remained fairly stable, while during winter, selection at the within-home range-scale switched from sites with high quality to sites with lower quality browse species which suggests depletion of high-quality species. Consistent with expectations from seasonal resource depletion, site fidelity (bimonthly home range overlap) was much lower in winter than in summer. 4. Coarse-scale habitat selection by moose as a function of forage variability revealed a scale-dependent trade-off between available browse quantity and browse quality. Moreover, resource depletion changed the winter selection criteria of free-ranging moose and we demonstrate how the behavioural response to such a dynamic process can be inferred from RSFs.