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Individual variation in behaviors such as aggression or boldness has the potential to influence the survival and reproductive success of individual animals. For example, individual variation in boldness means that some individuals will be more explorative and risk-tolerant than others, which could allow them to be more active in the face of danger or disturbance. Individual variation in boldness has been studied in many species, but fewer studies have examined patterns of boldness in mated pairs of animals. For many seabird species and some songbirds, mates tend to have similar levels of boldness and other personality characteristics. This suggests that coordination and decreased conflict between mated parents may allow for more effective care of young, leading to increased reproductive success. Boldness in male song sparrows (Melospiza melodia) has been shown to be both variable among individuals and repeatable for individuals, but similar work has not been conducted on females of the species. To determine if song sparrows within mated pairs have similar, different, or unrelated levels of boldness, we conducted Carolina wren (Thryothorus ludovicianus) alarm call playback experiments, simulating the presence of a predator, on the territories of 30 song sparrow pairs. At 24 of these territories, we conducted at least two playback trials to determine if boldness is repeatable for males and females. In a second study of the ability of song sparrow pairs to coordinate their boldness responses, we used playback experiments of song sparrow alarm calls to investigate whether males can distinguish between the calls of a stranger and their mate. Boldness and male response to a mate versus stranger were measured as the average distance from the playback speaker and the number of calls produced by each subject bird. Boldness, as measured by both distance from the speaker and number of calls, was repeatable for individuals and variable among individuals for males and females. M

As urban development continues to dominate landscapes across the United States, wildlife species are changing their behavior in many ways. Urban animals are often bolder, or less fearful of new stimuli, than rural animals. While natural selection and adaptation can drive behavioral changes in urban animals, other factors, such as learning or habituation, can also lead to behavioral modifications. Habituation may play an important role, if urban individuals learn through repeated exposure that humans do not represent a threat. To determine how repeated human exposure affects song sparrow (Melospiza melodia) populations in an urban and rural habitat, I measured boldness as flight initiation distance (FID), after repeated exposure to a nonthreatening human subject. FID measures the distance between human and bird, at which the bird flies away in fear of disturbance or predation. I collected FID data on male song sparrows in an urban and rural environment over 5 consecutive days. I found that after 5 days of successive trials, FID was individually repeatable in both urban and rural populations. Urban birds had consistently lower FID than their rural counterparts. I also found that FID went down over 5 days of repeated trials in the rural populations, but not the urban. These results suggest that habituation can occur quickly in rural birds and account for the greater boldness we typically see in urban populations.

Urbanization is a critical form of environmental change that can affect the physiology and behavior of wild animals and, notably, birds. One behavioral difference between birds living in urban and rural habitats is that urban males show elevated boldness or territorial aggression in response to simulated social challenge. This pattern has been described in several populations of song sparrow, Melospiza melodia. Such behavioral differences must be underpinned by differences in the brain, yet little work has explored how urbanization and neural function may be interrelated. We explored the relationship between urbanization and neural activation within a network of brain regions, collectively called the social behavior network, which contributes to the regulation of territorial aggression. Specifically, we captured free-living, territorial male song sparrows by playing them conspecific songs for 6-11 min, and then collected their brains. We estimated recent neural activation, as indicated by the immediate early gene FOS, and measured levels of a neuropeptide, arginine vasotocin (AVT), which is involved in the regulation of social behavior. Based on previous studies we expected urban males, which are generally more territorially aggressive, to have lower FOS expression in a node of the social behavior network implicated in regulating territoriality, the lateral septum (LS). Additionally, we expected urban males to have lower AVT expression in a brain region involved in the regulation of sociality, the medial bed nucleus of the stria terminalis (BSTm). We found that, compared to rural males, urban male song sparrows did have lower FOS expression in the LS. This pattern suggests that lower neural activation in the LS could contribute to behavioral adjustments to urbanization in male song sparrows. Additionally, counter to our predictions, urban male song sparrows had higher AVT-like immunoreactivity in the BSTm. Future work building upon these findings is needed to determ

Urbanization is a critical form of environmental change that can affect the physiology and behavior of wild animals and, notably, birds. One behavioral difference between birds living in urban and rural habitats is that urban males show elevated boldness or territorial aggression in response to simulated social challenge. This pattern has been described in several populations of song sparrow, Melospiza melodia. Such behavioral differences must be underpinned by differences in the brain, yet little work has explored how urbanization and neural function may be interrelated. We explored the relationship between urbanization and neural activation within a network of brain regions, collectively called the social behavior network, which contributes to the regulation of territorial aggression. Specifically, we captured free-living, territorial male song sparrows by playing them conspecific songs for 6-11 min, and then collected their brains. We estimated recent neural activation, as indicated by the immediate early gene FOS, and measured levels of a neuropeptide, arginine vasotocin (AVT), which is involved in the regulation of social behavior. Based on previous studies we expected urban males, which are generally more territorially aggressive, to have lower FOS expression in a node of the social behavior network implicated in regulating territoriality, the lateral septum (LS). Additionally, we expected urban males to have lower AVT expression in a brain region involved in the regulation of sociality, the medial bed nucleus of the stria terminalis (BSTm). We found that, compared to rural males, urban male song sparrows did have lower FOS expression in the LS. This pattern suggests that lower neural activation in the LS could contribute to behavioral adjustments to urbanization in male song sparrows. Additionally, counter to our predictions, urban male song sparrows had higher AVT-like immunoreactivity in the BSTm. Future work building upon these findings is needed to determ

Urban areas are a unique and growing habitat type. Animals living in this novel habitat are faced with new challenges, but may also encounter novel opportunities. Though urban animals have been observed to differ from their rural counterparts in a variety of behavioral and physiological traits, little is known about the specific features of urban areas that drive these differences and whether they are adaptive. Understanding this process is important from a conservation perspective and also to gain insight into how animals colonize novel habitats more generally. Using song sparrows (Melospiza melodia), a native songbird commonly found in urban areas, I explored responses to urbanization and the drivers and consequences of these responses with an eye toward understanding whether song sparrows had successfully adapted to urban habitats (Chapter I). I began by comparing body condition and levels of corticosterone, a hormone associated with energy management and the stress response in birds, between urban and rural populations (Chapter II). There was more variation across years than between habitats, suggesting that a variable environmental factor common to both habitats is the primary driver of these traits. I then compared territorial aggression levels and tested the effect of food availability on aggression (Chapter III). Fed rural birds and all urban birds had higher aggression levels than unfed rural birds, indicating that territorial aggression is related to resource availability in this species and that urban habitats may be perceived as more desirable. Finally, I looked for differences in reproductive timing and success and for relationships between reproductive success and aggression (Chapter IV). Higher reproductive success in urban populations, coupled with differences in the timing of successful nests between habitats, suggest differences in predation risk and predator community structure between habitats. In Chapter V, I synthesize my major findings and sug

Understanding the characteristics of high-quality avian habitat is critical for guiding salt marsh management and restoration. Existing insights into salt marsh avian habitat are often based on the composition of marsh vegetation, e.g., individual plant species cover. This study investigated whether the spatial configuration of marsh surface cover (e.g., patch number, density, size, shape complexity and compactness, degree of dissection of the landscape, variation and repetition of cover type, and the variance within these metrics) is a useful, additional indicator of avian habitat quality for the Alameda Song Sparrow (Melospiza melodia pusillula), a non-migratory California Species of Special Concern endemic to southern San Francisco Bay. M. m. pusillula density during the breeding seasons of 2002 through 2005 was estimated at 82 observation points in 10 marsh sites within the bird’s geographic range. The mean bird density index (overall mean: 5.61 birds detected per hectare of marsh) was not significantly different among marshes of different ages. We mapped the vegetation zones, open water, and upland areas within each marsh site using high resolution aerial photographs and automated classification analysis. We quantified the configuration of surface cover around each bird observation point by 31 metrics. Bird density index was best modeled by a multiple linear regression containing positive relationships with the metrics Mean Core Area Index and Patch Core Area Coefficient of Variation (R 2= 0.210, p < 0.0001). Qualitatively, this model suggested that M. m. pusillula abundance during the breeding season was greatest in marsh areas with compact patches that spanned a variety of patch sizes from moderate-to-large, uninterrupted by other cover. We conclude that configuration-based vegetation pattern analysis could usefully complement more customary composition-based habitat assessments to aid wetland habitat research, management, and restoration.

Emberizid sparrows (emberizidae) have played a prominent role in the study of avian vocal communication and social behavior. We present here brain transcriptomes for three emberizid model systems, song sparrow Melospiza melodia, white-throated sparrow Zonotrichia albicollis, and Gambel's white-crowned sparrow Zonotrichia leucophrys gambelii. Each of the assemblies covered fully or in part, over 89% of the previously annotated protein coding genes in the zebra finch Taeniopygia guttata, with 16,846, 15,805, and 16,646 unique BLAST hits in song, white-throated and white-crowned sparrows, respectively. As in previous studies, we find tissue of origin (auditory forebrain versus hypothalamus and whole brain) as an important determinant of overall expression profile. We also demonstrate the successful isolation of RNA and RNA-sequencing from post-mortem samples from building strikes and suggest that such an approach could be useful when traditional sampling opportunities are limited. These transcriptomes will be an important resource for the study of social behavior in birds and for data driven annotation of forthcoming whole genome sequences for these and other bird species.

Emberizid sparrows (emberizidae) have played a prominent role in the study of avian vocal communication and social behavior. We present here brain transcriptomes for three emberizid model systems, song sparrow Melospiza melodia, white-throated sparrow Zonotrichia albicollis, and Gambel's white-crowned sparrow Zonotrichia leucophrys gambelii. Each of the assemblies covered fully or in part, over 89% of the previously annotated protein coding genes in the zebra finch Taeniopygia guttata, with 16,846, 15,805, and 16,646 unique BLAST hits in song, white-throated and white-crowned sparrows, respectively. As in previous studies, we find tissue of origin (auditory forebrain versus hypothalamus and whole brain) as an important determinant of overall expression profile. We also demonstrate the successful isolation of RNA and RNA-sequencing from post-mortem samples from building strikes and suggest that such an approach could be useful when traditional sampling opportunities are limited. These transcriptomes will be an important resource for the study of social behavior in birds and for data driven annotation of forthcoming whole genome sequences for these and other bird species.

In birds, male song has been extensively studied, but female vocalizations have received little attention. Females of several North American species produce a unique vocalization, the nest departure call (NDC), upon leaving nests. Producing NDCs has costs due to acoustical properties that make nests easy to locate by predators. Thus, NDCs must also have benefits that balance or outweigh costs, and females should modulate call production as costs and benefits change. We explored whether female song sparrows, Melospiza melodia, adjust calling rate to reflect differential costs and benefits of calling induced by male presence, male quality (measured by body mass and song complexity), nest predator presence and nest height. Results suggest that calls benefit females by promoting male nest guarding and that females display adaptive plasticity in call production. Specifically, calling rate increased when the male was present, and male nest guarding increased when females gave an NDC. Females called less in the presence of a model nest predator, probably because the perceived costs of predator attraction outweighed the benefits of male recruitment. Conversely, females with heavier mates called more, perhaps because the efficacy of male nest guarding increases with mass. In addition, females called more from elevated nests in the presence of the predator and decreased calling later in the day. Male song complexity failed to predict calling rate, suggesting that this sexually selected trait does not reflect direct benefits gained by producing an NDC. Plasticity in calling probably exists because context-appropriate communication elevates fitness, whereas contextual mistakes in the decision to communicate result in fitness declines. © 2014 The Association for the Study of Animal Behaviour.

Understanding the characteristics of high-quality avian habitat is critical for guiding salt marsh management and restoration. Existing insights into salt marsh avian habitat are often based on the composition of marsh vegetation, e.g., individual plant species cover. This study investigated whether the spatial configuration of marsh surface cover (e.g., patch number, density, size, shape complexity and compactness, degree of dissection of the landscape, variation and repetition of cover type, and the variance within these metrics) is a useful, additional indicator of avian habitat quality for the Alameda Song Sparrow (Melospiza melodia pusillula), a non-migratory California Species of Special Concern endemic to southern San Francisco Bay. M. m. pusillula density during the breeding seasons of 2002 through 2005 was estimated at 82 observation points in 10 marsh sites within the bird's geographic range. The mean bird density index (overall mean: 5.61 birds detected per hectare of marsh) was not significantly different among marshes of different ages. We mapped the vegetation zones, open water, and upland areas within each marsh site using high resolution aerial photographs and automated classification analysis. We quantified the configuration of surface cover around each bird observation point by 31 metrics. Bird density index was best modeled by a multiple linear regression containing positive relationships with the metrics Mean Core Area Index and Patch Core Area Coefficient of Variation (R 2= 0.210, p < 0.0001). Qualitatively, this model suggested that M. m. pusillula abundance during the breeding season was greatest in marsh areas with compact patches that spanned a variety of patch sizes from moderate-to-large, uninterrupted by other cover. We conclude that configuration-based vegetation pattern analysis could usefully complement more customary composition-based habitat assessments to aid wetland habitat research, management, and restoration.

Extensive habitat loss and degradation have resulted in decreases in populations of tidal marsh breeding birds in the San Francisco Estuary in the past 150 years. We conducted point count surveys and nest monitoring in tidal marsh habitat at China Camp State Park from 1996 through 2007 to assess bird abundance, species richness and reproductive success over time. We found overall species richness at China Camp to be significantly lower than that of other San Pablo Bay tidal marshes, but also to be increasing during the study period. We present relative density indices and confirm breeding for three focal species that are of conservation concern: San Pablo song sparrow (Melospiza melodia samuelis), California black rail (Laterallus jamaicensis coturniculus), and California clapper rail (Rallus longirostris obsoletus). Song sparrows were observed at higher densities at China Camp than at other San Pablo Bay sites. There was no apparent trend in song sparrow density during the study period at China Camp, in contrast with the rest of San Pablo Bay, which displayed a significant decline. We determined song sparrow nest survival probability using the method described by Mayfield (1975) and found that nest survival at China Camp varied markedly among years, from >30% in 1996 to 6% in 2006, with no discernible trend. The main causes of nest failure were predation and tidal flooding. Song sparrow nests were found predominantly in gumplant (Grindelia stricta) and pickleweed (Sarcocornia pacifica) and less commonly in saltgrass (Distichlis spicata). Clapper rails nested exclusively in pickleweed and black rail nests were found mainly in gumplant and less commonly in saltgrass. China Camp’s expansive ancient marsh supports endemic and bird species of conservation concern, while serving as a reference site for tidal marsh studies in San Francisco Bay. The long-term monitoring of tidal marsh bird breeding parameters at China Camp has the potential to provide early detection of

Extensive habitat loss and degradation have resulted in decreases in populations of tidal marsh breeding birds in the San Francisco Estuary in the past 150 years. We conducted point count surveys and nest monitoring in tidal marsh habitat at China Camp State Park from 1996 through 2007 to assess bird abundance, species richness and reproductive success over time. We found overall species richness at China Camp to be significantly lower than that of other San Pablo Bay tidal marshes, but also to be increasing during the study period. We present relative density indices and confirm breeding for three focal species that are of conservation concern: San Pablo song sparrow (Melospiza melodia samuelis), California black rail (Laterallus jamaicensis coturniculus), and California clapper rail (Rallus longirostris obsoletus). Song sparrows were observed at higher densities at China Camp than at other San Pablo Bay sites. There was no apparent trend in song sparrow density during the study period at China Camp, in contrast with the rest of San Pablo Bay, which displayed a significant decline. We determined song sparrow nest survival probability using the method described by Mayfield (1975) and found that nest survival at China Camp varied markedly among years, from >30% in 1996 to 6% in 2006, with no discernible trend. The main causes of nest failure were predation and tidal flooding. Song sparrow nests were found predominantly in gumplant (Grindelia stricta) and pickleweed (Sarcocornia pacifica) and less commonly in saltgrass (Distichlis spicata). Clapper rails nested exclusively in pickleweed and black rail nests were found mainly in gumplant and less commonly in saltgrass. China Camp’s expansive ancient marsh supports endemic and bird species of conservation concern, while serving as a reference site for tidal marsh studies in San Francisco Bay. The long-term monitoring of tidal marsh bird breeding parameters at China Camp has the potential to provide early detection of

The San Francisco Bay estuary is highly urbanized, but it supports the largest remaining extent of tidal salt marshes on the west coast of North America as well as a diverse native bird community. San Francisco Bay tidal marshes are occupied by more than 113 bird species that represent 31 families, including five subspecies from three families that we denote as tidal-marsh obligates. To better identify the niche of bird species in tidal marshes, we present a review of functional groups based on foraging guilds and habitat associations. Foraging guilds describe the method by which species obtain food from tidal marshes, while habitat associations describe broad areas within the marsh that have similar environmental conditions. For example, the ubiquitous song sparrows (Alameda Melospiza melodia pusillula, Suisun M. m. maxillaris, and San Pablo M. m. samuelis) are surface-feeding generalists that consume prey from vegetation and the ground, and they are found across the entire marsh plain into the upland–marsh transition. In contrast, surface-feeding California black rails (Laterallus jamaicensis coturniculus) are cryptic, and generally restricted in their distribution to the mid- and high-marsh plain. Although in the same family, the endangered California clapper rail (Rallus longirostris obsoletus) has become highly specialized, foraging primarily on benthic fauna within marsh channels when they are exposed at low tide. Shorebirds such as the black-necked stilt (Himantopus mexicanus) typically probe in mud flats to consume macroinvertebrate prey, and are generally restricted to foraging on salt pans within the marsh plain, in ponds, or on mud flats during transitional stages of marsh evolution. The abundance and distribution of birds varies widely with changing water depths and vegetation colonization during different stages of restoration. Thus, tidal-marsh birds represent a rich and diverse community in bay marshes, with niches that may be distinguished by the foo

The San Francisco Bay estuary is highly urbanized, but it supports the largest remaining extent of tidal salt marshes on the west coast of North America as well as a diverse native bird community. San Francisco Bay tidal marshes are occupied by more than 113 bird species that represent 31 families, including five subspecies from three families that we denote as tidal-marsh obligates. To better identify the niche of bird species in tidal marshes, we present a review of functional groups based on foraging guilds and habitat associations. Foraging guilds describe the method by which species obtain food from tidal marshes, while habitat associations describe broad areas within the marsh that have similar environmental conditions. For example, the ubiquitous song sparrows (Alameda Melospiza melodia pusillula, Suisun M. m. maxillaris, and San Pablo M. m. samuelis) are surface-feeding generalists that consume prey from vegetation and the ground, and they are found across the entire marsh plain into the upland–marsh transition. In contrast, surface-feeding California black rails (Laterallus jamaicensis coturniculus) are cryptic, and generally restricted in their distribution to the mid- and high-marsh plain. Although in the same family, the endangered California clapper rail (Rallus longirostris obsoletus) has become highly specialized, foraging primarily on benthic fauna within marsh channels when they are exposed at low tide. Shorebirds such as the black-necked stilt (Himantopus mexicanus) typically probe in mud flats to consume macroinvertebrate prey, and are generally restricted to foraging on salt pans within the marsh plain, in ponds, or on mud flats during transitional stages of marsh evolution. The abundance and distribution of birds varies widely with changing water depths and vegetation colonization during different stages of restoration. Thus, tidal-marsh birds represent a rich and diverse community in bay marshes, with niches that may be distinguished by the foo

The immunocompetence handicap hypothesis proposes that testosterone (T)-dependent sexual signals are honest indicators of male health or genetic quality because only high-quality males are able to withstand the obligate effects of T-induced immunosuppression. In birds, the basic assumption that T suppresses immune function is equivocal, and the physiological mechanisms underlying T-induced immunosuppression remain to be investigated. We explored the proximate pathways of T-induced immunosuppression in song sparrows (Melospiza melodia) by treating captive nonbreeding males with different androgens and measuring several components of acquired immune function. Males implanted with T suppressed cell-mediated and humoral immune responses compared to males implanted with 5proportional to-dihydrotestosterone (DHT), dehydroepiandrosterone, or control ( empty) implants. Furthermore, T treatment increased plasma levels of corticosterone and decreased body mass and fat stores in relation to other treatments. The failure of DHT to depress immune function suggests that T-induced immunosuppression does not occur through a direct pathway because both T and DHT bind to androgen receptors on target cells. Instead, we outline indirect pathways that are likely responsible for suppression of the avian immune system that include stress-induced immunosuppression, aromatization to estrogen, and alterations in energy allocation that constrain expenditures toward immune system activation.

The prevalence and intensity of parasitic infection often increases in animals when they are reproducing. This may be a consequence of increased rates of parasite transmission due to reproductive effort. Alternatively, endocrine changes associated with reproduction can lead to immunosuppression. Here we provide support for a third potential mechanism: reduced immunocompetence as a consequence of adaptive reallocation of resources in times of increased energetic demand. In captive zebra finches Taeniopygia guttata, reproductive effort was manipulated through brood size. Enhanced effort was found to affect the production of antibodies towards sheep red blood cells. In addition, activity of zebra finches was manipulated independently of parental care. Experimentally increased daily workloads in activity reward schedules also suppressed antibody production. Thus, we show that not just the reproductive state, but the increased activity that accompanies reproduction is associated with immunocompetence. This mechanism may be sufficient to explain the increased parasitism observed in reproducing animals. We suggest that reduced immunocompetence as a consequence of increased reproductive effort may be an important pathway for the life history cost of reproduction.

President Roosevelt writes to naturalist Edward Howe Forbush that he has just read Forbush's report from last year on the destruction of birds in New England. Roosevelt asks if Forbush has noticed a change in numbers this spring or summer. Roosevelt has not noticed a difference, with many types of birds "as plentiful as ever," noting he has been observing birds in Oyster Bay for 31 years. Only quail and woodcock numbers seem diminished.

John Lewis Childs sends President Roosevelt details regarding various species of birds and their behaviors. Childs admits he was wrong, and the president right, about the nesting habits of the Long-eared vs. Short-eared owl. Childs describes a trip wherein he showed John Burroughs the nesting sites of a Brown Thrasher, and a Spotted Sandpiper, both incubating eggs. Childs compliments Roosevelt on the accuracy of his observations and description of a flock of passenger pigeons at Pine Knot, especially considering how busy he must be.