The Cerulean Warbler (Dendroica cerulea) is a Neotropical migrant bird species which is experiencing severe population declines. This study fills in gaps in the information available concerning Cerulean Warbler territoriality and breeding in a site in southeastern Indiana. During the summers of 2002 and 2003, 51 territories were mapped, with an average territory size of 0.21 hectares. Differences existed between territories and random sites for canopy cover, slope, canopy height, number of trees, diameter at breast height (DBH), the number of trees between 3 - 7.9 cm DBH, the number of trees between 8 - 14.9 cm DBH, the number of trees between 15 - 22.9 cm DBH, and the number of trees > 38 cm DBH. Nest productivity was very low in the study area, suggesting that Big Oaks National Wildlife Refuge was a sink population of Cerulean Warblers during the two years of the study. / Department of Biology

The Cerulean Warbler is a small migratory that is currently considered a species of special concern. Cerulean Warbler response to two types of silvicultural treatments in southern Indiana was assessed by comparing relative abundance and territory data collected during the two years prior to tree harvest to data collected one year immediately following the harvest. ArcGIS was then used to identify spatial attributes of territories that appeared to be important to the species. Lastly, a method of sampling canopy openings by using a Global Positioning System (GPS) unit was introduced. The results suggest that Cerulean Warblers were not negatively affected by the harvest and that moderate prescribed silviculture treatments that create small canopy openings can benefit the species. / Immediate Cerulean warbler response to silviculture in southern Indiana -- Effects of silviculture on spatial characteristics of Cerulean warbler territories -- A method of sampling canopy openings associated with Cerulean warbler territories. / Department of Biology

Preservation of large tracts of habitat is often recommended for long-term population viability of area-sensitive species. Large tracts may not always be available. Smaller patches, though not able to contain a viable population individually, may contribute to overall regional population viability if within the small patches pairs could successfully reproduce. By definition, area-sensitive species should have a minimum patch size threshold of habitat below which they will not likely reproduce. Two potential causes for positive relationships between patch size and production are inverse relationships between patch size and brood parasitism and patch size and food availability. My objectives were (1) to determine the minimum patch size thresholds of reproductive success for golden-cheeked warblers (Dendroica chrysoparia), black-and-white warblers (Mniotilta varia), and white-eyed vireos (Vireo griseus); (2) to determine whether thresholds for occupancy, territory establishment by males, or pairing success were indicative of thresholds of reproduction; (3) to determine whether the proportion of pairs fledging brown-headed cowbird (Molothrus ater) young was related to patch size, and (4) to determine the affects of patch size on food availability (i.e., arthropod abundance). The Vickery index of reproductive activity was used to determine reproductive activity of each male or pair and to quantify parasitism occurrences. I collected arthropods using branch clipping to assess the relationship between patch size and arthropod abundance. I found minimum patch size thresholds of reproductive success for golden-cheeked and black-and-white warblers, but not for white-eyed vireos. Minimum patch size of reproductive success was between 15 and 20.1 ha. Minimum patch size thresholds for occupancy, territory establishment by males, and pair formation were not consistent with thresholds for reproductive success. I found no relationships between patch size and cowbird parasitism or patch size and arthropod biomass. Conservation practices for target species based on thresholds of occupancy, territory establishment, or pair formation may not address issues of reproduction. The ability to identify thresholds of reproductive success for target species could be useful in conservation and management in multiple ways including setting goals for retention and restoration of a target species’ habitat patch size.

Preservation of large tracts of habitat is often recommended for long-term population viability of area-sensitive species. Large tracts may not always be available. Smaller patches, though not able to contain a viable population individually, may contribute to overall regional population viability if within the small patches pairs could successfully reproduce. By definition, area-sensitive species should have a minimum patch size threshold of habitat below which they will not likely reproduce. Two potential causes for positive relationships between patch size and production are inverse relationships between patch size and brood parasitism and patch size and food availability. My objectives were (1) to determine the minimum patch size thresholds of reproductive success for golden-cheeked warblers (Dendroica chrysoparia), black-and-white warblers (Mniotilta varia), and white-eyed vireos (Vireo griseus); (2) to determine whether thresholds for occupancy, territory establishment by males, or pairing success were indicative of thresholds of reproduction; (3) to determine whether the proportion of pairs fledging brown-headed cowbird (Molothrus ater) young was related to patch size, and (4) to determine the affects of patch size on food availability (i.e., arthropod abundance). The Vickery index of reproductive activity was used to determine reproductive activity of each male or pair and to quantify parasitism occurrences. I collected arthropods using branch clipping to assess the relationship between patch size and arthropod abundance. I found minimum patch size thresholds of reproductive success for golden-cheeked and black-and-white warblers, but not for white-eyed vireos. Minimum patch size of reproductive success was between 15 and 20.1 ha. Minimum patch size thresholds for occupancy, territory establishment by males, and pair formation were not consistent with thresholds for reproductive success. I found no relationships between patch size and cowbird parasitism or patch size and arthropod biomass. Conservation practices for target species based on thresholds of occupancy, territory establishment, or pair formation may not address issues of reproduction. The ability to identify thresholds of reproductive success for target species could be useful in conservation and management in multiple ways including setting goals for retention and restoration of a target species’ habitat patch size.

Mating behavior has a profound impact on reproductive success and the resulting genetic structure of offspring. Extra-pair copulation is a widely observed behavior within avian species. This study explored the genetic effects of mating behaviors of Prothonotary warblers, Protonotaria citrea (Parulidae), using co-dominant microsatellite markers. Prothonotary warblers are migratory songbirds that build nests in cavities, commonly found in wetland habitats. A set of artificial nest boxes were initiated by Dr Robert Reilly in 2002 in Dutch Gap, Chesterfield County Virginia, USA, a tidal tributary off the James River. From this population, 28 nest boxes were surveyed yielding 47 adults and 110 offspring. All individuals were genotyped and the multilocus genotypes were used to identify parentage. Using paternity exclusion, 27.2% of offspring were identified as resulting from extra-pair mating. Surprisingly, an additional 11.8% of offspring were classified as genetically unrelated to the resident female, being presumably the result of an egg dumping from a female not caught at the nest box. The vast majority of nest boxes, 82.1% in this study, had at least one offspring that was a result of a mating behavior outside of the social pair. There was also positive spatial autocorrelation in extra-pair paternity suggesting that these matings are not randomly distributed across the sampling landscape. These data show that Protonotaria citrea engage in both extra-pair copulations and nest parasitism. Whether this is normal mating behavior for the species, or something that is unique to this population is unknown.

Many organisms use both vegetation structure and social cues in selecting habitats. Many species of songbirds use the presence of breeding conspecifics as a social cue and sign of habitat quality, and can be induced to settle in unoccupied habitats by artificially broadcasting breeding song, a process referred to as “conspecific attraction”. In our study, we tested response to conspecific attraction a low-density population of the threatened Golden-winged Warbler (Vermivora chrysoptera) in Highland County, VA. Response to broadcast song was observed, with a modest increase in mean male abundance at survey points within 250 meters of treatments, and mean abundance decreasing outside of this range throughout the study area. Recruitment to conspecific playback was lower in our study than observed in previous research on high-density systems. Our results suggest more research is needed of the effectiveness of conspecific attraction in low density species and that its use should be tailored to the spatial and demographic conditions of the managed population.

As Neotropical migratory birds, Prothonotary Warblers are exposed to parasites in both tropical and temperate regions and may act as dispersal agents between geographic areas. This study identifies the prevalence of Haemosporidia, West Nile Virus (WNV), and St. Louis Encephalitis virus (SLEV) in this species. A total of 71.6% of captured Prothonotary Warblers were infected with Haemosporidia during the 2008 breeding season, and infection prevalence increased throughout the season. This temporal change in prevalence is likely due to infection relapse and transmission of new infections. No correlations between reproductive effort and infection status were observed, nor were any associations between infection prevalence and nest box location identified. WNV and SLEV were present in 37.5% and 6.3% of sampled Prothonotary Warblers, respectively. These results warrant more detailed analyses of pathogen transmission dynamics in this population, physiological mechanisms that affect infection susceptibility, and spatial and temporal trends in infection that may exist.

Oak wilt is a fatal disease of oaks caused by the fungus Ceratocystis fagacearum. Loss or degradation of habitat due to the disease may negatively affect the federally endangered golden-cheeked warbler (Setophaga chrysoparia). To assess the impact of oak wilt on golden-cheeked warblers, I investigated its influence on habitat selection and quality. I used remote sensing to estimate the amount of potential golden-cheeked warbler habitat currently affected by oak wilt, to predict the amount of potential habitat likely to be affected in the near future, and to assess the current probability of warbler occupancy in areas affected by oak wilt historically. I also quantified vegetative characteristics to assess overstory vegetation and regeneration in areas affected by the disease. I found proportional occupancy and territory density in unaffected areas to be, respectively, 3.5 and 1.8 times that of affected areas. Pairing success was 27% lower for territories containing oak wilt but fledging success was not affected. I estimated that 6.9% of potential golden-cheeked warbler habitat and 7.7% of the total area within my study region was affected by oak wilt in 2008. By 2018, I predicted that 13.3% of potential golden-cheeked warbler habitat and 16.0% of the study region would be affected by the disease. Using historical imagery, I found that areas affected by oak wilt in the past are less likely to be classified as current potential warbler habitat than areas never affected by the disease. I found no differences between the understory vegetation of affected and unaffected areas but that oaks were more common in the overstory than in the understory, suggesting that species composition in affected areas may shift in the years following an outbreak of the disease. My results suggest that the presence of oak wilt negatively influences habitat selection and quality for golden-cheeked warblers, likely due to reduced canopy cover in susceptible oak species. Additionally, oak wilt frequently occurs in golden-cheeked warbler habitat and will continue to spread into warbler habitat in the coming years. Future management efforts should address the threat oak wilt poses to golden-cheeked warblers by incorporating applicable preventative measures.

Several species of North American migratory songbirds reportedly experience seasonal diet shifts involving a shift from an insect diet during the breeding season to one incorporating fruits during migration and non-breeding periods but the extent to which dietary plasticity occurs in migratory songbirds is poorly quantified. Thus, I used stable carbon (ä13C) and nitrogen (ä15N) isotope analyses to evaluate the timing and extent of frugivory throughout the annual life cycle of 16 species of migratory songbirds, representing wide ranges in body size and reported diets. Birds were sampled during spring and fall migration at the Delta Marsh Bird Observatory in 2003. To investigate dietary patterns, I sampled multiple tissues (muscle, liver, whole blood, claws, bone collagen, feathers) as these represent different periods of diet integration due to varying elemental turnover rates.<p>Assuming that relatively low ä15N values represent a fruit diet and relatively high ä15N values represent an insect diet, I expected tissues representing fall migration (liver, blood, and muscle from fall-captured birds) and winter (greater coverts and claws from spring-captured birds) to have lower ä15N values than tissues representing spring migration (liver, blood, and muscle from spring-captured birds) and summer (tail feathers and claws from fall-captured birds) when fruits are presumed to be less common in songbird diets. Based on blood and claw ä15N values, there was no isotopic segregation of species I classified a priori as insectivores or omnivores. For most species, tissue ä15N values showed either no seasonal change or a shift opposite to my prediction (e.g., ä15N values higher in fall birds compared to spring birds). Boreal fruit ä15N values were lower than those for insects; however, ä15N values of agricultural fruits overlapped both boreal fruit and insect values suggesting that food web baselines did not conform to a simple (single) linear trophic-enrichment model. In Yellow-rumped Warblers (Dendroica coronata), within-tissue seasonal comparisons for liver, muscle and blood indicated a fruit diet during fall and winter and an insect diet during spring and summer; claws and feathers of birds captured in spring (representing winter diet) had unexpectedly high ä15N values. Diet-tissue isotopic discrimination factors associated with both a fruit diet and insect diet were taken from the literature and used to correct stable isotope values of tissues to putative diet because, currently, little is known about the nature of factors influencing discrimination factors to be used in simple linear dietary mixing models. There were differences in tissue ä13C and ä15N values depending on which discrimination factor was used. Based on mixing model results for tissue ä15N values, a higher proportion of insects vs. fruits was detected in the diet of Yellow-rumped Warblers for all tissues except muscle and claws. <p>My interpretations are contingent on the fact that the available natural history information, on which guild classifications were based, was correct and that elemental turnover rates and discrimination factors used were accurate. However, much uncertainty remains about the appropriate diet-tissue isotopic discrimination factors corresponding to fruit and insect diets. Due to extensive natural variability of stable nitrogen isotope values in food sources, possible anthropogenic influences and a lack of knowledge of the metabolic processes that can potentially affect stable isotope values, I caution against using stable isotope analysis alone to track frugivory in temperate North American migratory songbirds. Future research should focus on captive studies aimed at determining and validating discrimination factors of various tissues, particularly claws and feathers, for birds feeding on varying proportions of fruits and insects. Additionally, more information on the dietary habits of these migratory songbirds is needed, as previous estimates of insectivory and frugivory in songbirds may not be accurate.

Several species of North American migratory songbirds reportedly experience seasonal diet shifts involving a shift from an insect diet during the breeding season to one incorporating fruits during migration and non-breeding periods but the extent to which dietary plasticity occurs in migratory songbirds is poorly quantified. Thus, I used stable carbon (ä13C) and nitrogen (ä15N) isotope analyses to evaluate the timing and extent of frugivory throughout the annual life cycle of 16 species of migratory songbirds, representing wide ranges in body size and reported diets. Birds were sampled during spring and fall migration at the Delta Marsh Bird Observatory in 2003. To investigate dietary patterns, I sampled multiple tissues (muscle, liver, whole blood, claws, bone collagen, feathers) as these represent different periods of diet integration due to varying elemental turnover rates.<p>Assuming that relatively low ä15N values represent a fruit diet and relatively high ä15N values represent an insect diet, I expected tissues representing fall migration (liver, blood, and muscle from fall-captured birds) and winter (greater coverts and claws from spring-captured birds) to have lower ä15N values than tissues representing spring migration (liver, blood, and muscle from spring-captured birds) and summer (tail feathers and claws from fall-captured birds) when fruits are presumed to be less common in songbird diets. Based on blood and claw ä15N values, there was no isotopic segregation of species I classified a priori as insectivores or omnivores. For most species, tissue ä15N values showed either no seasonal change or a shift opposite to my prediction (e.g., ä15N values higher in fall birds compared to spring birds). Boreal fruit ä15N values were lower than those for insects; however, ä15N values of agricultural fruits overlapped both boreal fruit and insect values suggesting that food web baselines did not conform to a simple (single) linear trophic-enrichment model. In Yellow-rumped Warblers (Dendroica coronata), within-tissue seasonal comparisons for liver, muscle and blood indicated a fruit diet during fall and winter and an insect diet during spring and summer; claws and feathers of birds captured in spring (representing winter diet) had unexpectedly high ä15N values. Diet-tissue isotopic discrimination factors associated with both a fruit diet and insect diet were taken from the literature and used to correct stable isotope values of tissues to putative diet because, currently, little is known about the nature of factors influencing discrimination factors to be used in simple linear dietary mixing models. There were differences in tissue ä13C and ä15N values depending on which discrimination factor was used. Based on mixing model results for tissue ä15N values, a higher proportion of insects vs. fruits was detected in the diet of Yellow-rumped Warblers for all tissues except muscle and claws. <p>My interpretations are contingent on the fact that the available natural history information, on which guild classifications were based, was correct and that elemental turnover rates and discrimination factors used were accurate. However, much uncertainty remains about the appropriate diet-tissue isotopic discrimination factors corresponding to fruit and insect diets. Due to extensive natural variability of stable nitrogen isotope values in food sources, possible anthropogenic influences and a lack of knowledge of the metabolic processes that can potentially affect stable isotope values, I caution against using stable isotope analysis alone to track frugivory in temperate North American migratory songbirds. Future research should focus on captive studies aimed at determining and validating discrimination factors of various tissues, particularly claws and feathers, for birds feeding on varying proportions of fruits and insects. Additionally, more information on the dietary habits of these migratory songbirds is needed, as previous estimates of insectivory and frugivory in songbirds may not be accurate.