Differentiation of subspecies and sexes of Beringian Dunlins using morphometric measures

Five subspecies of Dunlins (Calidris alpina) that breed in Beringia are potentially sympatric during the non‐breeding season. Studying their ecology during this period requires techniques to distinguish individuals by subspecies. Our objectives were to determine (1) if five morphometric measures (body mass, culmen, head, tarsus, and wing chord) differed between sexes and among subspecies (C. a. actites, arcticola, kistchinski, pacifica, and sakhalina), and (2) if these differences were sufficient to allow for correct classification of individuals using equations derived from discriminant function analyses. We conducted analyses using morphometric data from 10 Dunlin populations breeding in northern Russia and Alaska, USA. Univariate tests revealed significant differences between sexes in most morphometric traits of all subspecies, and discriminant function equations predicted the sex of individuals with an accuracy of 83–100% for each subspecies. We provide equations to determine sex and subspecies of individuals in mixed subspecies groups, including the (1) Western Alaska group of arcticola and pacifica (known to stage together in western Alaska) and (2) East Asia group of arcticola, actites, kistchinski, and sakhalina (known to winter together in East Asia). Equations that predict the sex of individuals in mixed groups had classification accuracies between 75% and 87%, yielding reliable classification equations. We also provide equations that predict the subspecies of individuals with an accuracy of 22–96% for different mixed subspecies groups. When the sex of individuals can be predetermined, the accuracy of these equations is increased substantially. Investigators are cautioned to consider limitations due to age and feather wear when using these equations during the non‐breeding season. These equations will allow determination of sexual and subspecies segregation in non‐breeding areas, allowing implementation of taxonomic‐specific conservation actions.     

Nest covering in plovers: How modifying the visual environment influences egg camouflage

Camouflage is one of the most widespread antipredator defences, and its mechanistic basis has attracted considerable interest in recent years. The effectiveness of camouflage depends on the interaction between an animal's appearance and its background. Concealment can therefore be improved by changes to an animal's own appearance, by behaviorally selecting an optimal background, or by modifying the background to better match the animal's own appearance. Research to date has largely focussed on the first of these mechanisms, whereas there has been little work on the second and almost none on the third. Even though a number of animal species may potentially modify their environment to improve individual‐specific camouflage, this has rarely if ever been quantitatively investigated, or its adaptive value tested. Kittlitz's plovers (Charadrius pecuarius) use material (stones and vegetation) to cover their nests when predators approach, providing concealment that is independent of the inflexible appearance of the adult or eggs, and that can be adjusted to suit the local surrounding background. We used digital imaging and predator vision modeling to investigate the camouflage properties of covered nests, and whether their camouflage affected their survival. The plovers' nest‐covering materials were consistent with a trade‐off between selecting materials that matched the color of the eggs, while resulting in poorer nest pattern and contrast matching to the nest surroundings. Alternatively, the systematic use of materials with high‐contrast and small‐pattern grain sizes could reflect a deliberate disruptive coloration strategy, whereby high‐contrast material breaks up the telltale outline of the clutch. No camouflage variables predicted nest survival. Our study highlights the potential for camouflage to be enhanced by background modification. This provides a flexible system for modifying an animal's conspicuousness, to which the main limitation may be the available materials rather than the animal's appearance.     

Contrasting genetic diversity and population structure among three sympatric Madagascan shorebirds: parallels with rarity,endemism, and dispersal

Understanding the relative contributions of intrinsic and extrinsic factors to population structure and genetic diversity is a central goal of conservation and evolutionary genetics. One way to achieve this is through comparative population genetic analysis of sympatric sister taxa, which allows evaluation of intrinsic factors such as population demography and life history while controlling for phylogenetic relatedness and geography. We used ten conserved microsatellites to explore the population structure and genetic diversity of three sympatric and closely related plover species in southwestern Madagascar: Kittlitz's plover (Charadrius pecuarius), white‐fronted plover (C. marginatus), and Madagascar plover (C. thoracicus). Bayesian clustering revealed strong population structure in the rare and endemic Madagascar plover, intermediate population structure in the white‐fronted plover, and no detectable population structure in the geographically widespread Kittlitz's plover. In contrast, allelic richness and heterozygosity were highest for the Kittlitz's plover, intermediate for the white‐fronted plover and lowest for the Madagascar plover. No evidence was found in support of the “watershed mechanism” proposed to facilitate vicariant divergence of Madagascan lemurs and reptiles, which we attribute to the vagility of birds. However, we found a significant pattern of genetic isolation by distance among populations of the Madagascar plover, but not for the other two species. These findings suggest that interspecific variation in rarity, endemism, and dispersal propensity may influence genetic structure and diversity, even in highly vagile species.     

Colored plastic and metal leg bands do not affect survival of Piping Plover chicks

ABSTRACT Leg bands are commonly used to mark shorebird chicks as young as 1‐d old, but little is known about the possible impacts of bands on survival of prefledging shorebirds. We used a mark‐recapture framework to assess the impact of bands and banding‐related disturbance on prefledging survival in a federally endangered population of Piping Plovers (Charadrius melodus) breeding in the Great Lakes region from 2000 to 2008. We banded approximately 96% of all surviving chicks hatched prior to fledging, typically between 5 and 15 d of age. We used a multistate approach in program MARK whereby individuals contributed data as unbanded chicks before capture (N= 1073) and as banded chicks afterward (N= 780). The cumulative probability of surviving through 24 d of age was 0.63 and did not differ between banded and unbanded chicks. In addition, we found a positive effect of banding‐related disturbance on survival up to 3 d following banding (β= 0.60 CI: 0.17–1.02), possibly due to increased postbanding vigilance on the part of chicks and adults. Our results indicate that banding has no detrimental effect on survival of Piping Plover chicks prior to fledging and that current capture and banding methods are appropriate for this endangered species.     

Shorebird predation of horseshoe crab eggs in Delaware Bay: species contrasts and availability constraints

1. Functional responses -- the relationship between resource intake rate and resource abundance -- are widely used in explaining predator-prey interactions yet many studies indicate that resource availability is crucial in dictating intake rates. 2. For time-stressed migrant birds refuelling at passage sites, correct decisions concerning patch use are crucial as they determine fattening rates and an individual's future survival and reproduction. Measuring availability alongside abundance is essential if spatial and temporal patterns of foraging are to be explained. 3. A suite of shorebird species stage in Delaware Bay where they consume horseshoe crab Limulus polyphemus eggs. Several factors including spawning activity and weather give rise to marked spatial and temporal variation in the abundance and availability of eggs. We undertook field experiments to determine and contrast the intake rates of shorebird species pecking for surface and probing for buried eggs. 4. Whether eggs were presented on the sand surface or buried, we demonstrate strong aggregative responses and rapid depletion (up to 80%). Depletion was greater at deeper depths when more eggs were present. No consistent give-up densities were found. Type II functional responses were found for surface eggs and buried eggs, with peck success twice as high in the former. Maximum intake rates of surface eggs were up to 83% higher than those of buried eggs. 5. Caution is needed when applying functional responses predicted on the basis of morphology. Our expectation of a positive relationship between body size and intake rate was not fully supported. The smallest species, semipalmated sandpiper, had the lowest intake rate but the largest species, red knot, achieved only the same intake rate as the mid-sized dunlin. 6. These functional responses indicate that probing is rarely more profitable than pecking. Currently, few beaches provide egg densities sufficient for efficient probing. Areas where eggs are deposited on the sand surface are critical for successful foraging and ongoing migration. This may be especially true for red knot, which have higher energetic demands owing to their larger body size yet appear to have depressed intake rates because they consume smaller prey than their body size should permit.     

Habituation and habitat changes can moderate the impacts of human disturbance on shorebird breeding performance

Disturbance by humans is widely expected to reduce the reproductive fitness of nesting birds if disturbance reduces nest attentiveness, and unattended eggs experience increased risk of predation or exposure to potentially lethal temperature extremes. Yet, relatively few studies have examined the physiological or behavioural mechanisms whereby disturbance influences reproductive fitness, or the extent to which the costs of disturbance may be reduced through habituation. We compared the behavioural responses, egg temperatures and reproductive success of shore-nesting white-fronted plovers Charadrius marginatus to disturbance at two breeding sites experiencing low versus high human recreational activity, respectively. Daytime nest attentiveness decreased with increasing experimental disturbance at both sites, but this relationship differed between sites; for any given level of disturbance, incubating birds at the more disturbed site had greater nest attentiveness. They achieved this through habituation, allowing a closer human approach before leaving the nest, and returning to the nest faster after a disturbance event. Despite lower average daytime nest attentiveness at the more disturbed site, incubation temperatures did not differ significantly between sites. Nest mortality, mostly by natural mammalian and corvid predators, was significantly lower at the site experiencing high recreational activity. However, chick mortality was significantly greater at the more disturbed site, most likely because of predation by domestic dogs. Chick mortality may have been increased by the habitation of chicks, whose escape responses were much reduced at the more disturbed site. Nonetheless, annual fecundity was substantially higher at the more disturbed site, showing that the overall reproductive fitness of wild birds is not always compromised by human disturbance and urbanization.     

Determinants of preferred intertidal feeding habitat for Eastern Curlew: A study at two spatial scales

Abstract Broadscale habitat use by Eastern Curlews (Numenius madagascariensis) in their non‐breeding range in eastern Australia was assessed using low tide surveys on feeding grounds, where 60 skilled volunteers made repeated counts of the birds on intertidal flats, across 41% (9500 ha) of the intertidal habitat within Moreton Bay, Australia. We analysed 32 defined sections of intertidal flat, of roughly equal area (mostly 200–400 ha), which varied greatly in their curlew density (2–47 birds per 100 ha) and also in substrate and other environmental features. Sites with the least resistant substrates had densities three times those with the most resistant substrates. Of 10 environmental characteristics measured for each site, substrate resistance was the best predictor of curlew density (r² = 0.45). Characteristics that were poor predictors included distance to the nearest roost, level of human disturbance and distance to urban settlement. For a finer‐scale assessment, microhabitat use and feeding behaviour were recorded during low tide within 12 intertidal flats, which varied in size (23–97 ha), substrate, topography and other features. Across all flats, curlews strongly preferred to feed relatively close (0–50 m) to the low‐water line. They fed on a variety of substrates (including sand, sandy‐mud, mud and seagrass) in broadly similar proportions to their occurrence in the habitat. There was a statistically significant preference for sand, although its magnitude was not strong. These results indicate that curlews select habitat most strongly at a between‐flat rather than within‐flat scale.     

Effects of water discharge on fledging time,growth, and survival of piping plovers on the Missouri River

River flow management and modification is a global issue, and its effects on river-dependent organisms are pervasive. Flow modification can directly affect avian species through mortality or habitat loss, but less is known about indirect and sublethal effects of flow modification on reproductive output in these species. Young birds are more vulnerable to predation between hatching and fledging than after flight is achieved, but tradeoffs must be made to balance growth and survival. Predation pressure appears to be a significant factor affecting the time to fledging in altricial birds, but less is known about this threat for precocial birds. Birds reaching fledging earlier should have greater rates of survival to migration because their predator escape repertoire includes flight at an earlier age. We evaluated the effect of varying outflows from the Gavins Point Dam on the growth, age at fledging, and survival of piping plover (Charadrius melodus) chicks on the Missouri River (2006–2009). The study was characterized by 2 relatively high flow years (2006 and 2009) and 2 relatively low flow years (2007 and 2008). We used success rate in recapturing chicks in capture–mark–recapture models as an index for fledging. We attempted to recapture all chicks (n = 1,099) by hand every 3–4 days throughout the season to acquire morphological measurements. Models indicated that as flows from the dam increased, age at fledging increased. We also found that increasing flows were associated with decreasing daily survival rates (β_flow = −2.401, 95% CI: −4.351 to −0.452). Flow was also negatively related to chick mass gain, but we found less evidence for an effect on wing-chord length. Increased flows covered wet-substrate foraging habitat, and likely affected plover reproductive output directly through chick survival and indirectly through decreased growth and increased fledging times. © The Wildlife Society, 2013     

Why is timing of bird migration advancing when individuals are not?

Recent advances in spring arrival dates have been reported in many migratory species but the mechanism driving these advances is unknown. As population declines are most widely reported in species that are not advancing migration, there is an urgent need to identify the mechanisms facilitating and constraining these advances. Individual plasticity in timing of migration in response to changing climatic conditions is commonly proposed to drive these advances but plasticity in individual migratory timings is rarely observed. For a shorebird population that has significantly advanced migration in recent decades, we show that individual arrival dates are highly consistent between years, but that the arrival dates of new recruits to the population are significantly earlier now than in previous years. Several mechanisms could drive advances in recruit arrival, none of which require individual plasticity or rapid evolution of migration timings. In particular, advances in nest-laying dates could result in advanced recruit arrival, if benefits of early hatching facilitate early subsequent spring migration. This mechanism could also explain why arrival dates of short-distance migrants, which generally return to breeding sites earlier and have greater scope for advance laying, are advancing more rapidly than long-distance migrants.