Extensive Use of Intertidal Habitat by Shorebirds Outside Protected Nesting Areas

Conservation of shorebirds throughout their breeding and migratory ranges has become a priority as shorebird populations decline globally. Along the North Atlantic Coast, management efforts have particularly focused on preserving nesting habitat for piping plovers (Charadrius melodus), which are protected under the Endangered Species Act. It is unclear whether these conservation measures suffice to protect foraging habitat for piping plovers and other shorebirds on stopover during migration along the Atlantic Flyway. To evaluate the extent to which conservation of piping plover nesting areas extends to all habitats used by plovers, and to determine whether these protections also benefited non-breeding migratory shorebirds in the region, we conducted weekly shorebird surveys, recording the number and locations of piping plovers and other species, during northward and southward migration on Fire Island and Westhampton Island, New York, USA, from 2014–2016. We used canonical correspondence analysis (CCA) to assess the degree of spatiotemporal overlap between breeding plovers, foraging plovers, and other migratory shorebirds that temporarily stage at the site. The spatiotemporal distribution of migratory shorebirds matched more closely with piping plovers seen during foraging than piping plovers observed tending nests and engaging in other breeding activities. Migratory shorebirds and foraging piping plovers were more abundant and frequent in wet intertidal zones outside of fenced-off nesting areas, which were not protected under current management regimes. Therefore, additional protection of piping plover foraging habitat could benefit plovers and migratory shorebirds that use similar feeding grounds during stopover on northward and southward migration. © 2020 The Wildlife Society.     

Red fox trap success is correlated with piping plover chick survival

Predation management is an important component of managing species of conservation concern. The piping plover (Charadrius melodus; plover), a disturbance-dependent and conservation-reliant shorebird that nests on sandy beaches and barrier islands on the Atlantic Coast, was listed under the United States Endangered Species Act in 1986, with habitat loss and predation stated as key causes of its decline. We evaluated the relationship between a suite of predators and plover chick survival from 2015–2018. We used a camera grid to establish indices of the abundance of 3 known chick predators: red fox (Vulpes vulpes), raccoon (Procyon lotor), and domestic cat. We used camera detections in a survival model to assess potential relationships between predator species detection and plover chick survival. Plover chick survival was negatively related with red fox detection, but not with detection of the other 2 predators. In addition to the correlation with red fox detections, chick survival was negatively related to high plover nesting density. Our results suggest that red foxes were predators of piping plover chicks during our study, likely augmented by other density-dependent sources of mortality. Targeted predator management could aid in conservation of piping plovers in this system as a short-term solution, but long-term recovery plans must also address habitat limitation.     

Range-Wide Piping Plover Survival: Correlated Patterns and Temporal Declines

ABSTRACT 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 (φ_AHY). 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 = 0.69–0.81) than for Great Lakes and Atlantic Coast populations (range = 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 chick ecology following landscape-level disturbance

Population declines of disturbance-dependent species due to suppression of natural disturbances are realized across ecosystems. The piping plover (Charadrius melodus; plover), a disturbance-dependent and conservation-reliant shorebird that nests on sandy beaches and barrier islands on the Atlantic Coast, was listed under the United States Endangered Species Act in 1986. In 2012, Hurricane Sandy landed on Fire and Westhampton islands, barrier island nesting sites for plovers in New York, USA. Hurricane Sandy was a natural disturbance in this system, creating abundant nesting habitat. The number of chicks produced by a pair, or a population, is a direct measure of reproductive output, and gaining a better understanding of productivity and chick behavior following large-scale habitat creation may improve plover habitat management and potentially species persistence. We evaluated the effects of landscape features on habitat selection, behavior, and survival of plover broods using logistic regression, generalized linear mixed effects models, and survival models. Plover broods selected flatter sites with less dense vegetation than available at random. Chick foraging rates were highest in moist substrates and were lower in areas of higher nesting plover density. Chick survival was greater for broods that hatched earlier in the breeding season and increased as chicks aged. Generally, providing access to sites with flatter, moist substrates will likely result in higher quality brood rearing habitat on the landscape. Ultimately, vegetation removal and habitat management may be needed to reduce plover nesting density and ensure sufficient habitat, which may in turn improve plover chick survival. Moreover, sustaining natural landscape disturbances such as those resulting from storms, and not taking actions to prevent hurricane-created overwash, will allow these landscape features to persist.     

Captive-rearing piping plovers: Developing techniques to augment wild populations

Techniques for captive-rearing and releasing piping plovers (Charadrius melodus) were developed using a surrogate species, killdeer (Charadrius vociferus). We compared captive- and parent-reared killdeer, and parent-reared piping plovers and determined that growth and behavior were similar. After surrogate trials determined that captive-rearing was feasible, we used the same methods to raise piping plover chicks from salvaged eggs. For captive-reared chick of both species, survival to fledging was higher than and behaviors similar to parent-reared chicks in the wild. Rearing techniques were fine-tuned, and ten piping plover fledglings were released to the wild. Based on our results, we developed recommendations for captive-rearing piping plovers using salvaged eggs to enhance productivity of small populations. Zoo Biol 16:461–477, 1997. © 1997 Wiley-Liss, Inc.     

Factors affecting piping plover hatching success on Long Island,New York

Low hatching success may limit progress towards reaching productivity goals for Atlantic Coast piping plover (Charadrius melodus) recovery, despite management strategies to protect eggs from predators and decrease human disturbance of birds on nests. We measured piping plover hatching success on Eastern Long Island beaches and identified the major causes of egg failure to better understand why eggs that were otherwise intact (not depredated or destroyed by tidal flooding) failed to hatch. We documented egg and nest fates, dissected contents of unhatched eggs to determine viability, and recorded human and predator activity near a subset of plover nests on Suffolk County Parks properties. The low hatching success we recorded (0.60) in 2006 and 2007 would require higher chick survival rates than are typically observed for piping plovers to meet recovery targets for productivity. Few eggs showed signs of poor viability and overall egg hatchability was comparable to other ground nesting birds. Most egg failure was due to either depredation at unexclosed nests or nest abandonment by adults. The best predictor of nest abandonment was the maximum number of red fox tracks (Vulpes vulpes) counted on nearby transects (β = −1.16, 95% CI: −2.0 to −0.3) and we found evidence that plovers abandoned eggs in response to predation risk (e.g., a fox circling a nest exclosure). Adults from abandoned nests may have deserted eggs or been depredated. In either case, intact and viable eggs were abandoned. Nest abandonment was not related to human activity near nests, which were buffered from human disturbance by symbolic string fencing. Our results suggest that depredation and nest abandonment (e.g., desertion or death of adults) due to predator disturbance, not human disturbance or poor egg viability, contributed to the low hatching success we recorded. Active predator removal in addition to modification of predator exclosure use and design may be necessary to prevent direct (egg depredation) and indirect (nest abandonment) negative effects of predators on hatching success. © 2010 The Wildlife Society.     

Age‐specific survival and recruitment of piping plovers Charadrius melodus in the Great Lakes region

Juvenile survival and age at first breeding (i.e. recruitment) are critical parameters affecting population dynamics in birds, but high levels of natal dispersal preclude measurement of these variables in most species. We used multi‐state capture–recapture models to measure age‐specific survival and recruitment probabilities of piping plovers Charadrius melodus in the Great Lakes region during 1993–2012. This federally endangered population is thoroughly monitored throughout its entire breeding range, minimizing concerns that measures of survival and recruitment are confounded by temporary or permanent emigration. First‐year survival (± SE) averaged 0.284 ± 0.019 from mean banding age (9 d) and 0.374 ± 0.023 from fledging age (23 d). Factors that increased first‐year survival during the pre‐fledging period (9–23 d) included earlier hatching dates, older age at banding, greater number of fledglings at a given site, and better body condition at time of banding. However, when chicks that died prior to fledging were excluded from analysis, only earlier hatching dates improved first‐year survival estimates. Females had a higher probability (0.557 ± 0.066) of initiating breeding at age one than did males (0.353 ± 0.052), but virtually all plovers began breeding by age three. Adult survival was reduced by increased hurricane activity on the southeast U.S. Atlantic coast where Great Lakes piping plovers winter and by higher populations of merlins Falco columbarius. Mean annual adult survival declined from 1993 to 2012, and did not differ between males and females. Enhanced body condition led to higher survival to fledge and early breeding led to improved first‐year survival; therefore, management actions focused on ensuring access to quality feeding habitat for growing young and protecting early nests may increase recruitment in this federally endangered population.     

A COMPLETE SPECIES CENSUS AND EVIDENCE FOR REGIONAL DECLINES IN PIPING PLOVERS

Abstract: Complete population estimates for widely distributed species are rarely possible. However, for the third time in 10 years, an International Piping Plover (Charadrius melodus) Breeding and Winter Census was conducted throughout the species range in 2001. Nearly 1,400 participants from 32 U.S. states and Puerto Rico; 9 Canadian provinces; St. Pierre and Miquelon, France; Cuba; and the Bahamas visited 2,244 sites covering 11,836 km of shoreline habitat. During the winter census, 2,389 piping plovers were observed at 33.5% of potentially occupied sites (n = 352). Of these, 56.8% had ≤10 birds present. The breeding census recorded 5,945 adults at 777 of 1,892 sites surveyed. More than 80% of sites with piping plovers present had ≤10 birds. Results indicated an 8.4% increase from 1991 but only a 0.2% increase since 1996. Regional trends suggest that since 1991, number of breeding birds increased on the Atlantic Coast by 78% (2,920 birds; 12.4% increase since 1996) and by 80% in the Great Lakes (72 birds; 50% increase since 1996). However, plovers declined 15% (2,953 birds; 10% decline since 1996) in Prairie Canada/U.S. northern Great Plains. Subregional trends since 1991 reflect a 32.4% decline in Prairie Canada (972 birds; 42.4% decline since 1996), a 2.5% decline in the U.S. northern Great Plains (1,981 birds; 24% increase since 1996), 5.5% decline in eastern Canada (481 birds; 14% increase since 1996), although a 66.2% increase on the U.S. Atlantic Coast (2,430 birds; 12% since 1996). While numbers were down in much of the U.S. northern Great Plains since 1996, an increase (460%, 1,048 birds; 67.7% increase since 1991) was detected on the Missouri River. Results from 3 complete species census efforts provide essential data for conservation planning and assessment and illustrate the utility of global censuses for species of concern.     

Nest survival of piping plovers at a dynamic reservoir indicates an ecological trap for a threatened population

In the past 60 years, reservoirs have reshaped riverine ecosystems and transformed breeding habitats used by the threatened piping plover (Charadrius melodus; hereafter plover). Currently, 29 % of the Northern Great Plains plover population nests at reservoirs that might function as ecological traps because reservoirs have more diverse habitat features and greater dynamics in water levels than habitats historically used by breeding plovers. We examined factors influencing daily survival rates (DSR) of 346 plover nests at Lake Sakakawea (SAK; reservoir) during 2006–2009 by evaluating multiple a priori models, and we used our best model to hindcast nest success of plovers during 1985–2009. Our observed and hindcast estimates of nest success were low compared to published estimates. Previous findings indicate that plovers prefer nest sites that are low relative to water level. We found that elevation of nests above the water level had a strong positive correlation with DSR because water levels of SAK typically increased throughout the nesting period. Habitat characteristics on the reservoir differ from those that shaped nest-site selection for plovers. Accordingly, extraordinary nest loss occurs there in many years, largely due to inundation of nests, and based on low fledging rates those losses were not compensated by potential changes in chick survival. Therefore, our example supports the concept of ecological traps in birds because it addresses quantitative assessments of habitat preference and productivity over 25 years (since species listing) and affects a large portion of the population.     

Piping plover habitat selection and nest success on natural,managed, and engineered sandbars

Loss of breeding habitat and nest predation have contributed to the decline of many shorebird species. The United States Army Corps of Engineers (USACE) initiated a piping plover (Charadrius melodus) habitat creation and augmentation program on the Missouri River in the summer of 2004. The USACE increased unvegetated sandbar habitat by depositing dredged material (engineered sandbars) and by clearing vegetation from existing sandbars (managed sandbars). We evaluated the effects of this increase in nesting and foraging habitat on habitat selection and nest daily survival rate (DSR) of piping plovers on Lewis and Clark Lake and the Gavins Point Reach of the Missouri River from 2005 to 2007 (n = 623 nests). Piping plovers selected engineered sandbars more often than expected based on area and selected natural and managed habitats less than expected based on area. Daily survival rate on engineered sandbars was significantly higher than on natural or managed sandbars (log odds: 2.50, 95% CI: 1.05–5.94). Thus, plovers' habitat selection may have increased their nesting success. Our results suggest that habitat augmentation may stave off declines in piping plover populations limited by insufficient habitat and low nesting success. © 2011 The Wildlife Society.     

Human disturbance and stage‐specific habitat requirements influence snowy plover site occupancy during the breeding season

Habitat use has important consequences for avian reproductive success and survival. In coastal areas with recreational activity, human disturbance may limit use of otherwise suitable habitat. Snowy plovers Charadrius nivosus have a patchy breeding distribution along the coastal areas on the Florida Panhandle, USA. Our goal was to determine the relative effects of seasonal human disturbance and habitat requirements on snowy plover habitat use. We surveyed 303 sites for snowy plovers, human disturbance, and habitat features between January and July 2009 and 2010. We made multiple visits during three different sampling periods that corresponded to snowy plover breeding: pre‐breeding, incubation, and brood‐rearing and used multi‐season occupancy models to examine whether human disturbance, habitat features, or both influenced site occupancy, colonization (probability of transition from an unoccupied site to an occupied site), and extinction (probability of transition from an occupied site to an unoccupied site). Snowy plover site occupancy and colonization was negatively associated with human disturbance and site extinction was positively associated with human disturbance. Interdune vegetation had a negative effect on occupancy and colonization, indicating that plovers were less likely to use areas with uniform, dense vegetation among dunes. Also, dune shape, beach debris, and access to low‐energy foraging areas influenced site occupancy, colonization, and extinction. Plovers used habitat based on beach characteristics that provided stage‐specific resource needs; however, human disturbance was the strongest predictor of site occupancy. In addition, vegetation plantings used to enhance dune rehabilitation may negatively impact plover site occupancy. Management actions that decrease human disturbance, such as symbolic fencing and signage, may increase the amount of breeding habitat available to snowy plovers on the Florida Panhandle and in other areas with high human activity. The specific areas that require this protection may vary across snowy plover life history stages.     

Characterization of 36 polymorphic microsatellite loci in the Kentish plover (Charadrius alexandrinus) including two sex‐linked loci and their amplification in four other Charadrius species

We isolated 45 new Kentish plover (Charadrius alexandrinus) microsatellite loci. These were tested for polymorphism in 42 Kentish plovers breeding in the Çukurova Delta, Turkey. Thirty‐six of the 45 loci were polymorphic with observed heterozygosity varying between 0.22 and 0.93. Genotypes of individuals of known sex indicated that two loci were sex‐linked (Calex‐26 is located on the Z chromosome and Calex‐31 on the W chromosome). Additionally, we tested all loci for amplification in four other species of Charadridae (Kittlitz's plover, Madagascar plover, three‐banded plover and white‐fronted plover). On average 34 loci amplified per species (range 29–36).     

Integrating spatial data and shorebird nesting locations to predict the potential future impact of global warming on coastal habitats: A case study on Farasan Islands,Saudi Arabia

One of the expected effects of the global warming is changing coastal habitats by accelerating the rate of sea level rise. Coastal habitats support large number of marine and wetland species including shorebirds (plovers, sandpipers and allies). In this study, we investigate how coastal habitats may be impacted by sea level rise in the Farasan Islands, Kingdom of Saudi Arabia. We use Kentish plover Charadrius alexandrinus – a common coastal breeding shorebird – as an ecological model species to predict the influence of sea level rise. We found that any rise of sea level is likely to inundate 11% of Kentish plover nests. In addition, 5% of the coastal areas of Farasan Islands, which support 26% of Kentish plover nests, will be flooded, if sea level rises by one metre. Our results are constrained by the availability of data on both elevation and bird populations. Therefore, we recommend follow-up studies to model the impacts of sea level rise using different elevation scenarios, and the establishment of a monitoring programme for breeding shorebirds and seabirds in Farasan Islands to assess the impact of climate change on their populations.     

Coastal saltpans are a good alternative breeding habitat for Kentish plover Charadrius alexandrinus when umbrella species are present

The loss and degradation of natural habitats in coastal areas worldwide has adversely affected many waterbird species, changing their breeding distribution and reducing their productivity. Anthropogenic habitats such as saltpans can provide alternative or complementary habitats for waterbirds and mitigate the increasing human impact on natural coastal habitats. Unvegetated linear paths between salt ponds are used by ground‐nesting waterbird species to breed but their linear structure may facilitate the detection of nests by predators. This negative effect may, however, be counterbalanced by the advantages of breeding in mixed colonies. To evaluate the importance and the risks of breeding in saltpans we used the Kentish plover Charadrius alexandrinus breeding in coastal saltpans of southern Portugal as a model species. Specifically, we assessed the role of nest‐site characteristics, predation and nesting proximity to species with aggressive antipredator behaviour (black‐winged stilt Himantopus himantopus) on their breeding success. Kentish plovers selected nest‐sites on the edges of paths, with about 20% of water around the nest, and a mean visibility of more than 72%; however nest‐site characteristics were not correlated with nesting success. Predation was the main cause of nest loss in saltpans (42%); carrion crows Corvus corone were responsible for most daylight nest predation (58%) and red fox (Vulpes vulpes; 73%) for night predation. An 8‐yr monitoring plan of a Kentish plover population showed a linear increase in their breeding success as their breeding season increasingly overlapped with that of the black‐winged stilt. An experiment with artificial nests showed a significant increase in the number of exposure days (7 to 12) when these nests were within close distance of black‐winged stilt nests. Overall, our results showed that saltpans are an important alternative breeding habitat for the Kentish plover, especially for the maintenance of mixed species colonies.     

Susceptibility of avian hosts to experimental Gymnophalloides seoi infection

To determine whether avian species are susceptible to infection with Gymnophalloides seoi (a human-infecting intestinal trematode), we exposed 7 species of birds with metacercariae obtained from oysters. The birds were necropsied at days 2, 4, and 6 postinfection (PI). The highest worm recovery at day 6 PI was obtained from the Kentish plover (Charadrius alexandrinus; mean = 56.0%), followed by the Mongolian plover (C. mongolus; 49.3%), and the grey plover (Pluvialis squatarola; 32.3%). In contrast, no mature worms were recovered from the great knot (Calidris tenuirostris), dunlin (C. alpina), black-tailed gull (Larus crassirostris), and mallard (Anas platyrhynchos). Among the plovers, the worms attained the greatest size at day 6 PI (254.1 x 190.4 microm) in the Kentish plover, with a significantly higher number of eggs in the uterus. The 3 species of plovers are highly susceptible to experimental G. seoi infection, suggesting that they could play a role as definitive hosts for these worms in nature.     

Modeling foraging behavior of piping plovers to evaluate habitat restoration success

Habitat restoration projects are often deemed successful based on the presence of the target species within the habitat; however, in some cases the restored habitat acts as an ecological trap and does not help to improve the reproductive success of the target species. Understanding wildlife–habitat relationships through precise measurements of animal behavior can identify critical resources that contribute to high quality habitat and improve habitat restoration practice. We evaluated the success of a restored piping plover (Charadrius melodus) breeding habitat in New Jersey, USA. We identified the major factors influencing foraging rates, compared foraging activity budgets over 3 yr at restored and natural habitats, and explored the potential of artificial tidal ponds as a viable restoration alternative. Adult foraging rates were higher in artificial pond and ephemeral pool habitats, during low tide, and after breeding activity ended. Adult foraging rates were impeded by the presence of people and vehicles within 50 m. Chick foraging rates were highest at artificial ponds and bay shores and lowest in dunes and on sand flats. Chick foraging rates were strongly hindered by the presence of corvids and the number of people within 50 m. In addition, at artificial tidal ponds, piping plovers spent more time foraging and less time engaged in defensive behaviors (vigilance, crouching, and fleeing) compared to other potential habitats. Our findings support the hypothesis that artificial tidal ponds are a valuable, perhaps superior, foraging habitat. Future beach restoration projects should include this feature to maximize habitat quality and restoration success. © 2011 The Wildlife Society.     

Impacts of climate on prey abundance account for fluctuations in a population of a northern wader at the southern edge of its range

Understanding the mechanisms by which climate change will affect animal populations is vital for adaptive management. Many studies have described changes in the timing of biological events, which can produce phenological mismatch. Direct effects on prey abundance might also be important, but have rarely been studied. We examine the likely importance of variation in prey abundance in driving the demographics of a European golden plover ( Pluvialis apricaria ) population at its southern range margin. Previous studies have correlated plover productivity with the abundance of their adult cranefly (Tipulidae) prey, and modelled the phenology of both plover breeding and cranefly emergence in relation to temperature. Our analyses demonstrate that abundance of adult craneflies is correlated with August temperature in the previous year. Correspondingly, changes in the golden plover population are negatively correlated with August temperature 2 years earlier. Predictions of annual productivity, based on temperature-mediated reductions in prey abundance, closely match observed trends. Modelled variation in annual productivity for a future scenario of increasing August temperatures predicts a significant risk of extinction of the golden plover population over the next 100 years, depending upon the magnitude of warming. Direct effects of climate warming upon cranefly populations may therefore cause northward range contractions of golden plovers, as predicted by climate envelope modelling. Craneflies are an important food source for many northern and upland birds, and our results are likely to have wide relevance to these other species. Research into the potential for habitat management to improve the resilience of cranefly populations to high temperature should be an urgent priority.     

Piping Plover Chick Foraging,Growth, and Survival in the Great Plains

Abstract: We tested the hypothesis that piping plover (Charadrius melodus) habitat quality and chick survival on the Missouri River, USA, were lower on a cold-water reservoir and downstream from a hypolimnetic (cold-water) release dam with diel water fluctuations (Garrison Dam) than downstream from an epilimnetic dam (Gavins Point Dam). Plovers in adjacent alkali wetlands provided an index to the maximum reproductive potential in the region. Chicks gained weight more rapidly in the alkali wetlands than on epilimnetic and hypolimnetic river reaches. Invertebrate numbers and biomass were higher in the wetlands and epilimnetic reach, but chick survival was lower on the epilimnetic reach. Thus, piping plovers adapted to a variety of prey densities, and other factors, likely predation, reduced survival rates in the epilimnetic reach. Temporal and spatial variability in site quality indices suggests the need for a regional management strategy with different strategies at each site. Managers can minimize effects of local fluctuations in resource abundance and predators by ensuring protection of or creating geographically dispersed habitat. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):682–687; 2008)     

Habitat use by Mountain Plovers in prairie dog colonies in northeastern New Mexico

ABSTRACT Mountain Plovers (Charadrius montanus) are grassland birds that often breed in close association with colonies of black‐tailed prairie dogs (Cynomys ludovicianus). However, not all colonies provide plover nesting habitat or habitat of equal quality, and the characteristics of colonies important for plovers remain poorly understood. Over two years, I used plover distribution surveys, territory mapping, and habitat sampling to study habitat use by plovers in prairie dog colonies in shortgrass prairie in northeastern New Mexico. My objective was to document important components of plover breeding habitat in colonies by comparing characteristics of used and unused habitats at three spatial scales: colony, territory, and nest‐site. I found evidence of plover breeding in 14 of 44 colonies in 2009 and 13 of 43 colonies in 2010. Based on logistic regression, the probability of a colony being occupied by plovers was positively associated with colony size, but negatively associated with mean vegetation height. Preference for larger colonies could relate to minimum habitat requirements, or a potential tendency of this species to nest in social clusters. Shorter vegetation height was strongly correlated with greater bare ground and lower forb/subshrub cover, all characteristics that may be related to plover predator avoidance and foraging microhabitat. At both the territory and nest‐site scale, areas used by plovers had shorter vegetation, more bare ground, and less forb/subshrub cover than unused areas. Nest sites were also more sloped, perhaps to reduce risk of flooding, and located further away from the nearest prairie dog burrow, perhaps to reduce risk of disturbance. Overall, my results show that plover use of prairie dog colonies was influenced by landscape and habitat features of colonies, and suggest that large colonies are particularly valuable because they are most likely to contain adequate areas with preferred habitat characteristics.     

Ecological constraints on breeding system evolution: the influence of habitat on brood desertion in Kentish plover

1. One of the fundamental insights of behavioural ecology is that resources influence breeding systems. For instance, when food resources are plenty, one parent is able to care for the young on its own, so that the other parent can desert and became polygamous. We investigated this hypothesis in the context of classical polyandry when females may have several mates within a single breeding season, and parental duties are carried out largely by the male. 2. We studied a precocial wader, the Kentish plover Charadrius alexandrinus, that exhibits variable brood care such that the chicks may be raised by both parents, only by the female or, more often, only by the male. The timing of female desertion varies: some females desert their brood at hatching of the eggs and lay a clutch for a new mate, whereas other females stay with their brood until the chicks fledge. Kentish plovers are excellent organisms with which to study breeding system evolution, as some of their close relatives exhibit classical polyandry (Eurasian dotterel Eudromias morinellus, mountain plover Charadrius montanus), whereas others are polygynous (northern lapwing Vanellus vanellus). 3. Kentish plovers raised their broods in two habitats in our study site in southern Turkey: saltmarsh and lakeshore. Food intake was higher on the lakeshore than in the saltmarsh as judged from feeding behaviour of chicks and adults. As the season proceeded and the saltmarsh dried out, the broods moved toward the lakeshore. 4. As the density of plovers increased on lakeshore, the parents spent more time defending their young, and female parents stayed with their brood longer on the lakeshore. 5. We conclude that the influence of food abundance on breeding systems is more complex than currently anticipated. Abundant food resources appear to have profound implications on spatial distribution of broods, and the social interactions between broods constrain female desertion and polyandry.