Abstract Habitat selection is a crucial process affecting space use and reproductive success of birds. In this thesis, I investigated spatial and behavioural aspects of nest spacing, brood stage space use, habitat selection and factors affecting reproductive success of breeding common goldeneye (Bucephala clangula) using two large and long-term observational data sets from individually marked females. In the nesting stage, I found that spatial nesting pattern of goldeneye females changed from one year to the next and also between spatial scales. However, increasing aggregation of nesting females decreased nesting success due to increasing rate of nest desertion and nest predation especially at small spatial scale. These results provide evidence of a density-dependent population process in the common goldeneye in terms of association between annual spatial dispersion of nesting females and annual nesting success. In the brood stage, the most important factor affecting habitat selection was the amount of food. However, safe nest sites and food requirements of ducklings were not usually met in the same patch and females with broods adjusted their space-use tactics according to these critical breeding resources. Spatial divergence of these two obligatory resources induced brood movements at various distances shortly after hatching. During movements, broods used different landscape elements such as patches, corridors and matrix in a flexible way without clear fitness consequences in terms of duckling survival. Goldeneye broods suffered heavy losses especially during the early brood stage. Increasing predation risk by northern pike (Esox lucius) decreased survival of young ducklings, but frequent total brood losses suggest that also other factors affected duckling survival. Environmental factors such as temperature or rain were not related to the survival of ducklings.

Master of Science / Department of Biology / David A. Haukos / Abstract–Duckling Survival Lesser scaup (scaup) populations have been experiencing continent-wide decline since the 1980s. It is important to have complete understanding of the critical factors influencing population change (e.g., duckling survival, nesting success, and health) to advance our understanding of population dynamics and improving species conservation. Duckling survival is a primary driver of scaup demography. I conducted a capture-mark-recapture study using Cormack-Jolly-Seber models in Program MARK to compute apparent daily survival and recapture probabilities for 3256 individually marked ducklings with 620 recaptures during 2010 to 2013. The most parsimonious survival model based on a priori hypotheses found that Julian hatch date squared was the most significant predictor of survival and consistent through all four years. Mass at hatch also was significant as a quadratic effect. Duckling survival to 30 days ranged from 29.0 to 80.0. During this study, stabilizing selection played a significant role in duckling survival, which indicates that there was trade-offs for selection of an optimal timing of hatch on survival and a cost associated with hatching too early or too late and being too heavy or to light. / Abstract–Nest-Site Selection There is a hierarchical process of behavioral and environmental processes that influence habitat selection, which inherently influences the survival and fitness of that individual and contributes to population growth. I investigated nest fate, spatial attributes, and all relationships between high and low-water levels with habitat attributes (distance to upland, distance to open water, nearest neighbor distance) of located nests using general linear models in SAS, t-tests in R, and Hot Spot Analysis in ArcGIS of 481 nests over eight years. In low-water years, successful nests (X̅ = 1153 m) were located 22.0% farther from upland than unsuccessful nests (X̅ = 944 m), but support for a similar relationship was lacking in high-water years. Successful nests were located 21.0% and 23.0% (i.e., 49 and 50 m) closer to conspecific nests than unsuccessful nests in low and high-water levels, respectively. In both high and low-water level years, clusters of nests initiated later in the season coincided with Hot Spots for nest fate (i.e., high-quality habitat patches, clusters of successful nests), whereas areas that tended to be selected first, evidenced by clusters of nests initiated earlier, tended to overlap with clusters of Cold Spots for nest fate. The core Hot Spot for nest fate was in the same spot in both water level conditions and located in flooded emergent vegetation in the heart of Lower Red Rock Lake furthest from any upland habitat. Three out of six Cold Spots for nest fate both in high and low-water years were located in emergent vegetation on the perimeter of Lower Red Rock Lake adjacent to uplands. Density-dependence seems to be a factor affecting late-nesting scaup females that are apparently cuing in on the reproductive performance of conspecifics when determining where to nest. Therefore, management actions focused on survival and reproductive success of scaup should consider managing water levels and habitat for later nesting scaup to increase adult survival and ultimately recruitment of ducklings. / Abstract – Blood Parasite Prevalence Blood parasites, per se, do not lead to direct mortality, but instead reduce the health of individual birds, which may ultimately lead to decreased reproductive success. Evidence has shown that presence of blood parasites can reduce fitness, body condition, and reproductive success of waterfowl. For many avian species, the cost of reproduction is manifested as a negative relationship between female breeding effort and breeding season survival, with trade-offs occurring when these adaptive choices become detrimental to future reproductive performance. Blood was drawn for parasite load determination from 112 individual adult scaup captured from 2011to 2012 via spotlighting and drive-trapping. Parasite prevalence was determined through blood assays that were created using a two-slide wedge technique. Relationships among seasonal heterophile:lymphocyte ratio (a proxy for health), body mass at time of capture (throughout pre-, during, and post-breeding periods), breeding status (females only), and Julian date of capture (date of capture) of capture with parasite prevalence were analyzed using linear (lm) regression models in R 2.15.2. The blood parasite infection rate was 5.0%, with prevalence differing by gender with 33.3% of males positive for blood parasites compared to 1.0% of females. The presence of blood parasites did not affect health, fitness, or breeding status of scaup. A quadratic relationship was found with body mass and date of capture, indicating that body mass increased from pre-breeding period to the breeding period and decreased significantly at the end of the summer during molt. A negative relationship between the heterophile:lymphocyte ratio of female scaup and date of capture (i.e., the health of scaup females was greatest during the pre-breeding period after which it consistently decreased until the molting period). A strong negative correlation between heterophile:lymphocyte ratio and body mass was found in both genders, which indicated that scaup in poor body condition were also in poor health at the end of the breeding season.

Birth and death processes and the extent of dispersal directly affect population dynamics. Knowledge of ecological factors that influence these processes provides insight into natural selection and understanding about changes in population size. King eiders (Somateria spectabilis) breed across the arctic region of North America and winter in polar oceanic waters of the western and eastern regions of the continent. Here I studied a local population of King Eiders at Karrak Lake, Nunavut, where I used analysis of naturally-occurring stable isotopes (13C, 15N) from feathers, in conjunction with banding data, to investigate the extent of dispersal among winter areas and the influence of winter area on subsequent breeding. In addition, I used capture-mark-recapture methods to (1) investigate the relative contributions of survival and recruitment probabilities to local population dynamics, and (2) to test hypotheses about the influence of specific ecological factors on those probabilities or their components, e.g., nest success, duckling survival. Isotopic data suggested that female King Eiders were not strongly philopatric to wintering areas between years. Individuals that wintered in western seas initiated nests earlier and had slightly larger clutch sizes during early nest initiation relative to females that wintered in the east. Female condition during incubation did not vary by winter area. Female King Eiders of known breeding age were at least 3-years-old before their first breeding attempt. Age of first successful breeding attempt did not appear to be influenced by body size. However, after reaching breeding age, larger females apparently experienced greater breeding propensity. Adult survival rate (1996-2002) was estimated as 0.87 and recapture probabilities varied with time and ranged from 0.31 to 0.67. There is no evidence of survival advantages related to larger size. Population growth for this local study area was high, estimated at 20%/year with larger females contributing more to the population growth than smaller females. With continued population growth, density-dependent effects on components of recruitment appeared to emerge; the proportion of the female population that nested successfully declined with increasing population size. The probability of breeding successfully did not correlate with Mayfield estimates of nest success. To gain insight into King Eider brood ecology I, respectively, monitored 111 and 46 individually-marked ducklings from broods of 23 and 11 radio-marked King Eiders during 2000 and 2001. Total brood loss accounted for 84% of all duckling mortality with most brood loss (77%) less than 2 days after hatch. Estimated apparent survival rates of ducklings to 22 days of age were 0.10 for those that remained with radio-marked females, 0.16 for all ducklings, including those that had joined other broods, and 0.31 for broods. Ducklings brooded by larger females experienced higher survival than those brooded by smaller females, and ducklings that hatched earlier in the breeding season survived at higher rates. Overland brood movements of 1 km or more occurred in both years, and survival was greatest for ducklings that dispersed from Karrak Lake to smaller ponds than on Karrak Lake itself, the central nesting area. Estimates of duckling survival, combined with relative contributions to the population by adults, suggest that ecological factors such as body size can influence population growth. Furthermore, low duckling survival and delayed maturity, emphasize the need of high adult survival for population growth to occur. These data, in combination with evidence of dispersal among wintering areas have helped contribute to a broader understanding of North American King Eider demographics.

Birth and death processes and the extent of dispersal directly affect population dynamics. Knowledge of ecological factors that influence these processes provides insight into natural selection and understanding about changes in population size. King eiders (Somateria spectabilis) breed across the arctic region of North America and winter in polar oceanic waters of the western and eastern regions of the continent. Here I studied a local population of King Eiders at Karrak Lake, Nunavut, where I used analysis of naturally-occurring stable isotopes (13C, 15N) from feathers, in conjunction with banding data, to investigate the extent of dispersal among winter areas and the influence of winter area on subsequent breeding. In addition, I used capture-mark-recapture methods to (1) investigate the relative contributions of survival and recruitment probabilities to local population dynamics, and (2) to test hypotheses about the influence of specific ecological factors on those probabilities or their components, e.g., nest success, duckling survival. Isotopic data suggested that female King Eiders were not strongly philopatric to wintering areas between years. Individuals that wintered in western seas initiated nests earlier and had slightly larger clutch sizes during early nest initiation relative to females that wintered in the east. Female condition during incubation did not vary by winter area. Female King Eiders of known breeding age were at least 3-years-old before their first breeding attempt. Age of first successful breeding attempt did not appear to be influenced by body size. However, after reaching breeding age, larger females apparently experienced greater breeding propensity. Adult survival rate (1996-2002) was estimated as 0.87 and recapture probabilities varied with time and ranged from 0.31 to 0.67. There is no evidence of survival advantages related to larger size. Population growth for this local study area was high, estimated at 20%/year with larger females contributing more to the population growth than smaller females. With continued population growth, density-dependent effects on components of recruitment appeared to emerge; the proportion of the female population that nested successfully declined with increasing population size. The probability of breeding successfully did not correlate with Mayfield estimates of nest success. To gain insight into King Eider brood ecology I, respectively, monitored 111 and 46 individually-marked ducklings from broods of 23 and 11 radio-marked King Eiders during 2000 and 2001. Total brood loss accounted for 84% of all duckling mortality with most brood loss (77%) less than 2 days after hatch. Estimated apparent survival rates of ducklings to 22 days of age were 0.10 for those that remained with radio-marked females, 0.16 for all ducklings, including those that had joined other broods, and 0.31 for broods. Ducklings brooded by larger females experienced higher survival than those brooded by smaller females, and ducklings that hatched earlier in the breeding season survived at higher rates. Overland brood movements of 1 km or more occurred in both years, and survival was greatest for ducklings that dispersed from Karrak Lake to smaller ponds than on Karrak Lake itself, the central nesting area. Estimates of duckling survival, combined with relative contributions to the population by adults, suggest that ecological factors such as body size can influence population growth. Furthermore, low duckling survival and delayed maturity, emphasize the need of high adult survival for population growth to occur. These data, in combination with evidence of dispersal among wintering areas have helped contribute to a broader understanding of North American King Eider demographics.

Population surveys indicate a declining trend in abundance for the scoter genus at the continental level. Little is known about changes in life history traits responsible for the recent population decline of white-winged scoters (<i>Melanitta fusca deglandi</i>, hereafter scoters). Therefore, I studied nesting and duckling ecology of scoters at Redberry Lake, Saskatchewan, Canada during summers 2000-2001 when I found 198 nests. To examine nest-site selection, I compared habitat features between successful nests, failed nests, and random sites. Discriminant function analysis differentiated habitat features, measured at hatch, between successful nests, failed nests, and random sites; lateral (r = 0.65) and overhead (r = 0.35) concealment were microhabitat variables most correlated with canonical discriminant functions. I also modeled daily survival rate (DSR) of nests as a function of year, linear and quadratic trends with nest age, nest initiation date, and seven microhabitat variables. Nest survival from a time constant model (i.e., Mayfield nest success estimate) was 0.35 (95% CL: 0.27, 0.43). Estimates of nest success were lower than those measured at Redberry Lake in the 1970s and 1980s. In addition to nest survival increasing throughout the laying period and stabilizing during incubation, nest survival showed positive relationships with nest concealment and distance to water, and a negative relationship with distance to edge. Considering these factors, a model-averaged estimate of nest survival was 0.24 (95% CL: 0.09, 0.42). I conclude that scoters selected nesting habitat adaptively because (1) successful sites were more concealed than failed sites, (2) nest sites (i.e., successful and failed) had higher concealment than random sites, and (3) nest sites were on islands where success is greater than mainland. I then estimated duckling and brood survival with Cormack-Jolly-Seber models, implemented in Program Mark, from observations of 94 and 664 individually marked adult hens and ducklings, respectively. I tested hypotheses about duckling survival and (1) hatch date, (2) initial brood size at hatch, (3) duckling size and body condition at hatch, (4) offspring sex, (5) maternal female size and body condition at hatch, and (6) weather conditions within one week of hatching. Most mortality occurred during the first six days of duckling age. Variation in both duckling and brood survival were best modeled with effects of hatch date and initial brood size, while effects of female condition, female size, duckling size, and duckling condition were inconsistent. Survival probability clearly decreased with advancing hatch date and increased with larger initial brood sizes. Effects of weather and offspring sex in 2001, the only year such information was collected, suggested survival was negatively related to poor weather, but sex of ducklings, beyond size-related differences (i.e., sexual-size dimorphism), was unimportant. Estimates of survival to 28 days of age (30-day period), whether for ducklings (0.016, 0.021) or broods (0.084, 0.138) in 2000 or 2001, respectively, are the lowest of published studies and first for scoter broods in North America. I suspect intense gull predation shortly after hatch had the largest influence on duckling survival. Further research is needed to ascertain if low nesting success and duckling survival as well as other life cycle components are limiting scoter populations locally and throughout the rest of their breeding range.

Population surveys indicate a declining trend in abundance for the scoter genus at the continental level. Little is known about changes in life history traits responsible for the recent population decline of white-winged scoters (<i>Melanitta fusca deglandi</i>, hereafter scoters). Therefore, I studied nesting and duckling ecology of scoters at Redberry Lake, Saskatchewan, Canada during summers 2000-2001 when I found 198 nests. To examine nest-site selection, I compared habitat features between successful nests, failed nests, and random sites. Discriminant function analysis differentiated habitat features, measured at hatch, between successful nests, failed nests, and random sites; lateral (r = 0.65) and overhead (r = 0.35) concealment were microhabitat variables most correlated with canonical discriminant functions. I also modeled daily survival rate (DSR) of nests as a function of year, linear and quadratic trends with nest age, nest initiation date, and seven microhabitat variables. Nest survival from a time constant model (i.e., Mayfield nest success estimate) was 0.35 (95% CL: 0.27, 0.43). Estimates of nest success were lower than those measured at Redberry Lake in the 1970s and 1980s. In addition to nest survival increasing throughout the laying period and stabilizing during incubation, nest survival showed positive relationships with nest concealment and distance to water, and a negative relationship with distance to edge. Considering these factors, a model-averaged estimate of nest survival was 0.24 (95% CL: 0.09, 0.42). I conclude that scoters selected nesting habitat adaptively because (1) successful sites were more concealed than failed sites, (2) nest sites (i.e., successful and failed) had higher concealment than random sites, and (3) nest sites were on islands where success is greater than mainland. I then estimated duckling and brood survival with Cormack-Jolly-Seber models, implemented in Program Mark, from observations of 94 and 664 individually marked adult hens and ducklings, respectively. I tested hypotheses about duckling survival and (1) hatch date, (2) initial brood size at hatch, (3) duckling size and body condition at hatch, (4) offspring sex, (5) maternal female size and body condition at hatch, and (6) weather conditions within one week of hatching. Most mortality occurred during the first six days of duckling age. Variation in both duckling and brood survival were best modeled with effects of hatch date and initial brood size, while effects of female condition, female size, duckling size, and duckling condition were inconsistent. Survival probability clearly decreased with advancing hatch date and increased with larger initial brood sizes. Effects of weather and offspring sex in 2001, the only year such information was collected, suggested survival was negatively related to poor weather, but sex of ducklings, beyond size-related differences (i.e., sexual-size dimorphism), was unimportant. Estimates of survival to 28 days of age (30-day period), whether for ducklings (0.016, 0.021) or broods (0.084, 0.138) in 2000 or 2001, respectively, are the lowest of published studies and first for scoter broods in North America. I suspect intense gull predation shortly after hatch had the largest influence on duckling survival. Further research is needed to ascertain if low nesting success and duckling survival as well as other life cycle components are limiting scoter populations locally and throughout the rest of their breeding range.

This study provides information on the rates of DNA synthesis, sites of DNA synthesis, and DNA content of the avian salt gland during the osmoticstressing (plasma membrane synthesis) and destressing (plasma membrane turnover) cycle, in an effort to better understand the relationship of cell turnover to the initial events in plasma membrane amplification, differentiation, and turnover. The rate of DNA synthesis increases 12-24 h after the onset of osmotic stress, is maximal at about 24 h of osmotic stress, and decreases thereafter in fully stressed and destressed glands. The maximum DNA and protein content, and the maximum protein/DNA ratio are obtained after about 3 days of stress. Autoradiograms show that at 24 h of stress 70-80% of DNA synthesis occurs in connective tissue cells and 20-30% in parenchymal cells, but by 6 days of stress, synthesis occurs about equally in these cell groups. Because destressing is characterized by a large decrease in plasma membrane and in glandular protein, but by little DNA turnover or loss, the loss of plasma membrane is likely due to some type of cell dedifferentiation rather than cell turnover.

Like life-history theory, wildlife management decisions are typically predicated on trade-offs between benefits associated with investing resources to achieve higher reproductive or survival rates versus costs or risks of achieving those goals. On the Canadian prairies, most waterfowl conservation resources are directed to policies and programs that seek to increase duck nesting success. Limited attention has focused on post-hatching life-cycle stages, yet, despite considerable recent work on duckling survival rates, many uncertainties remain concerning how abiotic and biotic factors affect duckling survival rates. The role of upland habitat characteristics may be important but has received limited attention. I evaluated hypothesized sources of variation in duckling survival for 617 mallard (Anas platyrhynchos) broods on 27 Canadian prairie-parkland sites, with emphasis on assessing effects of managed and remnant natural upland habitats. I contrasted suites of a priori and post hoc exploratory models that incorporated effects of landscape, weather, female and brood-related variables to explain variation in duckling survival rates. Survival was lower for ducklings that used areas with high proportions of semi-permanent wetlands, as well as for broods that travelled farther overland. Exploratory analyses revealed further that survival of ducklings was negatively related to the amount of managed hayland. In contrast, duckling survival was positively associated with the amount managed grassland. There was no evidence of trade-offs between benefits of managing habitat to enhance duck nesting success versus costs in terms of lower subsequent duckling survival.<p> I also addressed unresolved questions about how birds balance costs and benefits of selecting habitats by determining the survival consequences of habitat choices made during brood-rearing. In theory, fitness should be higher in preferred habitats, but this assumption is rarely tested. Fitness consequences (i.e., duckling survival) of habitat selection patterns were determined at landscape and local scales using logistic regression and information-theoretic model selection techniques. Best-approximating landscape-level models indicated that mallard females selected brood-rearing areas with a high proportion of wetland and perennial upland habitats, but duckling survival was not related to habitat selection patterns at this scale. At finer spatial scales, females selected brood-rearing areas with high proportions of wetland habitats, but, contrary to expectation, duckling survival was lower when females raised their broods in these areas. Females avoided areas with abundant perennial cover and wetlands with little vegetative cover and, consistent with prediction, duckling survival was higher when females selected areas with low perennial cover. Thus, females did not consistently select brood-rearing habitats that conferred the highest fitness benefits. Rather, the relationship between habitat selection and duckling survival depended on spatial scale and habitats considered.

Like life-history theory, wildlife management decisions are typically predicated on trade-offs between benefits associated with investing resources to achieve higher reproductive or survival rates versus costs or risks of achieving those goals. On the Canadian prairies, most waterfowl conservation resources are directed to policies and programs that seek to increase duck nesting success. Limited attention has focused on post-hatching life-cycle stages, yet, despite considerable recent work on duckling survival rates, many uncertainties remain concerning how abiotic and biotic factors affect duckling survival rates. The role of upland habitat characteristics may be important but has received limited attention. I evaluated hypothesized sources of variation in duckling survival for 617 mallard (Anas platyrhynchos) broods on 27 Canadian prairie-parkland sites, with emphasis on assessing effects of managed and remnant natural upland habitats. I contrasted suites of a priori and post hoc exploratory models that incorporated effects of landscape, weather, female and brood-related variables to explain variation in duckling survival rates. Survival was lower for ducklings that used areas with high proportions of semi-permanent wetlands, as well as for broods that travelled farther overland. Exploratory analyses revealed further that survival of ducklings was negatively related to the amount of managed hayland. In contrast, duckling survival was positively associated with the amount managed grassland. There was no evidence of trade-offs between benefits of managing habitat to enhance duck nesting success versus costs in terms of lower subsequent duckling survival.<p> I also addressed unresolved questions about how birds balance costs and benefits of selecting habitats by determining the survival consequences of habitat choices made during brood-rearing. In theory, fitness should be higher in preferred habitats, but this assumption is rarely tested. Fitness consequences (i.e., duckling survival) of habitat selection patterns were determined at landscape and local scales using logistic regression and information-theoretic model selection techniques. Best-approximating landscape-level models indicated that mallard females selected brood-rearing areas with a high proportion of wetland and perennial upland habitats, but duckling survival was not related to habitat selection patterns at this scale. At finer spatial scales, females selected brood-rearing areas with high proportions of wetland habitats, but, contrary to expectation, duckling survival was lower when females raised their broods in these areas. Females avoided areas with abundant perennial cover and wetlands with little vegetative cover and, consistent with prediction, duckling survival was higher when females selected areas with low perennial cover. Thus, females did not consistently select brood-rearing habitats that conferred the highest fitness benefits. Rather, the relationship between habitat selection and duckling survival depended on spatial scale and habitats considered.