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Behavioural plasticity modulates temperature-related constraints on foraging time for a montane mammal

Contemporary climate change is altering temperature profiles across the globe. Increasing temperatures can reduce the amount of time during which conditions are suitable for animals to engage in essential activities, such as securing food. Behavioural plasticity, the ability to alter behaviour in response to the environment, may provide animals with a tool to adjust to changes in the availability of suitable thermal conditions. The extent to which individuals can alter fitness‐enhancing behaviours, such as food collection, to proximately buffer variation in temperature, however, remains unclear. Even less well understood are the potential performance advantages of flexible strategies among endotherms. We examined the degree to which individuals altered rates of food collection in response to temperature, and two potential benefits, using the American pika ( Ochotona princeps ), a temperature‐sensitive, food‐hoarding mammal, as a model. From July–September 2013–2015, we used motion‐activated cameras and in situ temperature loggers to examine pika food‐caching activity for 72 individuals across 10 sites in the central Rocky Mountains, USA. We quantified % nitrogen by cache volume as a metric of cache quality, and the number of events during which pikas were active in temperatures ≥25°C as a measure of potential thermoregulatory stress. We found a strong negative effect of temperature on the rate at which pikas cached food. Individual responses to temperature varied substantially in both the level of food‐collecting activity and in the degree to which individuals shifted activity with warming temperature. After accounting for available foraging time, individuals that exhibited greater plasticity collected a comparable amount of nitrogen, while simultaneously experiencing fewer occasions in which temperatures eclipsed estimated thermal tolerances. By varying food‐collection norms of reaction, individuals were able to plastically respond to temperature‐driven reductions in foraging time. Through this increased flexibility, individuals amassed food caches of comparable quality, while minimizing exposure to potentially stressful thermal conditions. Our results suggest that, given sufficient resource quality and availability, plasticity in foraging activity may help temperature‐limited endotherms adjust to climate‐related constraints on foraging time.

Wyoming

Top-down effect of repatriating bald eagles hinder jointly recovering competitors

1. The recovery of piscivorous birds around the world is touted as one of the great conservation successes of the 21st century, but for some species, this success was short-lived. Bald eagles, ospreys, and great blue herons began repatriating Voyageurs National Park, USA, in the mid-20th century. However, after 1990, only eagles continued their recovery, while osprey and heron recovery failed for unknown reasons. 2. We aimed to evaluate whether top-down effects of bald eagles, and bottom-up effects of inclement weather, habitat quality, and fish resources contributed to the failed recovery of ospreys and herons in a protected area. 3. We quantified the relative influence of top-down and bottom-up factors on nest colonization, persistence (i.e., nest reuse) and success for ospreys, and occurrence and size of heronries using 26 years (1986-2012) of spatially-explicit monitoring data coupled with multi-response hierarchical models and Bayesian variable selection approaches. 4. Bald eagles were previously shown to recover faster due to intensive nest protection and management. Increased numbers of eagles were associated with a reduction in the numbers of osprey nests, their nesting success, and heronry size; while higher local densities of nesting eagles deterred heronries nearby. We found little evidence of bottom-up limitations on the failed recovery of herons and ospreys. 5. We present a conservation conundrum: bald eagles are top predators and a flagship species of conservation that have benefited from intensive protection, but this likely hindered the recovery of ospreys and herons. Returning top predators, or rewilding, is widely promoted as a conservation strategy for top-down ecosystem recovery, but managing top predators in isolation of jointly recovering species can halt or reverse ecosystem recovery. Previous studies warn of the potential consequences of ignoring biotic interactions amongst recovering species, but we go further by quantifying how these interactions contributed to failed recoveries via impacts on the nesting demography of jointly recovering species. Multi-species management is paramount to realizing the ecosystem benefits of top predator recovery.

Minnesota

Native forage mediates influence of irrigated agriculture on migratory behaviour of elk

Ungulates migrate to maximize nutritional intake when forage varies seasonally. Populations of ungulates often include both migratory and non-migratory individuals, but the mechanisms driving individual differences in migratory behaviour are not well-understood. We quantified associations between hypothesized drivers of partial migration and the likelihood of migration for individual ungulates that experienced a range of environmental conditions and anthropogenic influences. We evaluated the effects of forage variation, conspecific density, and human land uses on migratory behaviour of 308 adult female elk in 16 herds across western Montana. We found irrigated agriculture on an individual's winter range reduced migratory behaviour, but individuals were more likely to migrate away from irrigated agricultural areas if better forage was available elsewhere or if they experienced high conspecific density on their winter range. When the forage available during the summer growing season varied predictably between years, elk were more likely to migrate regardless of whether they had access to irrigated agriculture. Our study shows that predictable availability of beneficial native forage can encourage migration even for ungulates with irrigated agriculture on their winter range. Perturbations that can affect the forage available to ungulates include wildfires, timber harvest, livestock grazing and changing weather patterns. If these or other disturbances negatively affect forage on summer ranges of migrants, or if they cause forage to vary unpredictably across space and time, our results suggest migratory behaviour may decline as a result.

Montana

Variation in selective regimes drives intraspecific variation in life-history traits and migratory behaviour along an elevational gradient

Comparative studies, across and within taxa, have made important contributions to our understanding of the evolutionary processes that promote phenotypic diversity. Trait variation along geographic gradients provides a convenient heuristic for understanding what drives and maintains diversity. Intraspecific trait variation along latitudinal gradients is well-known, but elevational variation in the same traits is rarely documented. Trait variation along continuous elevational gradients, however, provides compelling evidence that individuals within a breeding population may experience different selective pressures. Our objectives were to quantify variation in a suite of traits along a continuous elevational gradient, evaluate whether individuals in the population experience different selective pressures along that gradient and quantify variation in migratory tendency along that gradient. We examined variation in a suite of 14 life-history, morphological and behavioural traits, including migratory tendency, of yellow-eyed juncos along a continuous 1000-m elevational gradient in the Santa Catalina Mountains of Arizona. Many traits we examined varied along the elevational gradient. Nest survival and nestling growth rates increased, while breeding season length, renesting propensity and adult survival declined, with increasing elevation. We documented the migratory phenotype of juncos (partial altitudinal migrants) and show that individual migratory tendency is higher among females than males and increases with breeding elevation. Our data support the paradigm that variation in breeding season length is a major selective pressure driving life-history variation along elevational gradients and that individuals breeding at high elevation pursue strategies that favour offspring quality over offspring quantity. Furthermore, a negative association between adult survival and breeding elevation and a positive association between nest survival and breeding elevation help explain both the downslope and reciprocal upslope seasonal migratory movements that characterize altitudinal migration in many birds. Our results demonstrate how detailed studies of intraspecific variation in suites of traits along environmental gradients can lend new insights into the evolutionary processes that promote diversification and speciation, the causes of migratory behaviour, and how animal populations will likely respond to climate change.

Journal of Animal Ecology

Sources of variation in maternal allocation in a long-lived mammal

1. Life history theory predicts allocation of energy to reproduction varies with maternal age, but additional maternal features may be important to the allocation of energy to reproduction. We aimed to characterize age‐specific variation in maternal allocation and assess the relationship between maternal allocation and other static and dynamic maternal features. Mass measurements of 531 mothers and pups were used with Bayesian hierarchical models to explain the relationship between diverse maternal attributes and both the proportion of mass allocated by Weddell seal mothers, and the efficiency of mass transfer from mother to pup during lactation as well as the weaning mass of pups. Our results demonstrated that maternal mass was strongly and positively associated with the relative reserves allocated by a mother and a pup's weaning mass but that the efficiency of mass transfer declines with maternal parturition mass. Birthdate was positively associated with proportion mass allocation and pup weaning mass, but mass transfer efficiency was predicted to be highest at the mean birthdate. The relative allocation of maternal reserves declined with maternal age but the efficiency of mass transfer to pups increases, suggestive of selective disappearance of poor‐quality mothers. These findings highlight the importance of considering multiple maternal features when assessing variation in maternal allocation.

Journal of Animal Ecology

Wildfire reveals transient changes to individual traits and population responses of a native bumble bee ( Bombus vosnesenskii )

1. Fire-induced changes in the abundance and distribution of organisms, especially plants, can alter resource landscapes for mobile consumers driving bottom-up effects on their population sizes, morphologies, and reproductive potential. We expect these impacts to be most striking for obligate visitors of plants, like bees and other pollinators, but these impacts can be difficult to interpret due to the limited information provided by forager counts in the absence of survival or fitness proxies. 2. Increased bumble bee worker abundance is often coincident with the pulses of flowers that follow recent fire. However, it is unknown if observed postfire activity is due to underlying population growth or a stable pool of colonies recruiting more foragers to abundant resource patches. This distinction is necessary for determining the net impact of disturbance on bumble bees: are there population-wide responses or do just a few colonies reap the rewards? 3. We estimated colony abundance before and after fire in burned and unburned areas using a genetic mark-recapture framework. We paired colony abundance estimates with measures of body size, counts of queens, and estimates of foraging and dispersal to assess changes in worker size, reproductive output, and landscape-scale movements. 4. Higher floral abundance following fire not only increased forager abundance, but also the number of colonies from which those foragers came. Importantly, despite a larger population size we also observed increased mean worker size. Two years following fire, queen abundance was higher in both burned and unburned sites, potentially due to the dispersal of queens from burned into unburned areas. The effects of fire were transient; within two growing seasons, worker abundance was substantially reduced across the entire sampling area and body sizes were similar between burned and unburned sites. 5. Our results reveal how disturbance can temporarily release populations from resource limitation, boosting the genetic diversity, body size, and reproductive output of populations. Given that the effects of fire on bumble bees acted indirectly through pulsed resource availability, it is likely our results are generalizable to other situations, such as habitat restorations, where resource density is enhanced within the landscape.

California

Mate fidelity improves survival and breeding propensity of a long‐lived bird

Evolutionary and behavioural ecologists have long been interested in factors shaping the variation in mating behaviour observed in nature. Although much of the research on this topic has focused on the consequences of mate choice and mate change on annual reproductive success, studies of a potential positive link between mate fidelity and adult demographic rates have been comparatively rare. This is particularly true for long‐lived birds with multi‐year, socially monogamous pair bonds. We used a 26‐year capture–mark–recapture dataset of 3,330 black brent Branta bernicla nigricans to test whether breeding with a familiar mate improved future breeding propensity and survival. We predicted that experienced breeders nesting with a new partner would have rates of survival similar to familiar pairs because long‐lived species avoid jeopardizing survival since their lifetime fitness is sensitive to this vital rate. In contrast, we expected that any costs of breeding with a new partner would be paid through skipping the subsequent breeding attempt. We found that unfamiliar pairs had lower subsequent breeding propensity than faithful partners. However, contrary to our expectations, individuals breeding with a new mate also suffered reduced survival. These results add to a small number of studies indicating that a positive relationship between mate retention and adult demographic rates may exist in a diverse array of avian species. Given these results, researchers should consider costs of mate change that extend beyond within‐season reproductive success to fully understand the potential adaptive basis for perennial social monogamy. We caution that if mate retention enhances survival prospects, improvements in annual reproductive success with pair‐bond length could be a secondary factor favouring perennial social monogamy, particularly in species with slower life‐history strategies. Furthermore, some cases where annual reproductive success does not improve with pair‐bond duration, yet multi‐year pair bonds are common, could be explained by benefits afforded by mate fidelity to adult vital rates.

Journal of Animal Ecology

A test of the Niche Variation Hypothesis in a ruminant herbivore

Despite the shared prediction that the width of a population's dietary niche expands as food becomes limiting, the Niche Variation Hypothesis (NVH) and Optimal Foraging Theory (OFT) offer contrasting views about how individuals alter diet selection when food is limited. Classical OFT predicts that dietary preferences do not change as food becomes limiting, so individuals expand their diets as they compensate for a lack of preferred foods. In contrast, the NVH predicts that among-individual variation in cognition, physiology or morphology create functional trade-offs in foraging efficiency, thereby causing individuals to specialize on different subsets of food as food becomes limiting. To evaluate (a) the predictions of the NVH and OFT and (b) evidence for physiological and cognitive-based functional trade-offs, we used DNA microsatellites and metabarcoding to quantify the diet, microbiome and genetic relatedness (a proxy for social learning) of 218 moose Alces alces across six populations that varied in their degree of food limitation. Consistent with both the NVH and OFT, dietary niche breadth increased with food limitation. Increased diet breadth of individuals—rather than increased diet specialization—was strongly correlated with both food limitation and dietary niche breadth of populations, indicating that moose foraged in accordance with OFT. Diets were not constrained by inheritance of the microbiome or inheritance of diet selection, offering support for the little-tested hypothesis that functional trade-offs in food use (or lack thereof) determine whether populations adhere to the predictions of the NVH or OFT. Our results indicate that both the absence of strong functional trade-offs and the digestive physiology of ruminants provide contexts under which populations should forage in accordance with OFT rather than the NVH. Also, because dietary niche width increased with increased food limitation, OFT and the NVH provide theoretical support for the notion that plant–herbivore interaction networks are plastic rather than static, which has important implications for understanding interspecific niche partitioning. Lastly, because population-level dietary niche breadth and calf recruitment are correlated, and because calf recruitment can be a proxy for food limitation, our work demonstrates how diet data can be employed to understand a populations' proximity to carrying capacity.

Colorado, Utah, Wyoming

The demographic contributions of connectivity versus local dynamics to population growth of an endangered bird

Conservation and management increasingly focus on connectivity, because connectivity driven by variation in immigration rates across landscapes is thought to be crucial for maintaining local population and metapopulation persistence. Yet, efforts to quantify the relative role of immigration on population growth across the entire range of species and over time have been lacking. We assessed whether immigration limited local and range‐wide population growth of the endangered snail kite Rostrhamus sociabilis in Florida, USA, over 18 years using multi‐state, reverse‐time modelling that accounts for imperfect detection of individuals and unobservable states. Demographic contributions of immigration varied depending on the dynamics and geographic position of the local populations, were scale‐dependent and changed over time. By comparing the relative contributions of immigration versus local demography for periods of significant change in local abundance, we found empirical evidence for a disproportionately large role of immigration in facilitating population growth of a centrally located population—a connectivity ‘hub’. The importance of connectivity changed depending of the spatial scale considered, such that immigration was a more important driver of population growth at small versus large spatial scales. Furthermore, the contribution of immigration was much greater during time periods when local population size was small, emphasizing abundance‐dependent rescue effects. Our findings suggest that efforts aimed at improving local breeding habitat will likely be most effective at increasing snail kite population growth. More broadly, our results provide much needed information on the role of connectivity for population growth, suggesting that connectivity conservation may have the greatest benefits when efforts focus on centrally located habitat patches and small populations. Furthermore, our results highlight that connectivity is highly dynamic over time and that interpreting the effects of connectivity at local scales may not transfer to region‐wide dynamics.

Florida

The influence of species life history and distribution characteristics on species responses to habitat fragmentation in an urban landscape

Fragmentation within urbanized environments often leads to a loss of native species diversity; however, variation exists in responses among-species and among-populations within species. We aimed to identify patterns in species biogeography in an urbanized landscape to understand anthropogenic effects on vertebrate communities and identify species that are more sensitive or resilient to landscape change. We investigated patterns in species richness and species responses to fragmentation in southern Californian small vertebrate communities using multispecies occupancy models and determined factors associated with overall commonness and sensitivity to patch size for 45 small vertebrate species both among and within remaining non-developed patches. In general, smaller patches had fewer species, with amphibian species richness being particularly sensitive to patch size effects. Mammals were generally more common, occurring both in a greater proportion of patches and a higher proportion of the sites within occupied patches. Alternatively, amphibians were generally restricted to larger patches but were more ubiquitous within smaller patches when occupied. Species range size was positively correlated with how common a species was across and within patches, even when controlling for only patches that fell within a species' range. We found sensitivity to patch size was greater for more fecund species and depended on where the patch occurred within a species' range. While all taxa were more likely to occur in patches in the warmer portions of their ranges, amphibians and mammals were more sensitive to fragmentation in these warmer areas as compared to the rest of their ranges. Similarly, amphibians occurred at a smaller proportion of sites within patches in drier portions of their ranges. Mammals occurred at a higher proportion of sites that were also in drier portions of their range while reptiles did not differ in their sensitivity to patch size by range position. We demonstrate that taxonomy, life history, range size and range position can predict commonness and sensitivity of species across this highly fragmented yet biodiverse landscape. The impacts of fragmentation on species communities within an urban landscape depend on scale, with differences emerging among and within species and populations.

California

Age-related reproductive performance of the Adélie penguin, a long-lived seabird exhibiting similar outcomes regardless of individual life-history strategy

Age-related variation in reproductive performance in long-lived iteroparous vertebrate species is common, with performance being influenced by within-individual processes, such as improvement and senescence, in combination with among-individual processes, such as selective appearance and disappearance. Few studies of age-related reproductive performance have compared the role of these drivers within a metapopulation, subject to varying degrees of resource competition. We accounted for within- and among-individual changes among known-aged Adélie penguins Pygoscelis adeliae during 17 years (1997-2013), at three clustered colonies of disparate size, to understand patterns in age-related reproductive success during early and late adulthood. Age at first reproduction (AFR) was lowest, and number of breeding attempts highest, at the largest colony. Regardless of AFR, success improved with early post-recruitment experience. For both oldest and youngest recruitment groups, peak performance occurred at the end of their reproductive life span indicating a possible cost of reproduction. Intermediate recruitment groups reached peak performance in their mid-reproductive life span and with intermediate breeding experience, before decreasing. Breeding success was lowest for the initial breeding attempt regardless of AFR, but we observed subsequent variation relative to recruitment age. Gaining experience by delaying recruitment positively influenced reproductive performance early in the reproductive life span and was most evident for the youngest breeders. Oldest recruits had the highest initial and peak breeding success. Differences in AFR resulted in trade-offs in reproductive life span or timing of senescence but not in the overall number of breeding attempts. Patterns differed as a function of colony size, and thus competition for resources. Early life improvement in performance at the larger colonies was primarily due to within-individual factors and at the largest colony, AFR. Regardless of colony size late-life performance was positively related to the age at last reproduction, indicating selective disappearance of lower performing individuals. These results highlight that different life-history strategies were equally successful, indicating that individuals can overcome potential trade-offs associated with early- and late-life performance. These results have important implications for understanding the evolution of life-history strategies responsible for driving population change.

Beaufort Island ,Ross Island

Drivers of site fidelity in ungulates

While the tendency to return to previously visited locations—termed ‘site fidelity’—is common in animals, the cause of this behaviour is not well understood. One hypothesis is that site fidelity is shaped by an animal's environment, such that animals living in landscapes with predictable resources have stronger site fidelity. Site fidelity may also be conditional on the success of animals’ recent visits to that location, and it may become stronger with age as the animal accumulates experience in their landscape. Finally, differences between species, such as the way memory shapes site attractiveness, may interact with environmental drivers to modulate the strength of site fidelity. We compared inter-year site fidelity in 669 individuals across eight ungulate species fitted with GPS collars and occupying a range of environmental conditions in North America and Africa. We used a distance-based index of site fidelity and tested hypothesized drivers of site fidelity using linear mixed effects models, while accounting for variation in annual range size. Mule deer Odocoileus hemionus and moose Alces alces exhibited relatively strong site fidelity, while wildebeest Connochaetes taurinus and barren-ground caribou Rangifer tarandus granti had relatively weak fidelity. Site fidelity was strongest in predictable landscapes where vegetative greening occurred at regular intervals over time (i.e. high temporal contingency). Species differed in their response to spatial heterogeneity in greenness (i.e. spatial constancy). Site fidelity varied seasonally in some species, but remained constant over time in others. Elk employed a ‘win-stay, lose-switch’ strategy, in which successful resource tracking in the springtime resulted in strong site fidelity the following spring. Site fidelity did not vary with age in any species tested. Our results provide support for the environmental hypothesis, particularly that regularity in vegetative phenology shapes the strength of site fidelity at the inter-annual scale. Large unexplained differences in site fidelity suggest that other factors, possibly species-specific differences in attraction to known sites, contribute to variation in the expression of this behaviour. Understanding drivers of variation in site fidelity across groups of organisms living in different environments provides important behavioural context for predicting how animals will respond to environmental change.

Journal of Animal Ecology

Genomic signatures of thermal adaptation are associated with clinal shifts of life history in a broadly distributed frog

Temperature is a critical driver of ectotherm life-history strategies, whereby a warmer environment is associated with increased growth, reduced longevity and accelerated senescence. Increasing evidence indicates that thermal adaptation may underlie such life-history shifts in wild populations. Single nucleotide polymorphisms (SNPs) and copy number variants (CNVs) can help uncover the molecular mechanisms of temperature-driven variation in growth, longevity and senescence. However, our understanding of these mechanisms is still limited, which reduces our ability to predict the response of non-model ectotherms to global temperature change. In this study, we examined the potential role of thermal adaptation in clinal shifts of life-history traits (i.e. life span, senescence rate and recruitment) in the Columbia spotted frog Rana luteiventris along a broad temperature gradient in the western United States. We took advantage of extensive capture–recapture datasets of 20,033 marked individuals from eight populations surveyed annually for 14–18 years to examine how mean annual temperature and precipitation influenced demographic parameters (i.e. adult survival, life span, senescence rate, recruitment and population growth). After showing that temperature was the main climatic predictor influencing demography, we used RAD-seq data (50,829 SNPs and 6,599 putative CNVs) generated for 352 individuals from 31 breeding sites to identify the genomic signatures of thermal adaptation. Our results showed that temperature was negatively associated with annual adult survival and reproductive life span and positively associated with senescence rate. By contrast, recruitment increased with temperature, promoting the long-term viability of most populations. These temperature-dependent demographic changes were associated with strong genomic signatures of thermal adaptation. We identified 148 SNP candidates associated with temperature including three SNPs located within protein-coding genes regulating resistance to cold and hypoxia, immunity and reproduction in ranids. We also identified 39 CNV candidates (including within 38 transposable elements) for which normalized read depth was associated with temperature. Our study indicates that both SNPs and structural variants are associated with temperature and could eventually be found to play a functional role in clinal shifts in senescence rate and life-history strategies in R. luteiventris . These results highlight the potential role of different sources of molecular variation in the response of ectotherms to environmental temperature variation in the context of global warming.

Journal of Animal Ecology

Threading the needle: How humans influence predator–prey spatiotemporal interactions in a multiple‐predator system

Perceived predation risk and the resulting antipredator behaviour varies across space, time and predator identity. Communities with multiple predators that interact and differ in their use of space, time of activity and hunting mode create a complex landscape for prey to avoid predation. Anthropogenic presence and disturbance have the potential to shift interactions among predators and prey and the where and when encounters occur. We examined how white-tailed deer Odocoileus virginianus fawn spatiotemporal antipredator behaviour differed along an anthropogenic disturbance gradient that had black bears Ursus americanus , coyotes Canis latrans , bobcats Lynx rufus and humans present. We quantified (a) spatial co-occurrence in species distributions, (b) temporal overlap across the diel cycle and (c) spatiotemporal associations between humans, bears, coyotes, bobcats, adult male deer and fawns. We also examined how deer vigilance behaviour changed across the anthropogenic disturbance gradient and survey duration. Anthropogenic disturbance influenced spatiotemporal co-occurrence across multiple scales, often increasing spatiotemporal overlap among species. In general, species’ spatial co-occurrence was neutral or positive in anthropogenically disturbed environments. Bears and fawns, coyotes and adult male deer, and bobcats and fawns all had higher temporal overlap in the agriculture-development matrix sites. In addition, factors that influenced deer vigilance (e.g. distance to forest edge and predator relative abundance) in the agriculture-development matrix sites did not in the forest matrix site. By taking into account the different antipredator behaviours that can be detected and the different scales these behaviours might occur, we were able to gain a more comprehensive picture of how humans reduce available niche space for wildlife, creating the neutral and positive spatiotemporal associations between species that studies have been seeing in more disturbed areas.

Pennsylvania

Examination of the interaction between age-specific predation and chronic disease in the Greater Yellowstone Ecosystem

Predators may create healthier prey populations by selectively removing diseased individuals. Predators typically prefer some ages of prey over others, which may, or may not, align with those prey ages that are most likely to be diseased. The interaction of age-specific infection and predation has not been previously explored and likely has sizable effects on disease dynamics. We hypothesize that predator cleansing effects will be greater when the disease and predation occur in the same prey age groups. We examine the predator cleansing effect using a model where both vulnerability to predators and pathogen prevalence vary with age. We tailor this model to chronic wasting disease (CWD) in mule deer and elk populations in the Greater Yellowstone Ecosystem, with empirical data from Yellowstone grey wolves and cougars. Model results suggest that under moderate, yet realistic, predation pressure from cougars and wolves independently, predators may decrease CWD outbreak size substantially and delay the accumulation of symptomatic deer and elk. The magnitude of this effect is driven by the ability of predators to selectively remove late-stage CWD infections that are likely the most responsible for transmission, but this may not be the age class they typically select. Thus, predators that select for infected young adults over uninfected juveniles have a stronger cleansing effect, and these effects are strengthened when transmission rates increase with increasing prey morbidity. There are also trade-offs from a management perspective—that is, increasing predator kill rates can result in opposing forces on prey abundance and CWD prevalence. Our modelling exploration shows that predators have the potential to reduce prevalence in prey populations when prey age and disease severity are considered, yet the strength of this effect is influenced by predators' selection for demography or body condition. Current CWD management focuses on increasing cervid hunting as the primary management tool, and our results suggest predators may also be a useful tool under certain conditions, but not necessarily without additional impacts on host abundance and demography. Protected areas with predator populations will play a large role in informing the debate over predator impacts on disease.

Wyoming

Using piecewise regression to identify biological phenomena in biotelemetry datasets

1. Technological advances in the field of animal tracking have greatly expanded the potential to remotely monitor animals, opening the door to exploring how animals shift their behavior over time or respond to external stimuli. A wide variety of animal-borne sensors can provide information on an animal’s location, movement characteristics, external environmental conditions, and internal physiological status. 2. Here, we demonstrate how piecewise regression can be used to identify the presence and timing of potential shifts in a variety of biological responses using GPS telemetry and other biologging data streams. Different biological latent states can be inferred by partitioning a time-series into multiple segments based on changes in modeled responses (e.g., their mean, variance, trend, degree of autocorrelation) and specifying a unique model structure for each interval. 3. We provide six example applications highlighting a variety of taxonomic species, data streams, timescales and biological phenomena. These examples include a short-term behavioural response (flee and return) by a trumpeter swan Cygnus buccinator following a GPS collar deployment; remote identification of parturition based on movements by a pregnant moose Alces alces ; a physiological response (spike in heart-rate) in a black bear Ursus americanus to a stressful stimulus(presence of a drone); a mortality event of a trumpeter swan signalled by changes in collar temperature and overall dynamic body acceleration; an unsupervised method for identifying the onset, return, duration and staging use of sandhill crane Antigone canadensis migration; and estimation of the transition between incubation and brood-rearing (i.e. hatching) for a breeding trumpeter swan. 4. We implement analyses using the MCP package in R, which provides functionality for specifying and fitting a wide variety of user-defined model structures in a Bayesian framework and methods for assessing and comparing models using information criteria and cross-validation measures.

Journal of Animal Ecology

Identifying mechanisms underlying individual body size increases in a changing, highly seasonal environment: The growing trout of West Brook

As air temperature increases, it has been suggested that smaller individual body size may be a general response to climate warming. However, for ectotherms inhabiting cold, highly seasonal environments, warming temperatures may increase the scope for growth and result in larger body size. In a long-term study of individual brook trout Salvelinus fontinalis and brown trout Salmo trutta inhabiting a small stream network, individual lengths increased over the course of 15 years. As size-selective gains and losses to the population acted to reduce body sizes and mean body size at first tagging in the autumn (<60 mm) were not observed to change substantially over time, the increase in body size was best explained by higher individual growth rates. For brook trout, increasing water temperatures during the spring (when both trout species accomplish most of their total annual growth) was the primary driver of growth rate for juvenile fish and the environmental factor which best explained increases in individual body size over time. For brown trout, by contrast, reduction in and subsequent elimination of juvenile Atlantic salmon Salmo salar midway through the study period explained most of the increases in juvenile growth and body size. In addition to these major trends, a considerable amount of interannual variation in trout growth and body size was explained by other abiotic (stream flow) and biotic (population density) factors with the direction and magnitude of these effects differing by season, age-class and species. For example, stream flow was the dominant growth rate driver for adult fish with strong positive effects in the summer and autumn, but flow variation could not explain increases in body size as we observed no trend in flow. Overall, our work supports the general contention that for high-latitude ectotherms, increasing spring temperatures associated with a warming climate can result in increased growth and individual body size (up to a point), but context-dependent change in other factors can substantially contribute to both interannual variation and longer-term effects.

Massachusetts

Disentangling direct and indirect effects of extreme events on coastal wetland communities

One of the primary ways in which climate change will impact coastal freshwater wetlands is through changes in the frequency, intensity, timing and distribution of extreme weather events. Disentangling the direct and indirect mechanisms of population- and community-level responses to extreme events is vital to predicting how species composition of coastal wetlands will change under future conditions. We extended static structural equation modelling approaches to incorporate system dynamics in a multi-year multispecies occupancy model to quantify the effects of extreme weather events on a coastal freshwater wetland system. We used data from an 8-year study (2009–2016) on St. Marks National Wildlife Refuge in Florida, USA, to quantify species-specific and community-level changes in amphibian and fish occupancy associated with two flooding events in 2012 and 2013. We examine how physical changes to the landscape, including potential changes in salinity and increased wetland connectivity, may have contributed to or exacerbated the effects of these extreme weather events on the biota of isolated coastal wetlands. We provide evidence that the primary effects of flooding on the amphibian community were through indirect mechanisms via changes in the composition of the sympatric fish community that may have had lethal (i.e. through direct predation) or non-lethal (i.e. through direct or indirect competitive interactions) effects. In addition, we have shown that amphibian species differed in their sensitivity to direct flooding effects and indirect changes in the fish community and wetland-specific conductance, which led to variable responses across the community. These effects led to the overall decline in amphibian species richness from 2009 to 2016, suggesting that wetland-breeding amphibian communities on St. Marks National Wildlife Refuge may not be resilient to predicted changes in coastal disturbance regimes because of climate change. Understanding both direct and indirect effects, as well as species interactions, is important for predicting the effects of a changing climate on individual species, communities and ecosystems.

Journal of Animal Ecology