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.

A SEROLOGIC SURVEY OF FRANCISELLA TULARENSIS EXPOSURE IN WILDLIFE ON THE ARCTIC COASTAL PLAIN OF ALASKA, USA.

Tularemia is an infectious zoonotic disease caused by one of several subspecies of Francisella tularensis bacteria. Infections by F. tularensis are common throughout the northern hemisphere and have been detected in more than 250 wildlife species. In Alaska, US, where the pathogen was first identified in 1938, studies have identified F. tularensis antibodies in a diverse suite of taxa, including insects, birds, and mammals. However, few such investigations have been conducted recently and knowledge about the current distribution and disease ecology of F. tularensis is limited, particularly in Arctic Alaska, an area undergoing rapid environmental changes from climate warming. To help address these information gaps and provide insights about patterns of exposure among wildlife, we assessed the seroprevalence of F. tularensis antibodies in mammals and tundra-nesting geese from the Arctic Coastal Plain of Alaska, 2014-17. With a commercially available slide agglutination test, we detected antibodies in 14.7% of all individuals sampled (n=722), with titers ranging from 1:20 to 1:320. We detected significant differences in seroprevalence between family groups, with Canidae (foxes, Vulpes spp.) and Sciuridae (Arctic ground squirrel, Spermophilus parryii) having the highest seroprevalence at 21.5% and 33.3%, respectively. Mean seroprevalence for Ursidae (polar bears, Ursus maritimus) was 13.3%, whereas Cervidae (caribou, Rangifer tarandus) had comparatively low seroprevalence at 6.5%. Antibodies were detected in all Anatidae species sampled, with Black Brant (Branta bernicla nigricans) having the highest seroprevalence at 13.6%. The detection of F. tularensis antibodies across multiple taxa from the Arctic Coastal Plain and its nearshore marine region provides evidence of exposure to this pathogen throughout the region and highlights the need for renewed surveillance in Alaska.

Interspecific exchange of avian influenza virus genes in Alaska: the influence of trans-hemispheric migratory tendency and breeding ground sympatry.

The movement and transmission of avian influenza viral strains via wild migratory birds may vary by host species as a result of migratory tendency and sympatry with other infected individuals. To examine the roles of host migratory tendency and species sympatry on the movement of Eurasian low-pathogenic avian influenza (LPAI) genes into North America, we characterized migratory patterns and LPAI viral genomic variation in mallards (Anas platyrhynchos) of Alaska in comparison with LPAI diversity of northern pintails (Anas acuta). A 50-year band-recovery data set suggests that unlike northern pintails, mallards rarely make trans-hemispheric migrations between Alaska and Eurasia. Concordantly, fewer (14.5%) of 62 LPAI isolates from mallards contained Eurasian gene segments compared to those from 97 northern pintails (35%), a species with greater inter-continental migratory tendency. Aerial survey and banding data suggest that mallards and northern pintails are largely sympatric throughout Alaska during the breeding season, promoting opportunities for interspecific transmission. Comparisons of full-genome isolates confirmed near-complete genetic homology (>99.5%) of seven viruses between mallards and northern pintails. This study found viral segments of Eurasian lineage at a higher frequency in mallards than previous studies, suggesting transmission from other avian species migrating inter-hemispherically or the common occurrence of endemic Alaskan viruses containing segments of Eurasian origin. We conclude that mallards are unlikely to transfer Asian-origin viruses directly to North America via Alaska but that they are likely infected with Asian-origin viruses via interspecific transfer from species with regular migrations to the Eastern Hemisphere.

Life-history attributes of Arctic-breeding birds drive uneven responses to environmental variability across different phases of the reproductive cycle.

Animals exhibit varied life-history traits that reflect adaptive responses to their environments. For Arctic-breeding birds, traits related to diet, egg nutrient allocation, clutch size, and chick growth are predicted to be under increasing selection pressure due to rapid climate change and increasing environmental variability across high-latitude regions. We compared four migratory birds (black brant [Branta bernicla nigricans], lesser snow geese [Chen caerulescens caerulescens], semipalmated sandpipers [Calidris pusilla], and Lapland longspurs [Calcarius lapponicus]) with varied life histories at an Arctic site in Alaska, USA, to understand how life-history traits help moderate environmental variability across different phases of the reproductive cycle. We monitored aspects of reproductive performance related to the timing of breeding, reproductive investment, and chick growth from 2011 to 2018. In response to early snowmelt and warm temperatures, semipalmated sandpipers advanced their site arrival and bred in higher numbers, while brant and snow geese increased clutch sizes; all four species advanced their nest initiation dates. During chick rearing, longspur nestlings were relatively resilient to environmental variation, whereas warmer temperatures increased the growth rates of sandpiper chicks but reduced growth rates of snow goose goslings. These responses generally aligned with traits along the capital-income spectrum of nutrient acquisition and altricial-precocial modes of chick growth. Under a warming climate, the ability to mobilize endogenous reserves likely provides geese with relative flexibility to adjust the timing of breeding and the size of clutches. Higher temperatures, however, may negatively affect the quality of herbaceous foods and slow gosling growth. Species may possess traits that are beneficial during one phase of the reproductive cycle and others that may be detrimental at another phase, uneven responses that may be amplified with future climate warming. These results underscore the need to consider multiple phases of the reproductive cycle when assessing the effects of environmental variability on Arctic-breeding birds.

SURVEY OF ARCTIC ALASKAN WILDLIFE FOR INFLUENZA A ANTIBODIES: LIMITED EVIDENCE FOR EXPOSURE OF MAMMALS.

Influenza A viruses (IAVs) are maintained in wild waterbirds and have the potential to infect a broad range of species, including wild mammals. The Arctic Coastal Plain of Alaska supports a diverse suite of species, including waterfowl that are common hosts of IAVs. Mammals co-occur with geese and other migratory waterbirds during the summer breeding season, providing a plausible mechanism for interclass transmission of IAVs. To estimate IAV seroprevalence and identify the subtypes to which geese, loons, Arctic foxes ( Vulpes lagopus), caribou ( Rangifer tarandus), and polar bears ( Ursus maritimus) are potentially exposed, we used a blocking enzyme-linked immunosorbent assay (bELISA) and a hemagglutination inhibition (HI) assay to screen for antibodies to IAVs in samples collected during spring and summer of 2012-16. Apparent IAV seroprevalence using the bELISA was 50.3% in geese (range by species: 46-52.8%), 9% in loons (range by species: 3-20%), and 0.4% in Arctic foxes. We found no evidence for exposure to IAVs in polar bears or caribou by either assay. Among geese, we estimated detection probability from replicate bELISA analyses to be 0.92 and also found good concordance (>85%) between results from bELISA and HI assays, which identified antibodies reactive to H1, H6, and H9 subtype IAVs. In contrast, the HI assay detected antibodies in only one of seven loon samples that were positive by bELISA; that sample had low titers to both H4 and H5 IAV subtypes. Our results provide evidence that a relatively high proportion of waterbirds breeding on the Arctic Coastal Plain are exposed to IAVs, although it is unknown whether such exposure occurs locally or on staging or wintering grounds. In contrast, seroprevalence of IAVs in concomitant Arctic mammals is apparently low.

Nitrogen dynamics in an Alaskan salt marsh following spring use by geese.

Lesser snow geese (Anser caerulescens caerulescens) and Canada geese (Branta canadensis) use several salt marshes in Cook Inlet, Alaska, as stopover areas for brief periods during spring migration. We investigated the effects of geese on nitrogen cycling processes in Susitna Flats, one of the marshes. We compared net nitrogen mineralization, organic nitrogen pools and production in buried bags, nitrogen fixation by cyanobacteria, and soil and litter characteristics on grazed plots versus paired plots that had been exclosed from grazing for 3 years. Grazed areas had higher rates of net nitrogen mineralization in the spring and there was no effect of grazing on organic nitrogen availability. The increased mineralization rates in grazed plots could not be accounted for by alteration of litter quality, litter quantity, microclimate, or root biomass, which were not different between grazed and exclosed plots. In addition, fecal input was very slight in the year that we studied nitrogen cycling. We propose that trampling had two effects that could account for greater nitrogen availability in grazed areas: litter incorporation into soil, resulting in increased rates of decomposition and mineralization of litter material, and greater rates of nitrogen fixation by cyanobacteria on bare, trampled soils. A path analysis indicated that litter incorporation by trampling played a primary role in the nitrogen dynamics of the system, with nitrogen fixation secondary, and that fecal input was of little importance.

Forage digestibility and intake by lesser snow geese: effects of dominance and resource heterogeneity.

We measured forage intake, digestibility, and retention time for 11 free-ranging, human-imprinted lesser snow geese (Chen caerulescens caerulescens) as they consumed underground stembases of tall cotton-grass (Eriophorum angustifolium) on an arctic staging area in northeastern Alaska. Geese fed in small patches ([Formula: see text]=21.5 m2) of forage that made up ≤3% of the study area and consisted of high-quality "aquatic graminoid" and intermediate-quality "wet sedge" vegetation types. Dominant geese spent more time feeding in aquatic graminoid areas (r=0.61), but less total time feeding and more time resting than subdominant geese. Subdominant geese were displaced to areas of wet sedge where cotton-grass was a smaller proportion of underground biomass. Geese metabolized an average of 48% of the organic matter in stembases and there was a positive correlation between dominance and organic matter metabolizability (r=0.61). Total mean retention time of forage was 1.37 h and dry matter intake was 14.3 g/h. Snow geese that stage on the coastal plain of the Beaufort Sea likely use an extensive area because they consume a large mass of forage and exploit habitats that are patchily distributed and make up a small percentage of the landscape. Individual variation in nutrient absorption may result from agonistic interactions in an environment where resources are heterogeneously distributed.