Climate change and commercial fishing practices codetermine survival of a long-lived seabird.

Understanding the environmental mechanisms that govern population change is a fundamental objective in ecology. Although the determination of how top-down and bottom-up drivers affect demography is important, it is often equally critical to understand the extent to which, environmental conditions that underpin these drivers fluctuate across time. For example, associations between climate and both food availability and predation risk may suggest the presence of trophic interactions that may influence inferences made from patterns in ecological data. Analytical tools have been developed to account for these correlations, while providing opportunities to ask novel questions regarding how populations change across space and time. Here, we combine two modeling disciplines-path analysis and mark-recapture-recovery models-to explore whether shifts in sea-surface temperatures (SSTs) influenced top-down (entanglement in fishing equipment) or bottom-up (forage fish production) population constraints over 60 years, and the extent to which these covarying processes shaped the survival of a long-lived seabird, the Royal tern. We found that hemispheric trends in SST were associated with variation in the amount of fish harvested along the Atlantic coast of North America and in the Caribbean, whereas reductions in forage fish production were mostly driven by shifts in the amount of fish harvested by commercial fisheries throughout the North Atlantic the year prior. Although the indirect (i.e., stock depletion) and direct (i.e., entanglement) impacts of commercial fishing on Royal tern mortality has declined over the last 60 years, increased SSTs during this time period has resulted in a comparable increase in mortality risk, which disproportionately impacted the survival of the youngest age-classes of Royal terns. Given climate projections for the North Atlantic, our results indicate that threats to Royal tern population persistence in the Mid-Atlantic will most likely be driven by failures to recruit juveniles into the breeding population.

A life-history spectrum of population responses to simultaneous change in climate and land use.

Climate and land use change are two of the primary threats to global biodiversity; however, each species within a community may respond differently to these facets of global change. Although it is typically assumed that species use the habitat that is advantageous for survival and reproduction, anthropogenic changes to the environment can create ecological traps, making it critical to assess both habitat selection (e.g. where species congregate on the landscape) and the influence of selected habitats on the demographic processes that govern population dynamics. We used a long-term (1958-2011), large-scale, multi-species dataset for waterfowl that spans the United States and Canada to estimate species-specific responses to climate and land use variables in a landscape that has undergone significant environmental change across space and time. We first estimated the effects of change in climate and land use variables on habitat selection and population dynamics for nine species. We then hypothesized that species-specific responses to environmental change would scale with life-history traits, specifically: longevity, nesting phenology and female breeding site fidelity. We observed species-level heterogeneity in the demographic and habitat selection responses to climate and land use change, which would complicate community-level habitat management. Our work highlights the importance of multi-species monitoring and community-level analysis, even among closely related species. We detected several relationships between life-history traits, particularly nesting phenology, and species' responses to environmental change. One species, the early-nesting northern pintail (Anas acuta), was consistently at the extreme end of responses to land use and climate predictors and has been a species of conservation concern since their population began to decline in the 1980s. They, and the blue-winged teal, also demonstrated a positive habitat selection response to the proportion of cropland on the landscape that simultaneously reduced abundance the following year, indicative of susceptibility to ecological traps. By distilling the diversity of species' responses to environmental change within a community, our methodological approach and findings will help improve predictions of community responses to global change and can inform multi-species management and conservation plans in dynamic landscapes that are based on simple tenets of life-history theory.

Context-dependent variation in persistence of host populations in the face of disease.

Research Highlight: Valenzuela-Sánchez, A., Azat, C., Cunningham, A. A., Delgado, S., Bacigalupe, L. D., Beltrand, J., Serrano, J. M., Sentenac, H., Haddow, N., Toledo, V., Schmidt, B. R., & Cayuela, H. (2022). Interpopulation differences in male reproductive effort drive the population dynamics of a host exposed to an emerging fungal pathogen. Journal of Animal Ecology, 00, 1- 12. https://doi.org/10.1111/1365-2656.13603. Understanding the nuances of population persistence in the face of a stressor can help predict extinction risk and guide conservation actions. However, the exact mechanisms driving population stability may not always be known. In this paper, Valenzuela-Sánchez et al. (2022) integrate long-term mark-recapture data, focal measurements of reproductive effort, a population matrix model and inferences on life-history variation to reveal differences in demographic response to disease in a susceptible frog species (Rhinoderma darwinii). Valenzuela-Sánchez et al. found that demographic compensation via recruitment explained the positive population growth rate in their high disease prevalence population whereas the low disease prevalence population did not compensate and thus had decreasing population growth. Compensatory recruitment was likely due to the high probability of males brooding, and the high number of brooded larvae in the high prevalence population compared to low prevalence and disease-free populations. Valenzuela-Sánchez et al. also document faster generation times in the high prevalence population, which may indicate a faster life history that may be contributing to the population's ability to compensate for reduced survival. Lastly, the authors find a positive relationship between disease prevalence and the proportion of juveniles in a given population that suggest that there may be a threshold for disease prevalence that triggers increased reproductive effort. Altogether, their study provides novel support for increased reproductive effort as the pathway for compensatory recruitment leading to increasing population growth despite strong negative effects of disease on adult survival. Their results also caution the overgeneralization of the effects of stressors (e.g. disease) on population dynamics, where context-dependent responses may differ among host populations of a given species.

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.

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.

Detrimental impacts of climate change may be exacerbated by density-dependent population regulation in blue mussels.

The climate on our planet is changing and the range distributions of organisms are shifting in response. In aquatic environments, species might not be able to redistribute poleward or into deeper water when temperatures rise because of barriers, reduced light availability, altered water chemistry or any combination of these. How species respond to climate change may depend on physiological adaptability, but also on the population dynamics of the species. Density dependence is a ubiquitous force that governs population dynamics and regulates population growth, yet its connections to the impacts of climate change remain little known, especially in marine studies. Reductions in density below an environmental carrying capacity may cause compensatory increases in demographic parameters and population growth rate, hence masking the impacts of climate change on populations. On the other hand, climate-driven deterioration of conditions may reduce environmental carrying capacities, making compensation less likely and populations more susceptible to the effects of stochastic processes. Here we investigate the effects of climate change on Baltic blue mussels using a 17-year dataset on population density. Using a Bayesian modelling framework, we investigate the impacts of climate change, assess the magnitude and effects of density dependence, and project the likelihood of population decline by the year 2030. Our findings show negative impacts of warmer and less saline waters, both outcomes of climate change. We also show that density dependence increases the likelihood of population decline by subjecting the population to the detrimental effects of stochastic processes (i.e. low densities where random bad years can cause local extinction, negating the possibility for random good years to offset bad years). We highlight the importance of understanding, and accounting for both density dependence and climate variation when predicting the impact of climate change on keystone species, such as the Baltic blue mussel.

Changes in climate and land use interact to create an ecological trap in a migratory species.

Anthropogenic landscape alteration and climate change can have multiscale and interrelated effects on ecological systems. Such changes to the environment can disrupt the connection between habitat quality and the cues that species use to identify quality habitat, which can result in an ecological trap. Ecological traps are typically difficult to identify without fine-scale information on individual survival and fitness, but this information is rarely available over large temporal and spatial scales. The Prairie Pothole Region (PPR) of the United States and Canada has undergone extensive changes in the latter half of the 20th century due to advancements in agricultural technologies, water management practices and climate change. Historically, the PPR has been a highly productive area for breeding waterfowl. While the overall trends for dabbling ducks in the PPR have exhibited increasing abundances since the late 1980s, some species, such as the northern pintail, have been declining in abundance. We used a long-term dataset of pintail counts across the PPR to separate count data into a demographic process and a habitat selection process using a hierarchical model. The hierarchical model provided an alternative way of identifying ecological traps in the absence of individual survival and fitness. Our model also allowed us to account for the indirect pathways by which climate and agriculture impact pintail through their additional contribution to wetland availability, which is a primary driver of pintail demography and habitat selection. Decoupling these processes allowed us to identify an ecological trap related to increasing cropland land cover, in which pintail selected for cropland over alternative nesting habitat, likely due to the similarities with productive native mixed-grass prairie. However, large proportions of cropland within a region resulted in fewer pintail the following year, likely due to nest failures from predation and agricultural practices. In addition, we identified several regions in Canada where this ecological trap is contributing significantly to mismatches between habitat selection and demographic processes.

Shifting Vital Rate Correlations Alter Predicted Population Responses to Increasingly Variable Environments.

Time series of vital rates are often used to construct "environment-blind" stochastic population projections and calculate the elasticity of population growth to increased temporal variance in vital rates. Here, we show that the utility of this widely used demographic tool is greatly limited by shifts in vital rate correlations that occur as environmental drivers become increasingly variable. The direction and magnitude of these shifts are unpredictable without environmentally explicit models. Shifting vital rate correlations had the largest fitness effects on life histories with short to medium generation times, potentially hampering comparative analyses based on elasticities to vital rate variance for a wide range of species. Shifts in vital rate correlations are likely ubiquitous in increasingly variable environments, and further research should empirically evaluate the life histories for which detailed mechanistic relationships between vital rates and environmental drivers are required for making reliable predictions versus those for which summarized demographic data are sufficient.

Reproductive success of a keystone herbivore is more variable and responsive to climate in habitats with lower resource diversity.

The effects of climate on wild populations are often channelled through species interactions. Population responses to climate variation can therefore differ across habitats, owing to variation in the biotic community. Theory predicts that consumer demography should be less variable and less responsive to climate in habitats with greater resource diversity. We tested these predictions using a long-term study of breeding lesser snow geese along the western coast of Hudson Bay, Manitoba, Canada. Reproductive success was measured in 22 years from 114 locations, in either coastal or inland habitat types. We used Bayesian analysis to estimate the response of reproductive success to climate in each habitat type, along with residual variation not explained by climate. We then quantified gosling diet composition in each habitat type to test the prediction that reproductive success would be less variable and more responsive to climate in habitats with lower resource diversity. Reproductive success responded positively to seasonal warmness, but this response was much stronger in inland habitats than in coastal habitats. Site- and year-level random effects were also three to five times more variable in inland habitats. Simultaneously, land cover diversity and gosling diet diversity were lower in inland habitats. Our study illustrates that spatial variation in resource diversity (and thus, species interactions) can have important effects on consumer responses to climate. In this system, climate change is expected to disproportionately increase the reproductive success of snow geese in vast inland habitats, potentially counteracting management efforts to reduce the abundance of this keystone herbivore.

Understanding the demographic drivers of realized population growth rates.

Identifying the demographic parameters (e.g., reproduction, survival, dispersal) that most influence population dynamics can increase conservation effectiveness and enhance ecological understanding. Life table response experiments (LTRE) aim to decompose the effects of change in parameters on past demographic outcomes (e.g., population growth rates). But the vast majority of LTREs and other retrospective population analyses have focused on decomposing asymptotic population growth rates, which do not account for the dynamic interplay between population structure and vital rates that shape realized population growth rates (λt=Nt+1/Nt) in time-varying environments. We provide an empirical means to overcome these shortcomings by merging recently developed "transient life-table response experiments" with integrated population models (IPMs). IPMs allow for the estimation of latent population structure and other demographic parameters that are required for transient LTRE analysis, and Bayesian versions additionally allow for complete error propagation from the estimation of demographic parameters to derivations of realized population growth rates and perturbation analyses of growth rates. By integrating available monitoring data for Lesser Scaup over 60 yr, and conducting transient LTREs on IPM estimates, we found that the contribution of juvenile female survival to long-term variation in realized population growth rates was 1.6 and 3.7 times larger than that of adult female survival and fecundity, respectively. But a persistent long-term decline in fecundity explained 92% of the decline in abundance between 1983 and 2006. In contrast, an improvement in adult female survival drove the modest recovery in Lesser Scaup abundance since 2006, indicating that the most important demographic drivers of Lesser Scaup population dynamics are temporally dynamic. In addition to resolving uncertainty about Lesser Scaup population dynamics, the merger of IPMs with transient LTREs will strengthen our understanding of demography for many species as we aim to conserve biodiversity during an era of non-stationary global change.

A life-history perspective on the demographic drivers of structured population dynamics in changing environments.

Current understanding of life-history evolution and how demographic parameters contribute to population dynamics across species is largely based on assumptions of either constant environments or stationary environmental variation. Meanwhile, species are faced with non-stationary environmental conditions (changing mean, variance, or both) created by climate and landscape change. To close the gap between contemporary reality and demographic theory, we develop a set of transient life table response experiments (LTREs) for decomposing realised population growth rates into contributions from specific vital rates and components of population structure. Using transient LTREs in a theoretical framework, we reveal that established concepts in population biology will require revision because of reliance on approaches that do not address the influence of unstable population structure on population growth and mean fitness. Going forward, transient LTREs will enhance understanding of demography and improve the explanatory power of models used to understand ecological and evolutionary dynamics.

Disentangling the effects of climate, density dependence, and harvest on an iconic large herbivore's population dynamics.

Understanding the relative effects of climate, harvest, and density dependence on population dynamics is critical for guiding sound population management, especially for ungulates in arid and semiarid environments experiencing climate change. To address these issues for bison in southern Utah, USA, we applied a Bayesian state-space model to a 72-yr time series of abundance counts. While accounting for known harvest (as well as live removal) from the population, we found that the bison population in southern Utah exhibited a strong potential to grow from low density (ß0 = 0.26; Bayesian credible interval based on 95% of the highest posterior density [BCI] = 0.19-0.33), and weak but statistically significant density dependence (ß1 = -0.02, BCI = -0.04 to -0.004). Early spring temperatures also had strong positive effects on population growth (Pfat1 = 0.09, BCI = 0.04-0.14), much more so than precipitation and other temperature-related variables (model weight > three times more than that for other climate variables). Although we hypothesized that harvest is the primary driving force of bison population dynamics in southern Utah, our elasticity analysis indicated that changes in early spring temperature could have a greater relative effect on equilibrium abundance than either harvest or. the strength of density dependence. Our findings highlight the utility of incorporating elasticity analyses into state-space population models, and the need to include climatic processes in wildlife management policies and planning.

Combined effects of climate, predation, and density dependence on Greater and Lesser Scaup population dynamics.

An understanding of species relationships is critical in the management and conservation of populations facing climate change, yet few studies address how climate alters species interactions and other population drivers. We use a long-term, broad-scale data set of relative abundance to examine the influence of climate, predators, and density dependence on the population dynamics of declining scaup (Aythya) species within the core of their breeding range. The state-space modeling approach we use applies to a wide range of wildlife species, especially populations monitored over broad spatiotemporal extents. Using this approach, we found that immediate snow cover extent in the preceding winter and spring had the strongest effects, with increases in mean snow cover extent having a positive effect on the local surveyed abundance of scaup. The direct effects of mesopredator abundance on scaup population dynamics were weaker, but the results still indicated a potentil interactive process between climate and food web dynamics (mesopredators, alternative prey, and scaup). By considering climate variables and other potential effects on population dynamics, and using a rigorous estimation framework, we provide insight into complex ecological processes for guiding. conservation and policy actions aimed at mitigating and reversing the decline of scaup.

Effects of exploitation on an overabundant species: the lesser snow goose predicament.

Invasive and overabundant species are an increasing threat to biodiversity and ecosystem functioning world-wide. As such, large amounts of money are spent each year on attempts to control them. These efforts can, however, be thwarted if exploitation is compensated demographically or if populations simply become too numerous for management to elicit an effective and rapid functional response. We examined the influence of these mechanisms on cause-specific mortality in lesser snow geese using multistate capture-reencounter methods. The abundance and destructive foraging behaviours of snow geese have created a trophic cascade that reduces (sub-) Arctic plant, insect and avian biodiversity, bestowing them the status of 'overabundant'. Historically, juvenile snow geese suffered from density-related degradation of their saltmarsh brood-rearing habitat. This allowed harvest mortality to be partially compensated by non-harvest mortality (process correlation between mortality sources: ρ = -0.47; 90% BCI: -0.72 to -0.04). Snow goose family groups eventually responded to their own degradation of habitat by dispersing to non-degraded areas. This relaxed the pressure of density dependence on juvenile birds, but without this mechanism for compensation, harvest began to have an additive effect on overall mortality (ρ = 0.60; 90% BCI: -0.06 to 0.81). In adults, harvest had an additive effect on overall mortality throughout the 42-year study (ρ = 0.24; 90% BCI: -0.59 to 0.67). With the aim of controlling overabundant snow geese, the Conservation Order amendment to the International Migratory Bird Treaty was implemented in February of 1999 to allow for harvest regulations that had not been allowed since the early 1900s (e.g. a spring harvest season, high or unlimited bag limits and use of electronic calls and unplugged shotguns). Although harvest mortality momentarily increased following these actions, the increasing abundance of snow geese has since induced a state of satiation in harvest that has driven harvest rates below the long-term average. More aggressive actions will thus be needed to halt the growth and spread of the devastating trophic cascade that snow geese have triggered. Our approach to investigating the impacts of population control efforts on cause-specific mortality will help guide more effective management of invasive and overabundant species world-wide.

Riding the wetland wave: Can ducks locate macroinvertebrate resources across the breeding season?

Food availability varies considerably over space and time in wetland systems, and consumers must be able to track those changes during energetically-demanding points in the life cycle like breeding. Resource tracking has been studied frequently among herbivores, but receives less attention among consumers of macroinvertebrates. We evaluated the change in resource availability across habitat types and time and the simultaneous density of waterfowl consumers throughout their breeding season in a high-elevation, flood-irrigated system. We also assessed whether the macroinvertebrate resource density better predicted waterfowl density across habitats, compared to consistency (i.e., temporal evenness) of the invertebrate resource or taxonomic richness. Resource density varied marginally across wetland types but was highest in basin wetlands (i.e., ponds) and peaked early in the breeding season, whereas it remained relatively low and stable in other wetland habitats. Breeding duck density was positively related to resource density, more so than temporal resource stability, for all species. Resource density was negatively related to duckling density, however. These results have the potential to not only elucidate mechanisms of habitat selection among breeding ducks in flood-irrigated landscapes but also suggest there is not a consequential trade-off to selecting wetland sites based on energy density versus temporal resource stability and that good-quality wetland sites provide both.

Niche opportunities and invasion dynamics in a desert annual community.

Although many factors influence the ability of exotics to invade successfully, most studies focus on only a few variables to explain invasion; attempts at theoretical synthesis are largely untested. The niche opportunities framework proposes that the demographic success of an invader is largely affected by the availability of resources and the abundance of its enemies. Here, we use a 31-year study from a desert ecosystem to examine the niche opportunities framework via the invasion of the annual plant Erodium cicutarium. While the invader remained rare for two decades, a decline in granivory combined with an ideal climate window created an opportunity for E. cicutarium to escape control and become the dominant annual plant in the community. We show that fluctuations in consumption and resources can create niche opportunities for invaders and highlight the need for additional long-term studies to track the influence of changing climate and community dynamics on invasions.

Predators, alternative prey and climate influence annual breeding success of a long-lived sea duck.

1. Perturbations to ecosystems have the potential to directly and indirectly affect species interactions, with subsequent impacts on population dynamics and the vital rates that regulate them. 2. The few long-term studies of common eider breeding ecology indicate that reproductive success is low in most years, interrupted by occasional boom years. However, no study has explicitly examined the drivers of long-term variation in reproductive success. 3. Here, we use encounter history data collected across 41 years to examine the effects of arctic foxes (a terrestrial nest predator), local abundance and spatial distribution of lesser snow geese (an alternative prey source), and spring climate on common eider nest success. 4. Eider nest success declined over the course of the study, but was also highly variable across years. Our results supported the hypothesis that the long-term decline in eider nest success was caused by apparent competition with lesser snow geese, mediated by shared predators. This effect persisted even following a large-scale exodus of nesting geese from the eider colony. Nest success was also lowest in years of low arctic fox index, presumably driven by prey switching in years of low small mammal availability. However, increased snow goose abundance appeared to buffer this effect through prey swamping. The effect of spring climate depended on the stage of the breeding season; cold and wet and warm and dry conditions in early spring were correlated with decreased nest success, whereas warm and wet conditions in late spring increased eider nest success. 5. These results underscore the significance of both trophic interactions and climate in regulating highly variable vital rates, which likely have important consequences for population dynamics and the conservation of long-lived iteroparous species.

Intra-guild interactions and projected impact of climate and land use changes on North American pochard ducks.

The co-occurrence of functionally similar species is very common in nature, and is often put forward as a basis for ecosystem resilience to disturbance. At the same time, competition between similar species is also considered a strong driver of community composition. However, environmental stochasticity can alter this prediction, either because competitive abilities depend on time-varying factors or because covariance in species' responses to environmental conditions masks the effect of competition. Interactions other than competition can also influence community dynamics but have received less attention. We used a simplified community of two sympatric duck species (redhead Aythya americana and canvasback A. valisineria) and a previously published analysis of 50 years of demographic data to parameterize a stochastic, density-dependent, stage-structured model. These ducks interact via nest parasitism (mostly of canvasback by redhead) in addition to competition for food resources, with consequences at the demographic level; these interactions are modulated by habitat availability (number of ponds in the study landscape). We found that if habitat availability decreased there was a high risk of quasi-extinction, and redheads, although initially able to maintain their numerical dominance, quickly became the least abundant species because they perform worse during droughts. If habitat availability increased, we found that the initially more rare canvasback would increase in relative abundance, albeit slowly. We interpret this as a shift from a community influenced by nest parasitism (which is detrimental to canvasback) to a community mostly driven by species-specific dynamics due to relaxation of resource limitation.

Integrated modeling of communities: parasitism, competition, and demographic synchrony in sympatric ducks.

Functionally similar species often co-occur within an ecosystem, and they can compete for or facilitate each other's access to resources. The coupled dynamics of such species play an important role in shaping biodiversity and an ecosystem's resilience to perturbations. Here we study two congeneric North American ducks: Redhead Aythya americana and Canvasback A. vaselineria. Both are largely sympatric during the breeding season, and in addition to competition, facultative parasitic egg-laying can lead to interspecific density dependence. Using multi-population integrated models, we combined capture-recovery data, population surveys, and age ratio data in order to simultaneously estimate the mechanistic drivers of fecundity, survival, and population dynamics for both species. Canvasback numbers positively affected Redhead fecundity, whereas Redhead numbers negatively affected Canvasback fecundity, as expected due to parasitism. This interaction was modulated by wetland habitat availability in a way that matched the observation that Redhead hens parasitize Canvasback nests under all conditions but exhibit typical nesting behavior more frequently during years with numerous ponds. Once these effects of density and habitat were statistically controlled for, we found high levels of interspecific synchrony in both fecundity and survival (respectively, 75% and 49% of remaining variation). Thus, both neutral and non-neutral mechanisms affected the dynamics of these functionally similar species. In this and other systems, our method can be used to test hypotheses about species coexistence and to gain insights into the demographic drivers of community dynamics.

Demographic response to perturbations: the role of compensatory density dependence in a North American duck under variable harvest regulations and changing habitat.

1. Most wild animal populations are subjected to many perturbations, including environmental forcing and anthropogenic mortality. How population size varies in response to these perturbations largely depends on life-history strategy and density regulation. 2. Using the mid-continent population of redhead Aythya americana (a North American diving duck), we investigated the population response to two major perturbations, changes in breeding habitat availability (number of ponds in the study landscape) and changes in harvest regulations directed at managing mortality patterns (bag limit). We used three types of data collected at the continental scale (capture-recovery, population surveys and age- and sex ratios in the harvest) and combined them into integrated population models to assess the interaction between density dependence and the effect of perturbations. 3. We observed a two-way interaction between the effects on fecundity of pond number and population density. Hatch-year female survival was also density dependent. Matrix modelling showed that population booms could occur after especially wet years. However, the effect of moderate variation in pond number was generally offset by density dependence the following year. 4. Mortality patterns were insensitive to changes in harvest regulations and, in males at least, insensitive to density dependence as well. We discuss potential mechanisms for compensation of hunting mortality as well as possible confounding factors. 5. Our results illustrate the interplay of density dependence and environmental variation both shaping population dynamics in a harvested species, which could be generalized to help guide the dual management of habitat and harvest regulations.