Mammal communities of primeval forests as sentinels of global change.

Understanding the drivers and consequences of global environmental change is crucial to inform predictions of effects on ecosystems. We used the mammal community of Bialowieza Forest, the last lowland near-primeval forest in temperate Europe, as a sentinel of global change. We analyzed changes in stable carbon (δ13 C) and nitrogen (δ15 N) isotope values of hair in 687 specimens from 50 mammal species across seven decades (1946-2011). We classified mammals into four taxonomic-dietary groups (herbivores, carnivores, insectivores, and bats). We found a significant negative trend in hair δ15 N for the mammal community, particularly strong for herbivores. This trend is consistent with temporal patterns in nitrogen deposition from (15 N depleted) industrial fertilizers and fossil fuel emissions. It is also in line with global-scale declines in δ15 N reported in forests and other unfertilized, non-urban terrestrial ecosystems and with local decreases in N foliar concentrations. The global depletion of 13 C content in atmospheric CO2 due to fossil fuel burning (Suess effect) was detected in all groups. After correcting for this effect, the hair δ13 C trend became non-significant for both community and groups, except for bats, which showed a strong decline in δ13 C. This could be related to an increase in the relative abundance of freshwater insects taken by bats or increased use of methane-derived carbon in food webs used by bats. This work is the first broad-scale and long-term mammal isotope ecology study in a near-primeval forest in temperate Europe. Mammal communities from natural forests represent a unique benchmark in global change research; investigating their isotopic temporal variation can help identify patterns and early detections of ecosystem changes and provide more comprehensive and integrative assessments than single species approaches.

Experimental evidence that EPA and DHA are dietary requirements in a migratory shorebird, but they do not affect muscle oxidative capacity.

Dietary n-3 long chain polyunsaturated fatty acids (LCPUFAs) are hypothesized to be natural doping agents in migratory shorebirds, enabling prolonged flight by increasing membrane fluidity and oxidative capacity of the flight muscles. Animals can obtain n-3 LCPUFAs from the diet or by conversion of dietary α-linolenic acid, 18:3 n-3. However, the capacity to meet n-3 LCPUFA requirements from 18:3 n-3 varies among species. Direct tests of muscle oxidative enhancement and fatty acid conversion capacity are lacking in marine shorebirds that evolved eating diets rich in n-3 LCPUFAs. We tested whether the presence and type of dietary fatty acids influence the fatty acid composition and flight muscle oxidative capacity in western sandpipers (Calidris mauri). Sandpipers were fed diets low in n-3 PUFAs, high in 18:3 n-3, or high in n-3 LCPUFAs. Dietary fatty acid composition was reflected in multiple tissues, and low intake of n-3 LCPUFAs decreased the abundance of these fatty acids in all tissues, even with a high intake of 18:3 n-3. This suggests that 18:3 n-3 cannot replace n-3 LCPUFAs, and dietary n-3 LCPUFAs are required for sandpipers. Flight muscle indicators of enzymatic oxidative capacity and regulators of lipid metabolism did not change. However, the n-3 LCPUFA diet was associated with increased FAT/CD36 mRNA expression, potentially benefitting fatty acid transport during flight. Our study suggests that flight muscle lipid oxidation is not strongly influenced by n-3 PUFA intake. The type of dietary n-3 PUFA strongly influences the abundance of n-3 LCPUFAs in the body and could still impact whole-animal performance.

Winter-ground microhabitat use by differently coloured phenotypes affects return rate in a long-distance migratory bird.

Migratory bird populations are declining globally at alarming rates. Non-breeding site conditions affect breeding populations, but generalising non-breeding habitat conditions over large spatial regions cannot address potential fine-scale differences across landscapes or local populations. Plumage characteristics can mediate the effects of environmental conditions on individual fitness. However, whether different phenotypes use distinctive non-breeding sites, and whether they respond to non-breeding site conditions differently remains largely unknown. Stable isotopes (δ13C, δ15N, δ2H) of inert tissues are useful to infer habitat characteristics and geographic origins where those tissues were grown. We collected winter-grown feathers from pied flycatchers (Ficedula hypoleuca) on their breeding grounds over several years from males whose dorsal plumage colouration ranged continuously from brown to black and assessed their stable isotope values as proxies of local habitat conditions. Based on feather δ2H profiles we found that browner males spent their non-breeding season in drier habitats than black males. Assignment to origin analysis shows potential regional non-breeding ground separation between differently coloured males. High within-individual repeatability of both δ13C and δ15N indicate the pied flycatcher males return yearly to similar areas. Blacker males were more likely to return to the breeding grounds after dry years compared with brown males. The opposite was found in wet years. Our study demonstrates that different phenotypes are exposed to different non-breeding site conditions which can differentially affect individual survivorship. This has important ramifications for population dynamics under predicted climate change scenarios where especially brown phenotype pied flycatcher males may be under a risk of decreasing.

Stable isotopes and a changing world.

The measurement of naturally occurring stable isotope ratios of the light elements (C, N, H, O, S) in animal tissues and associated organic and inorganic fractions of associated environments holds immense potential as a means of addressing effects of global change on animals. This paper provides a brief review of studies that have used the isotope approach to evaluate changes in diet, isotopic niche, contaminant burden, reproductive and nutritional investment, invasive species and shifts in migration origin or destination with clear links to evaluating effects of global change. This field has now reached a level of maturity that is impressive but generally underappreciated and involves technical as well as statistical advances and access to freely available R-based packages. There is a need for animal ecologists and conservationists to design tissue collection networks that will best answer current and anticipated questions related to the global change and the biodiversity crisis. These developments will move the field of stable isotope ecology toward a more hypothesis driven discipline related to rapidly changing global events.

Anthropogenic and Climatic Drivers of Long-Term Changes of Mercury and Feeding Ecology in Arctic Beluga (Delphinapterus leucas) Populations.

We assessed long-term changes in the feeding ecology and mercury (Hg) accumulation in Eastern High Arctic-Baffin Bay beluga (Delphinapterus leucas) using total Hg and stable isotope (δ13C, δ15N) assays in teeth samples from historical (1854-1905) and modern (1985-2000) populations. Mean δ13C values in teeth declined significantly over time, from -13.01 ± 0.55‰ historically to -14.41 ± 0.28‰ in 2000, while no consistent pattern was evident for δ15N due to high individual variability within each period. The temporal shift in isotopic niche is consistent with beluga feeding ecology changing in recent decades to a more pelagic and less isotopically diverse diet or an ecosystem wide change in isotope profiles. Mercury concentrations in modern beluga teeth were 3-5 times higher on average than those in historical beluga. These results are similar to the long-term trends of Hg and feeding ecology reported in other beluga populations and in other Arctic marine predators. Similar feeding ecology shifts across regions and species indicate a consistent increased pelagic diet response to climate change as the Arctic Ocean progressively warmed and lost sea ice. Previously, significant temporal Hg increase in beluga and other Arctic animals was attributed solely to direct inputs of anthropogenic Hg from long-range sources. Recent advances in understanding the Arctic marine Hg cycle suggest an additional, complementary possibility─increased inputs of terrestrial Hg of mixed anthropogenic-natural origin, mobilized from permafrost and other Arctic soils by climate warming. At present, it is not possible to assign relative importance to the two processes in explaining the rise of Hg concentrations in modern Arctic marine predators.

Stable Mercury Trends Support a Long-Term Diet Shift Away from Marine Foraging in Salish Sea Glaucous-Winged Gulls over the Last Century.

Marine predators are monitored as indicators of pollution, but such trends can be complicated by variation in diet. Glaucous-winged gulls (Larus glaucescens) have experienced a dietary shift over the past century, from mainly marine to including more terrestrial/freshwater inputs, with unknown impacts on mercury (Hg) trends. We examined 109-year trends in total mercury (THg) and methylmercury (MeHg) concentrations in glaucous-winged gull feathers (1887-1996) from the Salish Sea. Adult flank feathers had higher MeHg concentrations than immature feathers, and males head feathers had higher THg concentrations than females. Overall, we found no evidence of a trend in feather MeHg or THg concentrations over time from 1887 to 1996. In the same individuals, δ15N, δ13C, and δ34S declined over time in gull feathers. In comparison, egg THg concentrations declined from 1970 to 2019 in two species of cormorants, likely reflecting decreases in local Hg sources. We conclude that diet shifts through time may have countered increased Hg deposition from long-range transport in glaucous-winged gulls. The lack of Hg trends over time in glaucous-winged gull feathers provides additional support that these gulls have decreased the amount of marine forage fish in their diet.

Using stable isotope (δ13C, δ15N) values from feces and breath to infer shorebird diets.

The use of stable isotopes of carbon (δ13C) and nitrogen (δ15N) from feces and breath offers potential as non-destructive tools to assess diets and nutrition. How stable isotope values derived from breath and feces compare with those from commonly used tissues, such as blood fractions and liver, remains uncertain, including understanding the metabolic routing of dietary nutrients. Here, we measured δ13C and δ15N from feces and δ13C of breath from captive Red-necked Stints (Calidris ruficollis) and 26 species of wild-caught migratory shorebirds (n = 259 individuals) and compared them against isotopic values from blood and feathers. For captive birds fed either cereal- or fish-based diets, differences in δ13C between feces and lipid-free diet were small, - 0.2 ± 0.5‰ and 0.1 ± 0.3‰, respectively, and differences in δ15N, - 0.7 ± 0.5‰ and - 0.5 ± 0.5‰, respectively. Hence, δ13C and δ15N values from feces can serve as proxies for ingested proteinaceous tissues and non-soluble carbohydrates because isotopic discrimination can be considered negligible. Stable isotope values in plasma and feces were strongly correlated in wild-caught shorebirds, indicating feces can be used to infer assimilated macronutrients. Breath δ13C was 1.6 ± 0.8‰ to 5.6 ± 1.2‰ lower than bulk food sources, and breath C derived from lipids was estimated at 47.5% (cereal) to 96.1% (fish), likely underlining the importance of dietary lipids for metabolism. The findings validate the use of stable isotope values of feces and breath in isotopic assays to better understand the dietary needs of shorebirds.

Breeding stage and tissue isotopic consistency suggests colony-level flexibility in niche breadth of an Arctic marine bird.

Organisms must overcome environmental limitations to optimize their investment in life history stages to maximize fitness. Human-induced climate change is generating increasingly variable environmental conditions, impacting the demography of prey items and, therefore, the ability of consumers to successfully access resources to fuel reproduction. While climate change effects are especially pronounced in the Arctic, it is unknown whether organisms can adjust foraging decisions to match such changes. We used a 9-year blood plasma δ13C and δ15N data set from over 700 pre-breeding Arctic common eiders (Somateria mollissima) to assess breeding-stage and inter-annual variation in isotopic niche, and whether inferred trophic flexibility was related to colony-level breeding parameters and environmental variation. Eider blood isotope values varied both across years and breeding stages, and combined with only weak relationships between isotopic metrics and environmental conditions suggests that pre-breeding eiders can make flexible foraging decisions to overcome constraints imposed by local abiotic conditions. From an investment perspective, an inshore, smaller isotopic niche predicted a greater probability to invest in reproduction, but was not related to laying phenology. Proximately, our results provide evidence that eiders breeding in the Arctic can alter their diet at the onset of reproductive investment to overcome increases in the energetic demand of egg production. Ultimately, Arctic pre-breeding common eiders may have the stage- and year-related foraging flexibility to respond to abiotic variation to reproduce successfully.

Migratory connectivity then and now: a northward shift in breeding origins of a long-distance migratory bird wintering in the tropics.

Temporal variation in the connectivity of populations of migratory animals has not been widely documented, despite having important repercussions for population ecology and conservation. Because the long-distance movements of migratory animals link ecologically distinct and geographically distant areas of the world, changes in the abundance and migratory patterns of species may reflect differential drivers of demographic trends acting over various spatial scales. Using stable hydrogen isotope analyses (δ2H) of feathers from historical museum specimens and contemporary samples obtained in the field, we provide evidence for an approximately 600 km northward shift over 45 years in the breeding origin of a species of songbird of major conservation concern (blackpoll warbler, Setophaga striata) wintering in the foothills of the eastern Andes of Colombia. Our finding mirrors predictions of range shifts for boreal-breeding species under warming climate scenarios and habitat loss in the temperate zone, and underscores likely drivers of widespread declines in populations of migratory birds. Our work also highlights the value of natural history collections to document the effects of global change on biodiversity.

Quantifying capital versus income breeding: New promise with stable isotope measurements of individual amino acids.

Capital breeders accumulate nutrients prior to egg development, then use these stores to support offspring development. In contrast, income breeders rely on local nutrients consumed contemporaneously with offspring development. Understanding such nutrient allocations is critical to assessing life-history strategies and habitat use. Despite the contrast between these strategies, it remains challenging to trace nutrients from endogenous stores or exogenous food intake into offspring. Here, we tested a new solution to this problem. Using tissue samples collected opportunistically from wild emperor penguins Aptenodytes forsteri, which exemplify capital breeding, we hypothesized that the stable carbon (δ13 C) and nitrogen (δ15 N) isotope values of individual amino acids (AAs) in endogenous stores (e.g. muscle) and in egg yolk and albumen reflect the nutrient sourcing that distinguishes capital versus income breeding. Unlike other methods, this approach does not require untested assumptions or diet sampling. We found that over half of essential AAs had δ13 C values that did not differ between muscle and yolk or albumen, suggesting that most of these AAs were directly routed from muscle into eggs. In contrast, almost all non-essential AAs differed in δ13 C values between muscle and yolk or between muscle and albumen, suggesting de novo synthesis. Over half of AAs that have labile nitrogen atoms (i.e. 'trophic' AA) had higher δ15 N values in yolk and albumen than in muscle, suggesting that they were transaminated during their routing into egg tissue. This effect was smaller for AAs with less labile nitrogen atoms (i.e. 'source' AA). Our results indicate that the δ15 N offset between trophic-source AAs (Δ15 Ntrophic-source ) may provide an index of the extent of capital breeding. The value of emperor penguin Δ15 NPro-Phe was higher in yolk and albumen than in muscle, reflecting the mobilization of endogenous stores; in comparison, the value of Δ15 NPro-Phe was similar across muscle and egg tissue in previously published data for income-breeding herring gulls Larus argentatus smithsonianus. Our results provide a quantitative basis for using AA δ13 C and δ15 N, and isotopic offsets among AAs (e.g. Δ15 NPro-Phe ), to explore the allocation of endogenous versus exogenous nutrients across the capital versus income spectrum of avian reproduction.

Tracing sources of carbon and hydrogen to stored lipids in migratory passerines using stable isotope (δ13C, δ2H) measurements.

Using measurements of naturally occurring stable isotopes in feathers to determine avian origin and migratory patterns is well established. However, isotopically determining nutritional origins of lipids, a major migratory fuel, has not been attempted. This study explores isotopic links between diet and stored lipids in captive white-throated sparrows (Zonotrichia albicollis) by providing isotopically distinct mixtures of carbohydrates/oils and drinking water and assessing the δ13C and δ2H values of stored lipid, breath CO2 (δ13C) and breath water vapour (δ2H). Stored lipid δ13C and δ2H values correlated with the isotopic values found in dietary carbohydrates/oils and drinking water treatments, respectively, indicating a clear traceable transfer of environmental dietary isotopic signals into body lipids. Dietary oils and carbohydrates contributed 80-82% of carbon and 44-46% of hydrogen, respectively, to stored lipids. Drinking water contributed 18-28% of hydrogen to stored lipids. Isotopic relationships were quantifiable using linear calibration algorithms which provide the basis for the construction of tissue isoscapes for migratory passerines. Breath CO2 δ13C values and breath water vapour δ2H values for fed and fasted birds reflected dietary sources. Breath CO2 δ13C values were higher for fasted birds than for fed birds by an average of 4.5‰ while breath water vapour δ2H values were lower for fasted birds by an average of 48.9‰. These results indicate that lipids and metabolites from their subsequent breakdown for fuel isotopically reflect dietary sources but complicate interpretation of such data, especially for wild migrating birds. Applications and limitations of these findings to the creation of "liposcapes" are examined.

Earlier and slower or later and faster: Spring migration pace linked to departure time in a Neotropical migrant songbird.

Migratory birds travel vast distances and the timing of migratory flights can affect survival and the ability to reproduce. For Neotropical migrant songbirds, early spring departure from wintering sites, early arrival to the breeding grounds and higher reproductive success have been related to the use of suitable habitats and environmental conditions during the non-breeding season. However, how migratory strategies are shaped by winter habitat choice is largely unknown due to the general inability to track birds from specific wintering habitats to stopovers or breeding destinations. We assessed how winter habitat (native forest vs. shade-grown coffee plantations) relates to spring departure date and migration pace in Swainson's Thrush Catharus ustulatus. We also determined the effect of departure date and total migration duration on the arrival date of birds detected near or within their breeding range. We used a novel application of Motus radiotelemetry arrays to track individuals from their wintering grounds in the Andes of South America along their migratory journey to North America. We found variation in migratory strategies between habitats, with birds wintering in native forest departing later than birds in coffee. We present isotopic evidence for native forest being of higher quality than shade-coffee for Swainson's Thrush and hypothesize that moister conditions in forest, as shown by stable isotope (δ13 C) analysis of thrush whole blood, provides favourable pre-migratory conditions allowing birds to delay departure from wintering grounds. Habitat, between-site and -year variation in departure date, suggests that birds made facultative adjustments to winter habitat quality and environmental conditions. Independent of habitat, birds that departed later migrated faster and this pattern was maintained along the migration route (n = 44). Migrating earlier and slower or later and faster was unlikely to result in significant differences in arrival time to breeding destinations. Our findings reveal underappreciated complexity in migratory decisions by long-distance migrants that contrast with the current paradigm of earlier departures and arrival from optimal habitats. The next step is to understand the relative fitness benefits of early versus late schedules or whether each strategy is an equally good response to experienced conditions.

Las aves migratorias viajan largas distancias y la fecha de sus vuelos migratorios puede afectar su supervivencia y habilidad para reproducirse. Para las aves migratorias Neotropicales canoras, salir más temprano de los sitios de invernada, llegar más temprano a los sitios de reproducción y tener mayor éxito reproductivo, está relacionado con el uso de hábitats y condiciones medioambientales adecuadas durante la época no reproductiva. Sin embargo, la forma en que las estrategias migratorias son moldeadas por la selección del hábitat invernal es en gran parte desconocida debido a la inhabilidad general de rastrear las aves desde hábitats de invernada específicos hasta los sitios de descanso o de reproducción. Determinamos la relación entre el hábitat de invierno (bosque nativo vs. plantaciones de café con sombra), la fecha de salida durante la migración de primavera y el número de días usados durante la migración por la mirla buchipecosa Catharus ustulatus. También determinamos el efecto de la fecha de salida y la duración total de la migración en la fecha de llegada de las aves detectadas cerca o dentro de su rango de reproducción. Usamos la novedosa red de radio telemetría Motus para rastrear individuos desde sus sitios de invernada en las montañas de los Andes en Sudamérica a lo largo de su viaje migratorio hasta Norte América. Encontramos variación en las estrategias migratorias entre hábitats, las aves invernando en bosque nativo salieron más tarde que las aves invernando en cafetales. Presentamos evidencia isotópica, que sugiere que el bosque nativo es de mejor calidad que las plantaciones de café con sombra. Hipotetizamos que condiciones más húmedas en el bosque, indicadas por análisis de isotopos estables (δ13 C) de la sangre de las mirlas, proveen condiciones favorables antes de la migración permitiendo a las aves retrasar su salida de los territorios de invernada. La variación en la fecha de salida entre hábitats, años y sitios, sugiere que las aves hicieron ajustes facultativos a la calidad del hábitat de invierno y a las condiciones medio ambientales. Independientemente del hábitat, las aves que salieron más tarde migraron más rápido, y este patrón fue mantenido a lo largo de la ruta migratoria (n = 44). Fue poco probable que migrar más temprano y más lento o más tarde y más rápido resultara en diferencias significativas en la fecha de llegada a los sitios de reproducción. Nuestros resultados revelan una subestimada complejidad en las decisiones migratorias de las aves y contrastan con el paradigma actual de salida y llegada más temprana en aves que usan hábitats óptimos. El siguiente paso es entender los beneficios relativos de migrar temprano versus tarde en el desempeño o si cada estrategia es una respuesta igualmente buena a las condiciones experimentadas.

Temporal patterns of foraging by silver-haired bats during migratory stopover revealed by isotopic analyses (δ13C) of breath CO2.

The extent to which migratory bats forage at stopover sites or while in migratory flight is poorly understood. Endogenous fat stores have lower δ13C values relative to the dietary substrates from which they were synthesized, and so, the fed versus fasted state of bats should be discernable by comparing their breath δ13C at capture to that after a known period of fasting. We captured silver-haired bats (Lasionycteris noctivagans) at a stopover site at Long Point, Ontario, Canada, during spring and fall migration. We collected breath samples at capture and after fasting in captivity for 12 h, providing a fasted-state δ13C value corresponding to metabolism of fat stores. We also collected and weighed fecal pellets produced while in captivity. Breath δ13C values at capture were positively correlated with mass of feces produced. During spring migration, δ13C values of breath CO2 at capture were low and similar to fasting values, but increased with date consistent with increased foraging at stopover and reliance on exogenous dietary nutrients as the season progressed. The opposite temporal pattern was found during fall migration. Our findings suggest that bats forage during migratory stopover when environmental conditions permit despite potential time trade-offs between feeding and travel, and the energy savings resulting from torpor during roosting. This study provides insight into the eco-physiology of bat migration and shows the importance of foraging habitat for migratory bats.

Mercury exposure to swallows breeding in Canada inferred from feathers grown on breeding and non-breeding grounds.

Aerial insectivorous birds such as swallows have been the steepest declining groups of birds in North America over the last 50 years but whether such declines are linked to contaminants has not been examined. We sampled feathers from five species of swallow at multiple locations to assess total mercury [THg] exposure for adults during the non-breeding season, and for juveniles on the breeding grounds. We assessed Hg exposure to juvenile birds in crop- and grass-dominated landscapes to determine if land-use practices influenced feather [THg]. We assayed feathers for stable isotopes (δ2H, δ13C, δ15N) as proxies for relative habitat use and diet to determine their potential influence on feather [THg]. Feather [THg] was highest in adult bank swallows (Riparia riparia) and purple martins (Progne subis) from Saskatchewan and adult cliff swallows (Petrochelidon pyrrhonota) from western regions, indicating differential exposure to Hg on the non-breeding grounds. Juvenile bank, barn (Hirundo rustica) and tree (Tachycineta bicolor) swallows had lower feather [THg] in crop-dominated landscapes than grass-dominated landscapes in Saskatchewan, potentially resulting from lower use of wetland-derived insects due to wetland drainage and intensive agriculture. Feather [THg] was related to juvenile feather stable isotopes for several species, suggesting complex interactions with diet and environmental factors. Many individuals had feather [THg] values >2 µg/g, a threshold at which deleterious effects may occur. Our findings indicate differential Hg exposure among species of swallow, regions and land-uses and highlight the need for additional research to determine dietary and finer-scale land-use impacts on individual species and populations.

Migratory monarchs that encounter resident monarchs show life-history differences and higher rates of parasite infection.

Environmental change induces some wildlife populations to shift from migratory to resident behaviours. Newly formed resident populations could influence the health and behaviour of remaining migrants. We investigated migrant-resident interactions among monarch butterflies and consequences for life history and parasitism. Eastern North American monarchs migrate annually to Mexico, but some now breed year-round on exotic milkweed in the southern US and experience high infection prevalence of protozoan parasites. Using stable isotopes (δ2 H, δ13 C) and cardenolide profiles to estimate natal origins, we show that migrant and resident monarchs overlap during fall and spring migration. Migrants at sites with residents were 13 times more likely to have infections and three times more likely to be reproductive (outside normal breeding season) compared to other migrants. Exotic milkweed might either attract migrants that are already infected or reproductive, or alternatively, induce these states. Increased migrant-resident interactions could affect monarch parasitism, migratory success and long-term conservation.

Tracking data and retrospective analyses of diet reveal the consequences of loss of marine subsidies for an obligate scavenger, the Andean condor.

Over the last century, marine mammals have been dramatically reduced in the world's oceans. We examined evidence that this change caused dietary and foraging pattern shifts of the Andean condor (Vultur gryphus) in Patagonia. We hypothesized that, after the decrease in marine mammals and the increase in human use of coastlines, condor diet changed to a more terrestrial diet, which in turn influenced their foraging patterns. We evaluated the diet by means of stable isotope analysis (δ13C, δ15N and δ34S) of current (last decade) and historical (1841-1933) feathers. We further evaluated the movement patterns of 23 condors using satellite tracking of individuals. Condors reduced their use of marine-derived prey in recent compared with historical times from 33 ± 13% to less than 8 ± 3% respectively; however, they still breed close to the coast. The average distance between the coast and nests was 62.5 km, but some nests were located close to the sea (less than 5 km). Therefore, some birds must travel up to 86 km from nesting sites, crossing over the mountain range to find food. The worldwide reduction in marine mammal carcasses, especially whales, may have major consequences on the foraging ecology of scavengers, as well as on the flux of marine inputs within terrestrial ecosystems.

Spring temperature, migration chronology, and nutrient allocation to eggs in three species of arctic-nesting geese: Implications for resilience to climate warming.

The macronutrients that Arctic herbivores invest in their offspring are derived from endogenous reserves of fat and protein (capital) that females build prior to the period of investment or from foods they consume concurrently with investment (income). The relative contribution from each source can be influenced by temporal and environmental constraints on a female's ability to forage on Arctic breeding areas. Warming temperatures and advancing Arctic phenology may alter those constraints. From 2011 to 2014, we examined relationships among spring temperature, timing of migration and reproduction, and the sources of nutrients females deposited in eggs for three sympatric species of geese that nested in northern Alaska. Compared to lesser snow geese (Anser caerulescens caerulescens) and greater white-fronted geese (Anser albifrons frontalis), black brant (Branta bernicla nigricans) were more likely to initiate follicle development during migration, resulting in fewer days between their arrival in the Arctic and the onset of incubation and requiring a relatively greater capital investment in eggs. Delaying follicle development until after their arrival in the Arctic provided snow geese and white-fronted geese an opportunity to forage near their nesting area and to deposit exogenous nutrients in eggs. With warmer spring temperatures, brant invested more capital in eggs, but snow geese invested less capital. Brant likely used capital to meet costs associated with earlier onset of follicle development when phenology was advanced, whereas snow geese used capital to compensate for poor foraging conditions during colder Arctic springs. Global warming is likely to reduce the quality of lower latitude marine habitats where brant acquire endogenous reserves and advancing Arctic phenology may increase their reliance on those reserves during reproduction. Near-term warming in northern Alaska may improve foraging conditions and favor the reproductive strategies of some herbivores such as snow geese and white-fronted geese that mainly invest Arctic nutrients in their offspring.

Tracing biogeochemical subsidies from glacier runoff into Alaska's coastal marine food webs.

Nearly half of the freshwater discharge into the Gulf of Alaska originates from landscapes draining glacier runoff, but the influence of the influx of riverine organic matter on the trophodynamics of coastal marine food webs is not well understood. We quantified the ecological impact of riverine organic matter subsidies to glacier-marine habitats by developing a multi-trophic level Bayesian three-isotope mixing model. We utilized large gradients in stable (δ13 C, δ15 N, δ2 H) and radiogenic (Δ14 C) isotopes that trace riverine and marine organic matter sources as they are passed from lower to higher trophic levels in glacial-marine habitats. We also compared isotope ratios between glacial-marine and more oceanic habitats. Based on isotopic measurements of potential baseline sources, ambient water and tissues of marine consumers, estimates of the riverine organic matter source contribution to upper trophic-level species including fish and seabirds ranged from 12% to 44%. Variability in resource use among similar taxa corresponded to variation in species distribution and life histories. For example, riverine organic matter assimilation by the glacier-nesting seabirds Kittlitz's murrelet (Brachyramphus brevirostris) was greater than that of the forest-nesting marbled murrelet (B. marmoratus). The particulate and dissolved organic carbon in glacial runoff and near surface coastal waters was aged (12100-1500 years BP 14 C-age) but dissolved inorganic carbon and biota in coastal waters were young (530 years BP 14 C-age to modern). Thus terrestrial-derived subsidies in marine food webs were primarily composed of young organic matter sources released from glacier ecosystems and their surrounding watersheds. Stable isotope compositions also revealed a divergence in food web structure between glacial-marine and oceanic sites. This work demonstrates linkages between terrestrial and marine ecosystems, and facilitates a greater understanding of how climate-driven changes in freshwater runoff have the potential to alter food web dynamics within coastal marine ecosystems in Alaska.

Assessing seasonal changes in animal diets with stable-isotope analysis of amino acids: a migratory boreal songbird switches diet over its annual cycle.

Tools to study seasonal changes in animal diets are needed to address a wide range of ecological questions. This is especially true of migratory animals that experience distinct environments where diets may be substantially different. However, tracking diets of individuals that move vast distances has proven difficult. Compound-specific isotope analysis has emerged as a valuable tool to study diets but has been little used to study dietary changes of migratory animals. Using this technique, we quantify seasonal variation in the annual diet of a migratory songbird (gray-cheeked thrush, Catharus minimus) and test the hypothesis that migrants change their diet in response to the energetic requirements of different periods of the annual cycle. By measuring δ13C and δ15N values of amino acids from feathers grown on the breeding grounds, blood formed during migration and claw grown on the wintering grounds, we found that migration is associated with greater consumption of fruit, compared to the breeding or wintering periods. This was confirmed by the lower trophic position of blood compared to feather and claw, by a decrease in the δ15N value of the source amino acid phenylalanine in blood as a function of days of stopover, and by the positive correlation between δ15N and δ13C values of phenylalanine in blood, and not in feather or claw. This study illustrates how isotopic analysis of amino acids can contribute to understand food webs, seasonal dietary changes and metabolic routing of nutrients in migratory animals.

Regional climate on the breeding grounds predicts variation in the natal origin of monarch butterflies overwintering in Mexico over 38 years.

Addressing population declines of migratory insects requires linking populations across different portions of the annual cycle and understanding the effects of variation in weather and climate on productivity, recruitment, and patterns of long-distance movement. We used stable H and C isotopes and geospatial modeling to estimate the natal origin of monarch butterflies (Danaus plexippus) in eastern North America using over 1000 monarchs collected over almost four decades at Mexican overwintering colonies. Multinomial regression was used to ascertain which climate-related factors best-predicted temporal variation in natal origin across six breeding regions. The region producing the largest proportion of overwintering monarchs was the US Midwest (mean annual proportion = 0.38; 95% CI: 0.36-0.41) followed by the north-central (0.17; 0.14-0.18), northeast (0.15; 0.11-0.16), northwest (0.12; 0.12-0.16), southwest (0.11; 0.08-0.12), and southeast (0.08; 0.07-0.11) regions. There was no evidence of directional shifts in the relative contributions of different natal regions over time, which suggests these regions are comprising the same relative proportion of the overwintering population in recent years as in the mid-1970s. Instead, interannual variation in the proportion of monarchs from each region covaried with climate, as measured by the Southern Oscillation Index and regional-specific daily maximum temperature and precipitation, which together likely dictate larval development rates and food plant condition. Our results provide the first robust long-term analysis of predictors of the natal origins of monarchs overwintering in Mexico. Conservation efforts on the breeding grounds focused on the Midwest region will likely have the greatest benefit to eastern North American migratory monarchs, but the population will likely remain sensitive to regional and stochastic weather patterns.