Nesting success is an important vital rate affecting the reproductive fitness of birds, and predation typically is the single most important factor affecting nesting success. Presumably, birds should nest in locations that maximize nest survival. If specific nest characteristics increase the probability that a nest will hatch, natural (phenotypic) selection could favour use of sites with these features, producing nonrandom patterns of nest site use. Alternatively, birds that are highly selective in nest site choices might be at a disadvantage if predators learn to forage preferentially in these locations and improve their efficiency in depredating nests; in this case, random nesting patterns could be favoured. Finally, it has been hypothesized that predation pressure can influence nest site selection patterns of entire bird communities. If predators develop a search image to hunt for bird nests, then nests that are most similar to each other, irrespective of species, should sustain higher mortality. To evaluate these hypotheses, I quantified nest site selection patterns of multiple species of ground-nesting dabbling ducks in areas where predation pressure was normally high, and compared these patterns to those on areas where predation was relaxed. Predation pressure was experimentally reduced by removing common predators of duck nests and females (mainly red foxes, coyotes, skunks and raccoons) on some study areas and not on others (controls). Predator removal and natural causes produced a 10-fold difference in duck nesting across study sites, allowing for investigation of effects of predation pressure on nest site selection of ducks. Coarse scale habitat selection patterns were similar to results reported in previous studies; blue-winged teal and northern shoveler were found more often in native grassland than in other habitat types, while gadwall and mallard nests occurred more frequently in shrub patches when compared with other habitat patches. A difference in nest site characteristics was observed between hatched and depredated nests for gadwall and northern shoveler but not for blue-winged teal and mallard. However, in all species, the nest site selection patterns were non-random. Thus, the process of nest predation did not shape patterns of nest site choice. Contrary to predictions, inter-specific overlap in nest site features was not related to predation pressure: nests that overlapped most with features of other species did not suffer higher predation, nor did inter-specific overlap in nest characteristics decrease during the nesting season. These findings were inconsistent with the hypothesis that community-level patterns of nest site use are differentiated as a result of predation pressure. Long-term work on nest site use by individually marked females of numerous ground-nesting bird species would be informative, as would experimental studies of other hypotheses about factors affecting nest site choices in birds.

Nesting success is an important vital rate affecting the reproductive fitness of birds, and predation typically is the single most important factor affecting nesting success. Presumably, birds should nest in locations that maximize nest survival. If specific nest characteristics increase the probability that a nest will hatch, natural (phenotypic) selection could favour use of sites with these features, producing nonrandom patterns of nest site use. Alternatively, birds that are highly selective in nest site choices might be at a disadvantage if predators learn to forage preferentially in these locations and improve their efficiency in depredating nests; in this case, random nesting patterns could be favoured. Finally, it has been hypothesized that predation pressure can influence nest site selection patterns of entire bird communities. If predators develop a search image to hunt for bird nests, then nests that are most similar to each other, irrespective of species, should sustain higher mortality. To evaluate these hypotheses, I quantified nest site selection patterns of multiple species of ground-nesting dabbling ducks in areas where predation pressure was normally high, and compared these patterns to those on areas where predation was relaxed. Predation pressure was experimentally reduced by removing common predators of duck nests and females (mainly red foxes, coyotes, skunks and raccoons) on some study areas and not on others (controls). Predator removal and natural causes produced a 10-fold difference in duck nesting across study sites, allowing for investigation of effects of predation pressure on nest site selection of ducks. Coarse scale habitat selection patterns were similar to results reported in previous studies; blue-winged teal and northern shoveler were found more often in native grassland than in other habitat types, while gadwall and mallard nests occurred more frequently in shrub patches when compared with other habitat patches. A difference in nest site characteristics was observed between hatched and depredated nests for gadwall and northern shoveler but not for blue-winged teal and mallard. However, in all species, the nest site selection patterns were non-random. Thus, the process of nest predation did not shape patterns of nest site choice. Contrary to predictions, inter-specific overlap in nest site features was not related to predation pressure: nests that overlapped most with features of other species did not suffer higher predation, nor did inter-specific overlap in nest characteristics decrease during the nesting season. These findings were inconsistent with the hypothesis that community-level patterns of nest site use are differentiated as a result of predation pressure. Long-term work on nest site use by individually marked females of numerous ground-nesting bird species would be informative, as would experimental studies of other hypotheses about factors affecting nest site choices in birds.

Mottled ducks (Anas fulvigula) are a non-migratory waterfowl species endemic to the western Gulf Coast, with a separate, genetically distinct subspecies (A. fulvigula fulvigula) occurring in peninsular Florida. Birds from Texas, Louisiana, and Florida were released in coastal South Carolina from 1975-1983, and banding data suggest an expanding population. I monitored 72 mottled duck nests and captured and radio-marked 196 pre-breeding and nesting females between 2010 and 2014 to study breeding ecology of these birds in the Ashepoo, Combahee, Edisto Rivers Basin. Nest success averaged 12% and varied with vegetation height and year. Indicated breeding pair surveys revealed breeding mottled ducks select managed wetland impoundments, predominately influenced by water depth. Future research should investigate ecology of nest predators of mottled ducks to devise successful habitat management strategies for breeding birds. Preliminary evidence suggests that managed wetland impoundments are important to breeding and brood rearing mottled ducks in coastal South Carolina.

Nesting and early life is a period of high mortality for many turtle species, so understanding how turtles select nest sites, and how those nest sites impact hatching success, may be important for successful species conservation. In this research, my objective was to 1) understand how the environment around potential nest sites (canopy, understory, and soil) influences gopher tortoise nest-site selection at burrows and 2) how that nest environment both directly (nest microclimate: temperature and moisture) and indirectly (nest characteristics: nest depth, distance from burrow, canopy and understory cover, percent clay in soil, and lay date) affects hatching success in naturally incubated tortoise nests. In the summers of 2022 and 2023, I conducted repeated searches at burrows to locate nests at the Jones Center at Ichauway and the Greenwood Ecological Reserve in southwestern Georgia. I collected soil samples, measured canopy and ground cover at gopher tortoise nest locations (n=132) and an equal number of comparison non-nest burrows. At nest sites, I also monitored temperature and moisture throughout incubation. To evaluate nest-site selection, I compared burrows with and without nests using multiple logistic regression to create a suite of five biologically relevant candidate models and compared models with Akaike's Information Criterion adjusted for small sample sizes. The top three models identified canopy cover and understory vegetation cover as the only significant predictors of nest presence at burrows, with tortoises in the sites nesting at burrows with lower understory and canopy cover. Furthermore, there was an interaction between the understory vegetation and canopy cover effects, where the effect of understory cover decreased as canopy cover increased. This suggests that the vegetation effect may be primarily driven by an avoidance of shade, whereby nests laid in burrows with high canopy cover were so shaded that understory vegetation had a weaker influence on nest-site selection. When tortoises nested in burrows with lower canopy cover, which was far more common than high canopy cover at our sites, they also avoided understory vegetation so that nest sites were least shaded. These results suggest that maintaining habitats with very open overstories may be most important for allowing gopher tortoises access to preferred nest sites. To quantify the direct and indirect effects of nest environment on hatching success, I built a structural equation model (SEM) in a Bayesian framework in which hatching success was affected by nest temperature and moisture, which were themselves affected by nest site characteristics. I found that nest microclimate could be predicted moderately well from characteristics of the nest environment (R2=0.25-0.49), with lay date influencing both temperature and moisture, vegetation affecting temperature, nest position influencing moisture and temperature variability, and percent clay in soil influencing moisture. Hatching success was highest at lower mean temperatures and moistures and at intermediate levels of temperature and moisture variability, but the ability of this model to predict hatching success was low (R2=0.10). I observed very high hatching success (87.5%) and, thus, eggs were generally receiving the conditions they needed to successfully develop and there was not much variation in hatching success to explain. This framework may be useful for investigating environmental causes of lower hatching success at less robust tortoise populations that may be experiencing low rates of natural hatching success. / Master of Science / Many turtle species experience high rates of mortality in early life, so understanding how turtles select areas to nest, and how those places impact hatching success, may be important for successful species conservation. In this research, my objective was to 1) understand how the environment around potential nest locations (vegetation and soil) influences where gopher tortoise place nests at burrows and 2) how the conditions of that nest location both directly and indirectly affect hatching success in natural gopher tortoise (Gopherus polyphemus) nests. In the summers of 2022 and 2023, I conducted repeated searches at burrows to locate nests at the Jones Center at Ichauway and the Greenwood Ecological Reserve in southwestern Georgia. I collected soil samples, measured canopy and vegetative ground cover at gopher tortoise nest locations and an equal number of burrows without nests. At nest sites, I also monitored temperature and moisture throughout incubation. To evaluate how tortoises chose nest locations, I created models to compare tortoise burrows with nests to burrows that were available for nesting, but where no nest was placed. The top three models identified canopy cover and understory vegetation cover as the only significant predictors of nest presence at burrows, with tortoises in my sites nesting at burrows with lower understory and canopy cover. Furthermore, there was an interaction between the understory vegetation and canopy cover effects, where the effect of understory cover decreased as canopy cover increased. This suggests that the vegetation effect may be primarily driven by an avoidance of shade, whereby nests laid in burrows with high canopy cover were so shaded that understory vegetation had a weaker influence on nest-site selection. When tortoises nested in burrows with lower canopy cover, which was far more common than high canopy cover at my sites, they also avoided understory vegetation so that nest sites were least shaded. These results suggest that maintaining habitats with open overstories may be most important for allowing gopher tortoises access to preferred nest sites. To quantify the direct and indirect effects of nest environment on hatching success, I built a structural equation model (SEM) in which hatching success was predicted by nest temperature and moisture, which were themselves predicted by nest-site characteristics. This allowed me to evaluate both the direct effects of nest temperature and soil and the indirect pathways by which nest environment may be influencing hatching success. I found that nest temperature and moisture could be predicted moderately well from characteristics of the nest environment (R2=0.25-0.49), with the date the nest was laid influencing both temperature and moisture, vegetation around the nest affecting temperature, nest position influencing moisture and temperature variability, and percent clay in soil influencing moisture. Hatching success was highest at lower mean temperatures and moistures and at intermediate levels of temperature and moisture variability, but the ability of this model to predict hatching success was low (R2=0.10). I observed very high hatching success (87.5%) and, thus, eggs were generally receiving the conditions they needed to successfully develop and there was not much variation in hatching success to explain. This framework may be useful for investigating environmental causes of lower hatching success at less robust tortoise populations that may be experiencing low rates of natural hatching success.

Master of Science / Division of Biology / David A. Haukos / The ring-necked pheasant (Phasianus colchicus) is a popular and economically important upland gamebird in Kansas. Population declines have stakeholders seeking methods to manage populations on agricultural lands. Cover crops planted during the breeding period may provide important resources pheasants require for survival and successful reproduction. I evaluated three cover crop mixes; a custom mix, commercial mix, a wildlife mix, and a chemical fallow control in three counties in western Kansas, during 2017 and 2018 to determine their potential as a management practice for increasing pheasant habitat. I tested the relative effects of spring cover crops on female pheasant survival, nest survival, nest-site selection, and mesocarnivore occupancy. Females pheasants (73) were captured via nightlighting during February – April and fitted with 15-g very-high-frequency radio collars and monitored by telemetry. I placed 58 camera traps on field edges and within cover crop treatments from April to September. Vegetation data were collected at nests and random points to assess nest-site selection and weekly random vegetation points were sampled within treatments. I used known fate and nest survival models in the package RMark interface in R to investigate adult and nest survival (R Core Team 2018). Adult breeding season survival was 0.57 (SE < 0.0001, CI = 0.5739 – 0.5740). Percent spring cover crop positively influenced adult survival (AICc wi = 0.450). Nest survival was 0.36 (SE < 0.001, CI = 0.3614 - 0.3614). Daily nest survival followed a pattern of high survival that gradually declined over the breeding season. Resource selection functions suggest female ring-necked pheasants selected vegetation between 5-7 dm at 50% VOR for nest sites (AICc wi = 0.97). Chi-square analyses suggest females selected Conservation Reserve Program (CRP) patches for nest sites more than expected during both years (2017 χ²₄ = 26.49, P < 0.001; 2018 χ²₄ = 9.80, P = 0.04). CRP supported 57% of nests and 56% of successful nests relative to other cover types. All three of the monitored nests in cover crops were depredated. Ring-necked pheasant occupancy was greatest on edges of treatments (ψ = 0.97, SE = 0.081) and influenced by proportion of the Chick Magnet seed mix (AICc wi = 0.68). Mesocarnivore occupancy was greatest on treatment edges with a constant occupancy of 0.99 (SE = 0.47, AICc wi = 0.66). Spring cover crops provide cover and foraging resources when the majority of agricultural practices are fallow. Spring cover crops do not provide sufficient vertical cover for nesting until after peak nesting occurs, especially during cooler than average winter and spring conditions such as 2018. However, there are tangible benefits of spring cover crops to other biological periods, such as adult female survival, and brood resources if placement of cover crops is targeted near quality nest habitat. My results indicates wheat is an ecological trap for nesting due to increased predation and destruction during harvest. Providing quality nest structure will reduce females nesting in wheat. Incorporation of spring cover crops is a beneficial wildlife management tool that can increase ring-necked pheasant habitat on the landscape.

Climate change is threatening biodiversity, causing populations and species to adapt, or otherwise, become extinct. Sea turtles have survived dramatic climate changes in the past, however, due to a history of intense human exploitation, and the current anthropogenic threats, their current resilience may be jeopardized. The main pursuits of this thesis were to i) evaluate the resistance of green turtles to predicted climate change impacts, using a globally significant rookery, in Poilão, Guinea-Bissau, as a case study; and ii) assess key population parameters to inform the conservation management of this resource. As the work developed I additionally had the opportunity to study the dynamics of an emerging disease in a juvenile foraging aggregation from Puerto Rico, which contributed to a broader understanding of resilience in this species. Specifically, I investigate the nest site selection behaviour of green turtles, their nesting environment, and the outcomes for their offspring, at Poilão, and apply this information to infer on the resilience of this population under future scenarios of climate change. I explore the connectivity established by the dispersal of post-hatchlings from Poilão, followed by their recruitment to foraging grounds, to set the geographical context of this major population. Lastly, I model the dynamics of Fibropapillomatosis, which affects juvenile green turtles globally, and examine the potential for disease recovery. The green turtle rookery in Poilão shows some resilience to expected climate change impacts. This significant population likely contributes to all juvenile foraging aggregations along the west coast of Africa, and to some extent to those in South America. Currently, green turtles are capable of recovery from Fibropapillomatosis, however, the incidence of disease may be enhanced by climate change.

The red-headed woodpecker is a primary cavity excavator found throughout the Eastern Deciduous Forest and parts of the Great Plains. Although widespread, the red-headed woodpecker is generally considered uncommon, with patchily distributed populations. Over the past 50 years, this species has experienced precipitous, range-wide declines. The red-headed woodpecker uses a variety of cover types to meet resource needs across the annual cycle, ranging from oak savannas and wetlands, to mature beech forests and urban parks. Given their apparent adaptability to such a wide range of habitats, causes of declines are perplexing. To understand and stem declines, recent studies have focused on quantifying this species’ habitat requirements. Most of these studies have focused on a single cover type, often open forests, during the breeding season. However, effective management requires comprehensive knowledge of red-headed woodpecker habitat needs in a variety of cover types across the annual cycle. My thesis seeks to address this knowledge gap. I studied red-headed woodpecker habitat selection during both the breeding and non-breeding seasons at Fort A.P. Hill (FAPH) in Caroline County, Virginia. FAPH contains a variety of cover types that red-headed woodpeckers are known to use for breeding and overwintering, including: wetlands, open forests and closed forests. In Chapter 1, I investigated winter roost-site selection of red-headed woodpeckers in a burned forest stand. My study was the first to quantify winter habitat selection for this species. My aim was to identify variables driving winter roost-site selection at two relevant spatial scales, and estimate their parameter weights using logistic regression. I found that red-headed woodpeckers preferred habitat around the roost tree that contained a higher basal area of snags and mast-producing trees. They differentially selected roost trees based on taxon and decay state. My results indicate that managers should adopt practices that promote snag generation and retention, and mature oak recruitment. In chapter 2, I focused on red-headed woodpecker breeding season habitat requirements, as many other studies have done. However, unlike other studies that investigated habitat selection within a single cover type, I found and characterized nest-sites in three distinct cover types (wetlands, closed forests, open forests). I used a comparative approach to identify cover-type specific nest-habitat thresholds at the landscape, patch and tree scales using boosted regression trees (BRT). Although models at the landscape scale inadequately discriminated between nest and available sites, models at the patch and tree scales achieved excellent discrimination ability. I found that red-headed woodpeckers are consistent in their preference for a number of habitat features at the patch (high medium/large snag density, open canopy) and tree (large diameter tree with less bark) scales, irrespective of cover type context. However, I also found cover-type specific habitat preferences at the patch scale, indicating red-headed woodpeckers are flexible in their selection of features surrounding the nest and responsive to the broader habitat context. My findings suggest that there are a number of habitat features that facilitate breeding for red-headed woodpeckers generally, and management that promotes these features will be effective in a variety of habitat contexts. They also indicate that management can be tailored to provide the most suitable habitat for each of these three commonly used cover types. Together, Chapters 1 and Chapter 2 expand our knowledge of this declining keystone species’ habitat needs across the full annual cycle. But there is still much to know. The choices of habitat selection have consequences. Determining the conditions under which habitat features function to either help or harm populations is a necessary next step. While not a part of this thesis, I am currently investigating the factors that drive nest success among cover types, linking breeding-site selection to population dynamics. I also have behavioral data that will help reveal the mechanisms that either facilitate or constrain the exploitation of food resources across cover types. Finally, the red-headed woodpecker is a facultative migrant that often shifts habitat associations to take advantage of seasonally available resources. Data from my point count surveys—conducted over two successive winter and summer seasons—allow for development of dynamic occupancy models. Modeling shifts in occupancy across seasons will show the habitat factors underlying seasonal shifts. Site-specific differences in colonization, extinction and frequency of occupancy, within seasons but across years, will further our understanding of what constitutes habitat quality for this species, in both the breeding and non-breeding seasons.

Three tern species, Roseate Tern (Sterna dougalli), Bridled Tern (Sterna anaethetus) and Crested Tern (Sterna bergii) co-bred at Baisha islet, east-north of Penghu, in the summer of 2006. Each species favored different kind of environment for nesting. All Crested Terns nested in the flat plane with vegetation; Roseate Terns in the cliff near the vegetation, and Bridled Tern primarily nested near the vertical rock or under a rocky roof. The Crested Tern had the highest nesting density, and Bridled Tern nested loosely. The hatching success of Roseate (75%) and Crested Tern (73%) were significantly higher than that of Bridled Tern (30%). Roseate and Crested Tern laid eggs synchronously and had apparently two wave of egg laying and the breeding performances between early- and late-laying period were quite different. The relationship between hatching success and nest-site characteristics of the three species was investigated. The results revealed that hatching success of Roseate Tern increased with the number of walls and neighbors. Early-laying nests (79%) and central nests (72%) were more successful than late (11%) and edge nests (45%) of Roseate Terns. The laying-period was also important factors affecting hatching success of Crested Tern. Overall, the low hatching success of the late-laying nests may be due to the change of environmental conditions and the losing advantage of group breeding in the late season. The low hatching success of Bridled Tern was considered owing to the asynchronous laying pattern, loosely built nesting and weak parental behaviors.

The aim of this study was to determine the habitat features affecting nest site selection and breeding success of the endangered Egyptian Vultures (Neophron percnopterus) breeding around the town of Beypazari. We searched and monitored nest sites in the study area (750 km2) for the years 2010 and 2011. The differences in terms of habitat features between nest sites and random points distributed along cliffs, and between successful and failed nest sites were investigated using both parametric approaches and machine learning methods with 21 habitat variables. The size of the Beypazari population of Egyptian Vultures was estimated to be 45 pairs. Seventeen nests in 2010 and 37 nests in 2011 were found and monitored. The breeding success of the population was estimated to be 100% in 2010 and 70% in 2011. Random Forests was the modeling technique with the highest accuracy and the modeling process chose 6 and 4 variables affecting nest site selection and breeding success of the species, respectively. Results showed that human impact was a potential factor governing the distribution of nest sites in the area and increased the probability of breeding failure as vultures clearly preferred to nest away from nearby villages, towns and roads, and nests on lower cliffs and nests that are close to the dump site (therefore the town center) was prone to failure. Utilization of elevation gradient and aspect showed trends similar to other populations of the species, with probability of nesting increasing at lower altitudes and for south facing cliffs. The overall results emphasize the potential conflict between human presence and the population of Egyptian Vultures in the area. Continuous monitoring of the nest sites and conservation activities towards raising public awareness are advised.

Many factors affect the productivity of songbirds. Which vegetation types the birds inhabit, nest-site characteristics, and adult behavior at the nest may affect predation and parasitism frequencies, fecundity, and nest survival and fledgling survival. All of these metrics determine reproductive success of individuals and may influence population persistence, especially for threatened and endangered species. My research investigated factors that affected these metrics for endangered black-capped vireos (Vireo atricapilla). Shrubland is considered high quality vireo habitat, with woodland vegetation types considered marginal. I located and monitored nests, conducted nest behavior observations, recorded behavior and predation at nests using video cameras, and resighted fledglings in shrubland, oak-juniper woodland, and deciduous woodland during the 2008–2010 breeding seasons. I monitored 302 black-capped vireo nests in 259 territories and resighted 350 fledglings with unique color combinations. Apparent nest success, nest survival, success of first nest attempts, parasitism and predation frequency, and fecundity did not differ statistically among vegetation types. Parasitism frequency was nearly twice as high in shrubland (22 percent) than in either woodland (12 percent in each) and varied by year. Nest-site characteristics differed among vegetation types, but nest survival was affected only by nest height and year; nests placed higher from the ground and nest attempts in 2008 and 2009 had lower survival. Fledgling survival was not affected by vegetation type or proximity of the nest to oak-juniper woodland. Nest behavior was not affected by vegetation characteristics, though nest attentiveness during incubation increased as average cover from 0 to 2 m increased. Females spent 80 percent more time on nests during incubation and 250 percent more time on nests during the nestling stage than males, but visitation was similar for each sex. Overall, the probability of nest success improved as male participation increased. My results emphasize the importance of male participation in determining the outcome of nests for species exhibiting bi-parental care. Furthermore, woodland habitats previously considered marginal may be good quality habitat in areas with large populations of black-capped vireos. Recognizing woodlands as non-typical, yet still suitable, habitat will allow managers to incorporate these vegetation types into management plans and recommendations for landowner conservation incentive programs.