The measurement of stress hormone levels in wild free-living animals is becoming an increasingly effective method for examining proximate mechanisms of animal behavior and the physiological impacts of human activities on wildlife. In these studies I measured plasma levels of the stress hormone corticosterone in the endangered red-cockaded woodpecker (Picoides borealis) to determine their role in the reproductive behavior of individuals in this species, and whether they are affected by human disturbance. In chapter one, I provide an introduction to the vertebrate stress response and I describe the natural history of the red-cockaded woodpecker. In chapter two, I compare corticosterone and reproductive hormone levels between breeding males and helper males to examine hormonal mechanisms of reproductive suppression in helper males. No hormonal differences existed between breeding and helper males. However, baseline corticosterone levels were lower in all males living in groups with two or more helper males, suggesting that male helpers reduce the workload of all other group members. In chapter three, I compare corticosterone levels between birds living in clusters subject to two different training restriction regimes on a military installation. Males living in clusters without training restrictions had lower baseline corticosterone than those living in clusters with training restrictions, suggesting that males habituate to chronic disturbance by downregulating baseline corticosterone levels. / Master of Science

Dispersal is a critical life-history component; it determines gene flow and has profound effects on population structure, demography, social systems, and population viability. To add to our knowledge of dispersal and, in particular, our understanding of the relationship between dispersal and inbreeding, I studied three aspects of the biology of the red-cockaded woodpecker: dispersal of breeding females; the costs, benefits, and frequency of inbreeding; and the effect of inbreeding on natal dispersal. Dispersal of breeding female red-cockaded woodpeckers is strongly associated with inbreeding avoidance and mate choice, weakly associated with site choice, and not found to be associated with social constraints. Estimates of mortality for non-dispersing and dispersing breeding females were 24 and 59 percent per year, respectively-rare evidence of the cost of breeding dispersal. Significant costs of close inbreeding were found. Closely related pairs (kinship coefficient greater than 0.1) had lower hatching success as well as lower survival and recruitment of fledglings than unrelated pairs. Moderately related pairs (kinship coefficient between 0 and 0.1) and moderately inbred individuals had increased hatching success, but did not produce more young. Despite documented costs of close inbreeding and a predictable spatial distribution of closely related males near the natal territory, female fledglings disperse a median of only two territories and a modal distance of one territory. Natal dispersal of females is affected by closely related males on the natal site but unaffected by closely related males or moderately related males that are off the natal site. / Master of Science

This project had 2 major objectives. The first objective was to assess how well the revised U.S. Fish and Wildlife Service Foraging Habitat Guidelines depict good quality habitat for the red-cockaded woodpecker (Picoides borealis) at Camp Lejeune, NC. To accomplish this, I used multiple linear and logistic regression to examine the relationships between fitness, habitat use, home range size, and habitat characteristics described in the guidelines. I assumed that habitat characteristics that confer quality were related to higher fitness, greater habitat use, and reduced home range size. To a large extent, the guidelines are validated. Red-cockaded woodpeckers responded favorably to habitat that mimics the historical, mature, and fire-maintained pine forests of the southeastern U.S., characterized by high densities of large pines, low densities of small and medium pines, and a lush herbaceous groundcover. Variables positively associated with habitat use and fitness were associated with reduced home range size, and those negatively associated with habitat use and fitness with increased home range size. Percent herbaceous groundcover was a significant regressor indicative of quality in every model. The second objective was to assess how well USFWS foraging partitions represent habitat used by red-cockaded woodpeckers. I conducted home range follows of 23 groups of red-cockaded woodpeckers and estimated the percentage of each home range encompassed by partitions of varying radii. The percentage of the actual home range included in the partition increased as a function of partition radius. The standard 800 m circular partition, on average, included 91% of the home range, but significant variation existed between groups. / Master of Science

One unresolved issue in the study of cooperative breeding is why individuals that delay their own reproduction assist others in raising young. Red-cockaded woodpecker helpers may again future indirect fitness benefits by increasing survival of breeders, who produce offspring related to the helper in the future. Breeder survival may be enhanced because of general benefits of group living, either because of the helper's own presence or because helping increases the production of fledglings. I demonstrate that breeder survival increases in the presence of helpers and additional young. Helpers may also reduce the workload of the breeder, and this may increase breeder survival as well. I show that breeders spend less time incubating and provisioning young when a helper is present. Helpers may also gain fitness benefits if once they become breeders, they are assisted by young they previously helped raise. Expected frequencies of reciprocal exchange of helping are low (2%). Slightly higher observed frequencies may be accounted for by preferential helping of kin and effects of territory quality. Reciprocity occurs no more often than expected among helpers unrelated to the young they help raise suggesting that young males do not preferentially helper former care-givers. i conclude that helpers do not gain fitness benefits from reciprocity. I examined the proximate causes of delayed breeding and helping behavior by measuring plasma testosterone (T) and prolactin (PRL) concentrations in female breeders, male breeders, and male helpers during different stages of the reproductive cycle. Among male breeders and helpers, T is low during the nonbreeding stage, peaks during copulation and declines during the incubation and nestling-provisioning stages. Helpers appear physiologically capable of reproducing: their T concentrations are equal those of breeders. Helpers unrelated to the breeding female have higher T than helper related to her. Sexual inactivity by male helpers is best explained by behavioral suppression. Female breeder, male breeder, and male helper PRL was equal and increased from the nonbreeding stage through the copulation and incubation stages. During the nestling provisioning stage, male breeder and male helper PRL declined, while female PRL continued to increase. I conclude that the physiological bases of helping behavior and parental behavior are the same. / Ph. D.

Cavity-excavating birds, such as woodpeckers, are ecosystem engineers and are often assumed to rely upon wood decay fungi to assist in softening the wood of potential excavation sites. Endangered red-cockaded woodpeckers (Picoides borealis; RCWs) are the only birds known to solely excavate through the sapwood and into the heartwood of living pine trees and generally take many years to complete their excavations. These birds may have developed a partnership with wood-inhabiting fungi to facilitate the excavation process. Past attempts to understand the complex relationships between cavity excavators and fungi relied on visual surveys of fruiting bodies, or evidence of decay, resulting in a one bird, one fungus paradigm. Using molecular methods, I investigated the relationships between RCWs and fungi, and found that the relationships between cavity-excavators and fungi involve multiple fungal species and are far more complex than previously imagined. Through a field survey, I showed that RCW excavations contain distinct communities of fungi, and propose two hypotheses to explain this result, (1) RCWs select trees with distinct fungal communities (tree selection hypothesis), or (2) RCWs promote distinct fungal communities via their excavations (bird facilitation hypothesis). By swabbing the birds, I found that RCWs carry fungal communities similar to those found in their completed excavations, demonstrating that RCWs may directly facilitate fungal dispersal during the excavation process. Through a test of the bird introduction hypothesis which implemented human-made experimental drilled cavity starts (incomplete excavations), half of which were inaccessible to the birds, I showed that RCW accessibility influences fungal community development in excavations. This experimental evidence demonstrates that the relationship between RCWs and fungal communities is a multipartite symbiosis may be mutualistic. Finally, by tracking fungal community development in experimental cavity starts through time, I also demonstrated that the fungal communities found in RCW excavations undergo succession, and that this process is influenced by the birds. The relationships described in this body of work provide the basis for future studies on cavity excavators and fungi, and also have implications for a diverse community of secondary cavity nesters, wood-inhabiting fungi, forest ecology, and the conservation of biodiversity. / Ph. D.

Cavity-nesting communities are structured by the creation of and competition for cavities as nest-sites. Viewing these communities as interconnected webs can help identify species interactions that influence community structure. This study examines cavity-nesting bird community interactions within the fire-maintained longleaf pine (Pinus palustris) ecosystem at Eglin Air Force Base, Florida. In chapter 1, I provide a background review of the ecology of my study system. In chapter 2, I use nest webs to depict the flow of cavity-creation and use at Eglin. I identified 2 webs into which most species could be placed. One web contained 6 species associated with pines. The second web contained 5 species associated with hardwoods. Red-cockaded woodpeckers (Picoides borealis) and northern flickers (Colaptes auratus) created most cavities used by other species within this community. In chapter 3, I describe snag densities and nest-site selection of the cavity-nesting bird community at Eglin. Large, mature pine snags were abundant, exceeding other reported densities for southern pine forests. Pine snags were heavily-used, despite the abundance of available red-cockaded woodpecker cavities in living pine. Hardwood snags accounted for 10% of nests found, and were used by 12 of 14 species. Diameters of nest-trees and available snags were below the range of optimal nest-snag diameters reported in other studies, indicating the need for site-specific snag management guidelines. In chapter 4, I combine a study of basic ecological principles with endangered species management to examine interactions within the cavity-nesting bird community at Eglin. I used a nest web to identify a potential indirect interaction between the red-cockaded woodpecker and large secondary cavity-nesters, mediated by the northern flicker. I used structural equation modeling to test a path model of this interaction. By experimentally manipulating cavity availability, I blocked links described in the model, confirming cavity creation and enlargement as mechanisms that influence this indirect relationship. I demonstrated that a red-cockaded woodpecker cavity-management technique could disrupt this indirect relationship by affecting northern flicker behavior, and provided an empirical example of how, in interactive ecological communities, single-species management can have indirect effects on non-target species. / Ph. D.