Over three consecutive breeding seasons I examined the breeding system of Eastern Kingbirds in Malheur National Wildlife Refuge in SE, Oregon. To assess genetic paternity, I optimized seven microsatellite markers and determined that extra pair paternity was common. Extra-pair young were present in approximately 60% of nests, representing 47% of young. I examined implications of extra-pair paternity from three perspectives. First, I examined the impact of extra-pair paternity on the opportunity for sexual selection within a socially monogamous system and identified correlates of male reproductive success. Some males were successful at the expense of others, and 24% of males did not sire any young. Thus, variance in male reproductive success was more than nine times greater than female variance, indicating that sexual selection operates strongly in this system. Extra-pair success was positively related to tarsus length and inversely dependent upon dawn song start time. Within-pair success was positively related to nest defense intensity during the incubation period. Second, I assessed the four genetic hypotheses proposed to explain female cooperation in extra-pair copulations. Extra-pair young were not distributed randomly among broods. Cuckolded within-pair males were not more closely related to females than were extra-pair sires. Pair wise comparisons indicated males with specific song and morphological traits were chosen as sires. Brood success (proportion of brood to fledge and recruit) was unrelated to thenumber of males who sired young in the brood. Extra-pair young were larger and heavier than their within-pair half-siblings. These results unambiguously support the 'good genes' hypothesis of female extra-pair mating. Last, I examined the influence of breeding date and parental condition (mother, social father and genetic sire) on offspring sex ratio. Given that male variance in reproductive success is much greater than that of females, I predicted the characters of genetic sires would be the most important factor influencing offspring sex ratio. Contrary to my prediction, male traits did not influence offspring sex ratio. However, more males hatched early in the season and females in better condition produced more sons. I also found that male nestling survival to the next breeding season was inversely related to date.

Eastern Kingbirds (Tyrannus tyrannus) breed from coast to coast in North America and build open-cup nests in trees. They have been extensively studied across most of their range and have only on occasion been documented to reuse a nest from a previous season. However, at Malheur National Wildlife Refuge (MNWR), located in southeastern Oregon, ~10 % of female Eastern Kingbirds reuse old nests of mainly American Robins (Turdus migratorius). In an attempt to address why nest reuse is so common at MNWR, I used artificial nests to evaluate two hypotheses as to why nest reuse is common in this breeding population. The first hypothesis states that Eastern Kingbirds reuse nests to save time and/or energy (TES) and the second one states nest reuse occurs because there is a shortage of suitable nest sites (NSS). I was able to reject the TES hypothesis because artificial nests provided no apparent reproductive benefits to Eastern Kingbirds, except that if a nest had failed it took less time to lay a replacement clutch after an initial failure if an artificial nest was used instead of building a new nest. A more reasonable explanation is that Eastern Kingbirds face a limited availability of suitable nest sites. With this in mind, I took vegetation measurements to address the hypothesis that Eastern Kingbirds make adaptive choices when selecting a nest site, in which case they would choose sites that increase their probability of breeding successfully. Successful nests, both natural and artificial, were placed higher in a tree and on a steeper angled nest branch than their failed counterparts. Those findings suggest that Eastern Kingbirds make adaptive choices when selecting a nest site.

This dissertation presents the results of a study that I undertook to better understand the breeding biology of Eastern Kingbirds (hereafter, kingbirds) at Malheur National Wildlife Refuge in southeastern Oregon from 2003 to 2009. Kingbirds are long-distance migratory songbirds that breed across much of North America. This species is socially monogamous but, via frequent extra-pair copulations, is genetically polygamous. Kingbirds exhibit relatively high breeding site fidelity, often returning to the same tree to nest in subsequent years. Both members of a pair provide parental care, but there are often specific duties performed by both male and female kingbirds. For example, males typically perform much of the vigilant nest defense that this species is well known for and contribute, to an extent, to the feeding of nestlings. Females, on the other hand, are entirely responsible for incubating and brooding. Beginning in 2002, most adults within the population of kingbirds at Malheur had been banded with a unique combination of three colored plastic leg bands and an aluminum USFWS band. Also, as many nests were located in each year as possible, and any young that survived to fledging age were banded with a unique combination of leg bands. Considerable effort was then spent each year to locate as many banded individuals as possible, which allowed me to document adult and juvenile survival. Annual survival rates of adult male and female kingbirds in the population at Malheur did not differ and were relatively high at approximately 0.65. Juvenile survival rate was approximately 0.29, indicating that slightly less than one-third of nestlings survive the interval between fledging and their first potential breeding season. Resighting probability was high for both sexes, although higher for adult males (0.94) than for adult female (0.84). The latter finding is consistent with the higher site fidelity of males than of females. Resighting probabilities for juveniles were much lower at 0.68 and 0.40 for males and females, respectively. Again, this was expected because natal site fidelity is typically much lower than breeding site fidelity. Compared to most other reports, resighting probability and return rates of juvenile kingbirds was high, presumably because the riparian habitats where kingbirds breed at Malheur function as an ecological island surrounded by, for kingbirds, unsuitable high desert habitat. Thus, unless they disperse very long distances, the only option for juvenile kingbirds is to begin nesting on the refuge. The collection of blood samples from birds has become an increasingly common practice in ornithology. The data that can be gained from these samples allow a number of interesting questions to be asked such as understanding the genetic mating system of a species, patterns of hormone secretion, and discerning migratory pathways via stable isotope analysis. The volume of blood collected is usually small and was assumed to cause no long-term negative effects on sampled individuals. However, few studies have rigorously examined the effect of blood sampling on survival. I used a multistate mark-recapture analysis to assess the effect blood sampling on annual survival of kingbirds by combining the annual survival data described above with whether or not individuals were subjected to blood sample collection. The results of this analysis indicated that blood sampling had no effect on annual survival rates of kingbirds. Whether or not this is the case for other species remains to be seen. However, my results support the assumption that when done correctly, blood sampling has little to no long-term negative effects on birds. Comparative analyses of many bird species show that testosterone secretion exhibits fairly predictable patterns among breeding birds of different mating systems. Monogamous species reach a peak during mate attraction and the period of female fertility which is then followed by a sharp decline when young are in the nest. By contrast, males of polygynous species tend to maintain higher levels of testosterone throughout the breeding season to, presumably, maximize opportunities for acquiring additional mates. Kingbirds are socially monogamous but cryptically polygamous. However, because of high rates of extra-pair paternity, variation in reproductive success among males is much higher than what is expected for a monogamous species, and, instead, is similar to what has been reported for polygynous species. Therefore from 2005 to 2009 I measured testosterone concentrations from the blood samples collected from male kingbirds to determine the breeding season profile of testosterone secretion in kingbirds and to understand the factors that influence testosterone variation among individuals. Contrary to expected for a monogamous species, the testosterone profile of kingbirds did not exhibit the brief peak in testosterone followed by a precipitous decline. Testosterone peaked early in the season, but declined very slowly as the nest cycle progressed. I attributed this gradual decline in testosterone to the cryptically polygynous nature of the kingbird extra-pair mating system. I also found substantial variation in testosterone concentration among male kingbirds and was able to identify several factors contributing to this variation. As expected, testosterone declined as the breeding season progressed (independent of stage in nest cycle) and nest density increased, while increases in testosterone were correlated with the number of fertile females within the population. This suggests that male kingbirds were capable of modulating testosterone concentrations to both cope with an increase in conspecific density, but also to ready themselves for times when extra-pair copulations were more likely.

We have been studying the population of Eastern Kingbirds breeding in riparian habitats in Malheur National Wildlife Refuge (eastern Oregon) since 2002. These efforts have created an ideal research environment wherein most adults in the population have been color banded and DNA sampled and, as part of other research projects, nearly all broods since 2003 have been paternity tested. I decided to use behavioral video recordings of parental nest behavior undertaken between 2003 and 2010 for two unrelated projects. First, I tested the effectiveness of video sampling nesting behavior (see below and chapter 2). Second, I tested whether male kingbirds were able to affect their level of paternal investment in accordance with their level or realized paternity (see below and chapter 3). Chapter 2 was split into three distinct questions: 1) are parental nesting behaviors repeatable, 2) is a one hour sample sufficient to capture variability in these behaviors, and 3) is the first hour of recording sufficient to capture variability in these behaviors. I found overwhelming evidence that the behaviors I measured were repeatable. This is truly important, for if repeatability was disproven, it would call into question the use of sampling throughout the field of animal behavior. I similarly found strong evidence that a one hour sample was sufficient to capture variability in parental behaviors. From this I was able to suggest that further sampling effort would be better spent increasing sample size rather than observation length. Testing whether the first hour of recording was sufficient to capture variability in parental behavior found more muddled results. While there generally was correlation between behavioral values in the first hour and those over a longer observation period, most behaviors were found to have significantly lower values in the first hour. I tested whether this was the result of a lingering observer effect or a natural effect of time of day and concluded that an observer effect was the more likely explanation. In chapter 3, I ran one of the more in depth and complicated tests for a relationship between paternal investment and realized paternity that I was able to find in the literature. I used the standard male feeding rate as a measure of male investment as well as a far more nuanced measure derived from the first Eigenvector of an analysis of six different paternal behaviors. These were both tested using Akaike's Information Criterion against a number of variables likely to affect parental behavior, including realized paternity. Ultimately, I found no evidence that kingbirds were able to affect paternal investment in response to lost paternity, and conclude that they likely had no means to assess realized paternity within the nest of their social mate. However, from the other parameters tested I was able to find that males increased their investment as brood size increased and as the female spent more time attending, but not feeding, the nestlings. Also, male investment decreased as territory density increased and flight feather length increased. Theory surrounding the variability of extrapair paternity in birds can be used to account for these results. In a denser population males are predicted to reduce paternal effort for one of two reasons: increased likelihood of paternity loss in his own nest and an increased investment in pursuing extrapair copulations of his own. Longer flight feathers are a sign of age and quality in kingbirds, so males with this trait are expected to spend more time pursuing extrapair copulations as they are more likely to be successful in this endeavor.