Many grassland songbird species exhibit sensitivity to patch size in North America’s fragmented prairie ecosystems, but the mechanisms explaining this area sensitivity are not well understood. I tested the effects of patch size and artificial conspecific location cues (song playback and decoys) on grassland songbird abundance in 23 northern tall-grass prairies in Manitoba, Canada, in 2010 and 2011. Richness and relative abundances increased with patch area; this effect was not explained by differences in local habitat structure, patch configuration, and adjacent matrix. Artificial cues elicited putative territory prospecting in small, previously unoccupied treatment patches from two focal species, Savannah Sparrow (Passerculus sandwichensis; n=3 treatment sites) and Le Conte’s Sparrow (Ammodramus leconteii; n=4 treatment sites), but not in control patches (n=3 for both focal species). Social information may influence the focal species’ settlement decisions, but the lack of permanent settlement response suggests social cues are unable to reverse their area sensitivity.

Badges of status are supposed to have insignificant production costs, so use costs are thought to be most important in ensuring signal reliability. Use costs arise from the use of the status signal in social interactions. Social experiences that arise from the use of inappropriate signals in social interactions may drive mechanisms that result in reduced fitness for inappropriate signalers. The role of social control, probing and familiarity in producing use costs was explored. There was no evidence that social control by dominants produced a cost for cheaters and no evidence that social control by subordinates produced a cost for inappropriate signaling by Trojans. Probing produced a cost for cheating when resource value was high but not when resource value was low. Familiarity had some effect on the cost and benefit of cheating but in patterns that were not predicted. Familiarity both eliminated a benefit of cheating and reduced a cost of cheating; therefore it is uncertain how familiarity affects honest signaling. The status signal of the receiver had no effect on the cost or benefits of cheating, and there was no evidence of punishment. Social experiences have the potential to affect signal development to produce a correspondence between signal and status. The effects of social experience on signal production were examined and there was little evidence that social experience influenced bib development. Neither aggression expressed nor aggression received was not predictive of bib size. Additionally, tests on the different measures of winning experience produced conflicting conclusions regarding the relationship between winning experience and bib development.

Behavioral traits can be remarkably flexible depending on the conditions in which they are expressed, yet, in spite of this flexibility, persistent differences between individuals appear to limit the potential expression of behaviors. For example, despite evidence that parents provide variable amounts of parental care in response to changing environmental conditions, they also differ in the overall level of care they provide. I used a behavioral reaction norm approach to study individual variation in parental care behavior in free-living house sparrows (Passer domesticus). I investigated the nature of this variation by studying the relationship between different forms of parental care, the biological basis of individual variation in care, and the effect of this variation in care on offspring. First, I found a positive covariance between nestling provisioning and nest defense. Parents that provided high levels of care in one context provided high levels of care in the other context, even after accounting for measures of offspring value. Second, I sought to identify the biological sources that create and maintain consistent individual differences in the level of care a parent provides. I found that the likelihood of feeding nestlings large food items was positively associated with genetic heterozygosity, but did not find evidence that nestling provisioning was influenced by additive genetic variation in this population. Parents hatched from larger eggs provisioned offspring at a higher rate than parents hatched from smaller eggs, but there was no effect of other conditions experienced in the nest on the level of care expressed as an adult. I also tested if differences in problem-solving ability were related to differences in parental care behavior. Although I found that problem-solving parents fledged more offspring than parents that could not solve the problem, parental care was not associated with any measure of problem-solving ability. Finally, I found that individual variation in parental care reaction norms predicted the growth rate, size, and immune response of nestlings, which in turn positively affected offspring survival and recruitment. My findings reveal factors maintaining individual differences in parental care behavior and offer new insights into the causes and consequences of individual variation.

Over the last 40 years North American grassland bird populations have declined more than any other bird guild. This trend is especially evident in Vermont, where species experiencing precipitous declines include the Savannah Sparrow (Passerculus sandwichensis) and Bobolink (Dolichonyx oryzivorus). These declines are linked to habitat loss due to reforestation and suburbanization as well as the intensification of grassland management. Modern grassland management includes earlier first-haying dates (late-May) and shorter intervals between haying events (35 days). These management practices have severe repercussions for songbird populations because 1) early-haying results in complete nest failure (99% Savannah Sparrow and 100% Bobolink nests), 2) the interval between the first and second haying is too short for birds to renest, and 3) intensively managed fields comprise a significant portion of the total available habitat (as much as 40%). In 2002-2006, I examined how hayfield and pasture management affected grassland songbird ecological and evolutionary behavior in the agricultural landscape of the Champlain Valley, Vermont and New York. I studied songbirds in four grassland management types: early-hayed fields harvested in late-May or early-June and again in mid-July; middle-hayed fields harvested in late-June or early-July; late-hayed fields harvested after 1 August; rotationally-grazed pastures, a matrix of small paddocks where cows are moved after the grass in a paddock is eaten to a low point. I addressed the following objectives: 1. Determined the annual productivity, survival, and recruitment of Bobolinks and Savannah Sparrows in the four treatment types. 2. Identified the effects of early-haying on the social and genetic mating systems of Savannah Sparrows. 3. Conducted a population viability analysis for Bobolinks and Savannah Sparrows nesting in the Champlain Valley, assessing sensitivities of life-history parameters and identifying effective management alternatives. This study provides information on how agricultural management affects the ecology, evolution, and viability of grassland birds. It will help inform landowners, managers, and law-makers about management practices and habitat requirements needed to sustain populations.

Master of Science / Biology / Alice Boyle / Dispersal is a behavior common to virtually all taxa with important consequences for gene flow, demography, and conservation. Mobile animals such as birds frequently engage in breeding dispersal, but the factors shaping this behavior are not well understood. In mid-continental grasslands, preliminary evidence suggested that Grasshopper Sparrows (Ammodramus savannarum) engaged in breeding dispersal within a single breeding season. This intriguing pattern shaped my research questions: (1) what are the patterns of within-season breeding dispersal in Grasshopper Sparrows? (2) why do some individual sparrows disperse, and others do not? and (3) what shapes settlement decisions following dispersal? I tested hypotheses based on spatial and temporal variation in nest predation, nest parasitism, and food availability. I studied Grasshopper Sparrows at 18 experimentally-managed watersheds with different fire and grazing regimes at Konza Prairie Biological Station during 2013-2015. To describe patterns, I combined re-sighting of 779 marked individuals, territory mapping, and radio-telemetry to quantify territory densities, turnover, and dispersal distances. To determine if nest predation or nest parasitism affected dispersal, I monitored the fate of 223 nests. I estimated food intake rates using plasma metabolites, and prey availability using sweep sampling. Densities of territorial Grasshopper Sparrows varied seasonally in management-specific ways. Turnover was remarkably high, with over half of territorial males being replaced each month. Over a third of males changed territories within-season, dispersing 0.1-9 km between breeding attempts. Dispersal decisions were related to past nest predation, but not nest parasitism. Dispersal likely yields fitness benefits, as sparrows that dispersed increased their chances of nest survival by 23% relative to site-faithful individuals. However, food availability did not affect settlement decisions. My study provides the first evidence of within-season breeding dispersal in Grasshopper Sparrows, and represents one of few tests of alternative hypotheses explaining dispersal decisions of songbirds. My results are consistent with a growing literature on the role of predation shaping dispersal, but suggest, somewhat surprisingly, that food is not important in post-dispersal habitat selection. High dispersal capacity coupled with adaptability to temporal and spatial change may be typical of grassland songbirds, implying that demographic studies and management decisions must consider their mobility for conservation.

Thesis (M.Sc. (Zoology)) -- University of Limpopo, 2012 / Sparrow-larks form a relatively small genus in the family Alaudidae and comprise only seven species distributed widely throughout Africa and parts of the Eurasian landmass. Sparrow-larks are unique amongst larks in that they are sexually dichromatic and exhibit biparental care. The chestnut-backed sparrow-lark Eremopterix leucotis is endemic to Africa with five subspecies recognized based on differences in plumage colouration. The five subspecies are distributed throughout the arid to semi-arid savannas of Africa with two subspecies (E. l. hoeschi and E. l. smithi) occurring in southern Africa. Despite their widespread occurrence and its interest for research on the evolution of characteristics in the family (e.g. being sexually dichromatic and exhibiting biparental care), very little is known of the biology and ecology of the Eremopterix larks. The chestnut-backed sparrow-lark is no exception and most of what we know of the species is based on incidental observations from a few nests. In an attempt to improve our knowledge of this interesting group of species, it was decided to study various aspects of the breeding biology and ecology, moult, vocalizations and geographical variation in the chestnutbacked sparrow-lark. The breeding biology of the chestnut-backed sparrow-lark was studied at Al3 farm (De Loskop) near Mogwadi in the Limpopo Province of South Africa from January 2008 to December 2010. Data collected during the study included: breeding seasonality, egg and clutch characteristics, duration of the incubation and nestling periods, nest-site characteristics, the roles and relative contribution of the sexes in the breeding cycle, nestling development, diet and nestling provisioning rate, and breeding success. Chestnut-backed sparrow-larks bred mostly during the dry season, which is from April to September in the study area. Nevertheless, the results revealed that breeding is bimodal with a main peak in breeding activity in late summer and autumn (March to April) and a second smaller peak in spring (September to October). The species showed geographical variation in clutch size with a mean of 1.88 eggs recorded in the study area as opposed to 1.00 recorded in the northern parts of its range. Egg dimensions compared well with measurements obtained from the Nest Record Card Scheme of the Animal Demography Unit, University of Cape Town, South Africa. The mean incubation period of 10.33 days recorded in this study compares favourably with that of other Eremopterix species viii (8–10 days), a genus with some of the shortest incubation periods amongst larks. The mean nestling period of 9.2 days (range: 8–10) in the study area was significantly less than the 10–12 days reported for populations in the northern range of the species, but it compares well with those of other sparrow-larks. Nest site characteristics, which were quantified within a 1 m2 quadrant with the nest as the centre, including nest dimensions, were consistent with those reported in the literature. Chestnut-backed sparrow-larks in the study area preferred to nest in areas with a high percentage of bare ground (median = 67.5%) and very little vegetation cover (median = 25%). Most nests faced in a southerly direction compared to nests in the north of the species’ range, which face in a north-easterly or easterly direction. The species’ preference to face the nests away from the midday sun most probably serves a thermoregulatory function to avoid excessive heat during the warmest parts of the day. Most nests (78.2%) had an apron varying in size from small and insignificant to large and well-developed. The functional significance of the apron remains a matter of conjecture and there was no association between breeding success and presence or absence of the apron. In addition, one pair constructed one nest with and another without an apron, suggesting that individual preference or characteristic is not a determinant factor in the construction of an apron. Both sexes took part in nest construction, incubation and feeding and brooding of nestlings. However, the relative contributions were not entirely symmetrical as males incubated a greater proportion (50.1%) of the time compared to females (43.1%), and the mean and median of male incubation shift lengths were longer than that of females, albeit not statistically significant. However, females made statistically significantly (P < 0.05) more nest visits to deliver food compared to males (54.6% vs. 45.5%). The average breeding success, estimated using Mayfield’s method, was 16.1% but there were inter-annual differences with the overall breeding success in 2010 being only 8.1% compared to 20.6% of 2008. Known causes of failure included nest depredation, flooding, starvation, nest abandonment and hatching failure. Statistical analysis of morphometric data of live specimens and museum study skins suggest that, in addition to being sexually dichromatic, chestnut-backed sparrowlarks also exhibit mild sexual size dimorphism. However, there was considerable overlap in these measurements between the sexes and as a result the biological significance of this sexual size dimorphism may be negligible. Nevertheless, the ix results show chestnut-backed sparrow-lark males tend to have longer wings and tails compared to females. This may be adaptive with respect to the extended display flights that males perform during the breeding season. Interestingly, the mean mass of breeding females in the study area was significantly more compared to males, whereas the SAFRING database, representing data recorded throughout the year, showed no significant differences in the mean mass between the sexes. The greater mass of breeding females may relate to physiological changes associated with the acquisition of resources and the development of structures and tissues associated with egg-laying and egg-production. Larks rely heavily on vocalizations to attract mates and advertise territories. Not surprisingly, the study revealed a rich vocal repertoire for the chestnut-backed sparrow-lark. The analysis of the vocalizations shows that chestnut-backed sparrowlarks have a display song performed by males, a sub-song sung by both sexes and various different calls used in different contexts, e.g. flight and alarm calls. The study also presents the first analysis and description of the vocalizations of nestlings. An interesting feature of the vocalizations of the chestnut-backed sparrow-larks was that they performed hetero-specific vocal mimicry, which was incorporated in the subsong. Moult is a relatively unknown aspect in the annual cycle of the majority of larks. Chestnut-backed sparrow-larks undergo post-breeding moult, which is an adaptation to reduce the conflict between moult and breeding as both activities have high energy demands. The moult study also showed that they undergo a partial moult in mid-winter, involving the inner-most secondaries and some of the contour feathers. The results of this study shed valuable light on the natural history of this species and contributed significantly to ornithology and our growing understanding of the biology and ecology of the family. The results can also form a basis for future inter- and intraspecific comparative studies. The study illustrates the importance of undertaking long term studies of species to account for inter-annual differences in various ecological parameters.

I investigated the effect of energy sector development and introduced crested wheatgrass (Agropyron cristatum) on grassland birds on Canadian Forces Base Suffield. I conducted point counts and mapped breeding territories in 2007 and 2008 for Savannah sparrows (Passerculus sandwichensis), chestnut-collared longspurs (Calcarius ornatus), and Spragues pipits (Anthus spragueii). I found that Savannah sparrows favored areas with taller vegetation, human disturbances and crested wheatgrass in both years. Longspurs used shorter vegetation and in were tolerant of disturbance. Crested wheatgrass was avoided by longspurs in both years. Pipit territories contained similar vegetation to longspurs, were sensitive to disturbance, and avoided placing territories in areas containing crested wheatgrass or trails in both years. Well sites, pipelines and junctions were not avoided by the three species. My research suggests that reducing the number of trails and the spread of crested wheatgrass will increase habitat availability for sensitive species of grassland birds. / Ecology

The diversity and abundance of the arthropod nest fauna of House Sparrows (Passer domesticus (L.)) and Tree Swallows (Tachycineta bicolor (Vieillot)) in southern Quebec were studied. Over 90,000 arthropods were extracted from the nests, including at least 71 species (50 families) of insects and at least 11 species (8 families) of mites. There were no significant differences in the species richness or diversity of nest arthropods from year to year (1997--1998), or from nest to nest within either host species. There was no significant difference in the overall species richness or diversity between House Sparrows and Tree Swallows. Cluster analyses of species richness and three diversity indices for all nests showed no clustering according to bird species. However, there were some significant differences in the abundance and diversity of particular arthropod taxa between the two bird species, with different ectoparasitic and non-ectoparasitic species dominant in the nests of each host species.

No description available.

abstract: In wild birds, the stress response can inhibit the activity of the innate immune system, which serves as the first line of defense against pathogens. By elucidating the mechanisms which regulate the interaction between stress and innate immunity, researchers may be able to predict when birds experience increased susceptibility to infections and can target specific mediators to mitigate stress-induced suppression of innate immune activity. Such elucidation is especially important for urban birds, such as the House Sparrow (Passer domesticus), because these birds experience higher pathogen prevalence and transmission when compared to birds in rural regions. I investigated the role of corticosterone (CORT) in stress-induced suppression of two measures of innate immune activity (complement- and natural antibody-mediated activity) in male House Sparrows. Corticosterone, the primary avian glucocorticoid, is elevated during the stress response and high levels of this hormone induce effects through the activation of cytosolic and membrane-bound glucocorticoid receptors (GR). My results demonstrate that CORT is necessary and sufficient for stress-induced suppression of complement-mediated activity, and that this relationship is consistent between years. Corticosterone, however, does not inhibit complement-mediated activity through cytosolic GR, and additional research is needed to confirm the involvement of membrane-bound GR. The role of CORT in stress-induced inhibition of natural antibody-mediated activity, however, remains puzzling. Stress-induced elevation of CORT can suppress natural antibody-mediated activity through the activation of cytosolic GR, but the necessity of this mechanism varies inter-annually. In other words, both CORT-dependent and CORT-independent mechanisms may inhibit natural antibody-mediated activity during stress in certain years, but the causes of this inter-annual variation are not known. Previous studies have indicated that changes in the pathogen environment or food availability can alter regulation of innate immunity, but further research is needed to test these hypotheses. Overall, my dissertation demonstrates that stress inhibits innate immunity through several mechanisms, but environmental pressures may influence this inhibitory relationship. / Dissertation/Thesis / Doctoral Dissertation Biology 2017