Episodes of high recruitment buffer against climate-driven mass mortality events in a North Pacific seabird population.

Longitudinal studies of marked animals provide an opportunity to assess the relative contributions of survival and reproductive output to population dynamics and change. Cassin's auklets are a long-lived seabird that maximizes annual reproductive effort in resource-rich years through a behaviour called double brooding, the initiation of a second breeding attempt following the success of the first during the same season. Our objective was to explore whether double brooding influenced population change by contributing a greater number of future recruits. We fit temporal symmetry models to 32 years of mark-recapture data of Cassin's auklets to infer the mechanisms underlying the observed variability in per capita recruitment rates. We found that periodic peaks in recruitment were explained by an increase in available nest sites, the proportion of the population double brooding 4 years prior, and spring upwelling conditions. Estimates of population change suggests a relatively stable population throughout the time series, attributable to a 'floating' demographic class of sexually mature individuals excluded from breeding by competition which quickly fill vacant sites following periods of low adult survival. Our results highlight the importance of recruitment in maintaining the population of a long-lived seabird periodically impacted by adverse environmental conditions.

Oceanographic drivers of winter habitat use in Cassin's Auklets.

Reduced prey abundance and severe weather can lead to a greater risk of mortality for seabirds during the non-breeding winter months. Resource patterns in some regions are shifting and becoming more variable in relation to past conditions, potentially further impacting survival and carryover to the breeding season. As animal tracking technologies and methods to analyze movement data have advanced, it has become increasingly feasible to draw fine-scale inference about how environmental variation affects foraging behavior and habitat use of seabirds during this critical period. Here, we used archival light-sensing tags to evaluate how interannual variation in oceanography affected the winter distribution of Cassin's Auklets from Southeast Farallon Island, California. Thirty-five out of 93 geolocators deployed from 2015 to 2017 were recovered and successfully recorded light-level data, from which geographic positions were estimated. Step-selection functions were applied to identify environmental covariates that best explained winter movement decisions and habitat use, revealing Cassin's Auklets dispersed farther from the colony during a winter with warm SST anomalies, but remained more centralized near the breeding colony during two average winters. Movement patterns were driven by avoidance of areas with higher sea surface temperatures and possible limits of dispersal from the breeding colony, and selection for areas with well-defined mesoscale fronts and cooler surface waters. Through multiple years of tagging and the application of step-selection functions, a robust and widely applied approach for analyzing animal movement in terrestrial species, we show how interannual differences in the movement patterns of a small seabird are driven by oceanographic variability across years. Understanding the winter habitat use of seabirds can help inform changes in population structure and measures of reproductive success, aiding managers in determining potential causes of breeding failures.

Increased reproductive investment associated with greater survival and longevity in Cassin's auklets.

Individuals increase lifetime reproductive output through a trade-off between investment in future survival and immediate reproductive success. This pattern may be obscured in certain higher quality individuals that possess greater reproductive potential. The Cassin's auklet (Ptychoramphus aleuticus) is a long-lived species where some individuals exhibit greater reproductive ability through a behaviour called double brooding. Here, we analyse 32 years of breeding histories from marked known-age auklets to test whether double brooding increases lifetime fitness despite the increased mortality and reduced lifespan higher reproductive effort would be expected to incur. Multistate mark-recapture modelling revealed that double brooding was strongly positively associated with higher annual survival and longevity. The mean (95% confidence interval) apparent survival was 0.69 (0.21, 0.91) for individuals that executed a single brood and 0.96 (0.84, 0.99) for those that double-brooded. Generalized linear mixed models indicated individuals that attempted multiple double broods over their lifetime were able to produce on average seven times as many chicks and live nearly 6 years longer than birds that never attempted a double brood. We found that high-quality individuals exhibited both increased reproductive effort and longevity, where heterogeneity in individual quality masked expected life-history trade-offs.

Maintenance of influenza A viruses and antibody response in mallards (Anas platyrhynchos) sampled during the non-breeding season in Alaska.

Prevalence of influenza A virus (IAV) infections in northern-breeding waterfowl has previously been reported to reach an annual peak during late summer or autumn; however, little is known about IAV infection dynamics in waterfowl populations persisting at high-latitude regions such as Alaska, during winter. We captured mallards (Anas platyrhynchos) throughout the non-breeding season (August-April) of 2012-2015 in Fairbanks and Anchorage, the two largest cities in Alaska, to assess patterns of IAV infection and antibody production using molecular methods and a standard serologic assay. In addition, we used virus isolation, genetic sequencing, and a virus microneutralization assay to characterize viral subtypes and to evaluate the immune response of mallards captured on multiple occasions through time. We captured 923 mallards during three successive sampling years: Fairbanks in 2012/13 and 2013/14, and Anchorage in 2014/15. Prevalence varied by age, season, and year/site with high and relatively stable estimates throughout the non-breeding season. Infected birds were detected in all locations/seasons except early-winter in Fairbanks during 2013/14. IAVs with 17 combinations of hemagglutinin (H1-5, H7-9, H11, H12) and neuraminidase (N1-6, N8, N9) subtypes were isolated. Antibodies to IAVs were detected throughout autumn and winter for all sampling locations and years, however, seroprevalence was higher among adults and varied among years. Mallards exhibited individual heterogeneity with regard to immune response, providing instances of both seroconversion and seroreversion to detected viral subtypes. The probability that an individual transitioned from one serostatus to another varied by age, with juvenile mallards having higher rates of seroconversion and seroreversion than adults. Our study provides evidence that a diversity of IAVs circulate in populations of mallards wintering at urban locations in Alaska, and we suggest waterfowl wintering at high-latitudes may play an important role in maintenance of viruses across breeding seasons.

Detection, prevalence, and transmission of avian hematozoa in waterfowl at the Arctic/sub-Arctic interface: co-infections, viral interactions, and sources of variation.

BACKGROUND: The epidemiology of avian hematozoa at high latitudes is still not well understood, particularly in sub-Arctic and Arctic habitats, where information is limited regarding seasonality and range of transmission, co-infection dynamics with parasitic and viral agents, and possible fitness consequences of infection. Such information is important as climate warming may lead to northward expansion of hematozoa with unknown consequences to northern-breeding avian taxa, particularly populations that may be previously unexposed to blood parasites. METHODS: We used molecular methods to screen blood samples and cloacal/oropharyngeal swabs collected from 1347 ducks of five species during May-August 2010, in interior Alaska, for the presence of hematozoa, Influenza A Virus (IAV), and IAV antibodies. Using models to account for imperfect detection of parasites, we estimated seasonal variation in prevalence of three parasite genera (Haemoproteus, Plasmodium, Leucocytozoon) and investigated how co-infection with parasites and viruses were related to the probability of infection. RESULTS: We detected parasites from each hematozoan genus in adult and juvenile ducks of all species sampled. Seasonal patterns in detection and prevalence varied by parasite genus and species, age, and sex of duck hosts. The probabilities of infection for Haemoproteus and Leucocytozoon parasites were strongly positively correlated, but hematozoa infection was not correlated with IAV infection or serostatus. The probability of Haemoproteus infection was negatively related to body condition in juvenile ducks; relationships between Leucocytozoon infection and body condition varied among host species. CONCLUSIONS: We present prevalence estimates for Haemoproteus, Leucocytozoon, and Plasmodium infections in waterfowl at the interface of the sub-Arctic and Arctic and provide evidence for local transmission of all three parasite genera. Variation in prevalence and molecular detection of hematozoa parasites in wild ducks is influenced by seasonal timing and a number of host traits. A positive correlation in co-infection of Leucocytozoon and Haemoproteus suggests that infection probability by parasites in one or both genera is enhanced by infection with the other, or that encounter rates of hosts and genus-specific vectors are correlated. Using size-adjusted mass as an index of host condition, we did not find evidence for strong deleterious consequences of hematozoa infection in wild ducks.

Transmission of influenza reflects seasonality of wild birds across the annual cycle.

Influenza A Viruses (IAV) in nature must overcome shifting transmission barriers caused by the mobility of their primary host, migratory wild birds, that change throughout the annual cycle. Using a phylogenetic network of viral sequences from North American wild birds (2008-2011) we demonstrate a shift from intraspecific to interspecific transmission that along with reassortment, allows IAV to achieve viral flow across successive seasons from summer to winter. Our study supports amplification of IAV during summer breeding seeded by overwintering virus persisting locally and virus introduced from a wide range of latitudes. As birds migrate from breeding sites to lower latitudes, they become involved in transmission networks with greater connectivity to other bird species, with interspecies transmission of reassortant viruses peaking during the winter. We propose that switching transmission dynamics may be a critical strategy for pathogens that infect mobile hosts inhabiting regions with strong seasonality.

A point mutation in the polymerase protein PB2 allows a reassortant H9N2 influenza isolate of wild-bird origin to replicate in human cells.

H9N2 influenza A viruses are on the list of potentially pandemic subtypes. Therefore, it is important to understand how genomic reassortment and genetic polymorphisms affect phenotypes of H9N2 viruses circulating in the wild bird reservoir. A comparative genetic analysis of North American H9N2 isolates of wild bird origin identified a naturally occurring reassortant virus containing gene segments derived from both North American and Eurasian lineage ancestors. The PB2 segment of this virus encodes 10 amino acid changes that distinguish it from other H9 strains circulating in North America. G590S, one of the 10 amino acid substitutions observed, was present in ~12% of H9 viruses worldwide. This mutation combined with R591 has been reported as a marker of pathogenicity for human pandemic 2009 H1N1 viruses. Screening by polymerase reporter assay of all the natural polymorphisms at these two positions identified G590/K591 and S590/K591 as the most active, with the highest polymerase activity recorded for the SK polymorphism. Rescued viruses containing these two polymorphic combinations replicated more efficiently in MDCK cells and they were the only ones tested that were capable of establishing productive infection in NHBE cells. A global analysis of all PB2 sequences identified the K591 signature in six viral HA/NA subtypes isolated from several hosts in seven geographic locations. Interestingly, introducing the K591 mutation into the PB2 of a human-adapted H3N2 virus did not affect its polymerase activity. Our findings demonstrate that a single point mutation in the PB2 of a low pathogenic H9N2 isolate could have a significant effect on viral phenotype and increase its propensity to infect mammals. However, this effect is not universal, warranting caution in interpreting point mutations without considering protein sequence context.

Trophic dynamics of shrinking Subarctic lakes: naturally eutrophic waters impart resilience to rising nutrient and major ion concentrations.

Shrinking lakes were recently observed for several Arctic and Subarctic regions due to increased evaporation and permafrost degradation. Along with lake drawdown, these processes often boost aquatic chemical concentrations, potentially impacting trophic dynamics. In particular, elevated chemical levels may impact primary productivity, which may in turn influence populations of primary and secondary consumers. We examined trophic dynamics of 18 shrinking lakes of the Yukon Flats, Alaska, that had experienced pronounced increases in nutrient (>200 % total nitrogen, >100 % total phosphorus) and ion concentrations (>100 % for four major ions combined) from 1985-1989 to 2010-2012, versus 37 stable lakes with relatively little chemical change over the same period. We found that phytoplankton stocks, as indexed by chlorophyll concentrations, remained unchanged in both shrinking and stable lakes from the 1980s to 2010s. Moving up the trophic ladder, we found significant changes in invertebrate abundance across decades, including decreased abundance of five of six groups examined. However, these decadal losses in invertebrate abundance were not limited to shrinking lakes, occurring in lakes with stable surface areas as well. At the top of the food web, we observed that probabilities of lake occupancy for ten waterbird species, including adults and chicks, remained unchanged from the period 1985-1989 to 2010-2012. Overall, our study lakes displayed a high degree of resilience to multi-trophic cascades caused by rising chemical concentrations. This resilience was likely due to their naturally high fertility, such that further nutrient inputs had little impact on waters already near peak production.

Estimating demographic parameters using a combination of known-fate and open N-mixture models.

Accurate estimates of demographic parameters are required to infer appropriate ecological relationships and inform management actions. Known-fate data from marked individuals are commonly used to estimate survival rates, whereas N-mixture models use count data from unmarked individuals to estimate multiple demographic parameters. However, a joint approach combining the strengths of both analytical tools has not been developed. Here we develop an integrated model combining known-fate and open N-mixture models, allowing the estimation of detection probability, recruitment, and the joint estimation of survival. We demonstrate our approach through both simulations and an applied example using four years of known-fate and pack count data for wolves (Canis lupus). Simulation results indicated that the integrated model reliably recovered parameters with no evidence of bias, and survival estimates were more precise under the joint model. Results from the applied example indicated that the marked sample of wolves was biased toward individuals with higher apparent survival rates than the unmarked pack mates, suggesting that joint estimates may be more representative of the overall population. Our integrated model is a practical approach for reducing bias while increasing precision and the amount of information gained from mark-resight data sets. We provide implementations in both the BUGS language and an R package.

Pronounced chemical response of Subarctic lakes to climate-driven losses in surface area.

Losses in lake area have been observed for several Arctic and Subarctic regions in recent decades, with unknown consequences for lake ecosystems. These reductions are primarily attributed to two climate-sensitive mechanisms, both of which may also cause changes in water chemistry: (i) increased imbalance of evaporation relative to inflow, whereby increased evaporation and decreased inflow act to concentrate solutes into smaller volumes; and (ii) accelerated permafrost degradation, which enhances sublacustrine drainage while simultaneously leaching previously frozen solutes into lakes. We documented changes in nutrients [total nitrogen (TN), total phosphorus (TP)] and ions (calcium, chloride, magnesium, sodium) over a 25 year interval in shrinking, stable, and expanding Subarctic lakes of the Yukon Flats, Alaska. Concentrations of all six solutes increased in shrinking lakes from 1985-1989 to 2010-2012, while simultaneously undergoing little change in stable or expanding lakes. This created a present-day pattern, much weaker or absent in the 1980s, in which shrinking lakes had higher solute concentrations than their stable or expanding counterparts. An imbalanced evaporation-to-inflow ratio (E/I) was the most likely mechanism behind such changes; all four ions, which behave semiconservatively and are prone to evapoconcentration, increased in shrinking lakes and, along with TN and TP, were positively related to isotopically derived E/I estimates. Moreover, the most conservative ion, chloride, increased >500% in shrinking lakes. Conversely, only TP concentration was related to probability of permafrost presence, being highest at intermediate probabilities. Overall, the substantial increases of nutrients (TN >200%, TP >100%) and ions (>100%) may shift shrinking lakes towards overly eutrophic or saline states, with potentially severe consequences for ecosystems of northern lakes.

Herbivores influence the growth, reproduction, and morphology of a widespread Arctic willow.

Shrubs have expanded in Arctic ecosystems over the past century, resulting in significant changes to albedo, ecosystem function, and plant community composition. Willow and rock ptarmigan (Lagopus lagopus, L. muta) and moose (Alces alces) extensively browse Arctic shrubs, and may influence their architecture, growth, and reproduction. Furthermore, these herbivores may alter forage plants in such a way as to increase the quantity and accessibility of their own food source. We estimated the effect of winter browsing by ptarmigan and moose on an abundant, early-successional willow (Salix alaxensis) in northern Alaska by comparing browsed to unbrowsed branches. Ptarmigan browsed 82-89% of willows and removed 30-39% of buds, depending on study area and year. Moose browsed 17-44% of willows and browsed 39-55% of shoots. Browsing inhibited apical dominance and activated axillary and adventitious buds to produce new vegetative shoots. Ptarmigan- and moose-browsed willow branches produced twice the volume of shoot growth but significantly fewer catkins the following summer compared with unbrowsed willow branches. Shoots on browsed willows were larger and produced 40-60% more buds compared to unbrowsed shoots. This process of shoot production at basal parts of the branch is the mechanism by which willows develop a highly complex "broomed" architecture after several years of browsing. Broomed willows were shorter and more likely to be re-browsed by ptarmigan, but not moose. Ptarmigan likely benefit from the greater quantity and accessibility of buds on previously browsed willows and may increase the carrying capacity of their own habitat. Despite the observed tolerance of willows to browsing, their vertical growth and reproduction were strongly inhibited by moose and ptarmigan. Browsing by these herbivores therefore needs to be considered in future models of shrub expansion in the Arctic.

Multi-trophic resilience of boreal lake ecosystems to forest fires.

Fires are the major natural disturbance in the boreal forest, and their frequency and intensity will likely increase as the climate warms. Terrestrial nutrients released by fires may be transported to boreal lakes, stimulating increased primary productivity, which may radiate through multiple trophic levels. Using a before-after-control-impact (BACI) design, with pre- and postfire data from burned and unburned areas, we examined effects of a natural fire across several trophic levels of boreal lakes, from nutrient and chlorophyll levels, to macroinvertebrates, to waterbirds. Concentrations of total nitrogen and phosphorus were not affected by the fire. Chlorophyll a levels were also unaffected, likely reflecting the stable nutrient concentrations. For aquatic invertebrates, we found that densities of three functional feeding groups did not respond to the fire (filterers, gatherers, scrapers), while two groups increased (shredders, predators). Amphipods accounted for 98% of shredder numbers, and we hypothesize that fire-mediated habitat changes may have favored their generalist feeding and habitat ecology. This increase in amphipods may, in turn, have driven increased predator densities, as amphipods were the most numerous invertebrate in our lakes and are commonly taken as prey. Finally, abundance of waterbird young, which feed primarily on aquatic invertebrates, was not affected by the fire. Overall, ecosystems of our study lakes were largely resilient to forest fires, likely due to their high initial nutrient concentrations and small catchment sizes. Moreover, this resilience spanned multiple trophic levels, a significant result for ecologically similar boreal regions, especially given the high potential for increased fires with future climate change.

e-Measures: insight into the challenges and opportunities of automating publicly reported quality measures.

Using electronic health records (EHR) to automate publicly reported quality measures is receiving increasing attention and is one of the promises of EHR implementation. Kaiser Permanente has fully or partly automated six of 13 the joint commission measure sets. We describe our experience with automation and the resulting time savings: a reduction by approximately 50% of abstractor time required for one measure set alone (surgical care improvement project). However, our experience illustrates the gap between the current and desired states of automated public quality reporting, which has important implications for measure developers, accrediting entities, EHR vendors, public/private payers, and government.

Individual heterogeneity in black brant survival and recruitment with implications for harvest dynamics.

We examined individual heterogeneity in survival and recruitment of female Pacific black brant (Branta bernicla nigricans) using frailty models adapted to a capture-mark-recapture context. Our main objectives were (1) to quantify levels of heterogeneity and examine factors affecting heterogeneity, and (2) model the effects of individual heterogeneity on harvest dynamics through matrix models. We used 24 years of data on brant marked and recaptured at the Tutakoke River colony, AK. Multievent models were fit as hidden Markov chain using program E-SURGE with an adequate overdispersion coefficient. Annual survival of individuals marked as goslings was heterogeneous among individuals and year specific with about 0.23 difference in survival between "high" (0.73)- and "low" (0.50)-quality individuals at average survival probability. Adult survival (0.85 ± 0.004) was homogeneous and higher than survival of both groups of juveniles. The annual recruitment probability was heterogeneous for brant >1-year-old; 0.56 (±0.21) and 0.31 (±0.03) for high- and low-quality individuals, respectively. Assuming equal clutch sizes for high- and low-quality individuals and that 80% of offspring were in the same quality class as the breeding female resulted in reproductive values about twice as high for high-quality individuals than low-quality individual for a given class of individuals producing differential contributions to population growth among groups. Differences in reproductive values greatly increased when we assumed high-quality individuals had larger clutch sizes. When we assumed that 50% of offspring were in the same quality class as their mothers and clutches were equal, differences in reproductive values between quality classes were greatly reduced or eliminated (breeders [BRs]). We considered several harvest scenarios using the assumption that 80% of offspring were in the same quality class as their mothers. The amount of compensation for harvest mortality declined as the proportion of high-quality individuals in the harvest increased, as differences in clutch sizes between groups decreased and as the proportion of BRs in the harvest increased. Synthesis and applications. Harvest at the same proportional level of the overall population can result in variable responses in population growth rate when heterogeneity is present in a population. λ was <1.0 under every scenario when harvest rates were >10%, and heterogeneity caused as much as +2% difference in growth rates at the highest levels of proportional harvest for low-quality individuals and the greatest differences in qualities between classes of individuals, a critical difference for a population with λ near 1.0 such as the brant. We observed less response in overall survival in the presence of heterogeneity because we did not observe heterogeneity in the annual survival of BRs. This analysis provides a comprehensive view of overall compensation at the population level and also constitutes the first example of a survival-recruitment model with heterogeneity. Individual heterogeneity should be more explicitly considered in harvest management of vertebrates.

Time-lagged variation in pond density and primary productivity affects duck nest survival in the Prairie Pothole Region.

The Prairie Pothole Region (PPR) is the primary breeding region for most species of North American dabbling ducks (Anas spp.). Conservation of these species is guided in part by knowledge of relationships between nest survival probability and habitat features. Positive relationships between duck nest survival and amount and configuration of herbaceous perennial vegetation have been observed in previous studies, but these 2- to 4-year studies might not have adequately characterized the temporal effect of wet-dry episodes on nest survival. Over an eight-year period, we studied nest survival of five species of ducks in the PPR relative to spatial and temporal variation in pond density, primary productivity, and hydrologic status of wetlands, soil, and vegetation on 52 study sites selected to span a gradient of spatial variation in proportion of herbaceous perennial vegetation and in number of wetland basins. We observed the fate of 12 754 nests. Consistent with past studies, 90% of nests that failed to hatch were destroyed by predators. Nest survival probability was positively related to current-year pond density and primary productivity, negatively related to pond density and primary productivity during the previous two years, and positively related to the number of wetland basins on the study site. Predicted relationships between nest survival and proportion or configuration of herbaceous perennial vegetation in the surrounding landscape were not supported. For mallard (Anas platyrhynchos), median estimated nest survival probability ranged from 0.02 (SE = 0.01) to 0.22 (SE = 0.02). Estimated nest survival was greatest on sites with numerous wetland basins that had transitioned from dry, unproductive conditions to wet, productive conditions in the previous 1-2 years. Our results were consistent with time-lagged responses of food webs to resource pulses in a broad array of ecosystems. Our study highlighted the importance of wetland basins and wet-dry episodes to duck nest survival in the PPR. Current habitat conservation efforts focus on landscapes with numerous wetland basins and a high proportion of herbaceous perennial vegetation. Our results suggest that future conservation efforts should focus on preserving high-density wetland complexes across as large a geographic extent as possible even in cropland-dominated landscapes.

Earlier migration timing, decreasing phenotypic variation, and biocomplexity in multiple salmonid species.

Climate-induced phenological shifts can influence population, evolutionary, and ecological dynamics, but our understanding of these phenomena is hampered by a lack of long-term demographic data. We use a multi-decade census of 5 salmonid species representing 14 life histories in a warming Alaskan stream to address the following key questions about climate change and phenology: How consistent are temporal patterns and drivers of phenology for similar species and alternative life histories? Are shifts in phenology associated with changes in phenotypic variation? How do phenological changes influence the availability of resource subsidies? For most salmonid species, life stages, and life histories, freshwater temperature influences migration timing--migration events are occurring earlier in time (mean = 1.7 days earlier per decade over the 3-5 decades), and the number of days over which migration events occur is decreasing (mean = 1.5 days per decade). Temporal trends in migration timing were not correlated with changes in intra-annual phenotypic variation, suggesting that these components of the phenotypic distribution have responded to environmental change independently. Despite commonalities across species and life histories, there was important biocomplexity in the form of disparate shifts in migration timing and variation in the environmental factors influencing migration timing for alternative life history strategies in the same population. Overall, adult populations have been stable during these phenotypic and environmental changes (λ ≈ 1.0), but the temporal availability of salmon as a resource in freshwater has decreased by nearly 30 days since 1971 due to changes in the median date of migration timing and decreases in intra-annual variation in migration timing. These novel observations advance our understanding of phenological change in response to climate warming, and indicate that climate change has influenced the ecology of salmon populations, which will have important consequences for the numerous species that depend on this resource.

Nest-site selection of passerines: effects of geographic scale and public and personal information.

Nest-site selection is an important determinant of individual fitness in birds. Understanding what information individuals use to choose nest sites is therefore important for understanding the evolution of nest-site selection, the dynamics of populations, and the conservation of species. We used five years of mark-recapture data for Mountain Bluebirds (Sialia currucoides) to examine how dispersal probability and nest-site selection vary with potential cues of nest-site quality. Dispersal distance between breeding seasons and nest-site selection were modeled as a function of personal reproductive success, conspecific density, conspecific reproductive success, and habitat type. Between years, the dispersal probability was related to personal reproductive success, not conspecific information, and individuals fledging fewer young dispersed longer distances. For dispersing individuals, the probability that a nest site was selected in year i was negatively related to distance from the nest site selected in year i - 1 for all age and sex classes, and positively related to conspecific density and reproductive success in year i - 1 for both second-year (SY) and after-hatch-year (AHY) females. However, nest-site selection in year i was more strongly related to conspecific density in year i- 1 for hatch-year (HY) females and was much more strongly related to the reproductive success of conspecifics in year i - 1 for AHY females. Nest-site selection of HY and AHY males was not consistently related to the metrics of conspecific information, but we suspect that relationships were obscured by competitive interactions. We found no evidence indicating that individuals respond differently to conspecific information at longer distances, suggesting that individuals limit dispersal to areas where they have prior knowledge. We predict that these patterns of nest-site selection will allow birds to loosely track nest-site quality and maintain an ideal free distribution, where average fitness is equal in all habitat types.

Factors contributing to process variance in annual survival of female greater sage-grouse in Montana.

Populations of Greater Sage-Grouse (Centrocercus urophasianus) have declined by 69-99% from historic levels, and information on population dynamics of these birds at a landscape scale is essential to informed management. We examined the relationships between hen survival and a suite of landscape-scale habitat and environmental conditions. We radio-marked 237 female Sage-Grouse and measured 426 vegetation plots during 2001-2004 at four sites in a 3200-km2 landscape in north-central Montana, USA. We used program MARK to model monthly survival rates for 11 seasonal intervals. There was strong support for the best-approximating model (AICc weight = 0.810), which indicated that (1) hen survival varied by season within years and by year within seasons, (2) nesting hens had higher nesting-season survival than non-nesting hens, and (3) individuals at one site had lower hunting-season survival than at other sites. We observed considerable variation in hen survival. Process variation was 0.255, with an expected range of annual survival of 0.12 to 1.0. The ratio of process to total variation was 0.999, indicating that observed variation was real and not attributable to sampling variation. We observed a nearly fourfold difference in maximum and minimum annual survival, ranging from 0.962 +/- 0.024 (mean +/- SE) for nesting hens in 2001-2002 to 0.247 +/- 0.050) for non-nesters in 2003-2004. Low annual survival in 2003 resulted from the compounded effects of a West Nile virus outbreak in August and a severe winter in 2003-2004. Increased hen mortality associated with severe winter weather contrasts with prior beliefs that Sage-Grouse populations are typically unaffected by winter weather conditions and underscores the importance of protecting winter sagebrush (Artemisia spp.) habitats.

Individual quality, survival variation and patterns of phenotypic selection on body condition and timing of nesting in birds.

Questions about individual variation in "quality" and fitness are of great interest to evolutionary and population ecologists. Such variation can be investigated using either a random effects approach or an approach that relies on identifying observable traits that are themselves correlated with fitness components. We used the latter approach with data from 1,925 individual females of three species of ducks (tufted duck, Aythya fuligula; common pochard, Aythya ferina; northern shoveler, Anas clypeata) sampled on their breeding grounds at Engure Marsh, Latvia, for over 15 years. Based on associations with reproductive output, we selected two traits, one morphological (relative body condition) and one behavioral (relative time of nesting), that can be used to characterize individual females over their lifetimes. We then asked whether these traits were related to annual survival probabilities of nesting females. We hypothesized quadratic, rather than monotonic, relationships based loosely on ideas about the likely action of stabilizing selection on these two traits. Parameters of these relationships were estimated directly using ultrastructural models embedded within capture-recapture-band-recovery models. Results provided evidence that both traits were related to survival in the hypothesized manner. For all three species, females that tended to nest earlier than the norm exhibited the highest survival rates, but very early nesters experienced reduced survival and late nesters showed even lower survival. For shovelers, females in average body condition showed the highest survival, with lower survival rates exhibited by both heavy and light birds. For common pochard and tufted duck, the highest survival rates were associated with birds of slightly above-average condition, with somewhat lower survival for very heavy birds and much lower survival for birds in relatively poor condition. Based on results from this study and previous work on reproduction, we conclude that nest initiation date and body condition covary with both reproductive and survival components of fitness. These associations lead to a positive covariance of these two fitness components within individuals and to the conclusion that these two traits are indeed correlates of individual quality.