Culling Versus Density Effects in Management of a Deer Population

Abstract: Wildlife managers often manipulate hunting regulations to control deer populations. However, few empirical studies have examined the level of hunting effort (hunter-days) required to limit population growth and demographic effects through harvesting of females. Moreover, the relative importance of density effects on population growth has not been quantified. We reconstructed a sika deer [Cervus nippon] population over a period of 12 years (1990–2001) using age- and sex-specific harvest data. Using cohort analysis, we analyzed population dynamics, focusing on 1) the relationship between hunting effort and hunting-induced mortality rate, 2) relative contributions of hunting mortality and recruitment of yearlings to annual changes in population growth rate, and 3) annual variation in recruitment rate. Population size increased until 1998 and declined thereafter. The population growth rate changed more in response to annual changes in recruitment rate than hunting mortality rate. Temporal variation in recruitment rate was not controlled by birth rate alone; direct density dependence, intensities of hunting mortality for fawns, and for females (≥2 yr of age), which accounted for the fawn survival rate, were required as factors to explain temporal variation. Density effects on the recruitment rate were not strong enough to regulate the population within the study period; high hunting mortality, with intensive female harvesting, was necessary to prevent population growth. Hunting effort was a good predictor of the hunting mortality rate, and female harvest had a negative effect on the recruitment rate through fawn survival. We suggest that >3,500 hunter-days and prioritization of female harvesting are required to prevent increases in this deer population.     

Building the Foundation for International Conservation Planning for Breeding Ducks across the U.S. and Canadian Border

We used publically available data on duck breeding distribution and recently compiled geospatial data on upland habitat and environmental conditions to develop a spatially explicit model of breeding duck populations across the entire Prairie Pothole Region (PPR). Our spatial population models were able to identify key areas for duck conservation across the PPR and predict between 62.1 – 79.1% (68.4% avg.) of the variation in duck counts by year from 2002 – 2010. The median difference in observed vs. predicted duck counts at a transect segment level was 4.6 ducks. Our models are the first seamless spatially explicit models of waterfowl abundance across the entire PPR and represent an initial step toward joint conservation planning between Prairie Pothole and Prairie Habitat Joint Ventures. Our work demonstrates that when spatial and temporal variation for highly mobile birds is incorporated into conservation planning it will likely increase the habitat area required to support defined population goals. A major goal of the current North American Waterfowl Management Plan and subsequent action plan is the linking of harvest and habitat management. We contend incorporation of spatial aspects will increase the likelihood of coherent joint harvest and habitat management decisions. Our results show at a minimum, it is possible to produce spatially explicit waterfowl abundance models that when summed across survey strata will produce similar strata level population estimates as the design-based Waterfowl Breeding Pair and Habitat Survey (r² = 0.977). This is important because these design-based population estimates are currently used to set duck harvest regulations and to set duck population and habitat goals for the North American Waterfowl Management Plan. We hope this effort generates discussion on the important linkages between spatial and temporal variation in population size, and distribution relative to habitat quantity and quality when linking habitat and population goals across this important region.     

Density dependence across multiple life stages in a temperate old-growth forest of northeast China

Recent studies on species coexistence suggest that density dependence is an important mechanism regulating plant populations. However, there have been few studies of density dependence conducted for more than one life-history stage or that control for habitat heterogeneity, which may influence spatial patterns of survival and mask density dependence. We explored the prevalence of density dependence across multiple life stages, and the effects of controlling for habitat heterogeneity, in a temperate forest in northeast China. We used generalized linear mixed-effects models to test for density-dependent mortality of seedlings and spatial point pattern analysis to detect density dependence for sapling-to-juvenile transitions. Conspecific neighbors had a negative effect on survival of plants in both life stages. At the seedling stage, we found a negative effect of conspecific seedling neighbors on survival when analyzing all species combined. However, in species-level analyses, only 2 of 11 focal species were negatively impacted by conspecific neighbors, indicating wide variation among species in the strength of density dependence. Controlling for habitat heterogeneity did not alter our findings of density dependence at the seedling stage. For the sapling-to-juvenile transition stage, 11 of 15 focal species showed patterns of local scale (<10 m) conspecific thinning, consistent with negative density dependence. The results varied depending on whether we controlled for habitat heterogeneity, indicating that a failure to account for habitat heterogeneity can obscure patterns of density dependence. We conclude that density dependence may promote tree species coexistence by acting across multiple life-history stages in this temperate forest.     

Quantifying density dependence in a bird population using human disturbance

Although density dependence has long been recognised as vital to population regulation, there have been relatively few studies demonstrating it spatially in wildlife populations, often due to the confounding effects of variation in habitat quality. We report on a study of woodlarks Lullula arborea, a species of European conservation concern, breeding on lowland heath in Dorset, England. We take the novel approach of utilising the birds’ response to human disturbance, which resulted in much of the variation in density but had no direct impact on demographic rates. Within years, in sites with greater density there were smaller mean chick masses, lower post-fledging survival, and higher rates of nestling mortality attributed to starvation. The effects on clutch size and fledging success were confounded by the area of grassland within a site. There was no effect on brood size. Density dependence also operated within sites between years: as density increased there were reductions in mean chick mass and post-fledging survival, while nestling mortality attributed to starvation increased. Density-dependent effects on clutch size were only weakly regulatory, whereas density-dependent starvation and post-fledging mortality rates contributed strongly to differences in overall breeding output. Heavier chicks (when 7 days old) were significantly more likely to fledge and less likely to starve. Broods with heavier chicks were more likely to supply recruits to the breeding population. Nestling mass was not a factor in survival in the immediate post-fledging period, suggesting that density-dependent processes act independently on this stage. We conclude that the number of birds per hectare of suitable habitat is a valid means of expressing density, and that habitat acts as a surrogate for food abundance through which density dependence operates on the woodlark population.     

Spatially varying density dependence drives a shifting mosaic of survival in a recovering apex predator (Canis lupus)

Understanding landscape patterns in mortality risk is crucial for promoting recovery of threatened and endangered species. Humans affect mortality risk in large carnivores such as wolves (Canis lupus), but spatiotemporally varying density dependence can significantly influence the landscape of survival. This potentially occurs when density varies spatially and risk is unevenly distributed. We quantified spatiotemporal sources of variation in survival rates of gray wolves (C. lupus) during a 21‐year period of population recovery in the Upper Peninsula of Michigan, USA. We focused on mapping risk across time using Cox Proportional Hazards (CPH) models with time‐dependent covariates, thus exploring a shifting mosaic of survival. Extended CPH models and time‐dependent covariates revealed influences of seasonality, density dependence and experience, as well as individual‐level factors and landscape predictors of risk. We used results to predict the shifting landscape of risk at the beginning, middle, and end of the wolf recovery time series. Survival rates varied spatially and declined over time. Long‐term change was density‐dependent, with landscape predictors such as agricultural land cover and edge densities contributing negatively to survival. Survival also varied seasonally and depended on individual experience, sex, and resident versus transient status. The shifting landscape of survival suggested that increasing density contributed to greater potential for human conflict and wolf mortality risk. Long‐term spatial variation in key population vital rates is largely unquantified in many threatened, endangered, and recovering species. Variation in risk may indicate potential for source‐sink population dynamics, especially where individuals preemptively occupy suitable territories, which forces new individuals into riskier habitat types as density increases. We encourage managers to explore relationships between adult survival and localized changes in population density. Density‐dependent risk maps can identify increasing conflict areas or potential habitat sinks which may persist due to high recruitment in adjacent habitats.     

Environmental drivers of habitat use by common bottlenose dolphins (Tursiops truncatus) in the Indian River Lagoon,Florida, USA

Anthropogenic impacts in estuarine systems can influence marine mammal habitat use, population dynamics, fitness, and mortality events. The objective was to examine habitat use among the resident common bottlenose dolphin (Tursiops truncatus) population inhabiting the Indian River Lagoon, Florida, and the influences of variation in environmental factors and prey availability in 2003–2015. We utilized photo-identification surveys, stratified random samples of prey, and environmental variables collected monthly. Kernel density estimation was used to determine the magnitude-per-unit area of dolphins across the IRL by wet and dry seasons each year, the values were used as a response variable in classification and regression tree analyses with fish community and environmental variables as predictors. Spatial patterns in dolphin density in the IRL were associated with salinity and dissolved oxygen levels, which are in part associated with freshwater discharges of nutrient and algae laden waters from the region's storm water management system. These findings isolate locations of concern for management of dolphin habitat, and anthropogenic drivers of dolphin distributions requiring further research.     

Density-dependent processes in the life history of fishes: evidence from laboratory populations of zebrafish Danio rerio

Population regulation is fundamental to the long-term persistence of populations and their responses to harvesting, habitat modification, and exposure to toxic chemicals. In fish and other organisms with complex life histories, regulation may involve density dependence in different life-stages and vital rates. We studied density dependence in body growth and mortality through the life-cycle of laboratory populations of zebrafish Danio rerio. When feed input was held constant at population-level (leading to resource limitation), body growth was strongly density-dependent in the late juvenile and adult phases of the life-cycle. Density dependence in mortality was strong during the early juvenile phase but declined thereafter and virtually ceased prior to maturation. Provision of feed in proportion to individual requirements (easing resource limitation) removed density dependence in growth and substantially reduced density dependence in mortality, thus indicating that 'bottom-up' effects act on growth as well as mortality, but most strongly on growth. Both growth and mortality played an important role in population regulation, with density-dependent growth having the greater impact on population biomass while mortality had the greatest impact on numbers. We demonstrate a clear ontogenic pattern of change in density-dependent processes within populations of a very small (maximum length 5 mm) fish, maintained in constant homogeneous laboratory conditions. The patterns are consistent with those distilled from studies on wild fish populations, indicating the presence of broad ontogenic patterns in density-dependent processes that are invariant to maximum body size and hold in homogeneous laboratory, as well as complex natural environments.     

Density-dependent habitat selection alters drivers of population distribution in northern Yellowstone elk

Although it is well established that density dependence drives changes in organismal abundance over time, relatively little is known about how density dependence affects variation in abundance over space. We tested the hypothesis that spatial trade-offs between food and safety can change the drivers of population distribution, caused by opposing patterns of density-dependent habitat selection (DDHS) that are predicted by the multidimensional ideal free distribution. We addressed this using winter aerial survey data of northern Yellowstone elk (Cervus canadensis) spanning four decades. Supporting our hypothesis, we found positive DDHS for food (herbaceous biomass) and negative DDHS for safety (openness and roughness), such that the primary driver of habitat selection switched from food to safety as elk density decreased from 9.3 to 2.0 elk/km². Our results demonstrate how population density can drive landscape-level shifts in population distribution, confounding habitat selection inference and prediction and potentially affecting community-level interactions.     

Patterns and causes of demographic variation in a harvested moose population: evidence for the effects of climate and density-dependent drivers

1.?Better understanding of the mechanisms affecting demographic variation in ungulate populations is needed to support sustainable management of harvested populations. While studies of moose Alces alces L. populations have previously explored temporal variation in demographic processes, managers responsible for populations that span large heterogeneous landscapes would benefit from an understanding of how demography varies across biogeographical gradients in climate and other population drivers. Evidence of thresholds in population response to manageable and un-manageable drivers could aid resource managers in identifying limits to the magnitude of sustainable change. 2.?Generalized additive models (GAMs) were used to evaluate the relative importance of population density, habitat abundance, summer and winter climatic conditions, primary production, and harvest intensity in explaining spatial variation in moose vital rates in Ontario, Canada. Tree regression was used to test for thresholds in the magnitudes of environmental predictor variables that significantly affected population vital rates. 3.?Moose population growth rate was negatively related to moose density and positively related to the abundance of mixed deciduous habitat abundant in forage. Calf recruitment was negatively related to a later start of the growing season and calf harvest. The ratio of bulls to cows was related to male harvest and hunter access, and thresholds were evident in predictor variables for all vital rate models. 4.?Findings indicate that the contributions of density-dependent and independent factors can vary depending on the scale of population process. The importance of density dependence and habitat supply to low-density ungulate populations was evident, and management strategies for ungulates may be improved by explicitly linking forest management and harvest. Findings emphasize the importance of considering summer climatic influences to ungulate populations, as recruitment in moose was more sensitive to the timing of vegetation green-up than winter severity. The efficacy of management decisions for harvested ungulates may require regional shifts in targets where populations span bioclimatic gradients. The use of GAMs in combination with recursive partitioning was demonstrated to be an informative analytical framework that captured nonlinear relationships common in natural processes and thresholds that are relevant to population management in diverse systems.     

Correlates of Mortality in an Exploited Wolf Population

Abstract We investigated the influence of habitat use on risk of death from hunting and trapping of 55 radiocollared gray wolves (Canis lupus) from an exploited insular population in Southeast Alaska, USA. We compared mortality rates for resident and nonresident wolves and used Cox proportional hazards regression to relate habitat composition within 100-m circular buffers around radiolocations to risk of death of resident and nonresident wolves. In addition, we included covariates representing distances to roads, logged stands, and lakes and streams in those analyses. We also compiled harvest data from 31 harvest units within the study area to compare densities of roads and distances from human settlements with rates of harvest. During our study 39 wolves died, of which 18 were harvested legally, 16 were killed illegally, and 5 died from natural causes. Legal and illegal harvest accounted for >87% of the mortality of radiocollared resident and nonresident wolves. Mean annual survival was 0.54 (SE = 0.17) for all wolves. Annual survival was 0.65 (SE = 0.17) for resident wolves and 0.34 (SE = 0.17) for nonresidents. Very few (19%) nonresident wolves survived to colonize vacant territories or join existing wolf packs. Roads, muskegs, and distances from lakes and streams were covariates positively associated with death of resident wolves. Clear-cuts were positively associated with risk of death of nonresident wolves. Rate of harvest increased with density of roads; however, road densities >0.9 km/km² had little additional effect on harvest rates. Harvest rates decreased with ocean distances from nearest towns or settlements. Roads clearly increased risk of death for wolves from hunting and trapping and contributed to unsustainable rates of harvest. Wildlife managers should consider effects of roads and other habitat features on harvest of wolves when developing harvest recommendations. They should expect substantial illegal harvest where wolf habitat is accessible to humans. Moreover, high rates of mortality of nonresident wolves exposed to legal and illegal harvest may reduce or delay successful dispersal, potentially affecting linkages between small disjunct wolf populations or population segments. We conclude that a combination of conservative harvest regulations and large roadless reserves likely are the most effective measures for conserving wolves where risks from human-caused mortality are high.     

Density‐dependent mortality in Pacific salmon: the ghost of impacts past?

Conservation biologists often ignore density dependence because at‐risk populations are typically small relative to historical levels. However, if populations are reduced as a result of impacts that lower carrying capacity, then density‐dependent mortality may exist at low population abundances. Here, we explore this issue in threatened populations of juvenile chinook salmon (Oncorhynchus tshawytscha). We followed the fate of more than 50 000 juvenile chinook in the Snake River Basin, USA to test the hypothesis that their survival was inversely associated with juvenile density. We also tested the hypotheses that non‐indigenous brook trout and habitat quality affect the presence or strength of density dependence. Our results indicate that juvenile chinook suffer density‐dependent mortality and the strength of density dependence was greater in streams in which brook trout were absent. We were unable to detect an effect of habitat quality on the strength of density dependence. Historical impacts of humans have greatly reduced population sizes of salmon, and the density dependence we report may stem from a shortage of nutrients normally derived from decomposing salmon carcasses. Cohorts of juvenile salmon may experience density‐dependent mortality at population sizes far below historical levels and recovery of imperiled populations may be much slower than currently expected.     

Experimental harvest regulations reveal that water availability during spring,not harvest,affects change in a waterfowl population

Population change is regulated by vital rates that are influenced by environmental conditions, demographic stochasticity, and, increasingly, anthropogenic effects. Habitat destruction and climate change threaten the future of many wildlife populations, and there are additional concerns regarding the effects of harvest rates on demographic components of harvested organisms. Further, many population managers strictly manage harvest of wild organisms to mediate population trends of these populations. The goal of our study was to decouple harvest and environmental variability in a closely monitored population of wild ducks in North America, where we experimentally regulated harvest independently of environmental variation over a period of 4 years. We used 9 years of capture–mark–recapture data to estimate breeding population size during the spring for a population of wood ducks in Nevada. We then assessed the effect of one environmental variable and harvest pressure on annual changes in the breeding population size. Climatic conditions influencing water availability were strongly positively related to population growth rates of wood ducks in our study system. In contrast, harvest regulations and harvest rates did not affect population growth rates. We suggest efforts to conserve waterfowl should focus on the effects of habitat loss in breeding areas and climate change, which will likely affect precipitation regimes in the future. We demonstrate the utility of capture–mark–recapture methods to estimate abundance of species which are difficult to survey and test the impacts of anthropogenic harvest and climate on populations. Finally, our results continue to add to the importance of experimentation in applied conservation biology, where we believe that continued experiments on nonthreatened species will be critically important as researchers attempt to understand how to quantify and mitigate direct anthropogenic impacts in a changing world.     

Reproductive strategies and body shape in the European pond turtle (Emys orbicularis) from contrasting habitats in Italy

Comparisons within and among populations offer important insights into variation in life-history traits and possible adaptive patterns to environmental conditions. We present the results of observed differences in body size, body shape and patterns of reproduction in four separate populations of the European pond turtle Emys orbicularis in central and southern Italy – coastal ( n =3) and mountainous ( n =1) sites and pond ( n =2) and canal ( n= 2) habitats – to determine whether phenotypic plasticity affects reproductive output. Although we did not find any significant latitudinal variation in body size, we observed significant differences in body shape between canal (rounded body shape) and pond (elongated body shape) systems and smaller size with rounded shape in the mountainous population. Reproductive output is similar among populations (median=5 eggs per clutch), whereas reproductive investment (relative clutch mass to maternal body mass) is higher in the mountain population (one clutch per year) than in coastal populations (two clutches per year), suggesting differential trade-offs between geographic locality, elevation and habitat type. Turtle shell shape and geographic location together affect reproductive output in E. orbicularis in Italy.     

Resource availability, breeding site selection, and reproductive success of red-winged blackbirds

Red-winged blackbirds are polygynous and show strong breeding site preferences, but it is unclear which environmental factors regulate their reproductive success and are ultimately responsible for shaping their patterns of habitat selection and their mating system. We evaluated the effect of variation in insect emergence rates on the reproductive success of male and female redwings nesting on replicate ponds. The number of male and female redwings that settled on a pond varied two- to three-fold among ponds, but was not related to insect emergence rates. Insect emergence rates had a positive effect on the number of nestlings successfully fledged by females, the number of nestlings fledged from male territories, and on the mass of nestlings at fledging. Typha stem density also varied widely among ponds, and was positively related to male and female settling density and mass of nestlings at fledging, but not to the number of nestlings fledged by females or males. We conclude that alternative breeding sites differ in their ability to support redwing reproduction, and that the availability of emerging odonates is an important environmental factor influencing the reproductive success of both male and female red-winged blackbirds. Received: 31 March 1997 / Accepted: 3 July 1997     

Density-dependent effects of mortality on the optimal body size to shift habitat: Why smaller is better despite increased mortality risk

Many animal species across different taxa change their habitat during their development. An ontogenetic habitat shift enables the development of early vulnerable-to-predation stages in a safe “nursery” habitat with reduced predation mortality, whereas less vulnerable stages can exploit a more risky, rich feeding habitat. Therefore, the timing of the habitat shift is crucial for individual fitness. We investigate the effect that size selectivity in mortality in the rich feeding habitat has on the optimal body size at which to shift between habitats using a population model that incorporates density dependence. We show that when mortality risk is more size dependent, it is optimal to switch to the risky habitat at a smaller rather than larger body size, despite that individuals can avoid mortality by staying longer in the nursery habitat and growing to safety in size. When size selectivity in mortality is high, large reproducing individuals are abundant and produce numerous offspring that strongly compete in the nursery habitat. A smaller body size at habitat shift is therefore favored because strong competition reduces growth potential. Our results reveal the interdependence among population structure, density dependence, and life history traits, and highlight the need for integrating ecological feedbacks in the study of life history evolution.     

Evolution of density- and patch-size-dependent dispersal rates

Based on a marginal value approach, we derive a nonlinear expression for evolutionarily stable (ES) dispersal rates in a metapopulation with global dispersal. For the general case of density-dependent population growth, our analysis shows that individual dispersal rates should decrease with patch capacity and-beyond a certain threshold-increase with population density. We performed a number of spatially explicit, individual-based simulation experiments to test these predictions and to explore further the relevance of variation in the rate of population increase, density dependence, environmental fluctuations and dispersal mortality on the evolution of dispersal rates. They confirm the predictions of our analytical approach. In addition, they show that dispersal rates in metapopulations mostly depend on dispersal mortality and inter-patch variation in population density. The latter is dominantly driven by environmental fluctuations and the rate of population increase. These conclusions are not altered by the introduction of neighbourhood dispersal. With patch capacities in the order of 100 individuals, kin competition seems to be of negligible importance for ES dispersal rates except when overall dispersal rates are low.     

Effect of habitat quality and phenotypic variation on abundance- and trait-based early warning signals of population collapses

Loss of resilience in population numbers in response to environmental perturbations may be predicted with statistical metrics called early warning signals (EWS) that are derived from abundance time series. These signals, however, have been shown to have limited success, leading to the development of trait-based EWS that are based on information collected from phenotypic traits such as body size. Experimental work assessing the efficacy of EWS under varying ecological and environmental factors are rare. In addition, disentangling how such warning signals are affected under varying ecological and environmental factors is key to their application in biological conservation. Here, we experimentally test how different rates of environmental forcing (i.e. warming) and varying ecological factors (i.e. habitat quality and phenotypic diversity) affected population stability and predictive power of early warning signals of population collapse. We analyzed population density and body size time series data from three phenotypically different populations of a protozoan ciliate Askenasia volvox in two levels of habitat quality subjected to three different treatments of warming (i.e. no warming, fast warming and slow warming). We then evaluated how well abundance- and trait-based EWS predicted population collapses under different levels of phenotypic diversity, habitat quality and warming treatments. Our results suggest that habitat quality and warming treatments had more profound effects than phenotypic diversity had on both population stability and on the performance of abundance-based signals of population collapse. In addition, trait-based EWS generally performed well, were reliable and more robust in forecasting population collapse than abundance-based EWS, regardless of variation in environmental and ecological factors. Our study points towards the development of a predictive framework that includes information from phenotypic traits such as body size as an indicator of loss of resilience of ecological systems in response to environmental perturbations.     

Individual‐based modelling of resource competition to predict density‐dependent population dynamics: a case study with white storks

Density regulation influences population dynamics through its effects on demographic rates and consequently constitutes a key mechanism explaining the response of organisms to environmental changes. Yet, it is difficult to establish the exact form of density dependence from empirical data. Here, we developed an individual‐based model to explore how resource limitation and behavioural processes determine the spatial structure of white stork Ciconia ciconia populations and regulate reproductive rates. We found that the form of density dependence differed considerably between landscapes with the same overall resource availability and between home range selection strategies, highlighting the importance of fine‐scale resource distribution in interaction with behaviour. In accordance with theories of density dependence, breeding output generally decreased with density but this effect was highly variable and strongly affected by optimal foraging strategy, resource detection probability and colonial behaviour. Moreover, our results uncovered an overlooked consequence of density dependence by showing that high early nestling mortality in storks, assumed to be the outcome of harsh weather, may actually result from density dependent effects on food provision. Our findings emphasize that accounting for interactive effects of individual behaviour and local environmental factors is crucial for understanding density‐dependent processes within spatially structured populations. Enhanced understanding of the ways animal populations are regulated in general, and how habitat conditions and behaviour may dictate spatial population structure and demographic rates is critically needed for predicting the dynamics of populations, communities and ecosystems under changing environmental conditions.     

Causes and consequences of winter mortality in fishes

Winter mortality has been documented in a large number of freshwater fish populations, and a smaller, but increasing, number of marine and estuarine fishes. The impacted populations include a number of important North American and European resource species, yet the sources of winter mortality remain unidentified in most populations where it has been documented. Among the potential sources, thermal stress and starvation have received the most research attention. Other sources including predation and pathogens have significant impacts but have received insufficient attention to date. Designs of more recent laboratory experiments have reflected recognition of the potential for interactions among these co-occurring stressors. Geographic patterns in winter mortality are, in some cases, linked to latitudinal clines in winter severity and variability. However, for many freshwater species in particular, the effects of local community structure (predators and prey) may overwhelm latitudinal patterns. Marine (and estuarine) systems differ from freshwater systems in several aspects important to overwintering fishes, the most important being the lack of isolating barriers in the ocean. While open population boundaries allow fish to adopt migration strategies minimizing exposure to thermal stresses, they may retard rates of evolution to local environments. Geographic patterns in the occurrence and causes of winter mortality are ultimately determined by the interaction of regional and local factors. Winter mortality impacts population dynamics through episodic depressions in stock size and regulation of annual cohort strength. While the former tends to act in a density-independent manner, the latter can be density dependent, as most sources of mortality tend to select against the smallest members of the cohort and population. Most stock assessment and management regimes have yet to explicitly incorporate the variability in winter mortality. Potential management responses include postponement of cohort evaluation (to after first winter of life), harvest restrictions following mortality events and habitat enhancement. Future research should place more emphasis on the ecological aspects of winter mortality including the influences of food-web structure on starvation and predation. Beyond illuminating an understudied life-history phase, studies of overwintering ecology are integral to contemporary issues in fisheries ecology including ecosystem management, habitat evaluation, and impacts of climate change.     

The effects of environmental heterogeneity on multivariate selection on reproductive traits in female great tits

Describing natural selection on phenotypic traits under varying environmental conditions is essential for a quantitative assessment of the scale at which adaptation might occur and of the impact of environmental variability on evolution. Here we analyzed patterns of multivariate selection via fecundity and viability on three reproductive traits (laying date, clutch size, and egg weight) in a population of great tits (Parus major). We quantified selection under different environmental conditions using (1) local variation in breeding density and (2) distinct areas of the population's habitat. We found that selection gradients were generally stronger for fecundity than for viability selection. We also found correlational selection acting on the combination of laying date and clutch size; this is the first documented evidence of such selection acting on these two traits in a passerine bird. Our analyses showed that both local breeding density and habitat significantly influenced selection patterns, hence favoring different patterns of reproductive investment at a small-scale relative to typical dispersal distances in this species. Canonical rotation of the nonlinear selection matrices yielded similar conclusions as traditional nonlinear selection analyses, and also showed that the main axes of selection and fitness surfaces varied over space within the population. Our results emphasize the importance of quantifying different forms of selection, and of including variation in environmental conditions at small scales to gain a better understanding of potential evolutionary dynamics in wild populations. This study suggests that the fitness landscape for this species is relatively rugged at scales relevant to the life histories of individual birds and their close relatives.