Energetic trade-offs in migration decision-making, reproductive effort and subsequent parental care in a long-distance migratory bird.

Migratory species trade-off long-distance movement with survival and reproduction, but the spatio-temporal scales at which these decisions occur are relatively unknown. Technological and statistical advances allow fine-scale study of animal decision-making, improving our understanding of possible causes and therefore conservation management. We quantified effects of reproductive preparation during spring migration on subsequent breeding outcomes, breeding outcomes on autumn migration characteristics and autumn migration characteristics on subsequent parental survival in Greenland white-fronted geese (Anser albifrons flavirostris). These are long-distance migratory birds with an approximately 50% population decline from 1999 to 2022. We deployed GPS-acceleration devices on adult females to quantify up to 5 years of individual decision-making throughout the annual cycle. Weather and habitat-use affected time spent feeding and overall dynamic body acceleration (i.e. energy expenditure) during spring and autumn. Geese that expended less energy and fed longer during spring were more likely to successfully reproduce. Geese with offspring expended more energy and fed for less time during autumn, potentially representing adverse fitness consequences of breeding. These behavioural comparisons among Greenland white-fronted geese improve our understanding of fitness trade-offs underlying abundance. We provide a reproducible framework for full annual cycle modelling using location and behaviour data, applicable to similarly studied migratory animals.

High-resolution recording of foraging behaviour over multiple annual cycles shows decline in old Adélie penguins' performance.

Age-related variation in foraging performance can result from both within-individual change and selection processes. These mechanisms can only be disentangled by using logistically challenging long-term, longitudinal studies. Coupling a long-term demographic data set with high-temporal-resolution tracking of 18 Adélie penguins (Pygoscelis adeliae, age 4-15 yrs old) over three consecutive annual cycles, we examined how foraging behaviour changed within individuals of different age classes. Evidence indicated within-individual improvement in young and middle-age classes, but a significant decrease in foraging dive frequency within old individuals, associated with a decrease in the dive descent rate. Decreases in foraging performance occurred at a later age (from 12-15 yrs old to 15-18 yrs old) than the onset of senescence predicted for this species (9-11 yrs old). Foraging dive frequency was most affected by the interaction between breeding status and annual life-cycle periods, with frequency being highest during returning migration and breeding season and was highest overall for successful breeders during the chick-rearing period. Females performed more foraging dives per hour than males. This longitudinal, full annual cycle study allowed us to shed light on the changes in foraging performance occurring among individuals of different age classes and highlighted the complex interactions among drivers of individual foraging behaviour.

Spatial consistency in drivers of population dynamics of a declining migratory bird.

Many migratory species are in decline across their geographical ranges. Single-population studies can provide important insights into drivers at a local scale, but effective conservation requires multi-population perspectives. This is challenging because relevant data are often hard to consolidate, and state-of-the-art analytical tools are typically tailored to specific datasets. We capitalized on a recent data harmonization initiative (SPI-Birds) and linked it to a generalized modelling framework to identify the demographic and environmental drivers of large-scale population decline in migratory pied flycatchers (Ficedula hypoleuca) breeding across Britain. We implemented a generalized integrated population model (IPM) to estimate age-specific vital rates, including their dependency on environmental conditions, and total and breeding population size of pied flycatchers using long-term (34-64 years) monitoring data from seven locations representative of the British breeding range. We then quantified the relative contributions of different vital rates and population structure to changes in short- and long-term population growth rate using transient life table response experiments (LTREs). Substantial covariation in population sizes across breeding locations suggested that change was the result of large-scale drivers. This was supported by LTRE analyses, which attributed past changes in short-term population growth rates and long-term population trends primarily to variation in annual survival and dispersal dynamics, which largely act during migration and/or nonbreeding season. Contributions of variation in local reproductive parameters were small in comparison, despite sensitivity to local temperature and rainfall within the breeding period. We show that both short- and long-term population changes of British breeding pied flycatchers are likely linked to factors acting during migration and in nonbreeding areas, where future research should be prioritized. We illustrate the potential of multi-population analyses for informing management at (inter)national scales and highlight the importance of data standardization, generalized and accessible analytical tools, and reproducible workflows to achieve them.

Multi-season climate projections forecast declines in migratory monarch butterflies.

Climate change poses a unique threat to migratory species as it has the potential to alter environmental conditions at multiple points along a species' migratory route. The eastern migratory population of monarch butterflies (Danaus plexippus) has declined markedly over the last few decades, in part due to variation in breeding-season climate. Here, we combined a retrospective, annual-cycle model for the eastern monarch population with climate projections within the spring breeding grounds in eastern Texas and across the summer breeding grounds in the midwestern U.S. and southern Ontario, Canada to evaluate how monarchs are likely to respond to climate change over the next century. Our results reveal that projected changes in breeding-season climate are likely to lead to decreases in monarch abundance, with high potential for overwintering population size to fall below the historical minimum three or more times in the next two decades. Climatic changes across the expansive summer breeding grounds will also cause shifts in the distribution of monarchs, with higher projected abundances in areas that become wetter but not appreciably hotter (e.g., northern Ohio) and declines in abundance where summer temperatures are projected to increase well above those observed in the recent past (e.g., northern Minnesota). Although climate uncertainties dominate long-term population forecasts, our analyses suggest that we can improve precision of near-term forecasts by collecting targeted data to better understand relationships between breeding-season climate variables and local monarch abundance. Overall, our results highlight the importance of accounting for the impacts of climate changes throughout the full-annual cycle of migratory species.

Modelling the responses of partially migratory metapopulations to changing seasonal migration rates: From theory to data.

Among-individual and within-individual variation in expression of seasonal migration versus residence is widespread in nature and could substantially affect the dynamics of partially migratory metapopulations inhabiting seasonally and spatially structured environments. However, such variation has rarely been explicitly incorporated into metapopulation dynamic models for partially migratory systems. We, therefore, lack general frameworks that can identify how variable seasonal movements, and associated season- and location-specific vital rates, can control system persistence. We constructed a novel conceptual framework that captures full-annual-cycle dynamics and key dimensions of metapopulation structure for partially migratory species inhabiting seasonal environments. We conceptualize among-individual variation in seasonal migration as two variable vital rates: seasonal movement probability and associated movement survival probability. We conceptualize three levels of within-individual variation (i.e. plasticity), representing seasonal or annual variation in seasonal migration or lifelong fixed strategies. We formulate these concepts as a general matrix model, which is customizable for diverse life-histories and seasonal landscapes. To illustrate how variable seasonal migration can affect metapopulation growth rate, demographic structure and vital rate elasticities, we parameterize our general models for hypothetical short- and longer-lived species. Analyses illustrate that elasticities of seasonal movement probability and associated survival probability can sometimes equal or exceed those of vital rates typically understood to substantially influence metapopulation dynamics (i.e. seasonal survival probability or fecundity), that elasticities can vary non-linearly, and that metapopulation outcomes depend on the level of within-individual plasticity. We illustrate how our general framework can be applied to evaluate the consequences of variable and changing seasonal movement probability by parameterizing our models for a real partially migratory metapopulation of European shags Gulosus aristotelis assuming lifelong fixed strategies. Given observed conditions, metapopulation growth rate was most elastic to breeding season adult survival of the resident fraction in the dominant population. However, given doubled seasonal movement probability, variation in survival during movement would become the primary driver of metapopulation dynamics. Our general conceptual and matrix model frameworks, and illustrative analyses, thereby highlight complex ways in which structured variation in seasonal migration can influence dynamics of partially migratory metapopulations, and pave the way for diverse future theoretical and empirical advances.

Enhancing monitoring and transboundary collaboration for conserving migratory species under global change: The priority case of the red kite.

Calls for urgent action to conserve biodiversity under global change are increasing, and conservation of migratory species in this context poses special challenges. In the last two decades the Convention on the Conservation of Migratory Species of Wild Animals (CMS) has provided a framework for several subsidiary instruments including action plans for migratory bird species, but the effectiveness and transferability of these plans remain unclear. Such laws and policies have been credited with positive outcomes for the conservation of migratory species, but the lack of international coordination and on-ground implementation pose major challenges. While research on migratory populations has received growing attention, considerably less emphasis has been given to integrating ecological information throughout the annual cycle for examining strategies to conserve migratory species at multiple scales in the face of global change. We fill this gap through a case study examining the ecological status and conservation of a migratory raptor and facultative scavenger, the red kite (Milvus milvus), whose current breeding range is limited to Europe and is associated with agricultural landscapes and restricted to the temperate zone. Based on our review, conservation actions have been successful at recovering red kite populations within certain regions. Populations however remain depleted along the southern-most edge of the geographic range where many migratory red kites from northern strongholds overwinter. This led us to a forward-looking and integrated strategy that emphasizes international coordination involving researchers and conservation practitioners to enhance the science-policy-action interface. We identify and explore key issues for conserving the red kite under global change, including enhancing conservation actions within and outside protected areas, recovering depleted populations, accounting for climate change, and transboundary coordination in adaptive conservation and management actions. The integrated conservation strategy is sufficiently general such that it can be adapted to inform conservation of other highly mobile species subject to global change.

Strong survival selection on seasonal migration versus residence induced by extreme climatic events.

Elucidating the full eco-evolutionary consequences of climate change requires quantifying the impact of extreme climatic events (ECEs) on selective landscapes of key phenotypic traits that mediate responses to changing environments. Episodes of strong ECE-induced selection could directly alter population composition, and potentially drive micro-evolution. However, to date, few studies have quantified ECE-induced selection on key traits, meaning that immediate and longer-term eco-evolutionary implications cannot yet be considered. One widely expressed trait that allows individuals to respond to changing seasonal environments, and directly shapes spatio-seasonal population dynamics, is seasonal migration versus residence. Many populations show considerable among-individual phenotypic variation, resulting in 'partial migration'. However, variation in the magnitude of direct survival selection on migration versus residence has not been rigorously quantified, and empirical evidence of whether seasonal ECEs induce, intensify, weaken or reverse such selection is lacking. We designed full annual cycle multi-state capture-recapture models that allow estimation of seasonal survival probabilities of migrants and residents from spatio-temporally heterogeneous individual resightings. We fitted these models to 9 years of geographically extensive year-round resighting data from partially migratory European shags Phalacrocorax aristotelis. We thereby quantified seasonal and annual survival selection on migration versus residence across benign and historically extreme non-breeding season (winter) conditions, and tested whether selection differed between females and males. We show that two of four observed ECEs, defined as severe winter storms causing overall low survival, were associated with very strong seasonal survival selection against residence. These episodes dwarfed the weak selection or neutrality evident otherwise, and hence caused selection through overall annual survival. The ECE that caused highest overall mortality and strongest selection also caused sex-biased mortality, but there was little overall evidence of sex-biased selection on migration versus residence. Our results imply that seasonal ECEs and associated mortality can substantially shape the landscape of survival selection on migration versus residence. Such ECE-induced phenotypic selection will directly alter migrant and resident frequencies, and thereby alter immediate spatio-seasonal population dynamics. Given underlying additive genetic variation, such ECEs could potentially cause micro-evolutionary changes in seasonal migration, and thereby cause complex eco-evolutionary population responses to changing seasonal environments.

Comparing abundance distributions and range maps in spatial conservation planning for migratory species.

Most spatial conservation planning for wide-ranging or migratory species is constrained by poor knowledge of species' spatiotemporal dynamics and is only based on static species' ranges. However, species have substantial variation in abundance across their range and migratory species have important spatiotemporal population dynamics. With growing ecological data and advancing analytics, both of these can be estimated and incorporated into spatial conservation planning. However, there is limited information on the degree to which including this information affects conservation planning. We compared the performance of systematic conservation prioritizations for different scenarios based on varying the input species' distributions by ecological metric (abundance distributions versus range maps) and temporal sampling resolution (weekly, monthly, or quarterly). We used the example of a community of 41 species of migratory shorebirds that breed in North America, and we used eBird data to produce weekly estimates of species' abundances and ranges. Abundance distributions at a monthly or weekly resolution led to prioritizations that most efficiently protected species throughout the full annual cycle. Conversely, spatial prioritizations based on species' ranges required more sites and left most species insufficiently protected for at least part of their annual cycle. Prioritizations with only quarterly species ranges were very inefficient as they needed to target 40% of species' ranges to include 10% of populations. We highlight the high value of abundance information for spatial conservation planning, which leads to more efficient and effective spatial prioritization for conservation. Overall, we provide evidence that spatial conservation planning for wide-ranging migratory species is most robust and efficient when informed by species' abundance information from the full annual cycle.

Modeling avian full annual cycle distribution and population trends with citizen science data.

Information on species' distributions, abundances, and how they change over time is central to the study of the ecology and conservation of animal populations. This information is challenging to obtain at landscape scales across range-wide extents for two main reasons. First, landscape-scale processes that affect populations vary throughout the year and across species' ranges, requiring high-resolution, year-round data across broad, sometimes hemispheric, spatial extents. Second, while citizen science projects can collect data at these resolutions and extents, using these data requires appropriate analysis to address known sources of bias. Here, we present an analytical framework to address these challenges and generate year-round, range-wide distributional information using citizen science data. To illustrate this approach, we apply the framework to Wood Thrush (Hylocichla mustelina), a long-distance Neotropical migrant and species of conservation concern, using data from the citizen science project eBird. We estimate occurrence and abundance across a range of spatial scales throughout the annual cycle. Additionally, we generate intra-annual estimates of the range, intra-annual estimates of the associations between species and characteristics of the landscape, and interannual trends in abundance for breeding and non-breeding seasons. The range-wide population trajectories for Wood Thrush show a close correspondence between breeding and non-breeding seasons with steep declines between 2010 and 2013 followed by shallower rates of decline from 2013 to 2016. The breeding season range-wide population trajectory based on the independently collected and analyzed North American Breeding Bird Survey data also shows this pattern. The information provided here fills important knowledge gaps for Wood Thrush, especially during the less studied migration and non-breeding periods. More generally, the modeling framework presented here can be used to accurately capture landscape scale intra- and interannual distributional dynamics for broadly distributed, highly mobile species.

Survival varies seasonally in a migratory bird: Linkages between breeding and non-breeding periods.

Migratory species form an important component of biodiversity; they link ecosystems across the globe, but are increasingly threatened by global environmental change. Understanding and mitigating threats requires knowledge of how demographic processes operate throughout the annual cycle, but this can be difficult to achieve when breeding and non-breeding grounds are widely separated. Our goal is to quantify the importance of variability in survival during the breeding and non-breeding seasons in determining variation in annual survival using a single population and, more broadly, the extent to which annual survival across species reflects variation in probability of surviving the migratory period. We use a 25-year dataset in which individuals of a long-distance migratory bird, the alpine swift Tachymarptis melba, were captured towards the beginning and end of each breeding season to estimate age- and season-specific survival probabilities and incorporate explicit estimation of the correlations in survival between age-classes and seasons. Monthly survival was higher during the breeding period than during the rest of the year and strongly affected by conditions in the breeding season; effects that remained apparent in the following non-breeding season, but not subsequently. Recruitment of juveniles was dependent on the timing of breeding, being higher if egg-laying commenced before the median date, and substantially lower if not. Across migratory bird species, variation in annual survival largely reflects variation in the probability of surviving the migratory period. Using a double-capture approach, even within a single season, provides valuable insights into the demography of migratory species, which will help understand the extent and impacts of the threats they face in a changing world.

Increased summer food supply decreases non-breeding movement in black-legged kittiwakes.

Individual condition at one stage of the annual cycle is expected to influence behaviour during subsequent stages, yet experimental evidence of food-mediated carry-over effects is scarce. We used a food supplementation experiment to test the effects of food supply during the breeding season on migration phenology and non-breeding behaviour. We provided an unlimited supply of fish to black-legged kittiwakes (Rissa tridactyla) during their breeding season on Middleton Island, Alaska, monitored reproductive phenology and breeding success, and used light-level geolocation to observe non-breeding behaviour. Among successful breeders, fed kittiwakes departed the colony earlier than unfed controls. Fed kittiwakes travelled less than controls during the breeding season, contracting their non-breeding range. Our results demonstrate that food supply during the breeding season affects non-breeding phenology, movement and distribution, providing a potential behavioural mechanism underlying observed survival costs of reproduction.

Spatial and temporal drivers of avian population dynamics across the annual cycle.

Untangling the spatial and temporal processes that influence population dynamics of migratory species is challenging, because changes in abundance are shaped by variation in vital rates across heterogeneous habitats and throughout the annual cycle. We developed a full-annual-cycle, integrated, population model and used demographic data collected between 2011 and 2014 in southern Indiana and Belize to estimate stage-specific vital rates of a declining migratory songbird, the Wood Thrush (Hylocichla mustelina). Our primary objective was to understand how spatial and temporal variation in demography contributes to local and regional population growth. Our full-annual-cycle model allowed us to estimate (1) age-specific, seasonal survival probabilities, including latent survival during both spring and autumn migration, and (2) how the relative contribution of vital rates to population growth differed among habitats. Wood Thrushes in our study populations experienced the lowest apparent survival rates during migration and apparent survival was lower during spring migration than during fall migration. Both mortality and high dispersal likely contributed to low apparent survival during spring migration. Population growth in high-quality habitat was most sensitive to variation in fecundity and apparent survival of juveniles during spring migration, whereas population growth in low-quality sites was most sensitive to adult apparent breeding-season survival. These results elucidate how full-annual-cycle vital rates, particularly apparent survival during migration, interact with spatial variation in habitat quality to influence population dynamics in migratory species.

Seasonal patterns and protection status of stopover hotspots for migratory landbirds in the eastern United States.

Migratory landbirds in North America are experiencing dramatic population declines. Although considerable research and conservation attention have been directed toward these birds' breeding and wintering grounds, far less is known about the areas used as stopover sites during migration. To address this knowledge gap, we used 5 years of weather surveillance radar data to map seasonal stopover densities of landbirds across the eastern United States during spring and autumn migration. We identified stopover hotspots covering 2.47 million ha that consistently support high densities of migratory landbirds in spring or autumn. However, only 16.7% of these sites are hotspots in both seasons. The distribution of hotspots is shifted eastward in autumn compared with spring. Deciduous forest is the most important habitat type in both seasons, with deciduous forest fragments embedded in broadly deforested regions having the highest probability of being hotspots. The concentration of birds in these forest fragments is stronger in spring, especially in the agricultural Midwest. We found generally higher stopover densities in protected areas than in unprotected areas in both seasons. Nonetheless, only one-third of identified stopover hotspots have some sort of protected status, and more than half of these protected hotspots are subject to extractive uses. A well-distributed network of well-protected stopover areas, complementing conservation efforts on the breeding and wintering grounds, is essential to sustaining healthy populations of migratory landbirds in North America.

Quantifying phenological landmarks of migration shows nonuniform use of the Caribbean by shorebirds.

Of the boreal- and Arctic-breeding North American shorebirds that migrate south through the Caribbean, most individuals continue farther south. However, for many species, some individuals remain beyond the southbound migration period (i.e., throughout the temperate winter and/or summer). This variation among individuals adds complexity to observation data, obscures migration patterns, and could prevent the examination of the use of different Caribbean regions by various shorebird species during migration and in the nonmigratory seasons. Here, we present a novel method that leverages a well-established statistical approach (generalized additive models) to systematically identify migration phenology even for complex passage migrant species with individuals that remain beyond migration. Our method identifies the active migration period using derivatives of a fitted GAM and then calculates phenology metrics based on quantiles of that migration period. We also developed indices to quantify oversummering and overwintering patterns with respect to migration. We analyzed eBird data for 16 North American shorebird species as they traveled South through the insular Caribbean, identifying separate migratory patterns for Cuba, Puerto Rico, Guadeloupe, Aruba, Bonaire, Curaçao, and Trinidad and Tobago. Our results confirm past reports and provide additional detail on shorebird migration in the Caribbean, and identify several previously unpublished regional patterns. Despite Puerto Rico being farther north and closer to continental North America, most species reached Puerto Rico later than other regions, supporting a long-standing hypothesis that migration strategy (transcontinental vs. transoceanic) leads to geographic differences in migration timing. We also found distinct patterns of migration curves, with some regions and species consistently having either symmetrical or skewed curves; these differences in migration curve shape reflect different migration processes. Our novel method proved reliable and adaptable for most species and serves as a valuable tool for identifying phenological patterns in complex migration data, potentially unlocking previously intractable data.

Influence of prey availability on habitat selection during the non-breeding period in a resident bird of prey.

BACKGROUND: For resident birds of prey in the temperate zone, the cold non-breeding period can have strong impacts on survival and reproduction with implications for population dynamics. Therefore, the non-breeding period should receive the same attention as other parts of the annual life cycle. Birds of prey in intensively managed agricultural areas are repeatedly confronted with unpredictable, rapid changes in their habitat due to agricultural practices such as mowing, harvesting, and ploughing. Such a dynamic landscape likely affects prey distribution and availability and may even result in changes in habitat selection of the predator throughout the annual cycle. METHODS: In the present study, we (1) quantified barn owl prey availability in different habitats across the annual cycle, (2) quantified the size and location of barn owl breeding and non-breeding home ranges using GPS-data, (3) assessed habitat selection in relation to prey availability during the non-breeding period, and (4) discussed differences in habitat selection during the non-breeding period to habitat selection during the breeding period. RESULTS: The patchier prey distribution during the non-breeding period compared to the breeding period led to habitat selection towards grassland during the non-breeding period. The size of barn owl home ranges during breeding and non-breeding were similar, but there was a small shift in home range location which was more pronounced in females than males. The changes in prey availability led to a mainly grassland-oriented habitat selection during the non-breeding period. Further, our results showed the importance of biodiversity promotion areas and undisturbed field margins within the intensively managed agricultural landscape. CONCLUSIONS: We showed that different prey availability in habitat categories can lead to changes in habitat preference between the breeding and the non-breeding period. Given these results we show how important it is to maintain and enhance structural diversity in intensive agricultural landscapes, to effectively protect birds of prey specialised on small mammals.

Migration ecology of western gray catbirds.

BACKGROUND: For many songbirds in North America, we lack movement details about the full annual cycle, notably outside the breeding season. Understanding how populations are linked spatially between breeding and overwintering periods (migratory connectivity) is crucial to songbird conservation and management. We assessed migratory connectivity for 2 breeding populations of Gray Catbirds (Dumetella carolinensis) west of and within the Rocky Mountains by determining migration routes, stopover sites, and overwintering locations. Additionally, we compared apparent annual survivorship for both populations. METHODS: We deployed 39 archival light-level geolocators and 21 Global Positioning System (GPS) tags on catbirds in the South Okanagan Valley, British Columbia, Canada, and 32 geolocators and 52 GPS tags in the Bitterroot River Valley, Montana, USA. These devices allowed us to determine migration routes, stopover sites, overwintering locations, and migratory connectivity. Migratory connectivity was quantified using Mantel's correlation. We used mark-recapture of colour banded catbirds in both sites to estimate apparent annual survivorship. RESULTS: We retrieved 6 geolocators and 19 GPS tags with usable data. Gray Catbirds from both populations passed through the Rocky Mountains eastward before heading south towards their overwintering locations in northeastern Mexico and Texas. Stopover sites during fall migration occurred primarily in Montana, Kansas, Oklahoma, and Arkansas. Overwintering locations spanned Texas and 5 states in northeastern Mexico. Individual catbirds used up to 4 distinct sites during the overwintering period. Catbirds separated by almost 500 km during the breeding season overlapped during the non-breeding season, suggesting weak migratory connectivity among western populations (Mantel's correlation = 0.013, P-value = 0.41). Catbird apparent annual survivorship estimates were higher in British Columbia (0.61 ± 0.06 females; 0.64 ± 0.05 males) than in Montana (0.34 ± 0.05 females; 0.43 ± 0.04 males), though the main driver of these differences remain unclear. CONCLUSIONS: Our results provide high precision geographic details during the breeding, migration, and overwintering phases of the annual cycle for western Gray Catbirds. Notably, we found that western catbirds followed the Central Flyway as opposed to the Pacific Flyway. We document that catbirds used multiple sites over winter, contrary to the popular belief that this phase of the annual cycle is stationary for most songbirds.

Demographic responses to climate change in a threatened Arctic species.

The Arctic is undergoing rapid and accelerating change in response to global warming, altering biodiversity patterns, and ecosystem function across the region. For Arctic endemic species, our understanding of the consequences of such change remains limited. Spectacled eiders (Somateria fischeri), a large Arctic sea duck, use remote regions in the Bering Sea, Arctic Russia, and Alaska throughout the annual cycle making it difficult to conduct comprehensive surveys or demographic studies. Listed as Threatened under the U.S. Endangered Species Act, understanding the species response to climate change is critical for effective conservation policy and planning. Here, we developed an integrated population model to describe spectacled eider population dynamics using capture-mark-recapture, breeding population survey, nest survey, and environmental data collected between 1992 and 2014. Our intent was to estimate abundance, population growth, and demographic rates, and quantify how changes in the environment influenced population dynamics. Abundance of spectacled eiders breeding in western Alaska has increased since listing in 1993 and responded more strongly to annual variation in first-year survival than adult survival or productivity. We found both adult survival and nest success were highest in years following intermediate sea ice conditions during the wintering period, and both demographic rates declined when sea ice conditions were above or below average. In recent years, sea ice extent has reached new record lows and has remained below average throughout the winter for multiple years in a row. Sea ice persistence is expected to further decline in the Bering Sea. Our results indicate spectacled eiders may be vulnerable to climate change and the increasingly variable sea ice conditions throughout their wintering range with potentially deleterious effects on population dynamics. Importantly, we identified that different demographic rates responded similarly to changes in sea ice conditions, emphasizing the need for integrated analyses to understand population dynamics.

Seasonal variation in the effects of artificial light at night on the occurrence of nocturnally migrating birds in urban areas.

Urban areas often contain large numbers of migratory bird species during seasonal migration, many of which are nocturnal migrants. How artificial light at night (ALAN) and urban landcover are associated with the diurnal occurrence of nocturnal migrants within urban areas across seasons has not been explored. Here, we use eBird bird occurrence information to estimate the seasonal species richness of nocturnally migrating passerines (NMP) within 333 well surveyed urban areas within the contiguous USA. We model the relationship between seasonal NMP species richness and ALAN, proportion of tree canopy cover, and proportion of impervious surface. NMP species richness reached its highest levels during spring and autumn migration and lowest during the winter and summer. Greater tree canopy cover was associated with higher NMP species richness during spring and autumn migration and the summer. A 10% increase in the proportion of tree canopy cover was associated with a 2.0% increase in NMP species richness during spring migration, a 1.8% increase during autumn migration, and a 0.9% increase during the summer. More impervious surface was associated with higher NMP species richness during the winter. A 10% increase in the proportion of impervious surface was associated with a 6.1-9.8% increase in NMP species richness. Higher ALAN was associated with lower NMP species richness during the winter and summer, and higher NMP species richness during spring and autumn migration. A 50% increase in ALAN was associated with a 3.0-3.6% decrease in NMP species richness during the winter, a 1.7% increase during spring migration, a 2.1% decrease during the summer, and a 5.0% increase during autumn migration. These findings highlight the variable effects of ALAN and urban landcover on the seasonal occurrence of NMP species in urban areas, the value of tree canopy cover during migration and the breeding season, and the importance of reducing ALAN during migration.

Migration distance and maternal resource allocation determine timing of birth in a large herbivore.

Birth timing is a key life-history characteristic that influences fitness and population performance. For migratory animals, however, appropriately timing birth on one seasonal range may be constrained by events occurring during other parts of the migratory cycle. We investigated how the use of capital and income resources may facilitate flexibility in reproductive phenology of migratory mule deer in western Wyoming, USA, over a 5-yr period (2015-2019). Specifically, we examined how seasonal interactions affected three interrelated life-history characteristics: fetal development, birth mass, and birth timing. Females in good nutritional condition at the onset of winter and those that migrated short distances had more developed fetuses (measured as fetal eye diameter in March). Variation in parturition date was explained largely by fetal development; however, there were up to 16 d of plasticity in expected birth date. Plasticity in expected birth date was shaped by income resources in the form of exposure to spring green-up. Although individuals that experienced greater exposure to spring green-up were able to advance expected birth date, being born early or late with respect to fetal development had no effect on birth mass of offspring. Furthermore, we investigated the trade-offs migrating mule deer face by evaluating support for existing theory that predicts that births should be matched to local peaks in resource availability at the birth site. In contrast to this prediction, only long-distance migrants that paced migration with the flush of spring green-up, giving birth shortly after ending migration, were able to match birth with spring green-up. Shorter-distance migrants completed migration sooner and gave birth earlier, seemingly trading off more time for offspring to grow and develop over greater access to resources. Thus, movement tactic had profound downstream effects on birth timing. These findings highlight a need to reconsider classical theory on optimal birth timing, which has focused solely on conditions at the birth site.

Full-annual demography and seasonal cycles in a resident vertebrate.

Wildlife demography is typically studied at a single point in time within a year when species, often during the reproductive season, are more active and therefore easier to find. However, this provides only a low-resolution glimpse into demographic temporal patterns over time and may hamper a more complete understanding of the population dynamics of a species over the full annual cycle. The full annual cycle is often influenced by environmental seasonality, which induces a cyclic behavior in many species. However, cycles have rarely been explicitly included in models for demographic parameters, and most information on full annual cycle demography is restricted to migratory species. Here we used a high-resolution capture-recapture study of a resident tropical lizard to assess the full intra-annual demography and within-year periodicity in survival, temporary emigration and recapture probabilities. We found important variation over the annual cycle and up to 92% of the total monthly variation explained by cycles. Fine-scale demographic studies and assessments on the importance of cycles within parameters may be a powerful way to achieve a better understanding of population persistence over time.