Direct and indirect effects of cougar predation on bighorn sheep fitness.

Predation has direct effects on prey population dynamics through mortality, and it can induce indirect effects through fear. The indirect effects of predation have been documented experimentally, but few studies have quantified them in nature so that their role in prey population dynamics remains controversial. Given the expanding or reintroduced populations of large predators in many areas, the quantification of indirect effects of predation is crucial. We sought to evaluate the direct and indirect fitness effects of intense cougar (Puma concolor) predation using 48 years of data on marked bighorn sheep (Ovis canadensis) on Ram Mountain, Alberta, Canada. We compared years of intense cougar predation with years with no or occasional cougar predation. We first quantified the effects of predation on neonatal, weaning, and overwinter lamb survival, three metrics potentially affected by direct and indirect effects. We then investigated the possible indirect effects of intense cougar predation on lamb production, female summer mass gain, and lamb mass at weaning. We found strong effects of cougar predation on lamb survival, lamb production, and seasonal mass gain of lambs and adult females. In years with high predation, neonatal, weaning, and overwinter lamb survival declined by 18.4%, 19.7% and 20.8%, respectively. Indirect effects included a 14.2% decline in lamb production. Female summer mass gain decreased by 15.6% and lamb mass at weaning declined by 8.0% in years of intense cougar predation. Our findings bring key insights on the impacts of predation on prey fitness by reporting moderate to large effects on recruitment and illustrate the importance of indirect effects of predation on population dynamics.

Vocal repertoire and individuality in the plains zebra (Equus quagga).

Acoustic signals are vital in animal communication, and quantifying them is fundamental for understanding animal behaviour and ecology. Vocalizations can be classified into acoustically and functionally or contextually distinct categories, but establishing these categories can be challenging. Newly developed methods, such as machine learning, can provide solutions for classification tasks. The plains zebra is known for its loud and specific vocalizations, yet limited knowledge exists on the structure and information content of its vocalzations. In this study, we employed both feature-based and spectrogram-based algorithms, incorporating supervised and unsupervised machine learning methods to enhance robustness in categorizing zebra vocalization types. Additionally, we implemented a permuted discriminant function analysis to examine the individual identity information contained in the identified vocalization types. The findings revealed at least four distinct vocalization types-the 'snort', the 'soft snort', the 'squeal' and the 'quagga quagga'-with individual differences observed mostly in snorts, and to a lesser extent in squeals. Analyses based on acoustic features outperformed those based on spectrograms, but each excelled in characterizing different vocalization types. We thus recommend the combined use of these two approaches. This study offers valuable insights into plains zebra vocalization, with implications for future comprehensive explorations in animal communication.

First dispersal records of the endangered banteng (Bos javanicus) in Thung Yai Naresuan West Wildlife Sanctuary, Thailand.

Banteng (Bos javanicus) is listed as an endangered species because of a global population decline of at least 50% over the last 25 years. The Western Forest Complex (WEFCOM) of Thailand has been identified as a priority site for banteng population recovery, and Huai Kha Keang Wildlife Sanctuary (HKK) is the most important source site for this species within the WEFCOM. We have provided evidence and discussed banteng dispersal from HKK to Thung Yai Naresuan West Wildlife Sanctuary (TYW). We sampled an area of 147 km2 in banteng habitat next to the border between HKK and TYW using camera traps. We divided the sampled area into four grid cells and placed camera traps during January to December 2022. We setup the camera traps near saltlicks and natural water sources, as important resources for banteng, to maximize capture probability. In total, 2835 trap days were obtained. Bantengs were captured in all seasons (RAI = 1.66), especially in dry dipterocarp forest, which contains the ground forage availability for banteng, and the low-slope area with elevation 600-700 m adjacent to the border between HKK and TYW. The results highlighted that banteng, which had never been reported in TYW before, appeared there for the first time. They most likely dispersed from the population source in HKK and settled in a habitat that is considered suitable for them. The habitat management and protection are significant for the future recovery of banteng populations in the TYW and the rest of protected areas in the WEFCOM.

Inter-annual variation in the microplastics abundance in feces of the Baird's tapir (Tapirus bairdii) from the Selva Maya, México.

Microplastics (MPs) are found in a wide range of ecosystems, from the Arctic to the deep ocean. However, there is no data on their presence in terrestrial mammals that inhabit the Selva Maya. The aim of this study is to detect the presence of MPs in the feces of the Baird's tapir (Tapirus bairdii) from the region of Calakmul, located in the Yucatan Peninsula, Mexico. We analyzed 129 fecal samples collected during 2017 and 2018, obtaining 57 and 72 samples during the rainy and dry seasons respectively. Sixty-eight percent of the samples contained 743 MPs with a mean of 19.3 ± 28.1 MPs/kg of dry weight (DW) feces in both years. An inter-annual variation in the average abundance of microplastic was observed during the two-year period (2017-2018), with a 72 % increase in these plastic particles in feces. Fourteen polymers were identified, with ethylene vinyl acetate (EVA), polypropylene (PP) and polyester (PES) being the most abundant during both years. Although the effects of MPs on the health of tapirs are not known, their presence is cause for concern. There is an urgent need for the implementation of appropriate plastic waste management programs in communities of the Selva Maya to diminish the consumption of MPs in species including humans where they pose a significant risk to health. ENVIRONMENTAL IMPLICATIONS: The use of plastics worldwide is increasing every day, so the presence of microplastics is and will continue to be a major environmental problem. It is known that contaminants can adhere to plastics, making them hazardous materials. Microplastics can contaminate remote areas such as Biosphere Reserves. Terrestrial species such as the tapir can ingest microplastics, putting their health at risk. Knowing the dispersion of microplastics is very important in order to manage them properly, taking into account their emission sources and type of polymer.

Proactive cursorial and ambush predation risk avoidance in four African herbivore species.

Most herbivores must balance demands to meet nutritional requirements, maintain stable thermoregulation and avoid predation. Species-specific predator and prey characteristics determine the ability of prey to avoid predation and the ability of predators to maximize hunting success. Using GPS collar data from African wild dogs, lions, impala, tsessebes, wildebeest and zebra in the Okavango Delta, Botswana, we studied proactive predation risk avoidance by herbivores. We considered predator activity level in relation to prey movement, predator and prey habitat selection, and preferential use of areas by prey. We compared herbivore behaviour to lion and wild dog activity patterns and determined the effect of seasonal resource availability and prey body mass on anti-predator behaviour. Herbivore movement patterns were more strongly correlated with lion than wild dog activity. Habitat selection by predators was not activity level dependent and, while prey and predators differed to some extent in their habitat selection, there were also overlaps, probably caused by predators seeking habitats with high prey abundance. Areas favoured by lions were used by herbivores more when lions were less active, whereas wild dog activity level was not correlated with prey use. Prey body mass was not a strong predictor of the strength of proactive predation avoidance behaviour. Herbivores showed stronger anti-predator behaviours during the rainy season when resources were abundant. Reducing movement when top predators are most active and avoiding areas with a high likelihood of predator use during the same periods appear to be common strategies to minimize predation risk. Such valuable insights into predator-prey dynamics are only possible when using similar data from multiple sympatric species of predator and prey, an approach that should become more prevalent given the ongoing integration of technological methods into ecological studies.

Climatically robust multiscale species distribution models to support pronghorn recovery in California.

We combined two climate-based distribution models with three finer-scale suitability models to identify habitat for pronghorn recovery in California now and into the future. We used a consensus approach to identify areas of suitable climate now and future for pronghorn in California. We compared the results of climate models from two separate hypotheses about their historical ecology in the state. Under the migration hypothesis, pronghorn were expected to be limited climatically by extreme cold in winter and extreme heat in summer; under the niche reduction hypothesis, historical pronghorn of distribution would have better represented the climatic limitations of the species. We combined occurrences from GPS collars distributed across three populations of pronghorn in the state to create three distinct habitat suitability models: (1) an ensemble model using random forests, Maxent, classification and regression Trees, and a generalized linear model; (2) a step selection function; and (3) an expert-driven model. We evaluated consensus among both the climate models and the suitability models to prioritize areas for, and evaluate the prospects of, pronghorn recovery. Climate suitability for pronghorn in the future depends heavily on model assumptions. Under the migration hypothesis, our model predicted that there will be no suitable climate in California in the future. Under the niche reduction hypothesis, by contrast, suitable climate will expand. Habitat suitability also depended on the methods used, but areas of consensus among all three models exist in large patches throughout the state. Identifying habitat for a species which has undergone extreme range collapse, and which has very fine scale habitat needs, presents novel challenges for spatial ecologists. Our multimethod, multihypothesis approach can allow habitat modelers to identify areas of consensus and, perhaps more importantly, fill critical knowledge gaps that could resolve disagreements among the models. For pronghorn, a better understanding of their upper thermal tolerances and whether historical populations migrated will be crucial to their potential recovery in California and throughout the arid Southwest.

White-tailed deer population dynamics in a multipredator landscape shaped by humans.

Large terrestrial mammals increasingly rely on human-modified landscapes as anthropogenic footprints expand. Land management activities such as timber harvest, agriculture, and roads can influence prey population dynamics by altering forage resources and predation risk via changes in habitat, but these effects are not well understood in regions with diverse and changing predator guilds. In northeastern Washington state, USA, white-tailed deer (Odocoileus virginianus) are vulnerable to multiple carnivores, including recently returned gray wolves (Canis lupus), within a highly human-modified landscape. To understand the factors governing predator-prey dynamics in a human context, we radio-collared 280 white-tailed deer, 33 bobcats (Lynx rufus), 50 cougars (Puma concolor), 28 coyotes (C. latrans), and 14 wolves between 2016 and 2021. We first estimated deer vital rates and used a stage-structured matrix model to estimate their population growth rate. During the study, we observed a stable to declining deer population (lambda = 0.97, 95% confidence interval: 0.88, 1.05), with 74% of Monte Carlo simulations indicating population decrease and 26% of simulations indicating population increase. We then fit Cox proportional hazard models to evaluate how predator exposure, use of human-modified landscapes, and winter severity influenced deer survival and used these relationships to evaluate impacts on overall population growth. We found that the population growth rate was dually influenced by a negative direct effect of apex predators and a positive effect of timber harvest and agricultural areas. Cougars had a stronger effect on deer population dynamics than wolves, and mesopredators had little influence on the deer population growth rate. Areas of recent timber harvest had 55% more forage biomass than older forests, but horizontal visibility did not differ, suggesting that timber harvest did not influence predation risk. Although proximity to roads did not affect the overall population growth rate, vehicle collisions caused a substantial proportion of deer mortalities, and reducing these collisions could be a win-win for deer and humans. The influence of apex predators and forage indicates a dual limitation by top-down and bottom-up factors in this highly human-modified system, suggesting that a reduction in apex predators would intensify density-dependent regulation of the deer population owing to limited forage availability.

Heavy metal and metalloid concentrations in red deer (Cervus elaphus) and their human health implications from One Health perspective.

The red deer is an ungulate and large game species. The contamination of the ecosystems by metal(loid)s may lead to the exposure of animals (as well as humans) through water and food resources. The direct contact of hunters and wild animal meat consumers with deer carcasses may be a potential contaminant source. This study aimed to determine the metal(loid)s' concentrations in the liver and kidney of red deer from two regions of Portugal (Idanha-a-Nova and Lousã), and to relate these with histopathologic lesions. Thirteen young male deer were submitted to metal(loid) determination (As, Cd, Co, Cr, Cu, Pb, and Zn) by inductively coupled plasma mass spectrophotometry (ICP-MS) and histopathology examination. Renal Cd (8.072 ± 5.766 mg/kg dw) and hepatic Pb (3.824 ± 6.098 mg/kg dw) mean values were high, considering the maximum values for consumption established by the European Commission. The hepatic mean value of Cu was significantly higher in Idanha-a-Nova (150.059 ± 33.321 mg/kg dw), and it is at the Cu toxicity limit considered for ruminants (150 mg/kg). The pollution induced by Panasqueira mines (Castelo Branco) may be a possible explanation for some of the findings, especially the higher values of hepatic Cu and Pb found in Idanha-a-Nova deer. These results have high importance under a One Health perspective, since they have implications in public health, and pose at risk the imbalance of animal populations and ecosystems.

First detection of Tetraparvovirus ungulate 1 in diseased cattle (Chinese Simmental) from Hunan province, China.

Tetraparvovirus is an emerging parvovirus infecting a variety of mammals and humans, and associated with human diseases including severe acute respiratory infection and acute encephalitis syndrome. In the present study, a Tetraparvovirus ungulate 1 (formerly known as bovine hokovirus) strain HNU-CBY-2023 was identified and characterized from diseased Chinese Simmental from Hunan province, China. The nearly complete genome of HNU-CBY-2023 is 5346 nt in size and showed genomic identities of 85-95.5% to the known Tetraparvovirus ungulate 1 strains from GenBank, indicating a rather genetic variation. Phylogenetic and genetic divergence analyses indicated that Tetraparvovirus ungulate 1 could be divided into two genotypes (I and II), and HNU-CBY-2023 was clustered into genotype II. This study, for the first time, identified Tetraparvovirus ungulate 1 from domestic cattle from mainland China, which will be helpful to understand the prevalence and genetic diversity of Tetraparvovirus ungulate 1.

Systematic reduction of natural enemies and competition across variable precipitation approximates buffelgrass invasiveness (Cenchrus ciliaris) in its native range.

Invasive grasses cause devastating losses to biodiversity and ecosystem function directly and indirectly by altering ecosystem processes. Escape from natural enemies, plant-plant competition, and variable resource availability provide frameworks for understanding invasion. However, we lack a clear understanding of how natural stressors interact in their native range to regulate invasiveness. In this study, we reduced diverse guilds of natural enemies and plant competitors of the highly invasive buffelgrass across a precipitation gradient throughout major climatic shifts in Laikipia, Kenya. To do this, we used a long-term ungulate exclosure experiment design across a precipitation gradient with nested treatments that (1) reduced plant competition through clipping, (2) reduced insects through systemic insecticide, and (3) reduced fungal associates through fungicide application. Additionally, we measured the interaction of ungulates on two stem-boring insect species feeding on buffelgrass. Finally, we measured a multiyear smut fungus outbreak. Our findings suggest that buffelgrass exhibits invasive qualities when released from a diverse group of natural stressors in its native range. We show natural enemies interact with precipitation to alter buffelgrass productivity patterns. In addition, interspecific plant competition decreased the basal area of buffelgrass, suggesting that biotic resistance mediates buffelgrass dominance in the home range. Surprisingly, systemic insecticides and fungicides did not impact buffelgrass production or reproduction, perhaps because other guilds filled the niche space in these highly diverse systems. For example, in the absence of ungulates, we showed an increase in host-specific stem-galling insects, where these insects compensated for reduced ungulate use. Finally, we documented a smut outbreak in 2020 and 2021, corresponding to highly variable precipitation patterns caused by a shifting Indian Ocean Dipole. In conclusion, we observed how reducing natural enemies and competitors and certain interactions increased properties related to buffelgrass invasiveness.

Genetic Analysis Reveals Dispersal Patterns of Japanese Serow in Two Different Habitats of a Mountainous Region.

Dispersal increases the costs of feeding and predation risk in the new environment and is reported to be biased toward habitats similar to the natal region in some mammals. The benefits and costs of dispersal often differ between sexes, and most mammals show male-biased dispersal in relation to a polygamous mating system. Japanese serow is generally a solitary and monogamous species. However, recent studies have shown that the sociality of serows on Mt. Asama differs between habitat types. In the mountain forests with low forage availability, solitary habits and social monogamy were observed, while, in alpine grasslands, female grouping and social polygyny were observed, which is probably due to abundant forage availability. We investigated the effects of habitat characteristics and sociality on the dispersal of serows using fecal and tissue samples from two different habitats on Mt. Asama. The Fst value between the two areas was significantly positive, and the mean relatedness within areas was significantly higher than that between areas, which suggests limited gene flow and natal habitat-biased dispersal. Bayesian clustering analysis showed unidirectional gene flow from forest to grassland, which was probably due to the high forage availability of the grassland. Analyses of the assignment index and mean relatedness did not show male-biased dispersal, even in the grassland, where serows were polygynous. Thus, polygyny in the grassland is not linked to male-biased dispersal. In summary, our study suggests that dispersal patterns in Japanese serows are affected by habitat rather than social differences.

Development of a high-density sub-species-specific targeted SNP assay for Rocky Mountain bighorn sheep (Ovis canadensis canadensis).

Due to their abundance and relative ease of genotyping, single nucleotide polymorphisms (SNPs) are a commonly used molecular marker for contemporary population genetic and genomic studies. A high-density and cost-effective way to type SNP loci is Allegro targeted genotyping (ATG), which is a form of targeted genotyping by sequencing developed and offered by Tecan genomics. One major drawback of this technology is the need for a reference genome and information on SNP loci when designing a SNP assay. However, for some non-model species genomic information from other closely related species can be used. Here we describe our process of developing an ATG assay to target 50,000 SNPs in Rocky Mountain bighorn sheep, using a reference genome from domestic sheep and SNP resources from prior bighorn sheep studies. We successfully developed a high accuracy, high-density, and relatively low-cost SNP assay for genotyping Rocky Mountain bighorn sheep that genotyped ~45,000 SNP loci. These loci were relatively evenly distributed throughout the genome. Furthermore, the assay produced genotypes at tens of thousands of SNP loci when tested on other mountain sheep species and subspecies.

Estimating Mineral Requirements of Wild Herbivores: Modelling Arctic Caribou (Rangifer tarandus granti) in Summer.

Mineral requirements are poorly described for most wildlife. Consequently, the role of forage minerals in movement and productivity are poorly understood for sedentary and migratory ungulates, such as reindeer and caribou (Rangifer tarandus). We applied estimates of maintenance, lactation, body mass change, and antler growth to production curves (body mass, daily intake, and milk yield) for female caribou to calculate their mineral requirements over summer. The total requirements (mg or g·d-1) were divided by the daily intake (kg·d-1) to estimate the minimum concentration of minerals required in the diet (mg or g·kg-1) to balance demand. The daily requirements (mg·d-1) of all minerals increased from parturition to the end of summer. The minimum dietary concentrations (mg·kg-1) of macro-minerals (Ca, P, Mg, Na, K) declined as food intake (kg·d-1) increased over summer. The minimum dietary concentrations (Fe, Mn, Cu, Zn) were heavily influenced by body mass gain, which increased through late lactation even though food intakes rose. Our modeling framework can be applied to other wild ungulates to assess the impacts of changing forage phenology, plant community compositions, or environmental disturbances on movement and productivity.

Linking African herbivore community enamel isotopes and environments: challenges, opportunities, and paleoecological implications.

Paleoenvironmental reconstructions of fossil sites based on isotopic analyses of enamel typically rely on data from multiple herbivore taxa, with the assumption that this dietary spectrum represents the community's isotopic range and provides insights into local or regional vegetation patterns. However, it remains unclear how representative the sampled taxa are of the broader herbivore community and how well these data correspond to specific ecosystems. Verifying these underlying assumptions is essential to refining the utility of enamel isotopic values for paleoenvironmental reconstructions. This study explores potential links between modern herbivore community carbon isotopic enamel spectra, biome types, and climate in sub-Saharan Africa. This region is one of the most comprehensively isotopically sampled areas globally and is of particular relevance to hominin evolution. Our extensive data compilation reveals that published enamel isotopic data from sub-Saharan Africa typically sample only a small percentage of the taxa documented at most localities and that some biome types (e.g., subtropical savannas) are dramatically overrepresented relative to others (e.g., forests) in these modern data sets. Multiple statistical analyses, including linear models and cluster analyses, revealed weak relationships of associated mammalian herbivore enamel isotopic values, biome type, and climate parameters. These results confound any simple assumptions about how community isotopic profiles map onto specific environments, highlighting the need for more precise strategic approaches in extending isotopic frameworks into the past for paleoecological reconstructions. Developing more refined modern analogs will ultimately allow us to more accurately characterize the isotopic spectra of paleo-communities and link isotopic dietary signatures to specific ecosystems.

Cougars, wolves, and humans drive a dynamic landscape of fear for elk.

To manage predation risk, prey navigate a dynamic landscape of fear, or spatiotemporal variation in risk perception, reflecting predator distributions, traits, and activity cycles. Prey may seek to reduce risk across this landscape using habitat at times and in places when predators are less active. In multipredator landscapes, avoiding one predator could increase vulnerability to another, making the landscape of fear difficult to predict and navigate. Additionally, humans may shape interactions between predators and prey, and induce new sources of risk. Humans can function as a shield, providing a refuge for prey from human-averse carnivores, and as a predator, causing mortality through hunting and vehicle collisions and eliciting a fear response that can exceed that of carnivores. We used telemetry data collected between 2017 and 2021 from 63 Global Positioning System-collared elk (Cervus canadensis), 42 cougars (Puma concolor), and 16 wolves (Canis lupus) to examine how elk habitat selection changed in relation to carnivores and humans in northeastern Washington, USA. Using step selection functions, we evaluated elk habitat use in relation to cougars, wolves, and humans, diel period (daytime vs. nighttime), season (summer calving season vs. fall hunting season), and habitat structure (open vs. closed habitat). The diel cycle was critical to understanding elk movement, allowing elk to reduce encounters with predators where and when they would be the largest threat. Elk strongly avoided cougars at night but had a near-neutral response to cougars during the day, whereas elk avoided wolves at all times of day. Elk generally used more open habitats where cougars and wolves were most active, rather than altering the use of habitat structure depending on the predator species. Elk avoided humans during the day and ~80% of adult female mortality was human caused, suggesting that humans functioned as a "super predator" in this system. Simultaneously, elk leveraged the human shield against wolves but not cougars at night, and no elk were confirmed to have been killed by wolves. Our results add to the mounting evidence that humans profoundly affect predator-prey interactions, highlighting the importance of studying these dynamics in anthropogenic areas.

Weather-dependent changes in habitat use by Alpine chamois.

BACKGROUND: Alterations in weather patterns due to climate change are accelerated in alpine environments, but mountains also provide a wide range of niches and potential refuge areas. In order to identify future critical habitat for mountain ungulates for effective protection, it is important to understand their spatial responses to changing weather conditions without movement constraints by human disturbance. METHODS: Using integrated step selection functions, we investigated fine-scale changes in seasonal habitat use in response to weather and time of day for 55 GPS-collared adult Alpine chamois in summer and 42 individuals in winter in a strictly protected area. RESULTS: Chamois reacted to increasing precipitation and wind speeds primarily by moving to lower elevations in summer and winter. However, reactions to high summer temperatures predominantly involved preferences for increasing tree cover density and northerly slopes. Snow depth had little effect on habitat choice, and southerly slopes were preferred in winter regardless of temperature. At night, chamois moved to steeper slopes and lower elevations than during daytime in both seasons, and to more open areas in summer. Steeper slopes were also preferred with increasing tree cover density. CONCLUSIONS: Chamois employ adaptive fine-scale adjustments in their habitat choice consistent with respect to efficient thermoregulation and protection from both weather extremes and predation risk in summer and winter. Movement responses to climate change are therefore expected to be far more complex than simple altitudinal changes in distribution. Particularly the role of forest cover must not be underestimated, as it appears to provide important thermal refuge habitat from high summer temperatures.

Evaluating movement-based methods for estimating the frequency and timing of parturition in mule deer.

BACKGROUND: Information on reproduction of harvested species such as mule deer (Odocoileus hemionus) is vital for conservation and management. Furthermore, parturition in ungulates may be detected using patterns of movement logged by GPS transmitters. Several movement-based methods have been developed to detect parturition in ungulates including the Peterson method, behavioral change point analysis (BCPA), rolling minimum convex polygons (rMCP), individual-based method (IBM), and population-based method (PBM). Our objectives were to (1) test the accuracy and the precision of each previously described method and (2) develop an improved method optimized for mule deer that incorporated aspects of the other methods. METHODS: We determined parturition timing and status for female mule deer fitted with GPS collars and implanted with vaginal implant transmitters (VITs). We used movement patterns before and after parturition to set movement thresholds for each movement-based method. Following model training, we used location and birth date data from an external dataset to test the effectiveness of each movement-based method. Additionally, we developed a novel method for detecting parturition called the analysis of parturition indicators (API). We used two regression analyses to determine the accuracy and precision of estimates generated by each method. RESULTS: The six methods we employed varied in accuracy, with the API, rMCP, and BCPA being most accurate. Precision also varied among methods, with the API, rMCP, and PBM generating the most precise estimates of parturition dates. The API and the rMCP performed similarly and better overall than any of the other existing methods. CONCLUSIONS: We found that movement-based methods could be used to accurately and precisely detect parturition in mule deer. Further, we determined that the API and rMCP methods had the greatest overall success at detecting parturition in mule deer. The relative success of the API and rMCP may be attributed to the fact that both methods use home range size to detect parturition and are validated using known parturition dates of collared deer. We present the API as an efficient method of estimating birth status and timing of parturition of mule deer fitted with GPS transmitters, as well as affirm the effectiveness of a previously developed method, rMCP.

Predation risk drives long-term shifts in migratory behaviour and demography in a large herbivore population.

Migration is an adaptive life-history strategy across taxa that helps individuals maximise fitness by obtaining forage and avoiding predation risk. The mechanisms driving migratory changes are poorly understood, and links between migratory behaviour, space use, and demographic consequences are rare. Here, we use a nearly 20-year record of individual-based monitoring of a large herbivore, elk (Cervus canadensis) to test hypotheses for changing patterns of migration in and adjacent to a large protected area in Banff National Park (BNP), Canada. We test whether bottom-up (forage quality) or top-down (predation risk) factors explained trends in (i) the proportion of individuals using 5 different migratory tactics, (ii) differences in survival rates of migratory tactics during migration and whilst on summer ranges, (iii) cause-specific mortality by wolves and grizzly bears, and (iv) population abundance. We found dramatic shifts in migration consistent with behavioural plasticity in individual choice of annual migratory routes. Shifts were inconsistent with exposure to the bottom-up benefits of migration. Instead, exposure to landscape gradients in predation risk caused by exploitation outside the protected area drove migratory shifts. Carnivore exploitation outside the protected area led to higher survival rates for female elk remaining resident or migrating outside the protected area. Cause-specific mortality aligned with exposure to predation risk along migratory routes and summer ranges. Wolf predation risk was higher on migratory routes than summer ranges of montane-migrant tactics, but wolf predation risk traded-off with heightened risk from grizzly bears on summer ranges. A novel eastern migrant tactic emerged following a large forest fire that enhanced forage in an area with lower predation risk outside of the protected area. The changes in migratory behaviour translated to population abundance, where abundance of the montane-migratory tactics declined over time. The presence of diverse migratory life histories maintained a higher total population abundance than would have been the case with only one migratory tactic in the population. Our study demonstrates the complex ways in which migratory populations change over time through behavioural plasticity and associated demographic consequences because of individuals balancing predation risk and forage trade-offs.

Reproductive tactics, birth timing and the risk-resource trade-off in an income breeder.

In variable environments, habitats that are rich in resources often carry a higher risk of predation. As a result, natural selection should favour individuals that balance allocation of time to foraging versus avoiding predation through an optimal decision-making process that maximizes fitness. The behavioural trade-off between resource acquisition and risk avoidance is expected to be particularly acute during gestation and lactation, when the energetic demands of reproduction peak. Here, we investigated how reproductive female roe deer adjust their foraging activity and habitat use during the birth period to manage this trade-off compared with non-reproductive juveniles, and how parturition date constrains individual tactics of risk-resource management. Activity of reproductive females more than doubled immediately following parturition, when energy demand is highest. Furthermore, compared with non-reproductive juveniles, they increased their exposure to risk by using open habitat more during daytime and ranging closer to roads. However, these post-partum modifications in behaviour were particularly pronounced in late-parturient females who adopted a more risk-prone tactic, presumably to compensate for the growth handicap of their late-born offspring. In income breeders, individuals that give birth late may be constrained to trade risk avoidance for foraging during peak allocation to reproduction, with probable consequences for individual fitness.