Do wintering conditions drive population trends in semipalmated sandpipers (Calidris pusilla)? Evidence from a corticosterone biomarker

Thesis (M.S.) University of Alaska Fairbanks, 2018Some of the most extreme long-distance migrants, Arctic-breeding shorebirds are disproportionately represented in tallies of declining species worldwide. For many shorebirds, including the semipalmated sandpiper (Calidris pusilla), the specific causes and mechanisms behind population declines have not been identified. Stressful conditions affecting birds during wintering are often implicated. Interactions between events and processes occurring in the disparate locations used throughout the annual cycle also may be critical in shaping both individual life histories and population demographics. The main objectives of my graduate research were: a) to examine whether semipalmated sandpipers wintering in specific locations incur differential levels of stress; and b) to test whether stressful conditions may carry over between different stages of an individual's life cycle. Using measurements of corticosterone (the primary avian stress hormone) deposited in winter-grown feathers, I examined the contribution of breeding season and fall migration to winter-incurred stress, and looked for evidence of carryover effects from wintering conditions to spring migration and subsequent reproductive performance. In Chapter 1, I compared the levels of stress exposure of 40 semipalmated sandpipers that bred at five Arctic sites and spent the austral summer in distinct regions (identified via light-sensing geolocators) across their tropical 'wintering' range. I found stress exposure varied by wintering region, and birds using locations along the Atlantic coast of northeastern South America and the Pacific coast of Central America had the highest feather corticosterone levels. I did not find evidence that carryover effects from the breeding season and/or fall migration influenced birds' physiology during winter. In Chapter 2, I investigated whether greater stress exposure during winter might subsequently affect birds during spring migration and/or breeding. I found that geolocator-tracked birds with increased stress levels delayed spring migration and initiated nests later. However, results for a larger dataset (including 254 birds breeding at seven sites across the North American Arctic) suggested low-stress birds nested later. It is possible the larger dataset included replacement clutches that could have confounded relationships with feather corticosterone, as only birds in better condition are likely to re-nest after clutch failure. In addition, I found evidence that stressful wintering conditions carryover to affect reproductive performance: females that accrued high levels of stress during wintering subsequently laid fewer eggs. In confirmed first nests, we found evidence for a clutch size-egg volume tradeoff, with high-stress females producing fewer offspring but potentially investing more in individual offspring. This research represents the first instance of the feather corticosterone technique being used to compare conditions across the wintering range of a calidrid shorebird and reveals specific wintering locations with high levels of stress exposure. This is also the first research that provides a mechanistic perspective on carryover effects between the wintering and breeding stages in a shorebird, through measurements of feather corticosterone. Finally, by showing that poor environmental conditions at wintering sites far from Arctic breeding areas may be detrimental to the reproductive performance of a species with declining populations, this research emphasizes the importance of considering full annual cycles in conservation and research efforts for migratory species

Effects of geolocators on hatching success, return rates, breeding movements, and change in body mass in 16 species of Arctic-breeding shorebirds

Additional Authors: Bruce Casler; Maureen Christie; Jonathan T. Coleman; Jesse R. Conklin; Willow B. English; H. River Gates; Olivier Gilg; Marie-Andrée Giroux; Ken Gosbell; Chris Hassell; Jim Helmericks; Andrew Johnson; Borgný Katrínardóttir; Kari Koivula; Jean-Francois Lamarre; Johannes Lang; David B. Lank; Nicolas Lecomte; Joe Liebezeit; Vanessa Loverti; Laura McKinnon; Clive Minton; David Mizrahi; Erica Nol; Veli-Matti Pakanen; Johanna Perz; Ron Porter; Jennie Rausch; Jeroen Reneerkens; Nelli Rönkä; Sarah Saalfeld; Nathan Senner; Benoît Sittler; Paul A. Smith; Kristine Sowl; Audrey Taylor; David H. Ward; Stephen YezerinacCitation: Weiser, E.L., R.B. Lanctot, S.C. Brown, J.A. Alves, P.F. Battley, R. Bentzen, J. Bêty, M.A. Bishop, M. Boldenow, L. Bollache, B. Casler, M. Christie, J.T. Coleman, J.R. Conklin, W.B. English, H.R. Gates, O. Gilg, M.-A. Giroux, K. Gosbell, C. Hassell, J. Helmericks, A. Johnson, B. Katrínardóttir, K. Koivula, E. Kwon, J.-F. Lamarre, J. Lang, D.B. Lank, N. Lecomte, J. Liebezeit, V. Loverti, L. McKinnon, C. Minton, D. Mizrahi, E. Nol, V.-M. Pakanen, J. Perz, R. Porter, J. Rausch, J. Reneerkens, N. Rönkä, S. Saalfeld, N. Senner, B. Sittler, P.A. Smith, K. Sowl, A. Taylor, D.H. Ward, S. Yezerinac, and B.K. Sandercock. 2016. Effects of geolocators on hatching success, return rates, breeding movements and change in body mass in 16 species of Arctic-breeding shorebirds. Movement Ecology 4: art12. URL: http://movementecologyjournal.biomedcentral.com/articles/10.1186/s40462-016-0077-6Background: Geolocators are useful for tracking movements of long-distance migrants, but potential negative effects on birds have not been well studied. We tested for effects of geolocators (0.8–2.0 g total, representing 0.1–3.9 % of mean body mass) on 16 species of migratory shorebirds, including five species with 2–4 subspecies each for a total of 23 study taxa. Study species spanned a range of body sizes (26–1091 g) and eight genera, and were tagged at 23 breeding and eight nonbreeding sites. We compared breeding performance and return rates of birds with geolocators to control groups while controlling for potential confounding variables. Result: We detected negative effects of tags for three small-bodied species. Geolocators reduced annual return rates for two of 23 taxa: by 63 % for semipalmated sandpipers and by 43 % for the arcticola subspecies of dunlin. High resighting effort for geolocator birds could have masked additional negative effects. Geolocators were more likely to negatively affect return rates if the total mass of geolocators and color markers was 2.5–5.8 % of body mass than if tags were 0.3–2.3 % of body mass. Carrying a geolocator reduced nest success by 42 % for semipalmated sandpipers and tripled the probability of partial clutch failure in semipalmated and western sandpipers. Geolocators mounted perpendicular to the leg on a flag had stronger negative effects on nest success than geolocators mounted parallel to the leg on a band. However, parallel-band geolocators were more likely to reduce return rates and cause injuries to the leg. No effects of geolocators were found on breeding movements or changes in body mass. Among-site variation in geolocator effect size was high, suggesting that local factors were important. Conclusions: Negative effects of geolocators occurred only for three of the smallest species in our dataset, but were substantial when present. Future studies could mitigate impacts of tags by reducing protruding parts and minimizing use of additional markers. Investigators could maximize recovery of tags by strategically deploying geolocators on males, previously marked individuals, and successful breeders, though targeting subsets of a population could bias the resulting migratory movement data in some species

Unexpected diversity in socially synchronized rhythms of shorebirds

The behavioural rhythms of organisms are thought to be under strong selection, influenced by the rhythmicity of the environment1, 2, 3, 4. Such behavioural rhythms are well studied in isolated individuals under laboratory conditions1, 5, but free-living individuals have to temporally synchronize their activities with those of others, including potential mates, competitors, prey and predators6, 7, 8, 9, 10. Individuals can temporally segregate their daily activities (for example, prey avoiding predators, subordinates avoiding dominants) or synchronize their activities (for example, group foraging, communal defence, pairs reproducing or caring for offspring)6, 7, 8, 9, 11. The behavioural rhythms that emerge from such social synchronization and the underlying evolutionary and ecological drivers that shape them remain poorly understood5, 6, 7, 9. Here we investigate these rhythms in the context of biparental care, a particularly sensitive phase of social synchronization12 where pair members potentially compromise their individual rhythms. Using data from 729 nests of 91 populations of 32 biparentally incubating shorebird species, where parents synchronize to achieve continuous coverage of developing eggs, we report remarkable within- and between-species diversity in incubation rhythms. Between species, the median length of one parent’s incubation bout varied from 1–19 h, whereas period length—the time in which a parent’s probability to incubate cycles once between its highest and lowest value—varied from 6–43 h. The length of incubation bouts was unrelated to variables reflecting energetic demands, but species relying on crypsis (the ability to avoid detection by other animals) had longer incubation bouts than those that are readily visible or who actively protect their nest against predators. Rhythms entrainable to the 24-h light–dark cycle were less prevalent at high latitudes and absent in 18 species. Our results indicate that even under similar environmental conditions and despite 24-h environmental cues, social synchronization can generate far more diverse behavioural rhythms than expected from studies of individuals in captivity5, 6, 7, 9. The risk of predation, not the risk of starvation, may be a key factor underlying the diversity in these rhythms

Landscape Change Detected over a Half Century in the Arctic National Wildlife Refuge Using High-Resolution Aerial Imagery

Rapid warming has occurred over the past 50 years in Arctic Alaska, where temperature strongly affects ecological patterns and processes. To document landscape change over a half century in the Arctic National Wildlife Refuge, Alaska, we visually interpreted geomorphic and vegetation changes on time series of coregistered high-resolution imagery. We used aerial photographs for two time periods, 1947–1955 and 1978–1988, and Quick Bird and IKONOS satellite images for a third period, 2000–2007. The stratified random sample had five sites in each of seven ecoregions, with a systematic grid of 100 points per site. At each point in each time period, we recorded vegetation type, microtopography, and surface water. Change types were then assigned based on differences detected between the images. Overall, 23% of the points underwent some type of change over the ~50-year study period. Weighted by area of each ecoregion, we estimated that 18% of the Refuge had changed. The most common changes were wildfire and postfire succession, shrub and tree increase in the absence of fire, river erosion and deposition, and ice-wedge degradation. Ice-wedge degradation occurred mainly in the Tundra Biome, shrub increase and river changes in the Mountain Biome, and fire and postfire succession in the Boreal Biome. Changes in the Tundra Biome tended to be related to landscape wetting, mainly from increased wet troughs caused by ice-wedge degradation. The Boreal Biome tended to have changes associated with landscape drying, including recent wildfire, lake area decrease, and land surface drying. The second time interval, after ~1982, coincided with accelerated climate warming and had slightly greater rates of change

Composition and Drivers of Gut Microbial Communities in Arctic-Breeding Shorebirds

Gut microbiota can have important effects on host health, but explanatory factors and pathways that determine gut microbial composition can differ among host lineages. In mammals, host phylogeny is one of the main drivers of gut microbiota, a result of vertical transfer of microbiota during birth. In birds, it is less clear what the drivers might be, but both phylogeny and environmental factors may play a role. We investigated host and environmental factors that underlie variation in gut microbiota composition in eight species of migratory shorebirds. We characterized bacterial communities from 375 fecal samples collected from adults of eight shorebird species captured at a network of nine breeding sites in the Arctic and sub-Arctic ecoregions of North America, by sequencing the V4 region of the bacterial 16S ribosomal RNA gene. Firmicutes (55.4%), Proteobacteria (13.8%), Fusobacteria (10.2%), and Bacteroidetes (8.1%) dominated the gut microbiota of adult shorebirds. Breeding location was the main driver of variation in gut microbiota of breeding shorebirds (R2 = 11.6%), followed by shorebird host species (R2 = 1.8%), and sampling year (R2 = 0.9%), but most variation remained unexplained. Site variation resulted from differences in the core bacterial taxa, whereas rare, lowabundance bacteria drove host species variation. Our study is the first to highlight a greater importance of local environment than phylogeny as a driver of gut microbiota composition in wild, migratory birds under natural conditions.publishedVersio

Composition and Drivers of Gut Microbial Communities in Arctic-Breeding Shorebirds

Gut microbiota can have important effects on host health, but explanatory factors and pathways that determine gut microbial composition can differ among host lineages. In mammals, host phylogeny is one of the main drivers of gut microbiota, a result of vertical transfer of microbiota during birth. In birds, it is less clear what the drivers might be, but both phylogeny and environmental factors may play a role. We investigated host and environmental factors that underlie variation in gut microbiota composition in eight species of migratory shorebirds. We characterized bacterial communities from 375 fecal samples collected from adults of eight shorebird species captured at a network of nine breeding sites in the Arctic and sub-Arctic ecoregions of North America, by sequencing the V4 region of the bacterial 16S ribosomal RNA gene. Firmicutes (55.4%), Proteobacteria (13.8%), Fusobacteria (10.2%), and Bacteroidetes (8.1%) dominated the gut microbiota of adult shorebirds. Breeding location was the main driver of variation in gut microbiota of breeding shorebirds (R2 = 11.6%), followed by shorebird host species (R2 = 1.8%), and sampling year (R2 = 0.9%), but most variation remained unexplained. Site variation resulted from differences in the core bacterial taxa, whereas rare, lowabundance bacteria drove host species variation. Our study is the first to highlight a greater importance of local environment than phylogeny as a driver of gut microbiota composition in wild, migratory birds under natural conditions.publishedVersio

Effects of leg flags on nest survival of four species of Arctic-breeding shorebirds

Marking wild birds is an integral part of many field studies. However, if marks affect the vital rates or behavior of marked individuals, any conclusions reached by a study might be biased relative to the general population. Leg bands have rarely been found to have negative effects on birds and are frequently used to mark individuals. Leg flags, which are larger, heavier, and might produce more drag than bands, are commonly used on shorebirds and can help improve resighting rates. However, no one to date has assessed the possible effects of leg flags on the demographic performance of shorebirds. At seven sites in Arctic Alaska and western Canada, we marked individuals and monitored nest survival of four species of Arctic-breeding shorebirds, including Semipalmated Sandpipers (Calidris pusilla), Western Sandpipers (C. mauri), Red-necked Phalaropes (Phalaropus lobatus), and Red Phalaropes (P. fulicarius). We used a daily nest survival model in a Bayesian framework to test for effects of leg flags, relative to birds with only bands, on daily survival rates of 1952 nests. We found no evidence of a difference in nest survival between birds with flags and those with only bands. Our results suggest, therefore, that leg flags have little effect on the nest success of Arctic-breeding sandpipers and phalaropes. Additional studies are needed, however, to evaluate the possible effects of flags on shorebirds that use other habitats and on survival rates of adults and chicks.acceptedVersio