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The Prairie Pothole Region of central Canada supports a diverse community of breeding waterbirds, but many species have declining populations and the demographic mechanisms driving the declines remain unknown. We conducted a 7-year field study during 1995–2001 to investigate the demographic performance of Marbled Godwits (Limosa fedoa) and Willets (Tringa semipalmata) breeding in managed wetlands near Brooks, Alberta. Mark-recapture analyses based on Cormack–Jolly– Seber models revealed that the annual rates of apparent survival for Marbled Godwits = 0.953 ± 0.012SE) and Willets ( = 0.861 ± 0.015SE) are among the highest rates of survivorship reported for any breeding or nonbreeding population of large-bodied shorebirds. Our estimates of life expectancy for males were comparable to longevity records in godwits (17.3 years ±5.8SE vs. 25–29+ years) and willets (7.7 ± 1.5SE vs. 10+ years). The two species both showed strong breeding site fidelity but differed in rates of mate fidelity. Pairs that reunited and males that switched mates usually nested 1.1–1.5 km. Returning pairs usually reunited in godwits (85%) but not in willets (28%), possibly because of species differences in adult survival or patterns of migration. Baseline estimates of annual survival for banded-only birds will be useful for evaluating the potential effects of new tracking tags or the environmental changes that have occurred during the past 20 years. Conservation strategies for large-bodied shorebirds should be focused on reduction of exposure to anthropogenic mortality because low rates of natural mortality suggest that losses to collisions at breeding sites or harvest at nonbreeding areas are likely to cause additive mortality. apparent survival, biometrics, demography, longevity, mark recapture, mate fidelity, shorebird, wader Applied ecology, Conservation ecology, Demography, Life history ecology, Population ecology, Zoology

Semipalmated Sandpipers (Calidris pusilla) are Arctic-breeding shorebirds that use staging sites in Atlantic Canada during their southbound migration to South America. The upper Bay of Fundy is recognized as a critical staging area, but hundreds of smaller sites outside the Bay of Fundy also host staging Semipalmated Sandpipers and have received comparatively little attention. Using the Motus Wildlife Tracking System, we tracked adult and juvenile Semipalmated Sandpipers and identified different staging strategies used by sandpipers tagged inside and outside the Bay of Fundy. Birds tagged in the Bay of Fundy remained there, while birds tagged along the Northumberland Strait, a tidal water body along the eastern coast of New Brunswick and northern Nova Scotia displayed multiple strategies. Most birds tagged along the Northumberland Strait used sites outside the Bay of Fundy exclusively, while a smaller proportion moved to the Bay of Fundy. Length of stay was shorter for birds using only sites outside the Bay of Fundy compared to birds using Bay sites or transferring between them. Choice of departure conditions also varied; birds using the Bay were more selective of specific wind conditions, favoring north and northwesterly winds. We found juvenile Semipalmated Sandpipers stayed 3 days longer in the region than adults and appeared to use a broader variety of sites. Our results highlight the importance of implementing conservation measures for smaller, lesser-known staging sites in Atlantic Canada, many of which are at an increased risk of loss and degradation due to climate change, coastal development, pollution, and human disturbance.

Droughts can affect invertebrate communities in wetlands, which can have bottom-up effects on the condition and survival of top predators. Shorebirds, key predators at coastal wetlands, have experienced widespread population declines and could be negatively affected by droughts. We explored, in detail, the effects of drought on multiple aspects of shorebird stopover and migration ecology by contrasting a year with average wet/dry conditions (2016) with a year with moderate drought (2017) at a major subarctic stopover site on southbound migration. We also examined the effects of drought on shorebird body mass during stopover across 14 years (historical: 1974–1982 and present-day: 2014–2018). For the detailed comparison of two years, in the year with moderate drought we documented lower invertebrate abundance at some sites, higher prey family richness in shorebird faecal samples, lower shorebird refuelling rates, shorter stopover durations for juveniles, and, for most species, a higher probability of making a subsequent stopover in North America after departing the subarctic, compared to the year with average wet/dry conditions. In the 14-year dataset, shorebird body mass tended to be lower in drier years. We show that even short-term, moderate drought conditions can negatively affect shorebird refuelling performance at coastal wetlands, which may carry-over to affect subsequent stopover decisions. Given shorebird population declines and predicted changes in the severity and duration of droughts with climate change, researchers should prioritize a better understanding of how droughts affect shorebird refuelling performance and survival. Supplementary Information The online version contains supplementary material available at 10.1007/s00442-021-05047-x.

The Bay of Fundy, Canada is a critical migratory stopover for Semipalmated Sandpipers (Calidris pusilla) during fall migration. Recent ecological changes combined with range-wide population declines indicate that a re-evaluation of habitat use and regional fidelity is required. The Bay of Fundy is divided into three regions, each of which contains multiple foraging and roost sites. In this study, we examined local movement and regional fidelity within and between years using radio-telemetry and field-readable markers. We deployed 194 radio-transmitters and 2295 field-readable flags on Semipalmated Sandpipers during migration in 2013-2014, and resighted these birds, along with additional birds flagged in 2012, between 2013 and 2016. Radio-tracked birds used on average 1.3 to 4.1 foraging and roost sites per day, with most making multiple daily movements, but only 2% made permanent moves between geographically distant regions within the bay during stopover. Additionally, 97% of birds returned to their original region in subsequent years. The use of multiple sites suggests Semipalmated Sandpipers are capable of adapting to dynamic conditions in the Bay of Fundy on a local scale, but the high regional fidelity suggests that their ability to adapt to changes occurring at a regional scale may be limited. Uncertainty about the capability of Semipalmated Sandpipers to flexibly move among different regions of the Bay of Fundy supports the need for conservation at both the site and regional level.

Effective conservation for listed migratory species requires an understanding of how drivers of population decline vary spatially and temporally, as well as knowledge of range-wide connectivity between breeding and nonbreeding areas. Environmental conditions distant from breeding areas can have lasting effects on the demography of migratory species, yet these consequences are often the least understood. Our objectives were to 1) evaluate associations between survival and extreme environmental disturbances at nonbreeding areas, including hurricanes, harmful algal blooms, and oil spills, and 2) estimate migratory connectivity between breeding and nonbreeding areas of midcontinental piping plovers (Charadrius melodus). We used capture and resighting data from 5067 individuals collected between 2002 and 2019 from breeding areas across the midcontinent, and nonbreeding areas throughout the Gulf of Mexico and southern Atlantic coasts of North America. We developed a hidden Markov multistate model to estimate seasonal survival and account for unobservable geographic locations. Hurricanes and harmful algal blooms were negatively associated with nonbreeding season survival, but we did not detect a similarly negative relationship with oil spills. Our results indicated that individuals from separate breeding areas mixed across nonbreeding areas with low migratory connectivity. Mixing among individuals in the nonbreeding season may provide a buffering effect against impacts of extreme events on any one breeding region. Our results suggest that understanding migratory connectivity and linking seasonal threats to population dynamics can better inform conservation strategies for migratory shorebirds.

Medium-distance migrant shorebirds that nest on beaches contend with human disturbance and habitat development on their breeding, migratory stopover and non-breeding sites. Conservation has focused on breeding areas because protection of nests is more tractable than protection of non-breeding habitat. As breeding ground management has encountered diminishing returns, interest in understanding threats in non-breeding areas has increased. However, robust estimates of non-breeding demographic rates (such as entry and persistence probabilities) and abundance are generally lacking, hindering the study of limiting or regulating factors. Estimating such rates is made more difficult by complex population dynamics at non-breeding sites. In the important non-breeding site of South Carolina, endangered Piping Plovers Charadrius melodus start arriving in July and some depart prior to December (the autumn-only population) while others remain through March (the wintering population, which in other locales has been shown to be closed in the winter months of December to February) or even until May. State uncertainty capture-mark-recapture models provide a means for estimating entry and persistence rates for such co-occurring populations. We estimated the proportion of the population entering the study area per survey (entry probability) and proportion remaining per survey (persistence rate) for both populations during the autumn, and abundance of the wintering population in the winter months, at four high-use sites in South Carolina in the non-breeding periods of 2006/7 and 2007/8, taking advantage of birds previously colour-ringed on the breeding grounds. We were able to make fairly precise estimates of entry and persistence rates with small sample sizes. Cumulative probability of entry was ~50% by the end of July and reached 95% for both populations by October. Estimated stopover duration for birds in the autumn-only population was 35 days in year 1 and 42 days in year 2. In the winter months (December to January), we estimated a wintering super-population size ± SE of 71±16 Piping Plovers in the first year and 75±16 Piping Plovers in the second year. If ringing programs on the breeding grounds continue, annual standardized resighting surveys in the nonbreeding period and mark-recapture models can provide robust estimates of entry and persistence rates and abundance. Long term monitoring programs can thus permit separation of demographic variance from sampling error, which is crucial for further research into population regulation and limiting factors. Our results suggest that habitat protection intended to benefit non-breeding Piping Plovers at our coastal sites should be in effect by late summer, as many birds are resident from July through the end of winter in March. This article is protected by copyright. All rights reserved.

Semipalmated Sandpiper (Calidris pusilla) populations have undergone significant declines at core nonbreeding sites in northeastern South America. Breeding populations have also declined in the eastern North American Arctic, but appear to be stable or increasing in the central and western Arctic. To identify vulnerable populations and sites, we documented the migratory connectivity of Semipalmated Sandpipers using light-level geolocators, deploying 250 at 8 Arctic sites across the species' breeding range from 2011 to 2015, plus 87 at a single wintering site in northeastern Brazil in 2013 and 2014. We recovered 59 units and resighted 7 more (26% return rate) on the breeding grounds, but none at the nonbreeding site. We recovered only ∼3% of units deployed in 2013 at eastern Arctic breeding sites, but recovered 33% of those deployed in 2015. Overall, birds with geolocators were 57% as likely to return as those carrying alphanumeric flags. Stopover durations at prairie sites (mean: 8.7 days southbou...

We compiled a >50‐year record of morphometrics for semipalmated sandpipers (Calidris pusilla), a shorebird species with a Nearctic breeding distribution and intercontinental migration to South America. Our data included >57,000 individuals captured 1972–2015 at five breeding locations and three major stopover sites, plus 139 museum specimens collected in earlier decades. Wing length increased by ca. 1.5 mm (>1%) prior to 1980, followed by a decrease of 3.85 mm (nearly 4%) over the subsequent 35 years. This can account for previously reported changes in metrics at a migratory stopover site from 1985 to 2006. Wing length decreased at a rate of 1,098 darwins, or 0.176 haldanes, within the ranges of other field studies of phenotypic change. Bill length, in contrast, showed no consistent change over the full period of our study. Decreased body size as a universal response of animal populations to climate warming, and several other potential mechanisms, are unable to account for the increasing and decreasing wing length pattern observed. We propose that the post‐WWII near‐extirpation of falcon populations and their post‐1973 recovery driven by the widespread use and subsequent limitation on DDT in North America selected initially for greater flight efficiency and latterly for greater agility. This predation danger hypothesis accounts for many features of the morphometric data and deserves further investigation in this and other species.

The objective of this study was to determine the implications of extreme tidal events on duration of stay and behavior of Semipalmated Sandpipers (Calidris pusilla) during migratory stopover in Cobequid Bay, Nova Scotia. This area is part of the Bay of Fundy and experiences the largest tidal range in the world. Radiotelemetry was used to monitor duration of stay of 30 adult and seven juvenile Semipalmated Sandpipers. Adults arriving in Cobequid Bay early in the migration period experienced a greater number of extreme high tides (> 15 m) that submerged preferred roost sites, and stayed on average 8.1 days longer than those that arrived later. Juvenile duration of stay was not significantly different from adults. When tides exceeded 15 m, Semipalmated Sandpipers engaged in over ocean flocking at high tide; however, this behavior was not observed when high tides were 13.6 m or less. These extra flights led to higher energy expenditure by early migrants, requiring an estimated 3.8 to 4.3-day increase in length of stay to reach the same mass as later migrants. In the future, predicted sea level rise could increase the frequency of extreme tidal amplitudes and result in greater energetic costs during Semipalmated Sandpiper stopover.

Aim: Parental care improves the survival of offspring and therefore has a major impact on reproductive success. It is increasingly recognized that coordinated biparental care is necessary to ensure the survival of offspring in hostile environments, but little is known about the influence of environmental fluctuations on parental cooperation. Assessing the impacts of environmental stochasticity, however, is essential for understanding how populations will respond to climate change and the associated increasing frequencies of extreme weather events. Here we investigate the influence of environmental stochasticity on biparental incubation in a cosmopolitan ground-nesting avian genus. Location: Global. Methods: We assembled data on biparental care in 36 plover populations (Charadrius spp.) from six continents, collected between 1981 and 2012. Using a space-for-time approach we investigate how average temperature, temperature stochasticity (i.e. year-to-year variation) and seasonal temperature variation during the breeding season influence parental cooperation during incubation. Results: We show that both average ambient temperature and its fluctuations influence parental cooperation during incubation. Male care relative to female care increases with both mean ambient temperature and temperature stochasticity. Local climatic conditions explain within-species population differences in parental cooperation, probably reflecting phenotypic plasticity of behaviour. Main conclusions: The degree of flexibility in parental cooperation is likely to mediate the impacts of climate change on the demography and reproductive behaviour of wild animal populations.

Long-distance migrants are assumed to be more time-limited during the pre-breeding season compared to the post-breeding season. Although breeding-related time constraints may be absent post-breeding, additional factors such as predation risk could lead to time constraints that were previously underestimated. By using an automated radio telemetry system, we compared pre- and post-breeding movements of long-distance migrant shorebirds on a continent-wide scale. From 2014 to 2016, we deployed radio transmitters on 1,937 individuals of 4 shorebird species at 13 sites distributed across North America. Following theoretical predictions, all species migrated faster during the pre-breeding season, compared to the post-breeding season. These differences in migration speed between seasons were attributable primarily to longer stopover durations in the post-breeding season. In contrast, and counter to our expectations, all species had higher airspeeds during the post-breeding season, even after accounting for seasonal differences in wind. Arriving at the breeding grounds in good body condition is beneficial for survival and reproductive success and this energetic constraint might explain why airspeeds are not maximised in the pre-breeding season. We show that the higher airspeeds in the post-breeding season precede a wave of avian predators, which could suggest that migrant shorebirds show predation-minimizing behaviour during the post-breeding season. Our results reaffirm the important role of time constraints during northward migration and suggest that both energy and predation-risk constrain migratory behaviour during the post-breeding season.

A recent study demonstrated that semipalmated sandpiper (Calidris pusilla) wing lengths have shortened from the 1980s to the present-day. We examined alternative and untested hypotheses for this change at an important stopover site, James Bay, Ontario, Canada. We evaluated morphometric patterns in wing length and bill length by age and sex, when possible, and assessed if wing shape has also changed during this time-period. We investigated patterns of morphological change in two additional Calidridine sandpipers, white-rumped sandpipers (Calidris fuscicollis) and least sandpipers (Calidris minutilla), to determine if shorter wing lengths are a widespread pattern in small sandpipers. We also examined allometric changes in wing and bill lengths to clarify if wing length declines were consistent with historical scaling relationships and indicative of a change in body size instead of only wing length change. We found that including sex and wing shape in analyses revealed important patterns in morphometric change for semipalmated sandpipers. Wing lengths declined for both sexes, but the magnitude of decline was smaller and not significant for males. Additionally, semipalmated sandpiper wings have become more convex, a shape that increases maneuverability in flight. Wing lengths, but not bill lengths, declined for most species and age classes, a pattern that was inconsistent with historical allometric scaling relationships. For juvenile semipalmated sandpipers, however, both bill and wing lengths declined according to historical scaling relationships, which could be a consequence of nutritional stress during development or a shift in the proportion of birds from smaller-sized, western breeding populations. Except for juvenile semipalmated sandpipers, we did not find evidence for an increase in the proportion of birds from different breeding populations at the stopover site. Given the wide, hemispheric distribution of these sandpipers throughout their annual cycles, our results, paired with those from a previous study, provide evidence for wide-spread reduction in wing lengths of Calidridine sandpipers since the 1980s. The shorter wing lengths and more convex wing shapes found in this study support the hypothesis that selection has favored more maneuverable wing morphology in small sandpipers.

Most of the known wintering areas of Piping Plovers (Charadrius melodus) are along the Atlantic and Gulf coasts of the United States and into Mexico, and in the Caribbean. However, 1066 threatened/endangered Piping Plovers were recently found wintering in The Bahamas, an area not previously known to be important for the species. Although representing about 27% of the birds counted during the 2011 International Piping Plover Winter Census, the location of their breeding site(s) was unknown. Thus, our objectives were to determine the location(s) of their breeding site(s) using molecular markers and by tracking banded individuals, identify spring and fall staging sites, and examine site fidelity and survival. We captured and color-banded 57 birds in January and February 2010 in The Bahamas. Blood samples were also collected for genetic evaluation of the likely subspecies wintering in The Bahamas. Band re-sightings and DNA analysis revealed that at least 95% of the Piping Plovers wintering in The Bahamas originated on the Atlantic coast of the United States and Canada. Re-sightings of birds banded in The Bahamas spanned the breeding distribution of the species along the Atlantic coast from Newfoundland to North Carolina. Site fidelity to breeding and wintering sites was high (88–100%). Spring and fall staging sites were located along the Atlantic coast of the United States, with marked birds concentrating in the Carolinas. Our estimate of true survival for the marked birds was 0.71 (95% CI: 0.61–0.80). Our results indicate that more than one third of the Piping Plover population that breeds along the Atlantic coast winters in The Bahamas. By determining the importance of The Bahamas to the Atlantic subspecies of Piping Plovers, future conservation efforts for these populations can be better focused on where they are most needed.

Listing a species under the Endangered Species Act (ESA) and developing a recovery plan requires U.S. Fish and Wildlife Service to establish specific and measurable criteria for delisting. Generally, species are listed because they face (or are perceived to face) elevated risk of extinction due to issues such as habitat loss, invasive species, or other factors. Recovery plans identify recovery criteria that reduce extinction risk to an acceptable level. It logically follows that the recovery criteria, the defined conditions for removing a species from ESA protections, need to be closely related to extinction risk. Extinction probability is a population parameter estimated with a model that uses current demographic information to project the population into the future over a number of replicates, calculating the proportion of replicated populations that go extinct. We simulated extinction probabilities of piping plovers in the Great Plains and estimated the relationship between extinction probability and various demographic parameters. We tested the fit of regression models linking initial abundance, productivity, or population growth rate to extinction risk, and then, using the regression parameter estimates, determined the conditions required to reduce extinction probability to some pre-defined acceptable threshold. Binomial regression models with mean population growth rate and the natural log of initial abundance were the best predictors of extinction probability 50 years into the future. For example, based on our regression models, an initial abundance of approximately 2400 females with an expected mean population growth rate of 1.0 will limit extinction risk for piping plovers in the Great Plains to less than 0.048. Our method provides a straightforward way of developing specific and measurable recovery criteria linked directly to the core issue of extinction risk. Published by Elsevier Ltd.

Large numbers of Semipalmated Sandpipers (Calidris pusilla) use the Bay of Fundy, Canada, as a stopover site during their southbound migration from breeding to wintering grounds. The species exhibits a geographic cline in bill length on the breeding grounds (larger in east). Based on a previous analysis of morphometric traits that detected a small temporal decline in bill length, it has been suggested that observed declines in the Semipalmated Sandpiper population staging in the Bay of Fundy were driven by a disproportionate loss from the eastern breeding population. New data collected in the Bay of Fundy has made it possible to reassess this suggestion. We compared new bill length data from the Bay of Fundy (2009–2014) to historical data (1981–1989, 1997–2007) to examine temporal changes in bill length at this stopover site. We also examined count data at a major roost site from 1981 to 2014 to assess temporal changes in migratory passage. Variation in median passage date is explained best by a combination of year and Arctic climate data. Birds are now migrating through the region later in the season; median passage date has advanced approximately 5 d from 1980s values. Our analyses suggest that while there was a small decline in bill length between the 1980s and early 2000s, it has since reversed and bills are now comparable to historical values. Given this reversal in mean bill length, coupled with population estimates that are still low relative to historical values, there is now little evidence to support the hypothesis that the decline between the 1980s and today in birds passing through the Bay of Fundy is linked to disproportionate losses of eastern breeders. We therefore suggest that population declines may be distributed more broadly across the breeding range of birds that use the Bay of Fundy as a stopover site.

Semipalmated Sandpipers (Calidris pusilla) are among the most common North American shorebirds. Breeding in Arctic North America, this species displays regional differences in migratory pathways and possesses longitudinal bill length variation. Previous investigations suggested that genetic structure may occur within Semipalmated Sandpipers and that three subspecies corresponding to western, central, and eastern breeding groups exist. In this study, mitochondrial control region sequences and nuclear microsatellite loci were used to analyze DNA of birds (microsatellites: n = 120; mtDNA: n = 114) sampled from seven North American locations. Analyses designed to quantify genetic structure and diversity patterns, evaluate genetic evidence for population size changes, and determine if genetic data support the existence of Semipalmated Sandpiper subspecies were performed. Genetic structure based only on the mtDNA data was observed, whereas the microsatellite loci provided no evidence of genetic differ...

Flooding can cause widespread nest failure and chick mortality in sandbar-and beach-nesting waterbirds, particularly when human activity has either altered natural hydrology or limited available nesting habitat. Such widespread reproductive failure could increase breeding-season dispersal, leading to the abandonment of established nesting sites. We examined how annual movement ψ varied by sex, reproductive success, and flooding in three Piping Plover (Charadrius melodus) breeding areas in Saskatchewan during 2002–2009, using a multistate capture-mark-recapture model in Program MARK (n = 782). On average, female Piping Plovers were twice as likely as males to disperse and both sexes were more likely to disperse following years of poor versus moderate reproductive success (minimum to maximum values: ψno-fledglings, 0.0054–0.0998 vs. ψtwo-fledglings, 0.0017–0.0607). In addition, breeding Piping Plovers exhibited higher dispersal following flood years, even in years of moderate reproductive success ...

The Semipalmated Sandpiper (Calidris pusilla) is a small, abundant shorebird that breeds primarily in sub-Arctic to mid-Arctic habitats across the Nearctic and winters principally along the northern and central coasts of South America. No subspecies have been described and little is known concerning their genetics. However, birds show a cline in bill length across the Arctic, with longest bills in the east and shortest in the west. There appear to be several ‘steps’ in the cline, suggesting a division into eastern, central and western breeding populations. Since females average longer bills than males in a breeding population, there is considerable overlap of bill lengths at migration staging areas. Based on bill length patterns and sightings and recoveries of marked individuals, most western breeders migrate south through the prairies, along with some birds from central Arctic populations. The remaining central Arctic breeders, and all eastern Arctic birds, migrate south through the north Atlan...

Counts of Semipalmated Sandpiper (Calidris pusilla) at some migratory stopover sites have shown pronounced declines over the last 35 years. Counts from breeding grounds avoid sources of bias that have proven troublesome for trends estimated from migration surveys. Published and unpublished data were reviewed to examine trends in densities of Semipalmated Sandpipers at breeding sites across Alaska and arctic Canada. Information was sparse, and some comparisons are tenuous because methods varied over time. Valid time series were obtained for 13 sites across the species' range. In Alaska, Semipalmated Sandpipers either increased or were at least stable at six sites and decreased at one site. Surveys at both sites in the central portion of the range suggested no change in abundance. In the eastern portion of the range, trends were variable: decreases were observed at two sites, a possible increase at one site, and no change at another. Thus, the species was generally increasing or stable in the west...

To conserve threatened species, managers require predictions about the effects of natural and anthropogenic factors on population growth that in turn require accurate estimates of survival, birth, and dispersal rates, and their correlation with natural and anthropogenic factors. For Piping Plovers (Charadrius melodus), fledging rate is often more amenable to management than adult survival, and population models can be used to estimate the productivity (young produced per breeding female) necessary to maintain or increase populations for given levels of survival. We estimated true survival and site fidelity of adult and subadult (from fledging to second year) Piping Plovers breeding in Saskatchewan using mark-resight data from 2002 to 2009. By estimating true survival rather than apparent survival (which is confounded with permanent emigration), we were able to provide more accurate projections of population trends. Average adult and subadult survival rates during our study were 0.80 and 0.57, respectively. Adult survival declined over time, possibly due in part to the loss of one breeding site to flooding. Average adult and subadult site fidelity were 0.86 and 0.46, respectively. Adult site fidelity declined during our study at two study sites, most strongly at the flooded site. Male and female Piping Plovers had similar survival rates, but males had greater site fidelity than females in some years. Based on our survival estimates, productivity needed for a stationary population was 0.75, a benchmark used for plover management on the Atlantic Coast, but not previously estimated for Prairie Canada. In stochastic simulations incorporating literature-based variation in survival rates, productivity needed for a stationary population increased to 0.86, still lower than that previously estimated for western populations. Mean productivity for our study sites ranged from 0.87 to 0.96 fledged young per pair. Our results suggest that fledging rates of Piping Plovers in Saskatchewan were sufficient to ensure a stationary or increasing population during our study period. However, large-scale habitat changes such as drought or anthropogenic flooding may lead to dispersal of breeding adults and possibly mortality that will increase the fledging rate needed for a stationary population. RESUMEN Para conservar especies amenazadas, se necesitan predicciones de los efectos de factores naturales y antropogenicos sobre el crecimiento de la poblacion, que a su vez requieren de estimaciones precisas de las tasas de supervivencia, reproduccion y la dispersion y su correlacion con factores naturales y antropogenicos. Para Charadrius melodus, la tasa de produccion de volantones es a menudo mas controlable que la supervivencia de adultos, y los modelos de poblacion pueden utilizarse para estimar la productividad (crias producidos por hembra reproductora) necesaria para mantener o aumentar las poblaciones, para un dado nivel de supervivencia. Estimamos la supervivencia real y la fidelidad al sitio de los adultos y subadultos (de la edad de volanton hasta su segundo ano) de C. melodus en Saskatchewan usando datos de marcaje y re-observacion de 2002 a 2009. Mediante la estimacion de la supervivencia real en vez de la supervivencia aparente (que se confunde con la emigracion permanente), pudimos hacer proyecciones mas precisas de las tendencias poblacionales. El promedio de las tasas de supervivencia de adultos y subadultos durante nuestro estudio fue de 0.80 y 0.57, respectivamente. La supervivencia de adultos se redujo con el tiempo, posiblemente debido en parte a la perdida de un lugar de reproduccion por inundacion. El promedio de la fidelidad al sitio de adultos y subadultos fue de 0.86 y 0.46, respectivamente. La fidelidad al sitio de los adultos disminuyo durante el estudio en dos sitios de estudio, con la mayor fuerza en el lugar inundado. Machos y hembras de C. melodus tuvieron tasas similares de supervivencia, pero los machos tuvieron una mayor fidelidad al lugar que las hembras en algunos anos. En base a nuestras estimaciones de supervivencia, la productividad necesaria para una poblacion estacionaria fue de 0.75, un punto de referencia utilizado para la gestion de C. melodus en la Costa Atlantica, pero que no se habia estimado anteriormente para las praderas de Canada. En las simulaciones estocasticas que incorporaron la variacion de las tasas de supervivencia basadas en la literatura, la productividad necesaria para mantener una poblacion estacionaria subio a 0.86, siendo todavia inferior a la previamente estimada para las poblaciones occidentales. El promedio de la productividad en nuestros sitios de estudio vario desde 0.87 hasta 0.96 volantones por pareja. Nuestros resultados sugieren que las tasas de produccion de volantones de C. melodus en Saskatchewan eran suficientes para asegurar una poblacion estacionaria o en aumento durante el periodo de estudio. Sin embargo, cambios a grande escala en el habitat, como la sequia o las inundaciones antropogenicas, pueden conducir a la dispersion de los adultos reproductores y posiblemente a mortalidad que requiere de una mayor tasa de produccion de volantones necesario para una poblacion estacionaria.

Resightings of uniquely marked birds from 2001 to 2008 were used to determine winter distributions of 4 breeding populations of a species at risk, the piping plover (Charadrius melodus). Although considerable overlap exists, a distinct pattern in winter distributions was evident. Birds originating from eastern Canada wintered farthest north compared to other populations. Most individuals from the United States Great Lakes were found in South Carolina and Georgia in winter, whereas birds from eastern Canada were found primarily in North Carolina. Although the great majority of birds marked in Prairie Canada were observed wintering in Texas, particularly southern Texas, breeding plovers from the United States Great Plains were more widely distributed on the gulf coast from Florida to Texas. Very few large-scale movements of individuals in winter were reported either within or between years. This study highlights the significance of geographic regions for eastern Canada, the United States Great Lakes, the United States Great Plains, and Prairie Canada populations, and demonstrates relatively high winter site fidelity. This information will help focus conservation efforts for specific breeding populations during the winter. © 2011 The Wildlife Society.

Age of first breeding is an important life history trait. Many Pipine Plovers (Charadrius melodus) do not breed as yearlings, but little information is available concerning the age of first breeding. From 2000 to 2006, we marked 991 chicks in three areas in Saskatchewan, Canada, and subsequently determined when 102 (49 females and 53 males) first bred. Females bred significantly earlier, on average, than males. More females (68%) bred as yearlings than did males (41%; P = 0.04), with most others first nesting when 2-yr old (29% of females and 50% of males). As expected from differences between the sexes in age of first breeding, younger females were more likely to pair with older males than were younger males with older females. Chicks that hatched early in the breeding season did not breed at an earlier age than those that hatched late in the year. Unlike older birds, juvenile Piping Plovers do not replace flight feathers during their first winter. As a result, 18 of 27 yearlings (67%) had worn outer primaries, whereas only one of 123 (1%) older birds had worn primaries. In addition, whereas 20 of 24 yearlings (83%) retained a few buff-tipped median coverts, none of 1 19 known older birds had such coverts. As a result, we were able to identify all yearlings by their worn primaries, buff-tipped median wing coverts, or both. Wing lengths of yearling Piping Plovers were 3% shorter than those of older birds, presumably due to wear. Because there is no evidence of differences in adult survival rates between the sexes and breeding habitat is available, we speculate that fewer yearling males than females breed because primary wear may reduce the ability of yearling males to perform aerial breeding displays.

Geographically isolated breeding populations of migratory shorebirds may be demographically connected through shared nonbreeding habitats. We used long-term (1998-2008) mark-recapture data on piping plovers (Charadrius melodus) collected from 7 separate studies located throughout North America to conduct a range-wide analysis of after hatch year apparent survival (WAHY). Our objectives were to compare concurrent survival estimates from disparate breeding sites and determine whether estimates followed similar trends or were correlated among breeding populations with shared wintering grounds. Average survival estimates were higher for Great Plains populations (range 5 0.69-0.81) than for Great Lakes and Atlantic Coast populations (range 5 0.56-0.71). Linear trend models indicated that apparent survival declined in 4 out of 7 populations, was unchanged in 3, and was generally highest among Great Plains populations. Based on a post hoc analysis, we found evidence of correlated year-to-year fluctuations in annual survival among populations wintering primarily along the southeastern United States Atlantic Coast and Gulf Coast. Our results indicate shared overwintering or stopover sites may influence annual variation in survival among geographically disparate breeding populations. Declines in piping plover survival are a cause for concern, and our results highlight the need for conservation efforts to include habitat used during the migratory and wintering periods.

Piping Plover (Charadrius melodus) is a migratory shorebird that is listed as endangered in Canada and the U.S. Great Lakes and as threatened throughout the rest of its breeding and winter range. We undertook a comprehensive molecular-genetic investigation to (1) address subspecific taxonomy, (2) characterize population genetic structure, and (3) infer past bottlenecks and demographic processes in this species. Analyses included individuals from 23 U.S. states and Canadian provinces and were based on mitochondrial DNA sequences (580 base pairs, n = 245) and 8 nuclear microsatellite loci (n = 229). Our findings provide support for separate Atlantic and Interior subspecies (C. m. melodus and C. m. circumcinctus, respectively). Birds from the Great Lakes region were allied with the Interior subspecies and should be referred to as C. m. circumcinctus. Population genetic analyses illustrated stronger genetic structure among Atlantic than among Interior birds, which may reflect reduced natal- and bree...

A hierarchical modelling approach was used to examine adult and age-specific survival in an 8-year study of breeding Semipalmated Sandpipers Calidris pusilla at La Perouse Bay, Canada. The survival of adult sandpipers was best described by a model with time dependence in local survival rate and probability of recapture. Annual variation in the local survival rate of adults was not correlated with nest success, timing of breeding or “return rates” and was not biased by an effect of first capture. Local survival rate of adult females (0.56, 95% c.l. = 0.51-0.61) was consistently lower than that of adult males (0.61, 95% c.l. = 0.56-0.66); these estimates were comparable with data from other shorebirds. The survival of returning young was best fitted by a model with both age and time dependence in local survival rate and probability of recapture. We evaluated our estimates of local survival rate with reference to patterns of breeding fidelity and philopatry in Semipalmated Sandpipers and other shorebirds.

Although Semipalmated Sandpipers Calidris pusilla are monogamous, with biparental incubation, most females (86–97%) deserted their broods to the care of their mates, 0–11 days (average 6) after their eggs hatched. Males left the brood an average of 8 days later, shortly before or after the chicks fledged. In several instances, females that deserted in one year remained with the chicks the next year, and vice versa. Females deserted chicks at nests that hatched later in the season at an earlier age than those had hatched earlier (r2> o.6). Since females appeared to have an energy deficit in at least some years, and suffered higher mortality rates than males during breeding, it is possible that females deserted broods in order to take advantage of better feeding conditions at migratory stopovers in northeastern North America early in the season. There was little evidence of higher nesting success or earlier hatching date in reuniting pairs, although if both members of a pair returned to the breeding area, 80% reunited. Increased survival of their mate may be most advantageous to males in ensuring that they obtain a female the following year.

We measured fresh body mass, total body fat, and fat-free dry mass (FFDM) of three species of Arctic-nesting calidrid sandpipers (Baird's Sandpiper [Calidris bairdii], hereafter “BASA”; Semipalmated Sandpiper [C. pusilla], hereafter “SESA”; and White-rumped Sandpiper [C. fuscicollis], hereafter “WRSA”) during spring stopovers in the Prairie Pothole Region (PPR) of North Dakota, and evaluated the contribution of stored fat to (1) energy requirements for migration to their Arctic-breeding grounds and (2) nutrient needs for reproduction. All spring migrant WRSA (n = 124) and BASA (n = 111), and all but 2 of 99 SESA we collected were ≥2 years old. Male and female BASA migrated through North Dakota concurrently, male SESA averaged earlier than females, and WRSA males preceded females. Fat indices (ratio of fat to FFDM) of male and female SESA and WRSA averaged approximately twice those of male and female BASA. Total body fat of male and female BASA increased with date in spring 1980, but not in 1981; slopes were similar for both sexes each year. Male and female SESA arrived lean in 1980 and 1981, and total body fat increased with date in both years, with similar slopes for all combinations of sex and year. Male and female WRSA arrived lean in 1980–1981 and 1981, respectively, and total body fat increased with date, whereas females arrived with fat reserves already acquired in 1980. Interspecific and sex differences in migration schedules probably contributed to variation in fat storage patterns by affecting maintenance energy costs and food availability. Estimated flight ranges of BASA suggest that few could have met their energy needs for migration to the breeding grounds exclusively from fat stored by the time of departure from North Dakota. Estimated flight ranges of SESA and WRSA, along with fresh body masses of both species when live-trapped on or near their breeding grounds in northern Canada, suggest that major parts of both populations stored adequate fat by departure from temperate mid-continental North America to meet their energy requirements for migration and part of their nutrient needs for reproduction. Dinámica de la Grasa en Chorlos que Nidifican en el Ártico durante la Primavera en el Área Continental Central de América del Norte

We surveyed the shoreline of a large terminal lake in southern Saskatchewan (Old Wives Lake) for sick or dead birds and estimated shorebird numbers during avian botulism outbreaks. Nearly double the number of shorebirds were present in 1999 compared with 1998. The most commonly observed shorebirds in both years were the Short-billed Dowitcher (Limnodromus griseus) and Long-billed Dowitcher (L. scolopaceus). Water levels were monitored, revealing a greater drawdown effect in 1998. Small species (e.g. Least Sandpiper (Calidris minutilla), Semipalmated Sandpiper (C. pusilla), and Sanderling (C. alba)) were much more common in 1998. Much of the data that exist on shorebird mortality during botulism kills is in the form of unpublished annual agency clean-up reports. These reports suggest that the American Avocet (Recurvirostra americana), yellowlegs (Tringa melanoleuca and T. flavipes), and the small Calidris sandpipers are the most commonly affected species. We suggest some biases inherent in these c...

Identification of relationships among geographically distinct populations of migratory species can provide an understanding of breeding and natal philopatry, migration pathways, and population mixing during winter. We used random amplified polymorphic DNA (RAPD) analyses to search for markers specific to difficult-to-differentiate shorebird species (e.g. long-billed dowitcher Limnodromus scolopaceus and short-billed dowitcher L. griseus) as well as geographically distinct breeding populations of Hudsonian godwits Limosa haemastica, red-necked phalaropes Phalaropus lobatus, semipalmated plovers Charadrius semipalmatus, dunlin Calidris alpina, pectoral sandpipers C. melanotos, semipalmated sandpipers C. pusilla and western sandpipers C. mauri. Markers clearly differentiated all shorebird species. Estimates of population differentiation varied greatly among species (FST = 0.095‐0.685) and correlated with interspecific variation in philopatry and geographical separation of breeding populations. We assigned individuals to putative breeding locales with greater certainty in welldifferentiated species than in poorly differentiated species. Our findings indicate specific phylogeographical structure varies among species, which has strong implications for conservation of habitats within migratory corridors. We suggest that RAPDs are useful in identifying geographical populations of migratory species and that molecular markers should be considered for tracking migratory birds throughout the annual cycle.

The breeding population of Semipalmated Sandpipers at La Perouse Bay, Churchill, Manitoba declined sharply during the 1980s. Using demographic information collected during eight summers of field work we built a stochastic matrix population model. The model successfully reconstructed the observed decline in the population during the course of the study. The predictions of the model agreed with the observed population level in 1993, 5 yr after the study was completed. Sensitivity analysis showed that the population trend was most sensitive to changes in adult survivorship and to immigration of adults onto the site. We ran simulations to consider five possible reasons for the decline: (1) low adult survivorship, (2) low fecundity, (3) delayed recruitment into the population, (4) low juvenile survivorship, and (5) insufficient immigration. For biologically reasonable values of these parameters, only adult survivorship and immigration rates could account for the observed decline, although juvenile survival also had some impact. These simulations also illustrate that sensitivity is a complex function of the demographic variables, and in particular, sensitivity to adult survival varies across the range of adult survival values that we considered. In addition, we used the model to explore the influence of annual variability in demographic parameters. It is known that the general effect of annual variation is to depress population trends. We showed that the sensitivity of the model to annual variability in survival is tied to the sensitivity of the model to changes in mean values of survival. With the empirical parameter values from this local population, we found that, when survival is high enough to promote stability of the population, the population is highly sensitive to the level of survival, and also to the amount of annual variability in survival. By contrast, the model is not very sensitive to fecundity and is correspondingly insensitive to annual variation in fecundity.

We used gut-content and stable-isotope techniques to determine diets of shorebirds staging at a prairie wetland complex. Stable-isotope ratios for carbon (13C/12C) and nitrogen (15N/14N) varied greatly within and among prey types and shorebirds, depending on location within the complex. Both dietary techniques suggested that Long-billed Dowitchers (Limnodromus scolopaceus) and Stilt Sandpipers (Calidris himantopus) ate mostly invertebrates, whereas Hudsonian (Limosa haemastica) and Marbled godwits (Limosa fedoa) ate mainly Potamogeton pectinatus tubers. In comparison, the stable-isotope technique indicated that godwits, especially juvenile Marbled Godwits, ate more invertebrates than is indicated by the gut-content analysis. The discrepancies between methods reflect the potential for bias in the application of these techniques. Researchers using stable isotopes to assess migratory shorebird diets should be aware of possible complications arising from isotopic variability within prey types, even over small geographic ranges. High isotopic variability at inland agro-wetland complexes might preclude reliable isotopic assessment of shorebird diets, especially long term. Rhizivory in godwits may be more common than is generally recognized, especially at inland stopovers during autumn migration in both North America and Eurasia, and should be factored into conservation initiatives for these species.

A female Semipalmated Sandpiper (Calidris pusilla) at least 16 years of age was observed at a nest near Churchill, Manitoba, which increases the previous longevity record for this species and ties that for the oldest small North American sandpiper. Mean annual adult survival rates since 1993 for Semipalmated Sandpipers in this area were not lower than those for previous years, despite considerable recent habitat destruction by Snow Geese (Chen caerulescens) and a continued decline in numbers of nesting sandpipers.

We examined the relationship between circulating levels of gonadal steroids (testosterone, progesterone, and estradiol) and breeding behavior in semipalmated sandpipers (Calidris pusilla) and red-necked phalaropes (Phalaropus lobatus) breeding sympatrically at La Pérouse Bay, 40 km east of Churchill, Manitoba. Semipalmated sandpipers are territorial and monogamous. Both parents incubate equally. Red-necked phalaropes are nonterritorial and polyandrous. Only male phalaropes care for eggs and young. Gonadal steroid hormone profiles were not reversed in the sex-role-reversed species. There was little difference in testosterone profiles between males of the territorial and nonterritorial species.