No description available.

Vocal learning is one of the most distinctive characteristics of the modern human species. Through the intricate interaction between vocal motor and auditory systems during early sensitive periods, humans spontaneously master the ability to speak and decode speech. Because vocal learning is so rare in vertebrates, songbirds (Oscines) are the primary model organisms used in studies of acoustic communication and vocal learning. The acquisition of songs in birds and speech in humans (learning of complex sounds with syntactic structures) exhibit similar developmental trajectories. Research on song learning has focused primarily on vocal production with limited emphasis on the role of auditory perception. While auditory tuning and sensorimotor feedback are indispensable for successful vocal learning and communication, how auditory tuning emerges at different levels of the neural processing hierarchy and how sensorimotor integration occurs in the brain during vocal learning is not fully understood. The neurobiology research described here thus focuses on auditory tuning and sensorimotor integration in vocal learning songbirds using multiple experimental approaches. In the first chapter, I describe peripheral auditory tuning in several songbird species. Using operant conditioning, I trained individual birds to report audible tones and assessed hearing thresholds over the 0.5 to 10 kHz frequency range. I also examined the relationship between song spectral energy and hearing by analyzing song frequency-power spectra and audiograms across species. I found that across songbird species, regardless of age, rearing condition, and sex, hearing range was similar: 0.5 to 8 kHz. Notably, the vocal energy in courtship song matches each species auditory sensitivity, indicating the coevolution of sensory and vocal motor systems. In the second chapter, I describe neuronal tuning in the auditory cortex (AC) of female songbirds. While male songbirds exhibit experience-dependent neuronal tuning in AC, the nature of AC tuning in females that do not sing has not been studied. I used in vivo acute electrophysiological recordings to examine neural responses to tones, ripple stimuli and songs. I compared neuronal firing patterns in female AC between different species groups and rearing conditions. My results suggest that higher-order auditory processing in female songbirds is conserved across species and that early song experience affects some aspects of tuning in the AC of females, suggesting that females exhibit experience-dependent changes in auditory tuning across development. In the final chapter, I examine a potential sensorimotor integration site, the caudal striatum (CSt), and its role in vocal learning. Auditory neurons in CSt suggest that the region may integrate auditory inputs and vocal motor commands to modulate sensorimotor learning. To study the effects of CSt lesions on song learning, I produced excitotoxic lesions in CSt across developmental stages. To label brain regions that project to or receive input from CSt, I injected anterograde or retrograde tracers into CSt. I also characterized the auditory tuning properties of CSt through electrophysiological recording. I found that CSt receives both dopaminergic and auditory projections but is not necessary for successful song learning. Electrophysiological data also show that auditory tuning properties of neurons in CSt are highly similar to neurons in other AC subregions, suggesting that CSt may be a sub-region of AC.

I examined the relationship of bird assemblages and species to habitat patterns over landscapes composed of a mosaic of habitat elements. I surveyed songbirds using a variation of the variable circular plot method during the 1994 and 1995 breeding seasons on 20 sites ranging in size from 50 to 72 ha on the Quantico Marine Corps Base, VA. Measures of community performance including species diversity, species richness, species equitability, and indices of relative abundance were calculated for each site. I determined the large-scale habitat characteristics of each site by analyzing coverages of each site from Quantico’s GIS database using FRAGSTATS. Landscape patches were defined using 2 different classification schemes to determine if both SAF cover type and generalized habitat classifications could be used to determine which large-scale habitat elements influence bird species and assemblages. I used stepwise multiple regression and stepwise logistic regression to determine which large-scale habitat measures and combinations thereof were associated with high and low measures of community performance. Diversity as measured by the Shannon-Wiener diversity index and the Simpson’s diversity index was positively related to the amount of high-contrast edge in a landscape in the SAF cover type based analysis. In the generalized habitat type based analysis, diversity was positively related to the number of different patch types per unit area in a landscape and negatively related to the percentage of hardwood forest in a landscape. The number of different patch types per unit area, the amount of contrast-weighted edge per unit area, and the percentage of mixed pine/hardwood forest in a landscape were selected most frequently as significant predictors of individual species relative abundance in both the SAF cover type and generalized habitat type based analyses. Habitat diversity was the most important factor influencing the large-scale selection of habitat by bird species on Quantico. With respect to individual species models, the 2 analyses yielded comparable results, and I believe that many of the common bird species occurring on Quantico can be managed according to either the SAF cover type classifications or the generalized habitat type classifications. / Master of Science

No description available.

Although grassland bird populations have steadily declined, little research has examined the effect of oil infrastructure on abundances of grassland songbirds. Even less research has identified mechanisms that explain observed effects. To evaluate this, I sampled abundance of 5 songbird species at oil well sites with different pump mechanisms, power sources, and activity levels; I also evaluated the effects of perch and road density and exotic vegetation, all of which are associated with oil development. Both Baird’s sparrows (Ammodramus bairdii) and Sprague’s pipits (Anthus spragueii) had lower abundances at all sites that contained oil infrastructure. The other 3 species, chestnut-collared longspurs (Calcarius ornatus), western meadowlarks (Sturnella neglecta), and Savannah sparrows (Passerculus sandwichensis), were relatively unaffected by oil wells, linear features, or exotic vegetation. Given that oil well sites negatively affected two species of concern, more research is needed to determine mitigation strategies. / October 2016

Grasslands are the most threatened terrestrial ecosystem in the world, and as they decline the species that depend on them also decline. Variable stocking rates and resting rangeland could be used by range managers to manipulate plant cover and create wildlife habitat. I used generalized linear mixed-models to evaluate how effects of rest on vegetation and songbirds varied based stocking rates previously applied from 2006-2014 in Grasslands National Park, Canada. My results indicate, in the mixed-grass prairie, succession is retrogressive following rest, and both vegetation and songbirds are resilient to grazing. Recovery of songbirds was linked to the recovery of habitat structure. Songbirds with flexible habitat requirements, such as Savannah and grasshopper sparrows, responded rapidly to livestock exclusion. The retrogressive nature of succession in mixed-grass prairie offers managers an opportunity to take chances when attempting to create wildlife habitat with livestock grazing, as risk of irreversible change is relatively low. / October 2016

A l’heure actuelle, il reste largement à étudier comment le codage sensoriel des signaux vocaux de communication contribue à leur détection et à leur reconnaissance. Peu d’études se sont, en effet, penchées sur le codage des vocalisations au niveau des régions auditives en fonction de l’individu qui les produit et du degré de familiarité avec cet individu. Dans ce cadre, les oiseaux chanteurs sont un bon modèle parce qu’ils utilisent des vocalisations pour interagir et reconnaître leurs congénères et qu’ils possèdent, de plus, un ensemble de régions auditives. Parmi ces régions, le nidopallium caudomedian (NCM), une aire auditive analogue du cortex auditif secondaire chez les mammifères, est actuellement considérée comme une région spécialisée dans le traitement des vocalisations (chants et cris) de l’espèce : les neurones du NCM répondent plus fortement aux vocalisations de l’espèce qu’à celles d’une autre espèce. À partir de là, parce que chez le diamant mandarin, le cri de distance permet aux individus, mâles ou femelles, de reconnaître leur partenaire sexuel, nous avons cherché à savoir si, chez les femelles comme chez les mâles, les neurones du NCM montraient une discrimination dans leurs réponses auditives entre le cri d’individus connus (parmi lesquels figurait le partenaire sexuel) et ceux d’individus inconnus et si ces réponses reflétaient le degré de familiarité de ces vocalisations. Les enregistrements de l’activité des neurones du NCM, chez des diamants mandarins vigiles (grâce à un système de télémétrie) ou anesthésiés, lors de la présentation de cris de distance, ont révélé, chez les femelles vivant en couple et ayant été familiarisées avec un autre couple de diamants mandarins, une plus forte augmentation de l’activité lors de la diffusion des cris d’individus connus, mâles ou femelles, qu’aux cris d’individus inconnus. Une telle augmentation n’a pas été, en outre, observée chez des femelles contrôles qui n’avaient jamais entendu ces mêmes cris auparavant. De plus, ils ont indiqué que le nombre de neurones montrant un fort degré de sélectivité ainsi que la quantité d’information portée par les trains de potentiels d’action étaient plus importants chez les femelles vivant en couple que chez les femelles contrôles. En revanche, chez les mâles, bien que la plupart des neurones montrait des réponses lors de la diffusion des cris, aucune différence n’a été mise en évidence entre les cris d’individus connus et ceux d’inconnus. Nous avons alors cherché à savoir comment, d’un point de vue acoustique, les cris de distance étaient représentés au sein du NCM. En se basant sur une étude comportementale ayant déterminé quelles étaient les caractéristiques acoustiques qui contribuaient à la reconnaissance de ces cris, nous avons cherché à savoir si les neurones du NCM étaient sensibles à ces mêmes caractéristiques acoustiques. Les résultats ont montré que, chez les femelles, la suppression de la fréquence fondamentale et la modification du timbre du cri du partenaire sexuel ou du propre cri de l’oiseau provoquaient une forte diminution des réponses au sein du NCM alors que, chez les mâles, les réponses variaient selon le paramètre modifié et le type de cri présenté. Nos résultats suggèrent donc que, chez le diamant mandarin, le NCM est impliqué dans le codage du cri de distance. Cependant, ils mettent en évidence des différences dans ce codage entre les mâles et les femelles. Chez les femelles, ce codage permet de discriminer entre les cris d’individus connus et ceux d’individus inconnus alors que chez les mâles, son rôle reste à être déterminé. Chez les femelles, l’expérience sociale au travers de la mémorisation des signaux de communication des individus peut donc façonner les propriétés fonctionnelles des neurones d’une aire auditive secondaire. Ces propriétés pourraient donc continuellement subir des changements pour s'adapter à l’environnement social de l’individu. / How sensory signals are encoded in the brain and whether their behavioural relevance affects their encoding are central questions in sensory neuroscience. Studies have consistently shown that behavioural relevance can change the neural representation of sounds in the auditory system, but what occurs in the context of natural acoustic communication where significance could be acquired through social interaction remains to be explored. The zebra finch, a highly social songbird species that forms lifelong pair bonds and uses a vocalization, the distance call, to identify its mate offers an opportunity to address this issue. One auditory area in the songbird telencephalon, the caudo-medial nidopallium (NCM) that is considered as being analogous to the secondary mammalian auditory cortex, has recently emerged as part of the neural substrate for sensory representation of species-specific vocalizations: the activation of NCM neurons is greatest when birds are exposed to conspecific song, as compared to heterospecific song or artificial stimuli. This led us to investigate whether, in the zebra finch, NCM neurons could contribute to the discrimination among vocalizations that differ in their degree of familiarity: calls produced by the mate, by familiar individuals (males or females), or by unfamiliar individuals (males or females). In females, behaviourally relevant calls, i.e. the mate’s call and familiar calls, evoked responses of greater magnitude than unfamiliar calls. This distinction between responses was seen both in multiunit recordings from awake freely moving mated females (using a telemetric system) and in single unit recordings from anesthetized mated females. In contrast, control females that had not heard them previously displayed response of similar magnitude to call stimuli. In addition, more cells showed highly selective responses in mated than in control females suggesting that experience-dependent plasticity in call-evoked responses resulted in enhanced discrimination of auditory stimuli. In males, as in females, call playback evoked robust auditory responses. However, neurons in males did not appear capable of categorizing the calls of individuals (males or females) as ‘‘familiar’’ or ‘‘unfamiliar’’. Then, we investigated how calls are represented in the NCM of zebra finches by assessing whether certain call-specific acoustic cues drove NCM neurons to a greater degree than others. Behavioural studies had previously identified call-specific acoustic cues that are necessary to elicit a vocal response from male and female zebra finches. Single-unit recordings indicated that NCM neurons in females were particularly sensitive to call modifications in the spectral domain: suppressing the fundamental frequency of call stimuli or modifying the relative energy levels of harmonics in call caused a marked decrease in response magnitude of NCM neurons. In males, NCM neurons also appear to be sensitive to call modifications in the spectral domain, however changes in magnitude of responses (increase or decrease) depended on the acoustic cue that had been modified.Our results provide evidence that the NCM is a telencephalic auditory region that contributes to the processing of the distance call, in females as well in males. However, how the distance call is processed and represented in the NCM appears to differ between males and females. In females, the NCM could be involved in dicrimination between call stimuli whereas, in males, its functional role in call-processing remains to be determined. Our results also suggest that, in females, social experience with the call of individuals, by affecting the degree to which neurons discriminated between these calls, may shape the functional properties of neurons in a telencephalic auditory area. The functional properties of auditory neurons may therefore change continuously to adapt to the social environment.

An undue focus on Temperate Zone oscines (songbirds or passerines) has led to a geographical bias in interpretation of song frmnction and evolution. This bias led initially to relatively simplistic theories of the ftmnction of bird song with vocalizations divided into 'songs' and 'calls'. Songs were complex, learned vocalizations, given by males in the breeding season, thnctioning in territory defence and mate attraction and stimulation. Calls, on the other hand, were simple innate vocalizations serving more immediate needs such as begging for food and raising an alarm. Female song, where it occurred, was considered an aberration. Further studies suggested that complex songs were associated with mate attraction functions while simpler songs were associated with territory defence. However it became apparent that the distinction between songs and calls was not nearly so clear-cut and the supposed connection between complexity and function in song was questioned. Moreover it was realised that female song could not be dismissed as a mere aberration. Another problem was the ftmnction of the dawn chorus, where research had failed to find a consistent, all-encompassing explanation. Since most studies had been done on Northern Hemisphere songbirds, it was becoming clear that the geographical focus needed to be broadened. The life histories of Northern Hemisphere TemperateZone songbirds are very different from those in many other regions. In contrast to the situation in this zone, maintenance of year-round territory, territory defence by both male and female, life-long social monogamy and extensive female vocalization are widespread in tropical, subtropical and Southern Hemisphere regions. Recently it has been suggested that more intensive studies of vocalizations in these regions might help clariQi some of these issues and consequently an endemic Australian passerine was chosen for the current study. The study focused on the vocalizationsof the grey butcherbird Cracticus torquatus Artamidae, which displays the life history features described above. The main study population was located in the Brisbane suburbs of Rainworth and Bardon. Additional data were gathered from other Brisbane suburbs and bushland sites within the city and at Lake Broadwater near Dalby, Queensland. Vocalizations were initially recorded electronically and analysed using Canary sound editing program. Vocalization data were supplemented using an aural recording method, which was independently checked for reliability. Behavioural data including posture during vocalizations and interactions with other birds were also gathered. Initial investigationsrevealed the existence of two main categories of vocalizations - those given by the family group during the day, all year round and those given at dawn by males during the breeding season. For the focal study populations, group vocalizations were studied throughout the year over several years, however the song given at dawn by males during the breeding season proved to be quite complex and three birds from three territories in the main study area were chosen as case studies. Recordings were made of the vocalizations of the three case study birds over three breeding seasons. Starting and fmishing times (with respect to civil twilight) were recorded in order to determine both changes in song bout duration and starting time throughout the breeding season. Additional birds from the same area, from the other Brisbane suburbs and from the bushland sites were studied to check the validity of conclusions drawn from the case studies. The results of the investigations revealed a vocalization structure that contrasted strongly with the simple picture of bird song drawn from study ofNorthernHemisphere Temperate Zone passerines. The vocalizations given during the day, often referred to as the 'song' of this species, were very different from the early dawn song given by the male during the breeding season. This latter appeared to be song sensu stricto according to the paradigms developed for Northern Hemisphere birds. The thytime vocalizations, however, fitted neither the classic definition of 'song' nor the classic definition of 'call'. This relatively long-term study revealed different starting time patterns and periodicity for thy vocalizations and male breeding season song. Day vocalizations commenced at a fairly constant time with respect to civil twilight throughoutthe year but breeding season song started progressively earlier from the beginning through to the middle of the season then progressively later till the end of the season. Relative finishing time of breeding song however remained constant so that the duration of breeding season song gradually increased then decreased paralleling the change in starting times. A consequence of the two distinct classes of vocalizations was that there were essentially two distinct 'dawn choruses'. One, consisting of group vocals, was sung all year round; the other was given by males singing 'breeding season' song. Since there was no reason the expect that a single function would necessarily be ascribed to both choruses, this raised the possibility that some of the confusion surrounding the ftinction and nature of the 'dawn chorus' originated from a failure to recognize the existence of two such choruses. Variation in time and space showed ifirther differences between the two vocalization classes. The breeding song of each male was distinctly different from that of his neighbours and there was a marked change in the repertoire of any individual from one year to the next. In marked contrast, daytime vocalization repertoires of neighbouring groups were virtually indistinguishable and changed little from year to year. These findings, together with information from recent literature, suggested that the two song classes had a different ontogeny, function and possibly evolution. It was proposed that territory declaration was the function of dawn singing by grey butcherbird family groups but that the function most consistent with adult male dawn song was attraction of females for extra-pair copulations. It was suggested that chorusing itself was to some extent an accidental by-product of the advantage to the individual or group of singing at dawn although a recently proposed function, the social dynamics function, could not be ruled out. Further differences from the Northern Hemisphere situation were detected in subsong. First, subsong was given by birds in their first year and also by adult males. The finding of subsong in adult males was not without precedent as it has been documented previously for a small number of passerines, especially those that change repertoire from year to year. Subsong in young (first year) birds, however, was unusual in that birds practised in small groups rather than in the complete isolation usually associated with subsong. Moreover they did not practise adult male song but instead practised group daytime vocalizations. It was suggested that it was important for birds to learn to sing in company for the important task of group territory defence. Further investigation of the literature and observations during the present study revealed similar vocalization classes and behaviours in other members of the Artamidae and other endemic Australasian taxa. These literature investigations also revealed that the possession two song vocalization classes was quite widespread although they tended to be restricted (but not exclusive) to males rather than found in males and females. These findings led to furtherresearch into the significanceofAustraliain the evolution of songbirds, the role of co-operative breeding in Australianpasserines, and finally to an hypothesis for a possible origin of male bird song. It is suggested that male song arose in a social environment where the male and female were in frequent vocal communication. If the tendency to seek extra-pair copulations (EPCs) and female choice had already been incorporated into the suite of passerine behaviours, it would be necessary to avoid the mate during such activities and the male would need to advertise with a signal distinct from group vocalizations. Early dawn, with poor light conditions, could be a favourable time for these activities. Thus it is proposed that the ancestral condition was with all group members singing most vocalisations, the intermediate situation was similar to that in the grey butcherbird and the 'advanced' condition was where female and other group member vocalizations (other than calls) have dropped out and only male song remains.

Grassland bird populations in North America are in steady decline. Despite declining faster and more consistently than any other group of birds, grassland songbird populations are relatively understudied and little is known about factors driving breeding-site philopatry and dispersal. Landscape and habitat composition may influence fidelity of grassland songbirds to a breeding area. As predicted by the theory of the <i>Ideal Free Distribution</i>, high-quality sites are likely to have a higher percentage of return breeders than low-quality sites because higher quality sites should have more or better-quality resources birds need for improved fitness. Using stable hydrogen isotope (äD) analysis, I approximated minimum fidelity rates of two grassland songbirds to two landscape (grass- vs crop-dominated landscapes) and two habitat (native grass vs planted grass) types. I hypothesized that grassland songbirds would return more readily to higher quality sites. For Spragues Pipit (<i>Anthus spragueii</i>), a habitat specialist, this would mean returning more readily to native grass habitat in grass dominated landscapes. I expected no difference in return rate of Savannah Sparrow (<i>Passerculus sandwichensis</i>), a habitat generalist, to either habitat or landscape. However, I found that the proportion of non-returning breeders was not influenced by landscape or habitat for either species. Furthermore, I examined attributes (distance from capture point to nearest crop and to the nearest road, as well as the percentage of native grass, planted grass, water and woody vegetation within landscape and territory buffers around the capture point) of the landscape and territory of each individual to determine if specific landscape or territory characteristics influenced their return rate to a breeding area. Neither species showed an affinity or aversion to any of the landscape or territory characteristics considered.<p> At a larger scale, geographical position within the breeding range may influence dispersal rates of migratory songbirds. Given that environmental factors often change in a clinal manner, central populations presumably experience the most favorable environmental conditions and peripheral populations the least favorable. Therefore, geographically peripheral locations likely occur in ecologically marginal or stressful conditions, resulting in higher dispersal rates of migratory birds. I examined the differences in dispersal rates of two grassland songbirds at two geographically distinct locations; one centrally located in the Spragues Pipit breeding range (Last Mountain Lake area, SK, Canada) and one at the periphery (Bowdoin National Wildlife Refuge, MT, USA). As expected, Spragues Pipits at the Bowdoin National Wildlife Refuge had a significantly larger dispersal rate than at the Last Mountain Lake area. Savannah Sparrow dispersal rates did not differ between locations at the more conservative outlier classification. These results provide some limited evidence that geographical position within the breeding range can influence dispersal rates.<p> Using äD analysis, I found that local amount-weighted growing-season deuterium in precipitation (äDp) at locations within grassland ecosystems differed from those long-term (45+ year) models described by stable hydrogen isotope ratio basemaps, illustrated in Hobson and Wassenaar (1997), Meehan et al. (2004) and Bowen et al. (2005). Therefore, I describe how äDp values were corrected from the long-term isoscape value predicted by Bowen et al. (2005). This method of determining year-specific local weighted growing-season äDp is an improvement upon the currently used Bowen et al. (2005) isoscape that is based on long-term precipitation patterns. To improve assignment of individuals to origins based on their äDf values, future research should incorporate year-to-year variation by applying year-specific corrections to the Bowen et al. (2005) isoscape.<p> More research is needed to determine the factors affecting the philopatry and dispersal of grassland songbirds in order to conserve them.

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.