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.

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.

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.

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.

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.

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.