Non-native grazers affect physiological and demographic responses of greater sage-grouse.

Non-native ungulate grazing has negatively impacted native species across the globe, leading to massive loss of biodiversity and ecosystem services. Despite their pervasiveness, interactions between non-native grazers and native species are not fully understood. We often observe declines in demography or survival of these native species, but lack understanding about the mechanisms underlying these declines. Physiological stress represents one mechanism of (mal)adaptation, but data are sparse. We investigated glucocorticoid levels in a native avian herbivore exposed to different intensities of non-native grazing in the cold desert Great Basin ecosystem, USA. We measured corticosterone, a glucocorticoid in feathers for a large sample (n = 280) of female greater sage-grouse (Centrocercus urophasianus) from three study areas in Northern Nevada and Southern Oregon with different grazing regimes of livestock and feral horses. We found that greater feral horse density was associated with higher corticosterone levels, and this effect was exacerbated by drought conditions. Livestock grazing produced similar results; however, there was more model uncertainty about the livestock effect. Subsequent nesting success was lower with increased feather corticosterone, but corticosterone levels were not predictive of other vital rates. Our results indicate a physiological response by sage-grouse to grazing pressure from non-native grazers. We found substantial among-individual variation in the strength of the response. These adverse effects were intensified during unfavorable weather events, highlighting the need to reevaluate management strategies in the face of climate change.

Zero-inflated count distributions for capture-mark-reencounter data.

The estimation of demographic parameters is a key component of evolutionary demography and conservation biology. Capture-mark-recapture methods have served as a fundamental tool for estimating demographic parameters. The accurate estimation of demographic parameters in capture-mark-recapture studies depends on accurate modeling of the observation process. Classic capture-mark-recapture models typically model the observation process as a Bernoulli or categorical trial with detection probability conditional on a marked individual's availability for detection (e.g., alive, or alive and present in a study area). Alternatives to this approach are underused, but may have great utility in capture-recapture studies. In this paper, we explore a simple concept: in the same way that counts contain more information about abundance than simple detection/non-detection data, the number of encounters of individuals during observation occasions contains more information about the observation process than detection/non-detection data for individuals during the same occasion. Rather than using Bernoulli or categorical distributions to estimate detection probability, we demonstrate the application of zero-inflated Poisson and gamma-Poisson distributions. The use of count distributions allows for inference on availability for encounter, as well as a wide variety of parameterizations for heterogeneity in the observation process. We demonstrate that this approach can accurately recover demographic and observation parameters in the presence of individual heterogeneity in detection probability and discuss some potential future extensions of this method.

Density-dependence produces spurious relationships among demographic parameters in a harvested species.

Harvest of wild organisms is an important component of human culture, economy, and recreation, but can also put species at risk of extinction. Decisions that guide successful management actions therefore rely on the ability of researchers to link changes in demographic processes to the anthropogenic actions or environmental changes that underlie variation in demographic parameters. Ecologists often use population models or maximum sustained yield curves to estimate the impacts of harvest on wildlife and fish populations. Applications of these models usually focus exclusively on the impact of harvest and often fail to consider adequately other potential, often collinear, mechanistic drivers of the observed relationships between harvest and demographic rates. In this study, we used an integrated population model and long-term data (1973-2016) to examine the relationships among hunting and natural mortality, the number of hunters, habitat conditions, and population size of blue-winged teal Spatula discors, an abundant North American dabbling duck with a relatively fast-paced life history strategy. Over the last two and a half decades of the study, teal abundance tripled, hunting mortality probability increased slightly ( < 0.02 ), and natural mortality probability increased substantially ( > 0.1 ) at greater population densities. We demonstrate strong density-dependent effects on natural mortality and fecundity as population density increased, indicative of compensatory harvest mortality and compensatory natality. Critically, an analysis that only assessed the relationship between survival and hunting mortality would spuriously indicate depensatory mortality due to multicollinearity between abundance, natural mortality and hunting mortality. Our findings demonstrate that models that only consider the direct effect of hunting on survival or natural mortality can fail to accurately assess the mechanistic impact of hunting on population dynamics due to multicollinearity among demographic drivers. This multicollinearity limits inference and may have strong impacts on applied management actions globally.

Estimating survival and adoption rates of dependent juveniles.

Population growth and fitness are typically most sensitive to adult survival in long-lived species, but variation in recruitment often explains most of the variation in fitness, as past selection has canalized adult survival. Estimating juvenile survival until age of independence has proven challenging, because marking individuals in this age class may directly affect survival. For Greater Sage-grouse, uniquely marking juveniles in the first days of life likely results in adverse effects to survival, detection of juveniles is not perfect, and females adopt juveniles from other parents. These challenges are encountered by researchers studying avian and mammalian species with similar life histories, yet methods do not exist that explicitly estimate all these components of the recruitment process. We propose a novel data collection method and demographic model to simultaneously estimate rates of detection, survival, and adoption of juvenile individuals. Using multiple cameras to film the beginning of juvenile activity on specific days, we obtained counts of juveniles associated with marked females. Increases of juveniles to broods provided information that enabled us to estimate rates of adoption that can be applied at the population level. Losses from broods informed apparent survival. These losses could be attributed to death, or they could be chicks that were adopted by other females. We found evidence that apparent survival of juveniles was influenced by localized weather patterns when chicks were young. Similarly, we found that young chicks were more susceptible to the adverse effect of attending females being flushed by an observer. Both of these patterns diminished quickly as chicks aged. We provide the first-ever estimates of interval-specific adoption rates. Our results suggest that researchers should be cautious when designing studies to estimate juvenile survival. More importantly, they provide insight into adoption, a behavior that has been known to exist for decades.

Long-term trends and drought: Spatiotemporal variation in juvenile sex ratios of North American ducks.

Sex ratios affect population dynamics and individual fitness, and changing sex ratios can be indicative of shifts in sex-specific survival at different life stages. While climate and landscape changes alter sex ratios of wild bird populations, long-term, landscape scale assessments of sex ratios are rare. Further, little work has been done to understand changes in sex ratios in avian communities. In this manuscript, we analyze long-term (1961-2015) data on five species of ducks across five broad climatic regions of the United States to estimate the effects of drought and long-term trends on the proportion of juvenile females captured at banding. As waterfowl have a 1:1 sex ratio at hatch, we interpret changes in sex ratios of captured juveniles as changes in sex-specific survival rates during early life. Seven of 12 species-region pairs exhibited evidence for long-term trends in the proportion of juvenile females at banding. The proportion of juvenile females at banding increased for duck populations in the western United States and typically declined for duck populations in the eastern United States. We only observed evidence for an effect of drought in two of the 12 species-region pairs, where the proportion of females declined during drought. As changes to North American landscapes and climate continue and intensify, we expect continued changes in sex-specific juvenile survival rates. More broadly, we encourage further research examining the mechanisms underlying long-term trends in juvenile sex ratios in avian communities.

A hierarchical model for jointly assessing ecological and anthropogenic impacts on animal demography.

The management of sustainable harvest of animal populations is of great ecological and conservation importance. Development of formal quantitative tools to estimate and mitigate the impacts of harvest on animal populations has positively impacted conservation efforts. The vast majority of existing harvest models, however, do not simultaneously estimate ecological and harvest impacts on demographic parameters and population trends. Given that the impacts of ecological drivers are often equal to or greater than the effects of harvest, and can covary with harvest, this disconnect has the potential to lead to flawed inference. In this study, we used Bayesian hierarchical models and a 43-year capture-mark-recovery dataset from 404,241 female mallards Anas platyrhynchos released in the North American midcontinent to estimate mallard demographic parameters. Furthermore, we model the dynamics of waterfowl hunters and habitat, and the direct and indirect effects of anthropogenic and ecological processes on mallard demographic parameters. We demonstrate that density dependence, habitat conditions and harvest can simultaneously impact demographic parameters of female mallards, and discuss implications for existing and future harvest management models. Our results demonstrate the importance of controlling for multicollinearity among demographic drivers in harvest management models, and provide evidence for multiple mechanisms that lead to partial compensation of mallard harvest. We provide a novel model structure to assess these relationships that may allow for improved inference and prediction in future iterations of harvest management models across taxa.

A niche for null models in adaptive resource management.

As global systems rapidly change, our collective ability to predict future ecological dynamics will become increasingly important for successful natural resource management. By merging stakeholder objectives with system uncertainty, and by adapting actions to changing systems and knowledge, adaptive resource management (ARM) provides a rigorous platform for making sound decisions in a changing world. Critically, however, applications of ARM could be improved by employing benchmarks (i.e., points of reference) for determining when learning is occurring through the cycle of monitoring, modeling, and decision-making steps in ARM. Many applications of ARM use multiple model-based hypotheses to identify and reduce systematic uncertainty over time, but generally lack benchmarks for gauging discovery of scientific evidence and learning. This creates the danger of thinking that directional changes in model weights or rankings are indicative of evidence for hypotheses, when possibly all competing models are inadequate. There is thus a somewhat obvious, but yet to be filled niche for including benchmarks for learning in ARM. We contend that carefully designed "ecological null models," which are structured to produce an expected ecological pattern in the absence of a hypothesized mechanism, can serve as suitable benchmarks. Using a classic case study of mallard harvest management that is often used to demonstrate the successes of ARM for learning about ecological mechanisms, we show that simple ecological null models, such as population persistence (Nt +1 = Nt ), provide more robust near-term forecasts of population abundance than the currently used mechanistic models. More broadly, ecological null models can be used as benchmarks for learning in ARM that trigger the need for discarding model parameterizations and developing new ones when prevailing models underperform the ecological null model. Identifying mechanistic models that surpass these benchmarks will improve learning through ARM and help decision-makers keep pace with a rapidly changing world.

Individual heterogeneity in fitness in a long-lived herbivore.

Heterogeneity in the intrinsic quality and nutritional condition of individuals affects reproductive success and consequently fitness. Black brant (Branta bernicla nigricans) are long-lived, migratory, specialist herbivores. Long migratory pathways and short summer breeding seasons constrain the time and energy available for reproduction, thus magnifying life-history trade-offs. These constraints, combined with long lifespans and trade-offs between current and future reproductive value, provide a model system to examine the role of individual heterogeneity in driving life-history strategies and individual heterogeneity in fitness. We used hierarchical Bayesian models to examine reproductive trade-offs, modeling the relationships between within-year measures of reproductive energy allocation and among-year demographic rates of individual females breeding on the Yukon-Kuskokwim Delta, Alaska, using capture-recapture and reproductive data from 1988 to 2014. We generally found that annual survival tended to be buffered against variation in reproductive investment, while breeding probability varied considerably over the range of clutch size-laying date combinations. We provide evidence for relationships between breeding probability and clutch size, breeding probability and nest initiation date, and an interaction between clutch size and initiation date. Average lifetime clutch size also had a weak positive relationship with apparent survival probability. Our results support the use of demographic buffering strategies for black brant. These results also indirectly suggest associations among environmental conditions during growth, fitness, and energy allocation, highlighting the effects of early growth conditions on individual heterogeneity, and subsequently, lifetime reproductive investment.

Mate fidelity improves survival and breeding propensity of a long-lived bird.

Evolutionary and behavioural ecologists have long been interested in factors shaping the variation in mating behaviour observed in nature. Although much of the research on this topic has focused on the consequences of mate choice and mate change on annual reproductive success, studies of a potential positive link between mate fidelity and adult demographic rates have been comparatively rare. This is particularly true for long-lived birds with multi-year, socially monogamous pair bonds. We used a 26-year capture-mark-recapture dataset of 3,330 black brent Branta bernicla nigricans to test whether breeding with a familiar mate improved future breeding propensity and survival. We predicted that experienced breeders nesting with a new partner would have rates of survival similar to familiar pairs because long-lived species avoid jeopardizing survival since their lifetime fitness is sensitive to this vital rate. In contrast, we expected that any costs of breeding with a new partner would be paid through skipping the subsequent breeding attempt. We found that unfamiliar pairs had lower subsequent breeding propensity than faithful partners. However, contrary to our expectations, individuals breeding with a new mate also suffered reduced survival. These results add to a small number of studies indicating that a positive relationship between mate retention and adult demographic rates may exist in a diverse array of avian species. Given these results, researchers should consider costs of mate change that extend beyond within-season reproductive success to fully understand the potential adaptive basis for perennial social monogamy. We caution that if mate retention enhances survival prospects, improvements in annual reproductive success with pair-bond length could be a secondary factor favouring perennial social monogamy, particularly in species with slower life-history strategies. Furthermore, some cases where annual reproductive success does not improve with pair-bond duration, yet multi-year pair bonds are common, could be explained by benefits afforded by mate fidelity to adult vital rates.

Long-term research and hierarchical models reveal consistent fitness costs of being the last egg in a clutch.

Maintenance of phenotypic heterogeneity in the face of strong selection is an important component of evolutionary ecology, as are the consequences of such heterogeneity. Organisms may experience diminishing returns of increased reproductive allocation as clutch or litter size increases, affecting current and residual reproductive success. Given existing uncertainty regarding trade-offs between the quantity and quality of offspring, we sought to examine the potential for diminishing returns on increased reproductive allocation in a long-lived species of goose, with a particular emphasis on the effect of position in the laying sequence on offspring quality. To better understand the effects of maternal allocation on offspring survival and growth, we estimated the effects of egg size, timing of breeding, inter- and intra-annual variation, and position in the laying sequence on gosling survival and growth rates of black brant Branta bernicla nigricans breeding in western Alaska from 1987 to 2007. We found that gosling growth rates and survival decreased with position in the laying sequence, regardless of clutch size. Mean egg volume of the clutch a gosling originated from had a positive effect on gosling survival (ß = 0.095, 95% CRI: 0.024, 0.165) and gosling growth rates (ß = 0.626, 95% CRI: 0.469, 0.738). Gosling survival (ß = -0.146, 95% CRI: -0.214, -0.079) and growth rates (ß = -1.286, 95% CRI: -1.435, -1.132) were negatively related to hatching date. These findings indicate substantial heterogeneity in offspring quality associated with their position in the laying sequence. They also potentially suggest a trade-off mechanism for females whose total reproductive investment is governed by pre-breeding state.

Estimating correlations among demographic parameters in population models.

Estimating correlations among demographic parameters is critical to understanding population dynamics and life-history evolution, where correlations among parameters can inform our understanding of life-history trade-offs, result in effective applied conservation actions, and shed light on evolutionary ecology. The most common approaches rely on the multivariate normal distribution, and its conjugate inverse Wishart prior distribution. However, the inverse Wishart prior for the covariance matrix of multivariate normal distributions has a strong influence on posterior distributions. As an alternative to the inverse Wishart distribution, we individually parameterize the covariance matrix of a multivariate normal distribution to accurately estimate variances (σ 2) of, and process correlations (ρ) between, demographic parameters. We evaluate this approach using simulated capture-mark-recapture data. We then use this method to examine process correlations between adult and juvenile survival of black brent geese marked on the Yukon-Kuskokwim River Delta, Alaska (1988-2014). Our parameterization consistently outperformed the conjugate inverse Wishart prior for simulated data, where the means of posterior distributions estimated using an inverse Wishart prior were substantially different from the values used to simulate the data. Brent adult and juvenile annual apparent survival rates were strongly positively correlated (ρ = 0.563, 95% CRI 0.181-0.823), suggesting that habitat conditions have significant effects on both adult and juvenile survival. We provide robust simulation tools, and our methods can readily be expanded for use in other capture-recapture or capture-recovery frameworks. Further, our work reveals limits on the utility of these approaches when study duration or sample sizes are small.

Changes in behavior are unable to disrupt a trophic cascade involving a specialist herbivore and its food plant.

Changes in ecological conditions can induce changes in behavior and demography of wild organisms, which in turn may influence population dynamics. Black brant (Branta bernicla nigricans) nesting in colonies on the Yukon-Kuskokwim Delta (YKD) in western Alaska have declined substantially (~50%) since the turn of the century. Black brant are herbivores that rely heavily on Carex subspathacea (Hoppner's sedge) during growth and development. The availability of C. subspathacea affects gosling growth rates, which subsequently affect pre- and postfledging survival, as well as size and breeding probability as an adult. We predicted that long-term declines in C. subspathacea have affected gosling growth rates, despite the potential of behavior to buffer changes in food availability during brood rearing. We used Bayesian hierarchical mixed-effects models to examine long-term (1987-2015) shifts in brant behavior during brood rearing, forage availability, and gosling growth rates at the Tutakoke River colony. We showed that locomotion behaviors have increased (ß = 0.05, 95% CRI: 0.032-0.068) while resting behaviors have decreased (ß = -0.024, 95% CRI: -0.041 to -0.007), potentially in response to long-term shifts in forage availability and brood density. Concurrently, gosling growth rates have decreased substantially (ß = -0.100, 95% CRI: -0.191 to -0.016) despite shifts in behavior, mirroring long-term declines in the abundance of C. subspathacea (ß = -0.191, 95% CRI: -0.355 to -0.032). These results have important implications for individual fitness and population viability, where shifts in gosling behavior putatively fail to mitigate long-term declines in forage availability.

Brood Size Affects Future Reproduction in a Long-Lived Bird with Precocial Young.

Estimation of trade-offs between current reproduction and future survival and fecundity of long-lived vertebrates is essential to understanding factors that shape optimal reproductive investment. Black brant geese (Branta bernicla nigricans) fledge more goslings, on average, when their broods are experimentally enlarged to be greater than the most common clutch size of four eggs. Thus, we hypothesized that the lesser frequency of brant clutches exceeding four eggs results, at least partially, from a future reduction in survival, breeding probability, or clutch size for females tending larger broods. We used an 8-year mark-recapture data set (Barker robust design) with 5 years of clutch and brood manipulations to estimate long-term consequences of reproductive decisions in brant. We did not find evidence of a trade-off between reproductive effort and true survival or future initiation date and clutch size. Rather, future breeding probability was maximized ( 0.92±0.03 [SE]) for manipulated females tending broods of four goslings and was lower for females tending smaller (one gosling; 0.63±0.09 [SE]) or larger (seven goslings; 0.52±0.15 [SE]) broods. Our results suggest that demographic trade-offs for female brant tending large broods may reduce the fitness value of clutches larger than four and, therefore, contribute to the paucity of larger clutches. The lack of a trade-off between reproductive effort and survival provides evidence that survival, to which fitness is most sensitive in long-lived animals, is buffered against temporal variation in brant.

Timing of precipitation in an arid environment: Effects on population performance of a large herbivore.

Climate models predict that shifts in temperature and precipitation patterns are likely to occur across the globe. Changing climate will likely have strong effects on arid environments as a result of increased temperatures, increasing frequency and intensity of droughts, and less consistent pulses of rainfall. Therefore, understanding the link between patterns of precipitation, temperature, and population performance of species occupying these environments will continue to increase in importance as climatic shifts occur within these natural ecosystems. We sought to evaluate how individual, maternal, population, and environmental, particularly temperature and precipitation, level factors influence population performance of a large herbivore in an arid environment. We used mule deer (Odocoileus hemionus) as a representative species and quantified juvenile survival to test hypotheses about effects of environmental factors on population performance. Precipitation events occurring in mid- to late-pregnancy (January-April) leading to spring green-up, as indexed by normalized difference in vegetation index, had the strongest positive effect on juvenile survival and recruitment. In addition, larger neonates had an increased probability of survival. Our findings indicate that timing and amount of precipitation prior to parturition have strong influences on maternal nutritional condition, which was passed on to young. These results have important implications for understanding how animal populations may benefit from timing of precipitation during spring and prior to parturition, especially in arid environments.

Weather, habitat composition, and female behavior interact to modify offspring survival in Greater Sage-Grouse.

Weather is a source of environmental variation that can affect population vital rates. However, the influence of weather on individual fitness is spatially heterogeneous and can be driven by other environmental factors, such as habitat composition. Therefore, individuals can experience reduced fitness (e.g., decreased reproductive success) during poor environmental conditions through poor decisions regarding habitat selection. This requires, however, that habitat selection is adaptive and that the organism can correctly interpret the environmental cues to modify habitat use. Greater Sage-Grouse (Centrocercus urophasianus) are an obligate of the sagebrush ecosystems of western North America, relying on sagebrush for food and cover. Greater Sage-Grouse chicks, however, require foods with high nutrient content (i.e., forbs and insects), the abundance of which is both temporally and spatially dynamic and related primarily to water availability. Our goal was to assess whether nest site selection and movements of broods by females reduced the negative effect of drought on offspring survival. As predicted, chick survival was negatively influenced by drought severity. We found that sage-grouse females generally preferred to nest and raise their young in locations where their chicks would experience higher survival. We also found that use of habitats positively associated with chick survival were also positively associated with drought severity, which suggests that females reduced drought impacts on their dependent young by selecting more favorable environments during drought years. Although our findings suggest that female nest site selection and brood movement rates can reduce the negative effects of drought on early offspring survival, the influence of severe drought conditions was not completely mitigated by female behavior, and that drought conditions should be considered a threat to Greater Sage-Grouse population persistence.

Ultimate regulation of fecundity in species with precocial young: declining marginal value of offspring with increasing brood size does not explain maximal clutch size in Black Brent geese.

Lack 18:125-128 (1967) proposed that clutch size in precocial species was regulated by nutrients available to females during breeding. Drent and Daan 68:225-252 (1980) proposed the individual optimization hypothesis, whereby individual state determines the optimal combination of breeding date and clutch size. Neither hypothesis accounts for variation in nutrients among females at the end of egg laying, strong right truncations in clutch size distributions, or the fact that many species with precocial young are determinate layers. One solution is that there is a maximum clutch size, above which the number of fledged young declines. We manipulated brood size in Black Brent geese to decouple brood size from maternal quality and produce broods larger than the natural maximum. We recaptured marked goslings to assess variation in prefledging survival as a function of brood size and we estimated relative prefledging survival of goslings using a Bayesian hierarchical approach. We considered effects of natural clutch size, brood size and their interaction on probability that we captured goslings at about 4 weeks of age. Prefledging survival declined with increasing brood size ([Formula: see text] = -0.53; 95% CI -0.91 to -0.16), while laid clutch size had little influence on prefledging survival ([Formula: see text] = -0.04; 95% CI -0.42 to 0.32). Despite declining per capita survival with increasing brood size, the most productive brood size was six goslings, which is greater than the typical maximum clutch size of five. Thus, reduced survival in large broods, by itself, is not the sole mechanism that limits maximum clutch size. We suggest elsewhere that incubation limitation and lower residual reproductive value for females tending larger broods may be other mechanisms limiting maximal clutch size in brent.

Fine-scale genetic structure among greater sage-grouse leks in central Nevada.

BACKGROUND: Mating systems that reduce dispersal and lead to non-random mating might increase the potential for genetic structure to arise at fine geographic scales. Greater sage-grouse (Centrocercus urophasianus) have a lek-based mating system and exhibit high site fidelity and skewed mating ratios. We quantified population structure by analyzing variation at 27,866 single-nucleotide polymorphisms in 140 males from ten leks (within five lek complexes) occurring in a small geographic region in central Nevada. RESULTS: Lek complexes, and to a lesser extent individual leks, formed statistically identifiable clusters in ordination analyses, providing evidence for fine-scale geographic genetic differentiation. Lek geography predicted genetic differentiation even at a small geographic scale, which could be sharpened by strong site fidelity. Relatedness was also higher among individuals within lek complexes (and leks), suggesting that reproductive skew, where few males participate in most of the successful matings, could also potentially contribute to genetic differentiation. Models incorporating a habitat resistance surface as a proxy for potentially reduced movement due to landscape features indicated that both geographic distance and habitat suitability (i.e. preferred habitat) predicted genetic structure, with no significant effect of man-made barriers to movement (i.e. power lines and roads). Finally, we illustrate how data sets containing fewer loci (<4000) had less statistical precision and failed to detect the full degree of genetic structure. CONCLUSION: Our results suggest that habitat features and lek site geography of sage-grouse shape fine scale genetic structure, and highlight how larger data sets can have increased precision and accuracy for quantifying ecologically relevant genetic structure over small geographic scales.

Evaluating vegetation effects on animal demographics: the role of plant phenology and sampling bias.

Plant phenological processes produce temporal variation in the height and cover of vegetation. Key aspects of animal life cycles, such as reproduction, often coincide with the growing season and therefore may inherently covary with plant growth. When evaluating the influence of vegetation variables on demographic rates, the decision about when to measure vegetation relative to the timing of demographic events is important to avoid confounding between the demographic rate of interest and vegetation covariates. Such confounding could bias estimated effect sizes or produce results that are entirely spurious. We investigated how the timing of vegetation sampling affected the modeled relationship between vegetation structure and nest survival of greater sage-grouse (Centrocercus urophasianus), using both simulated and observational data. We used the height of live grasses surrounding nests as an explanatory covariate, and analyzed its effect on daily nest survival. We compared results between models that included grass height measured at the time of nest fate (hatch or failure) with models where grass height was measured on a standardized date - that of predicted hatch date. Parameters linking grass height to nest survival based on measurements at nest fate produced more competitive models, but slope coefficients of grass height effects were biased high relative to truth in simulated scenarios. In contrast, measurements taken at predicted hatch date accurately predicted the influence of grass height on nest survival. Observational data produced similar results. Our results demonstrate the importance of properly considering confounding between demographic traits and plant phenology. Not doing so can produce results that are plausible, but ultimately inaccurate.

Carryover effects and climatic conditions influence the postfledging survival of greater sage-grouse.

Prebreeding survival is an important life history component that affects both parental fitness and population persistence. In birds, prebreeding can be separated into pre- and postfledging periods; carryover effects from the prefledging period may influence postfledging survival. We investigated effects of body condition at fledging, and climatic variation, on postfledging survival of radio-marked greater sage-grouse (Centrocercus urophasianus) in the Great Basin Desert of the western United States. We hypothesized that body condition would influence postfledging survival as a carryover effect from the prefledging period, and we predicted that climatic variation may mediate this carryover effect or, alternatively, would act directly on survival during the postfledging period. Individual body condition had a strong positive effect on postfledging survival of juvenile females, suggesting carryover effects from the prefledging period. Females in the upper 25th percentile of body condition scores had a postfledging survival probability more than twice that (Φ = 0.51 ± 0.06 SE) of females in the bottom 25th percentile (Φ = 0.21 ± 0.05 SE). A similar effect could not be detected for males. We also found evidence for temperature and precipitation effects on monthly survival rates of both sexes. After controlling for site-level variation, postfledging survival was nearly twice as great following the coolest and wettest growing season (Φ = 0.77 ± 0.05 SE) compared with the hottest and driest growing season (Φ = 0.39 ± 0.05 SE). We found no relationships between individual body condition and temperature or precipitation, suggesting that carryover effects operated independently of background climatic variation. The temperature and precipitation effects we observed likely produced a direct effect on mortality risk during the postfledging period. Conservation actions that focus on improving prefledging habitat for sage-grouse may have indirect benefits to survival during postfledging, due to carryover effects between the two life phases.

Spatial variation in life-history trade-offs results in an ideal free distribution in Black Brant Geese.

Ideal free distribution theory predicts that individuals distribute themselves so fitness is equal among patches. In this paper we evaluate all components of adult fitness to assess the hypothesis that individuals distribute themselves among seven brood-rearing areas so that trade-offs among different life history traits result in equal mean fitness among individuals using different areas. We used estimates of vital rates (clutch size, nest survival, pre-fledging survival, post-fledging survival, juvenile survival, and breeding probability) to estimate brood-rearing-area-specific per capita recruitment rates and survival for adult females. We summed brood-rearing-area-specific per capita recruitment and adult survival to calculate brood-rearing-area-specific estimates of lambda. We found little variation in lambda among brood-rearing areas and lifetime fitness implications of changing brood-rearing area were negligible (< 1% brood-rearing area mean fitness). We conclude that adult female Black Brant distribute themselves in an ideal free manner, resulting in equal fitness among females using these areas.