Urbanization is viewed as a major threat to global biodiversity because of its role in the loss and fragmentation of low-lying, productive habitats associated with coastal plains and river valleys. My study examines the effects of urbanization on the movements and distribution of songbirds in Calgary, Alberta, Canada. I conducted playback and translocation experiments to assess the permeability of small-scale (e.g. transportation corridors, rivers) and large-scale (e.g. multi-lane expressways, areas of urban development) features of the urban landscape, respectively. I then used these empirical data to parameterize spatially explicit models and determine functional landscape connectivity across the study area. Finally, using point surveys conducted at 183 sites across the urban matrix, I examined the role of land cover type, local vegetation characteristics, landscape-level forest cover, and isolation from natural features on the distribution of songbirds. In 563 playback trials involving the responses of 2241 birds, I found that the size of the gap in vegetation was the most important determinant of movement across linear features; the likelihood of movement sharply decreasing as the gap in vegetation exceeded 30 m. The results of 176 translocation trials provided further evidence of the barrier effect of gaps. Multiple gaps, in particular, constrained the movements of both yellow warblers (Dendroica petechia) and black-capped chickadees (Poecile atricapillus). The bird surveys revealed that natural forest stands played a critical role in sustaining regional avian diversity in the study area. Moreover, functional distance to the nearest forested natural area or water body often explained more variation in the probability of occurrence of focal species than straight-line distance, suggesting that barriers identified from the permeability experiments may have affected not only the movements of songbirds but their settlement patterns as well. Taken together, my results suggest that preserving a functionally connected network of natural areas is vital to conserving avian biodiversity in cities. My research describes novel methodologies for characterizing the composition and configuration of highly heterogeneous and fragmented landscapes. It also provides a unique examination of the link between the movement behaviour of individual birds and population-level distribution patterns within this context. / Ecology

Rapid increases in plasma androgens are generally associated with short-term aggressive challenges in many breeding vertebrates. However, some animals such as song sparrows (Melospiza melodia) are aggressive year-round, even during the non-breeding season, when gonads are regressed and systemic testosterone (T) levels are non-detectable. In contrast, levels of the prohormone dehydroepiandrosterone (DHEA) are elevated year-round in the plasma and brain. The local conversion of brain DHEA to potent androgens may be critical in regulating non-breeding aggression. 3β-hydroxysteroid dehydrogenase/Δ4-Δ5 isomerase (3β-HSD) catalyzes DHEA conversion to androstenedione (AE) and the cofactor NAD⁺ assists in this transformation. In this thesis, I asked whether brain 3β-HSD activity is regulated by social encounters in seasonally breeding male songbirds. In Experiment 1, I looked at the long-term seasonal regulation of brain 3β-HSD activity. 3β-HSD activity was highest in the non-breeding season compared to the breeding season and molt. In Experiment 2, I hypothesized that brain 3β-HSD activity is rapidly regulated by short-term social encounters during the non-breeding season. A 30 min social challenge increased aggressive behavior. Without exogenous NAD⁺, there was ~355% increase in 3β-HSD activity in the caudal telencephalon and ~615% increase in the medial central telencephalon compared to controls (p<0.05). With exogenous NAD⁺, there was no effect of social challenge on 3β-HSD activity. These data suggest that endogenous cofactors play a critical role in the neuroendocrine response to social challenges. The increase in brain DHEA conversion to AE during social challenges may be a mechanism to rapidly increase local androgens in the non-breeding season, when there are many costs of systemic T.

This study shows the importance of the basic spectro-temporal song parameters in territorial contests between male songbirds. Chiffchaff and willow warbler were selected as model species for their phylogenetic and ecological similarity, strong territorial behaviour, but very different singing styles. We found that bigger males of both species sing lower pitched songs and that males of both species adjust their behaviour according to song pitch of their rival simulated by playback. Thus, it seems that both species use song pitch to acoustically assess the body size of their rivals. We also show that song length and syllable rate are important features of the song, probably signalling short-term motivation to escalate territorial conflict.

Rapid increases in plasma androgens are generally associated with short-term aggressive challenges in many breeding vertebrates. However, some animals such as song sparrows (Melospiza melodia) are aggressive year-round, even during the non-breeding season, when gonads are regressed and systemic testosterone (T) levels are non-detectable. In contrast, levels of the prohormone dehydroepiandrosterone (DHEA) are elevated year-round in the plasma and brain. The local conversion of brain DHEA to potent androgens may be critical in regulating non-breeding aggression. 3β-hydroxysteroid dehydrogenase/Δ4-Δ5 isomerase (3β-HSD) catalyzes DHEA conversion to androstenedione (AE) and the cofactor NAD⁺ assists in this transformation. In this thesis, I asked whether brain 3β-HSD activity is regulated by social encounters in seasonally breeding male songbirds. In Experiment 1, I looked at the long-term seasonal regulation of brain 3β-HSD activity. 3β-HSD activity was highest in the non-breeding season compared to the breeding season and molt. In Experiment 2, I hypothesized that brain 3β-HSD activity is rapidly regulated by short-term social encounters during the non-breeding season. A 30 min social challenge increased aggressive behavior. Without exogenous NAD⁺, there was ~355% increase in 3β-HSD activity in the caudal telencephalon and ~615% increase in the medial central telencephalon compared to controls (p<0.05). With exogenous NAD⁺, there was no effect of social challenge on 3β-HSD activity. These data suggest that endogenous cofactors play a critical role in the neuroendocrine response to social challenges. The increase in brain DHEA conversion to AE during social challenges may be a mechanism to rapidly increase local androgens in the non-breeding season, when there are many costs of systemic T. / Science, Faculty of / Zoology, Department of / Graduate

Song production in songbirds is controlled by parts of the brain known as the song control regions (SCRs). During spring, gonads increase in size, males sing to attract mates, and SCRs become larger. This neuroplasticity is controlled by the change in day length and increased plasma testosterone (T) levels. Plasma T can be reduced by stress through the production of corticosterone (CORT), through the production of beta-endorphin, or through direct effects on the testes via the nervous system. We determined the T, estradiol, and CORT hormonal profiles of wild House Finches by capturing and sampling blood from the finches every season for two years. To track SCR neuroplasticity in the wild, we also measured the volume of two specific SCRs, the HVC and RA, every season. We then examined the effects of stress on the finch endocrine system in the wild by performing a 30-minute restraint stress protocol once every season and took blood samples before and after the restraint. To determine whether stress and/or CORT affect neuroplasticity in SCRs, we captured male house finches during winter and brought them into captivity. They were allowed to acclimate to captivity for one month on short days (8L:16D) before we transferred them to long days (16L:8D) and restraint stressed half the birds. We measured their gonads, plasma T and CORT levels, volumes of the HVC and RA, and the number of new neurons in the HVC. HVC volumes were smaller in stressed than non-stressed birds, while RA volumes did not differ. There was no difference in number of new neurons or estimated total number of neurons in the HVC between control and restrained birds. Because the HVC is involved in song production, it is possible that stress negatively impacts singing behavior and reproductive success in House Finches. Future work should address how natural stressors may affect neuroplasticity in birds.

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Background noise is ubiquitous and can impair acoustic communication and influence signaling behavior in animals. Despite evidence demonstrating myriad effects of anthropogenic noise on animal communication, precisely how natural background noise influences communication and behavior remains unclear. Yet, natural sources of background noise, such as rushing rivers or crashing ocean surf, share similar power spectra to sources of anthropogenic noise and can occur at high amplitudes, potentially masking acoustic signals. To investigate the effects of water-generated noise on songbird behavior, we experimentally broadcast landscape-level playbacks of ocean surf and river noise in coastal California, USA, and riparian habitat in Idaho, USA, respectively. In Chapter 1, we conducted a conspecific playback experiment examining how territorial defense behaviors of lazuli buntings (Passerina amoena) and spotted towhees (Pipilo maculatus) vary in response to broadcast water noise. We also incorporated cicada noise from a serendipitous Okanagana spp. emergence as a biotic source influencing lazuli bunting behavior. Both species produce songs that share substantial spectral overlap with low-frequency, water-generated noise, and lazuli bunting song shares an additional high-frequency overlap with cicada calls. Thus, there is potential for background acoustic conditions to mask conspecific signals. We found that detection and discrimination of conspecific playback occurred more slowly for both species as sound level increased. Lazuli buntings also exhibited divergent flight behaviors in response to high- and low-frequency acoustic sources, both dependent and independent of sound level. In Chapter 2, we investigated how amplitude and frequency of water-generated noise influences spectral and temporal song characteristics in six songbird species. We recorded individuals defending territories across 37 sites, with each site representing one of four acoustic environments: naturally quiet ‘controls’, naturally noisy ‘positive controls’ adjacent to the ocean or a whitewater river, ‘phantom’ playback sites with continuous broadcast of low-frequency water noise, and ‘shifted’ playback sites with continuous broadcast of high-frequency water noise. We predicted that all individuals exposed to ‘positive control’, ‘phantom’, or ‘shifted’ noise would adjust song structure, but the magnitude of signal modification would be larger in noisier locations and the type of modification would depend on the spectral profile of the acoustic environment. No two species altered songs in precisely the same way. However, song structure of all six species varied with amplitude and/or frequency of background noise. Together our results demonstrate that natural noise can impair agonistic behaviors and influence vocal structure. These findings suggest that the natural acoustic environment shapes acoustic communication, highlighting natural soundscapes as an under-appreciated axis of the environment.

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