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Burrowing species rely on subterranean and subnivean sites to fulfill important life-history and behavioral processes, including predator avoidance, thermoregulation, resting, and reproduction. For these species, burrow architecture can affect the quality and success of such processes, since characteristics like tunnel width and chamber depth influence access by predators, thermal insulation, and energy spent digging. Wolverines (Gulo gulo) living in Arctic tundra environments dig burrows in snow during winter for resting sites and reproductive dens, but there are few published descriptions of such burrows. We visited 114 resting burrows and describe associated architectural characteristics and non-snow structure. Additionally, we describe characteristics of 15 reproductive den sites that we visited during winter and summer. Although many resting burrows were solely excavated in snow, most incorporated terrain structures including cliffs, talus, river shelf ice, thermokarst caves, and stream cutbanks. Burrows typically consisted of a single tunnel leading to a single chamber, though some burrows had multiple entrances, branching tunnels, or both. Tunnels in resting burrows were shorter than those in reproductive dens, and resting chambers were typically located at the deepest part of the burrow. Reproductive dens were associated with snowdrift-forming terrain features such as streambeds, cutbanks on lake edges, thermokarst caves, and boulders. Understanding such characteristics of Arctic wolverine resting and reproductive structures is critical for assessing anthropogenic impacts as snowpack undergoes climate-driven shifts.

Hibernation is associated with long lifespan: on average, hibernating mammals live 15% longer than nonhibernators of equivalent mass. We investigated how survival varies with sex, season, and the deployment of biologgers in arctic ground squirrels (Urocitellus parryii (Richardson, 1825)), a widely distributed northern hibernator. The duration of hibernation in arctic ground squirrels differs markedly by sex: females hibernate 30% longer each year than males, a behavioural trait that could positively affect female survival. Additionally, males engage in aggressive territorial and food cache defense in spring and fall, which may decrease survival in this sex. From 13 years of mark–recapture data, we estimated apparent survival of arctic ground squirrels in Arctic Alaska (USA) using Cormack–Jolly–Seber models in program MARK. We found that females had higher annual survival ([Formula: see text] (95% confidence interval (CI): 0.469, 0.913)) than males ([Formula: see text] (95% CI: 0.416, 0.670)), with a maximum observed lifespan (10 years) that exceeded that of males (6 years). We also show that biologger use and implantation did not significantly impact survival. Quantifying basic arctic ground squirrel demographics from this well-studied population illustrates how sex-specific hibernation parameters may influence lifespan differences in male and female arctic ground squirrels and provides support for the safety of biologging devices.

Background Migrations in temperate systems typically have two migratory phases, spring and autumn, and many migratory ungulates track the pulse of spring vegetation growth during a synchronized spring migration. In contrast, autumn migrations are generally less synchronous and the cues driving them remain understudied. Our goal was to identify the cues that migrants use in deciding when to initiate migration and how this is updated while en route. Methods We analyzed autumn migrations of Arctic barren-ground caribou (Rangifer tarandus) as a series of persistent and directional movements and assessed the influence of a suite of environmental factors. We fitted a dynamic-parameter movement model at the individual-level and estimated annual population-level parameters for weather covariates on 389 individual-seasons across 9 years. Results Our results revealed strong, consistent effects of decreasing temperature and increasing snow depth on migratory movements, indicating that caribou continuously update their migratory decision based on dynamic environmental conditions. This suggests that individuals pace migration along gradients of these environmental variables. Whereas temperature and snow appeared to be the most consistent cues for migration, we also found interannual variability in the effect of wind, NDVI, and barometric pressure. The dispersed distribution of individuals in autumn resulted in diverse environmental conditions experienced by individual caribou and thus pronounced variability in migratory patterns. Conclusions By analyzing autumn migration as a continuous process across the entire migration period, we found that caribou migration was largely related to temperature and snow conditions experienced throughout the journey. This mechanism of pacing autumn migration based on indicators of the approaching winter is analogous to the more widely researched mechanism of spring migration, when many migrants pace migration with a resource wave. Such a similarity in mechanisms highlights the different environmental stimuli to which migrants have adapted their movements throughout their annual cycle. These insights have implications for how long-distance migratory patterns may change as the Arctic climate continues to warm.

To increase the probability of detecting odour plumes, and so increase prey capture success, when winds are stable central place foraging seabirds should fly crosswind to maximize the round-trip distance covered. At present, however, there is no empirical evidence of this theoretical prediction. Here, using an extensive GPS tracking dataset, we investigate, for the first time, the foraging movements of Bulwer's petrels (Bulweria bulwerii) in the persistent North Atlantic trade winds. To test the hypotheses that, in stable winds, petrels use crosswind to maximize both the distance covered and the probability of detecting olfactory cues, we combine state-space models, generalized additive models and Gaussian plume models. Bulwer's petrels had the highest degree of selectivity for crosswinds documented to date, often leading to systematic zig-zag flights. Crosswinds maximized both the distance travelled and the probability of detecting odour plumes integrated across the round-trip (rather than at any given point along the route, which would result in energetically costly return flight). This evidence suggests that petrels plan round-trip flights at departure, integrating expected costs of homeward journeys. Our findings, which are probably true for other seabirds in similar settings, further highlight the critical role of wind in seabird foraging ecology., This work was funded by the Fundação para a Ciência e a Tecnologia (FCT, Portugal) and the European Regional Development Fund through the projects: UIDB/04292/2020 and UIDP/04292/2020, granted to MARE; UIDP/50017/2020, UIDB/50017/2020 and LA/P/0094/2020, granted to CESAM; and PD/BD/135537/2018 awarded to F.V.

Climate change is rapidly altering the composition and availability of snow, with implications for snow-affected ecological processes, including reproduction, predation, habitat selection, and migration. How snowpack changes influence these ecological processes is mediated by physical snowpack properties, such as depth, density, hardness, and strength, each of which is in turn affected by climate change. Despite this, it remains difficult to obtain meaningful snow information relevant to the ecological processes of interest, precluding a mechanistic understanding of these effects. This problem is acute for species that rely on particular attributes of the subnivean space, for example depth, thermal resistance, and structural stability, for key life-history processes like reproduction, thermoregulation, and predation avoidance. We used a spatially explicit snow evolution model to investigate how habitat selection of a species that uses the subnivean space, the wolverine, is related to snow depth, snow density, and snow melt on Arctic tundra. We modeled these snow properties at a 10 m spatial and a daily temporal resolution for 3 years, and used integrated step selection analyses of GPS collar data from 21 wolverines to determine how these snow properties influenced habitat selection and movement. We found that wolverines selected deeper, denser snow, but only when it was not undergoing melt, bolstering the evidence that these snow properties are important to species that use the Arctic snowpack for subnivean resting sites and dens. We discuss the implications of these findings in the context of climate change impacts on subnivean species.

Significant reductions in sea ice in the Pacific Arctic have occurred over the last 2 decades. Comparing the results of similarly conducted studies, but from different time periods, can increase our understanding of how marine mammals are responding to this change. We modeled the habitat selection and movement behavior of juvenile bearded seals Erignathus barbatus tagged with satellite transmitters in the Bering, Chukchi, and Beaufort seas during 2014-2018, a period of rapid decline in sea ice cover. We compared our results to an earlier study of juvenile bearded seals tagged in the Chukchi Sea during 2004-2009, a period of relatively stable sea ice coverage, and found differences. Seals in the earlier period strongly selected habitat near the ice edge and intermediate ice concentrations (50-60%) in both winter and spring. Seals in the later period strongly selected habitat away from the ice edge, showed no selection for any ice concentration in winter, and weakly selected low ice concentrations in spring (

Human modification of landscapes includes extensive addition of linear features, such as roads and transmission lines. These can alter animal movement and space use and affect the intensity of interactions among species, including predation and competition. Effects of linear features on animal movement have seen relatively little research in avian systems, despite ample evidence of their effects in mammalian systems and that some types of linear features, including both roads and transmission lines, are substantial sources of mortality. Here, we used satellite telemetry combined with step-selection functions designed to explicitly incorporate the energy landscape (el-SSFs) to investigate the effects of linear features and habitat on movements and space use of a large soaring bird, the golden eagleAquila chrysaetos, during migration. Our sample consisted of 32 adult eagles tracked for 45 spring and 39 fall migrations from 2014-2017. Fitted el-SSFs indicated eagles had a strong general preference for south-facing slopes, where thermal uplift develops predictably, and that these areas are likely important aspects of migratory pathways. el-SSFs also revealed that roads and railroads affected movement during both spring and fall migrations, but eagles selected areas near roads to a greater degree in spring compared to fall and at higher latitudes compared to lower latitudes. During spring, time spent near linear features often occurred during slower-paced or stopover movements, perhaps in part to access carrion produced by vehicle collisions. Regardless of the behavioral mechanism of selection, use of these features could expose eagles and other soaring species to elevated risk via collision with vehicles and/or transmission lines. Linear features have been previously documented to affect the ecology of terrestrial species (e.g., large mammals) by modifying individuals’ movement patterns; our work shows these effects on movement extend to avian taxa.

In populations across many taxa, a large fraction of sexually mature individuals do not breed but are attempting to enter the breeding population. Such individuals, often referred to as "floaters," can play critical roles in the dynamics and stability of these populations and buffer them through periods of high adult mortality. Floaters are difficult to study, however, so we lack data needed to understand their roles in the population ecology and conservation status of many species. Here, we analyzed satellite telemetry data with a newly developed mechanistic space use model based on an Ornstein-Uhlenbeck process to help overcome the paucity of data in studying the differential habitat selection and space use of floater and territorial golden eagles Aquila chrysaetos. Our sample consisted of 49 individuals tracked over complete breeding seasons across 4 years, totaling 104 eagle breeding seasons. Modeling these data mechanistically was required to disentangle key differences in movement and particularly to separate aspects of movement driven by resource selection from those driven by use of a central place. We found that floaters generally had more expansive space use patterns and larger home ranges, as well as evidence that they partition space with territorial individuals seemingly on fine scales through differential habitat and resource selection. Floater and territorial eagle home ranges overlapped markedly, suggesting that floaters use the interstices between territories. Furthermore, floater and territorial eagles differed in how they selected for uplift variables, key components of soaring birds' energy landscape, with territorial eagles apparently better able to find and use thermal uplift. We also found relatively low individual heterogeneity in resource selection, especially among territorial individuals, suggesting a narrow realized niche for breeding individuals, which varied from the level of among-individual variation present during migration. This work furthers our understanding of floaters' potential roles in the population ecology of territorial species and suggests that conserving landscapes occupied by territorial eagles also protects floaters.

The complexity of real food webs involves uncertainty in data and in underlying ecological processes, and modeling approaches deal with these challenges differently. Generalized modeling provides a linear stability analysis without narrow specification of all processes, and conventional dynamical systems models approximate functional forms to discuss trajectories in phase space. This study compares results and ecological interpretations from both methods in four-species ecological networks at steady state. We find that a specific (dynamical systems) model only provides a subset of stability data from the generalized model, which spans many plausible dynamic scenarios, allowing for conflicting results. Nevertheless, both approaches reveal that fixed points become stable when nutrient flows to predators are fettered and even more when the basal growth rate approaches a maximum. The specific model identifies a distinct ecosystem response to bottom-up forcing, the enrichment of lower trophic levels. Enrichment stabilizes a fixed point when basal species are in a resource-deprived environment but destabilizes it if resources become more abundant. The generalized model provides less specific information since infinitely many paths of enrichment are hypothetical. Nevertheless, generalized modeling of ecological systems is a powerful technique that enables a meta analysis of these uncertain complex systems.

Longitudinal studies of marked animals provide an opportunity to assess the relative contributions of survival and reproductive output to population dynamics and change. Cassin's auklets are a long-lived seabird that maximizes annual reproductive effort in resource-rich years through a behaviour called double brooding, the initiation of a second breeding attempt following the success of the first during the same season. Our objective was to explore whether double brooding influenced population change by contributing a greater number of future recruits. We fit temporal symmetry models to 32 years of mark-recapture data of Cassin's auklets to infer the mechanisms underlying the observed variability in per capita recruitment rates. We found that periodic peaks in recruitment were explained by an increase in available nest sites, the proportion of the population double brooding 4 years prior, and spring upwelling conditions. Estimates of population change suggests a relatively stable population throughout the time series, attributable to a 'floating' demographic class of sexually mature individuals excluded from breeding by competition which quickly fill vacant sites following periods of low adult survival. Our results highlight the importance of recruitment in maintaining the population of a long-lived seabird periodically impacted by adverse environmental conditions.

Life history variation is thought to be mainly a result of energetic trade-offs among fitness components; however, detecting these trade-offs in natural populations has yielded mixed results. Individual quality and environmental variation may mask expected relationships among fitness components because some higher quality individuals may be able to acquire more resources and invest more in all functions. Thus, life history variation may be more affected by variation in individual quality than varying strategies to resolve energetic trade-offs, e.g. costs of reproduction. Here, we investigated whether variation in female quality or costs of reproduction is a larger factor in shaping differences in life history trajectories by assessing the relationship between survival and individual reproductive performance using a 32-year longitudinal data set of repeated reproductive measurements from 273 individually marked, known-aged female grey seals (Halichoerus grypus) from the Sable Island breeding colony. We defined individual reproductive performance using two traits: reproductive frequency (a female’s probability of breeding) and provisioning performance (provisions given to young measured by offspring mass), computed using mixed effects models separately for (1) all reproductive events, and (2) an age-class specific reproductive investment. Individual differences contributed a large portion of the variance in reproductive traits, with individuals displaying a range in individual reproductive frequencies from 0.45 to 0.94, and a range of average pup weaning masses from 34.9 kg to 61.8 kg across their lifetime. We used a Cormack-Jolly-Seber open-population model to estimate the effect of these reproductive performance traits on adult survival probability. Our approach estimated a positive relationship between reproductive performance and survival, where individuals that consistently invest well in their offspring survive longer. The best supported model estimated survival as a function of age-class specific provisioning performance, where late-life performance was quite variable and had the greatest impact on survival, possibly indicating individual variation in senescence. There was no evidence to support a trade-off in reproductive performance and survival at the individual level. These results suggest that in grey seals, individual quality is a stronger driver in life history variation than varying strategies to mitigate trade-offs among fitness components.

An individuals size when young may be an important source individual variation in lifetime reproductive performance, as size effects on ontogenetic development can have cascading physiological and behavioral consequences throughout life. Here, we explored how natal size influences subsequent reproductive performance in grey seals (Halichoerus grypus) using mark-recapture and repeated measures reproductive data on a marked sample of 363 females that were measured for standard length at roughly 4 weeks of age and eventually recruited to the Sable Island breeding colony. Two reproductive traits were considered: provisioning performance (mass of weaned offspring), modeled using linear mixed effects models; and reproductive frequency (the rate at which a female returns to breed), modeled using mixed-effects multistate mark-recapture models. After accounting for female age, experience, and offspring sex, we found a positive association between natal length and our measures of reproductive performance. Mothers with the longest natal lengths produced pups nearly 8 kg heavier and were 20% more likely to breed in a given year than mothers with the shortest natal lengths. Because correlation in individual body length between natal and adult life stages is weak (animals that were longer than average as pups do not always grow to be longer than average adults), the covariation between natal length and future reproductive performance reported here may act more as a carry-over effect from the size advantages afforded in the juvenile stage that allow for greater adult performance, rather than a physical trait that is maintained throughout life.

Seasonal resource pulses can have enormous impacts on species interactions. In marine ecosystems, air-breathing predators often drive their prey to deeper waters. However, it is unclear how ephemeral resource pulses such as near-surface phytoplankton blooms alter the vertical trade-off between predation avoidance and resource availability in consumers, and how these changes cascade to the diving behaviour of top predators. We integrated data on Weddell seal diving behaviour, diet stable isotopes, feeding success and mass gain to examine shifts in vertical foraging throughout ice break-out and the resulting phytoplankton bloom each year. We also tested hypotheses about the likely location of phytoplankton bloom origination (advected or producedin situwhere seals foraged) based on sea ice break-out phenology and advection rates from several locations within 150 km of the seal colony. In early summer, seals foraged at deeper depths resulting in lower feeding rates and mass gain. As sea ice extent decreased throughout the summer, seals foraged at shallower depths and benefited from more efficient energy intake. Changes in diving depth were not due to seasonal shifts in seal diets or horizontal space use and instead may reflect a change in the vertical distribution of prey. Correspondence between the timing of seal shallowing and the resource pulse was variable from year to year and could not be readily explained by our existing understanding of the ocean and ice dynamics. Phytoplankton advection occurred faster than ice break-out, and seal dive shallowing occurred substantially earlier than local break-out. While there remains much to be learned about the marine ecosystem, it appears that an increase in prey abundance and accessibility via shallower distributions during the resource pulse could synchronize life-history phenology across trophic levels in this high-latitude ecosystem.

Audio playback of vocalizations by conspecifics is commonly used to elicit calls when surveying birds of prey. Methods for call surveys vary widely in their use of silent listening periods, and usually range from 3–15 min in length. We aimed to refine this approach for detecting Great Horned Owls (Bubo virginianus) in Arctic Alaska, which is the northernmost limit of their breeding range. We used two playback protocols: protocol 1 entailed uninterrupted playback, whereas protocol 2 interspersed silent listening periods with playback during 12-min surveys. In playback surveys consisting of 166 point counts during the 2017 and 2018 breeding seasons, the probability of detecting a Great Horned Owl was 0.46 (95% CI ¼ 60.09) with protocol 1 and 0.35 (95% CI ¼ 60.12) with protocol 2 (P ¼ 0.18). The probability of detection rose with the length of the playback: of all owls detected during the 12-min surveys, 23% (95% CI ¼ 66.4%) responded within the first 3 min, 52 6 7.6% within the first 6 min, and 80 6 6.1% within 9 min. Including silent listening periods was not necessary for detecting Great Horned Owls during call surveys. We found no correlation between probability of detection and either cloud cover or wind speed (P ¼ 0.60 and P ¼ 0.28, respectively). However, we found a negative correlation between temperature and probability of detection (P ¼ 0.02). From these surveys, we calculated the density of Great Horned Owls in the Middle Fork Koyukuk Valley, Alaska (approximately 67.589°N, 149.789°W) was 4.2 6 2.6 owls/km2 during the winters of 2017 and 2018, which represents the first estimate of density at the northern breeding limit of the species.

The Arctic is changing rapidly, introducing new challenges and opportunities for animals. Rising air temperatures are driving dramatic shifts in habitat quality and availability, including increased shrub abundance, reduced pond area, and reduced springtime snow cover (Hinzman et al. 2005). In some cases, these changes have net positive effects for particular species, as exemplified by northward range expansions of moose and beavers following shrubification (Tape et al. 2016, 2018).

Bering–Chukchi–Beaufort Sea bowhead whales are born at about 4.2 m in length and 1000 kg, in lead systems along the NW Alaskan coast. Bowheads in the other northern stocks are also born in ice-covered seas. Maximum body lengths (standard measure) can reach 19 m. Regarding coloration: the skin is gray on neonates and young calves, and black on yearlings and adults. The size and shape of the chin patch is variable and does not change through life. After adulthood age, the specific areas of the skin lose pigment and turn white around the eye, the peduncle, genital groove, and base of the pectoral limb. Growth is rapid from birth to age 1, followed by a 3- to 4-year growth hiatus. Following the hiatus, the increase in body length is slow to sexual maturity at ~25 years and ~13.5 m in length. The Inupiat recognize several growth forms associated with age; these include aġvaaq (neonate), ingutuq (yearling), qairaliq (postyearling), along with several names for large adults. Allometry of the head is marked and likely associated with extreme development of the baleen rack. The head is very large in relation to body—perhaps the largest in Mammalia. The blubber is among the thickest of any cetacean (to ~38 cm). Body mass is similar to right whales of similar length; large bowheads can reach 100 metric tons or larger. Longevity can reach 200 or more years. The age of physical maturity is unclear, and adults may increase in length through life. Their ability to navigate ice-covered seas and break through ice to breathe is singular among baleen whales.

Home range dynamics and movement are central to a species’ ecology and strongly mediate both intra- and interspecific interactions. Numerous methods have been introduced to describe animal home ranges, but most lack predictive ability and cannot capture effects of dynamic environmental patterns, such as the impacts of air and water flow on movement.Here, we develop a practical, multi-stage approach for statistical inference into the behavioral mechanisms underlying how habitat and dynamic energy landscapes—in this case how airflow increases or decreases the energetic efficiency of flight—shape animal home ranges based around central places. We validated the new approach using simulations, then applied it to a sample of 12 adult golden eaglesAquila chrysaetostracked with satellite telemetry.The application to golden eagles revealed effects of habitat variables that align with predicted behavioral ecology. Further, we found that males and females partition their home ranges dynamically based on uplift. Specifically, changes in wind and sun angle drove differential space use between sexes, especially later in the breeding season when energetic demands of growing nestlings require both parents to forage more widely.This method is easily implemented using widely available programming languages and is based on a hierarchical multistate Ornstein-Uhlenbeck space use process that incorporates habitat and energy landscapes. The underlying mathematical properties of the model allow straightforward computation of predicted utilization distributions, permitting estimation of home range size and visualization of space use patterns under varying conditions.

In populations of many taxa, a large fraction of sexually mature individuals do not breed but are attempting to enter the breeding population. Such individuals, often referred to as “floaters” play critical roles in determining dynamics and stability of these populations. Floaters are difficult to study, however, so we lack data on the roles they play in population ecology and conservation status of many species. Here, we paired satellite telemetry and a new mechanistic space use model based on an Ornstein-Uhlenbeck process to study the differential habitat selection and space use of floater and territorial golden eaglesAquila chrysaetos. Our sample consisted of 49 individuals tracked over complete breeding seasons across four years, totalling 104 eagle breeding seasons. Modeling these data with the new mechanistic approach was required to parse key differences in movement and separate aspects of resource selection from central place behavior. We found that floaters generally had more expansive space use patterns and larger home ranges, partitioning space with territorial individuals seemingly on fine scales through differential habitat and resource selection. Floater and territorial eagle home ranges overlapped markedly, suggesting floaters use the interstices between territories. Further, floater and territorial eagles differed in how they selected for uplift variables, key components of soaring birds’ energy landscape, with territorial eagles apparently better able to find and use thermal uplift. We also found relatively low individual heterogeneity in resource selection, especially among territorial individuals, suggesting a narrow realized niche for breeding individuals. This work furthers our understanding of floaters’ potential roles in population ecology of territorial species, as well as suggests that conserving landscapes occupied by territorial eagles also protects floaters.

Long-distance migrations by ungulate species are a globally imperiled natural phenomenon and conservation of them requires monitoring population vital rates. Satellite telemetry tracking is widely used for understanding the spatial distribution and movement of animals, especially migratory animals in remote environments. Recently, analytical methods have been developed to infer parturition events from movement data in multiple species that calve in isolation, but to date such methods have not been tested on animals that both migrate and spatially aggregate during calving. We applied two movement-based methods developed to infer parturition in nonmigratory woodland caribou (Rangifer tarandus caribou (Gmelin, 1788)) to 241 reproductive seasons spanning 6 years of GPS data from migratory barren-ground caribou (Rangifer tarandus granti J.A. Allen, 1902). We compared results from both methods to data from aerial surveys of collared females during the calving period. We found that each movement-based method had ∼80% overall accuracy to identify calving events, with interannual variation ranging from 61% to 100%. When we considered instances when the two analytical methods agreed on parturition outcome, the accuracy increased to 89% with an annual range of 73%–100%. Using these methods, we identified marked interannual differences in peak calving dates and higher parturition rates than previously reported for this caribou herd. The successful application of these analyses to a migratory, gregarious ungulate suggests a broader applicability of the methodology.

We investigated whether inferences about harbor seal (Phoca vitulina Linnaeus, 1758) whisker shed status and growth rates based on whisker morphology could improve dietary time series from whisker isotopic profiles. We identified morphometric characteristics of Alaskan harbor seal whiskers, including a smooth root section (SRS) and a bumpy section. The SRS was longer for spring versus fall-collected whiskers and matched the length of fully grown, shed whiskers. Thus, SRS length can differentiate whisker shed and growth status, and we used it to determine the sequence of whisker shedding by cohort in summer-captured harbor seals. The mean interbump length (IBL) correlated with whisker length, potentially providing a proxy for whisker growth rate. We compared carbon isotope ratios (expressed as δ13C values) along the three longest whiskers from 10 harbor seals, using two approaches for converting whisker position to date: (1) a standard growth rate constant based on captive studies and (2) individually adjusted growth rates based on whisker morphology. Intraindividual patterns of whisker δ13C values became more synchronous when expressed by deposition date rather than by position; however, adjusting growth rates based on IBL did not improve synchrony further. These findings suggest that whisker morphology can contribute whisker growth rate and shedding information for dietary reconstruction.

Reduced prey abundance and severe weather can lead to a greater risk of mortality for seabirds during the non-breeding winter months. Resource patterns in some regions are shifting and becoming more variable in relation to past conditions, potentially further impacting survival and carryover to the breeding season. As animal tracking technologies and methods to analyze movement data have advanced, it has become increasingly feasible to draw fine-scale inference about how environmental variation affects foraging behavior and habitat use of seabirds during this critical period. Here, we used archival light-sensing tags to evaluate how interannual variation in oceanography affected the winter distribution of Cassin's Auklets from Southeast Farallon Island, California. Thirty-five out of 93 geolocators deployed from 2015 to 2017 were recovered and successfully recorded light-level data, from which geographic positions were estimated. Step-selection functions were applied to identify environmental covariates that best explained winter movement decisions and habitat use, revealing Cassin's Auklets dispersed farther from the colony during a winter with warm SST anomalies, but remained more centralized near the breeding colony during two average winters. Movement patterns were driven by avoidance of areas with higher sea surface temperatures and possible limits of dispersal from the breeding colony, and selection for areas with well-defined mesoscale fronts and cooler surface waters. Through multiple years of tagging and the application of step-selection functions, a robust and widely applied approach for analyzing animal movement in terrestrial species, we show how interannual differences in the movement patterns of a small seabird are driven by oceanographic variability across years. Understanding the winter habitat use of seabirds can help inform changes in population structure and measures of reproductive success, aiding managers in determining potential causes of breeding failures.

Individual variation in reproductive ability is a key component of natural selection within populations, driving the evolution of life histories and population responses to changing environmental conditions. Evidence that population density affects individual-level fitness in wild populations is limited, particularly for long-lived animals, which are difficult to observe on a biologically relevant scale. We tested for individual heterogeneity in reproductive performance in female grey seals (Halichoerus grypus) using 35 yr of mark-resighting data at Sable Island, Canada (43.93° N 59.91° W). We used Bayesian generalized linear mixed-effect models and multistate open robust design mark-resight models to investigate whether population size negatively influences individual reproductive performance. We measured reproductive performance in two ways: reproductive frequency (the probability of returning to the island to breed) and annual provisioning performance (the probability of successfully weaning a pup given a female bred). Sighting histories of 1,655 known-aged females with a total of 22,961 pupping events were used for analysis. After accounting for effects of female age, parity, and random year effects, we found that both provisioning performance and reproductive frequency demonstrated a strong, positive correlation with population size. Among-individual variance in reproductive traits and responses to population size indicated considerable heterogeneity in overall reproductive performance. As population size grew, "robust" females increased their reproductive performance more than their more "frail" conspecifics in both reproductive traits, resulting in an amplification of differences among individuals. Consequently, simulations from posterior distributions revealed a large fitness consequence of heterogeneity in this population, with "frail" individuals having 47.1% fewer successful pups than more "robust" females (mean reproductive output ± SD: 9.12 ± 3.77 pups for frail individuals, 16.97 ± 2.94 for robust individuals). Repeatability of overall reproductive performance across environments indicates individual quality may be more influential to lifetime reproductive success than costs associated with reproductive investment. This quantification of relative fitness and its dynamics is crucial to understanding broad evolutionary processes in natural populations.

Introduced plants can positively affect population viability by augmenting the diet of native herbivores, but can negatively affect populations if they are subpar or toxic resources. In organisms with complex life histories, such as insects specializing on host plants, the impacts of a novel host may differ across life stages, with divergent effects on population persistence. Most research on effects of novel hosts has focused on adult oviposition preference and larval performance, but adult preference may not optimize offspring performance, nor be indicative of host quality from a demographic perspective. We compared population growth rates of the Baltimore checkerspot butterfly, Euphydryas phaeton, on an introduced host, Plantago lanceolata (English plantain), and the native host Chelone glabra (white turtlehead). Contrary to the previous findings suggesting that P. lanceolata could be a population sink, we found higher population growth rates (λ) on the introduced than the native host, even though some component parameters of λ were higher on the native host. Our findings illustrate the importance of moving beyond preference-performance studies to integrate vital rates across all life stages for evaluating herbivore-host plant relationships. Single measures of preference or performance are not sufficient proxies for overall host quality nor do they provide insights into longer term consequences of novel host plant use. In our system, in particular, P. lanceolata may buffer checkerspot populations when the native host is limiting, but high growth rates could lead to crashes over longer time scales.

The home-range concept is central in animal ecology and behavior, and numerous mechanistic models have been developed to understand home range formation and maintenance. These mechanistic models usually assume a single, contiguous home range. Here we describe and implement a simple home-range model that can accommodate multiple home-range centers, form complex shapes, allow discontinuities in use patterns, and infer how external and internal variables affect movement and use patterns. The model assumes individuals associate with two or more home-range centers and move among them with some estimable probability. Movement in and around home-range centers is governed by a two-dimensional Ornstein-Uhlenbeck process, while transitions between centers are modeled as a stochastic state-switching process. We augmented this base model by introducing environmental and demographic covariates that modify transition probabilities between home-range centers and can be estimated to provide insight into the movement process. We demonstrate the model using telemetry data from sea otters (Enhydra lutris) in California. The model was fit using a Bayesian Markov Chain Monte Carlo method, which estimated transition probabilities, as well as unique Ornstein-Uhlenbeck diffusion and centralizing tendency parameters. Estimated parameters could then be used to simulate movement and space use that was virtually indistinguishable from real data. We used Deviance Information Criterion (DIC) scores to assess model fit and determined that both wind and reproductive status were predictive of transitions between home-range centers. Females were less likely to move between home-range centers on windy days, less likely to move between centers when tending pups, and much more likely to move between centers just after weaning a pup. These tendencies are predicted by theoretical movement rules but were not previously known and show that our model can extract meaningful behavioral insight from complex movement data.

Shifts in the distribution of benthivorous predators provide an indication of underlying environmental changes in benthic-mediated ecosystems. Spectacled Eiders (Somateria fischeri) are benthivorous sea ducks that spend the nonbreeding portion of their annual cycle in the Bering, Chukchi, Beaufort, and East Siberian seas. Sea ducks generally molt in biologically productive areas with abundant prey. If the distribution of eiders at molting areas matches prey abundance, spatial shifts may indicate changes in environmental conditions in the Arctic. We used a randomization procedure to test for shifts in the distribution of satellite telemetry locations received from Spectacled Eiders in the 1990s and 2008–2011 within 4 late-summer, ice-free molting areas: Indigirka–Kolyma, northern Russia; Ledyard Bay, eastern Chukchi Sea; Norton Sound, northeastern Bering Sea; and Mechigmenskiy Gulf, northwestern Bering Sea. We also tested for interannual and interdecadal changes in dive depth required to reach pre...

Group living is a widespread behavior thought to be an evolutionary adaptation for reducing predation risk. Many group-living species, however, spend a portion of their life cycle as dispersed individuals, suggesting that the costs and benefits of these opposing behaviors vary temporally. Here, we evaluated mechanistic hypotheses for explaining individual dispersion as a tactic for reducing predation risk at reproduction (i.e., birthing) in an otherwise group-living animal. Using simulation analyses parameterized by empirical data, we assessed whether dispersion increases reproductive success by (i) increasing predator search time, (ii) reducing predator encounter rates because individuals are inconspicuous relative to groups, or (iii) eliminating the risk of multiple kills per encounter. Simulations indicate that dispersion becomes favorable only when detectability increases with group size and there is risk of multiple kills per encounter. This latter effect, however, is likely the primary mechanism driving females to disperse at reproduction because group detectability effects are presumably constant year-round. We suggest that the risk of multiple kills imposed by highly vulnerable offspring may be an important factor influencing dispersive behavior in many species, and conservation strategies for such species will require protecting sufficient space to allow dispersion to effectively reduce predation risk.

The stability of ecological networks of varying topologies and predator-prey relationships is explored by applying the concept of generalized modeling. The effects of omnivory, complexity, enrichment, number of top predators, and predatory response are discussed. The degree of omnivory plays a large role in governing web stability at steady state. Complexity as measured from connectance and network size is not a perfect indicator of stability; large, highly connected webs can be just as stable as smaller, less connected ones. Learning behavior as expressed in Holling's type III predatory response is stabilizing for food webs and provides exceptions to the paradox of enrichment for some topologies.

Animals can respond to dynamic environments through phenological plasticity of life history events; however, changes in one part of the annual cycle can diminish the success of subsequent life history events. Our aims were to determine the associations between reproduction and moult phenology across years and to quantify phenological plasticity across varying environmental conditions. We conducted demographic surveys of 4,252 flipper-tagged Weddell seals (Leptonychotes weddellii) in the Ross Sea, Antarctica during four austral summers. At each sighting, seals were assigned a moult code based on the visible presence of new fur and the start date of each animal’s moult was back-calculated. Reproductive success and parturition dates were obtained for the breeding season prior to and following the moult. We found that successful reproduction delayed moult by 16 days relative to non-parturient females. Phenology of the intervening moult was indicative of previous reproductive dynamics but not predictive of subsequent reproductive outcomes. Across years, moult phenology varied by about two weeks and covaried strongly with sea ice break-out timing for all reproductive categories. Our findings suggest these polar mammals have some flexibility within the annual cycle that allows adjustment of moult phenology to fluctuating environmental conditions without compromising future reproductive success.

AstractLong distance migration can increase lifetime fitness, but can be costly, incurring increased energetic expenses and higher mortality risks. Stopover and other en route behaviors allow animals to rest and replenish energy stores and avoid or mitigate other hazards during migration. Some animals, such as soaring birds, can subsidize the energetic costs of migration by extracting energy from flowing air. However, it is unclear how these energy sources affect or interact with behavioral processes and stopover in long-distance soaring migrants. To understand these behaviors and the effects of processes that might enhance use of flight subsidies, we developed a flexible mechanistic model to predict how flight subsidies drive migrant behavior and movement processes. The novel modelling framework incorporated time-varying parameters informed by environmental covariates to characterize a continuous range of behaviors during migration. This model framework was fit to GPS satellite telemetry data collected from a large soaring and opportunist foraging bird, the golden eagle (Aquila chrysaetos), during migration in western North America. Fitted dynamic model parameters revealed a clear circadian rhythm in eagle movement and behavior, which was directly related to thermal uplift. Behavioral budgets were complex, however, with evidence for a joint migrating/foraging behavior, resembling a slower paced fly-and-forage migration, which could facilitate efficient refueling while still ensuring migration progress. In previous work, ecological and foraging conditions are normally considered to be the key aspects of stopover location quality, but taxa that can tap energy sources from moving fluids to drive migratory locomotion, such as the golden eagle, may pace migration based on both foraging opportunities and available flight subsidies.

For migrating animals, realized migration routes and timing emerge from hundreds or thousands of movement decisions made along migration routes. Local weather conditions along migration routes continually influence these decisions, and even relatively small changes in en route weather may cumulatively result in major shifts in migration patterns. Here, we analysed satellite tracking data to score a discrete navigation decision by a large migratory bird as it navigated a high-latitude, 5000 m elevation mountain range to understand how those navigational decisions changed under different weather conditions. We showed that wind conditions in particular areas along the migration pathway drove a navigational decision to reroute a migration; conditions encountered predictably resulted in migrants routing either north or south of the mountain range. With abiotic conditions continuing to change globally, simple decisions, such as the one described here, might additively emerge into new, very different migration routes.