Density-independent Processes Research Articles

Logistic-growth models measuring density feedback are sensitive to population declines, but not fluctuating carrying capacity.

Analysis of long-term trends in abundance of animal populations provides insights into population dynamics. Population growth rates are the emergent interplay of inter alia fertility, survival, and dispersal. However, the density feedbacks operating on some vital rates ("component feedback") can be decoupled from density feedbacks on population growth rates estimated using abundance time series ("ensemble feedback"). Many of the mechanisms responsible for this decoupling are poorly understood, thereby questioning the validity of using logistic-growth models versus vital rates to infer long-term population trends. To examine which conditions lead to decoupling, we simulated age-structured populations of long-lived vertebrates experiencing component density feedbacks on survival. We then quantified how imposed stochasticity in survival rates, density-independent mortality (catastrophes, harvest-like removal of individuals) and variation in carrying capacity modified the ensemble feedback in abundance time series simulated from age-structured populations. The statistical detection of ensemble density feedback from census data was largely unaffected by density-independent processes. Long-term population decline caused from density-independentmortalitywas the main mechanism decoupling the strength of component versus ensemble density feedbacks. Our study supports the use of simple logistic-growth models to capture long-term population trends, mediated by changes in population abundance, when survival rates are stochastic, carrying capacity fluctuates, and populations experience moderate catastrophic mortality over time.

Density regulation amplifies environmentally induced population fluctuations.

Density-dependent regulation is ubiquitous in population dynamics, and its potential interaction with environmental stochasticity complicates the characterization of the random component of population dynamics. Yet, this issue has not received attention commensurate with its relevance for descriptive and predictive modeling of population dynamics. Here we use a Bayesian modeling approach to investigate the contribution of density regulation to population variability in stochastic environments. We analytically derive a formula linking the stationary variance of population abundance/density under Gompertz regulation in a stochastic environment with constant variance to the environmental variance and the strength of density feedback, to investigate whether and how density regulation affects the stationary variance. We examine through simulations whether the relationship between stationary variance and density regulation inferred analytically under the Gompertz model carries over to the Ricker model, widely used in population dynamics modeling. The analytical decomposition of the stationary variance under stochastic Gompertz dynamics implies higher variability for strongly regulated populations. Simulation results demonstrate that the pattern of increasing population variability with increasing density feedback found under the Gompertz model holds for the Ricker model as well, and is expected to be a general phenomenon with stochastic population models. We also analytically established and empirically validated that the square of the autoregressive parameter of the Gompertz model in AR(1) form represents the proportion of stationary variance due to density dependence. Our results suggest that neither environmental stochasticity nor density regulation can alone explain the patterns of population variability in stochastic environments, as these two components of temporal variation interact, with a tendency for density regulation to amplify the magnitude of environmentally induced population fluctuations. This finding has far-reaching implications for population viability. It implies that intense intra-specific resource competition increases the risk of environment-driven population collapse at high density, making opportune harvesting a sensible practice for improving the resistance of managed populations such as fish stocks to environmental perturbations. The separation of density-dependent and density-independent processes will help improve population dynamics modeling, while providing a basis for evaluating the relative importance of these two categories of processes that remains a topic of long-standing controversy among ecologists.

Correction to: Strength of correlation between wildlife collision data and hunting bags varies among ungulate species and with management scale

Most European ungulate species are increasing in numbers and expanding their range. For the management and monitoring of these species, 64% of European countries rely on indirect proxies of abundance (e.g., hunting bag statistics). With increasing ungulate numbers, data on ungulate-vehicle collisions (UVC) may provide an important and inexpensive, complementary data source. Currently, it is unclear how bag statistics compare with UVC. A direct comparison of these two indices is important because both are used in ungulate management. We evaluated the relationship between UVC and ungulate hunting bags across bioclimatic, regional, and local scales, using five time lags (t−3 to t+1) for the five most common wild ungulate species in Sweden. For all species, hunting bags and UVC correlated positively, but correlation strength and time lags varied across scales and among species. The two indices correlated most strongly at the local management scale. Correlation between both indices was strong for the smaller deer species and wild boar, in particular, but much weaker for moose where we found the best fit using a 2-year time lag. For the other species, indices from the same year correlated best. We argue that the reason for moose data behaving differently is that, in Sweden, moose are formally managed using a 3-year time plan, while the other species are not. Accordingly, moose hunting bags are influenced more strongly by density-independent processes than bags of the other species. Consequently, the mismatch between the two indices may generate conflicting conclusions for management depending on the method applied.

Strength of correlation between wildlife collision data and hunting bags varies among ungulate species and with management scale

Most European ungulate species are increasing in numbers and expanding their range. For the management and monitoring of these species, 64% of European countries rely on indirect proxies of abundance (e.g., hunting bag statistics). With increasing ungulate numbers, data on ungulate-vehicle collisions (UVC) may provide an important and inexpensive, complementary data source. Currently, it is unclear how bag statistics compare with UVC. A direct comparison of these two indices is important because both are used in ungulate management. We evaluated the relationship between UVC and ungulate hunting bags across bioclimatic, regional, and local scales, using five time lags (t−3 to t+1) for the five most common wild ungulate species in Sweden. For all species, hunting bags and UVC correlated positively, but correlation strength and time lags varied across scales and among species. The two indices correlated most strongly at the local management scale. Correlation between both indices was strong for the smaller deer species and wild boar, in particular, but much weaker for moose where we found the best fit using a 2-year time lag. For the other species, indices from the same year correlated best. We argue that the reason for moose data behaving differently is that, in Sweden, moose are formally managed using a 3-year time plan, while the other species are not. Accordingly, moose hunting bags are influenced more strongly by density-independent processes than bags of the other species. Consequently, the mismatch between the two indices may generate conflicting conclusions for management depending on the method applied.

Host nutrition mediates interactions between plant viruses, altering transmission and predicted disease spread

Interactions among co-infecting pathogens are common across host taxa and can affect infectious disease dynamics. Host nutrition can mediate these among-pathogen interactions, altering the establishment and growth of pathogens within hosts. It is unclear, however, how nutrition-mediated among-pathogen interactions affect transmission and the spread of disease through populations. We manipulated the nitrogen (N) and phosphorus (P) supplies to oat plants in growth chambers and evaluated interactions between two aphid-vectored Barley and Cereal Yellow Dwarf Viruses: PAV and RPV. We quantified the effect of each virus on the other's establishment, within-plant density, and transmission. Co-inoculation significantly increased PAV density when N and P supplies were low and tended to increase RPV density when N supply was high. Co-infection increased PAV transmission when N and P supplies were low and tended to increase RPV transmission when N supply was high. Despite the parallels between the effects of among-pathogen interactions on density and transmission, changes in virus density only partially explained changes in transmission, suggesting that virus density-independent processes contribute to transmission. A mathematical model describing the spread of two viruses through a plant population, parameterized with empirically derived transmission values, demonstrated that nutrition-mediated among-pathogen interactions could affect disease spread. Interactions that altered transmission through virus density-independent processes determined overall disease dynamics. Our work suggests that host nutrition alters disease spread through among-pathogen interactions that modify transmission.

Nonlinearity in interspecific interactions in response to climate change: Cod and haddock as an example.

Climate change has profound ecological effects, yet our understanding of how trophic interactions among species are affected by climate change is still patchy. The sympatric Atlantic haddock and cod are co-occurring across the North Atlantic. They compete for food at younger stages and thereafter the former is preyed by the latter. Climate change might affect the interaction and coexistence of these two species. Particularly, the increase in sea temperature (ST) has been shown to affect distribution, population growth and trophic interactions in marine systems. We used 33-year long time series of haddock and cod abundances estimates from two data sources (acoustic and trawl survey) to analyse the dynamic effect of climate on the coexistence of these two sympatric species in the Arcto-Boreal Barents Sea. Using a Bayesian state-space threshold model, we demonstrated that long-term climate variation, as expressed by changes of ST, affected species demography through different influences on density-independent processes. The interaction between cod and haddock has shifted in the last two decades due to an increase in ST, altering the equilibrium abundances and the dynamics of the system. During warm years (ST over ca. 4°C), the increase in the cod abundance negatively affected haddock abundance while it did not during cold years. This change in interactions therefore changed the equilibrium population size with a higher population size during warm years. Our analyses show that long-term climate change in the Arcto-Boreal system can generate differences in the equilibrium conditions of species assemblages.

Population Trends and Urbanization: Simulating Density Effects Using a Local Regression Approach

Density-dependent population growth regulates long-term urban expansion and shapes distinctive socioeconomic trends. Despite a marked heterogeneity in the spatial distribution of the resident population, Mediterranean European countries are considered more homogeneous than countries in other European regions as far as settlement structure and processes of metropolitan growth are concerned. However, rising socioeconomic inequalities among Southern European regions reflect latent demographic and territorial transformations that require further investigation. An integrated assessment of the spatio-temporal distribution of resident populations in more than 1000 municipalities (1961–2011) was carried out in this study to characterize density-dependent processes of metropolitan growth in Greece. Using geographically weighted regressions, the results of our study identified distinctive local relationships between population density and growth rates over time. Our results demonstrate that demographic growth rates were non-linearly correlated with other variables, such as population density, with positive and negative impacts during the first (1961–1971) and the last (2001–2011) observation decade, respectively. These findings outline a progressive shift over time from density-dependent processes of population growth, reflecting a rapid development of large metropolitan regions (Athens, Thessaloniki) in the 1960s, to density-dependent processes more evident in medium-sized cities and accessible rural regions in the 2000s. Density-independent processes of population growth have been detected in the intermediate study period (1971–2001). This work finally discusses how a long-term analysis of demographic growth, testing for density-dependent mechanisms, may clarify the intrinsic role of population concentration and dispersion in different phases of the metropolitan cycle in Mediterranean Europe.

Source water variability as a driver of rockfish recruitment in the California Current Ecosystem: implications for climate change and fisheries management

Elucidating connections between ocean climate variability and change and recruitment of juvenile fishes to adult populations is critical for understanding variability in stock–recruit dynamics. Recruitment to adult rockfish populations in the California Current Ecosystem (CCE) is highly variable, leading to short- and long-term changes in abundance, productivity, forage availability, and potential fisheries yield. We used regional ocean model output, oceanographic data, and a 34-year time series of pelagic juvenile rockfish to investigate the interaction between changes in CCE source waters, as reflected by physical water mass properties, and recruitment variability. Specifically, variability of “spiciness” on upper water isopycnals explains a substantial fraction of the variation in pelagic juvenile rockfish abundance. High rockfish abundances correspond to cooler, fresher waters with higher dissolved oxygen (i.e., “minty”) conditions, indicative of Pacific subarctic water. By contrast, years of low rockfish abundance are associated with warmer, more saline, and more oxygen-deficient (i.e., “spicy”) conditions, reflecting waters of subtropical or equatorial origin. Transport and source waters in the CCE are key factors determining density-independent processes and subsequent recruitment to adult populations.

Density-independent and density-dependent factors affecting spatio-temporal dynamics of Atlantic cod (Gadus morhua) distribution in the Gulf of Maine

Abstract Due to strong spatial interplays between intraspecific interactions and environmental forcing, both density-dependent and density-independent processes can affect spatio-temporal dynamics of fish populations in a spatially explicit fashion. To this end, this study investigated the underlying mechanisms of spatio-temporal dynamics of Atlantic cod (Gadus morhua) in the Gulf of Maine (GoM). Based on the data from the Northeast Fisheries Science Center (NEFSC) bottom-trawl surveys in spring and fall from 1982 to 2013, empirical cumulative distribution function (ECDF) curves and geographic distribution indices were used to examine the species–environment and abundance–occupancy relationship, respectively. Then, a variable-coefficient generalized additive model was constructed to quantify the simultaneous effects of environmental variables and population size on the spatio-temporal dynamics of cod distribution. Area occupied remained relatively high through the late 1990s, but underwent a pronounced contraction into the western GoM (WGoM) for the reminder of the time-series. The model results suggest that the spatio-temporal dynamics of GoM cod have been driven by complex interactions of density-dependent and density-independent factors over the past three decades. Better knowledge of these dynamics can improve our understanding of the causality of abundance–occupancy and species–environment relationships and help to reduce error estimates for survey-based indices.

A Natural History of Change in Native Bees Associated with Lowbush Blueberry in Maine

More than 120 native bee species have been documented in Maine since 1930 in association with the native plant Vaccinium angustifolium (Lowbush Blueberry). We report 3 studies in commercial Lowbush Blueberry fields: (1) a survey of diversity in Osmia (mason bees) and closely related Megachile (leaf-cutter bees) using trap nests in 93 fields from 1990 to 2012, (2) a 29-year study of a native bee community, and (3) an examination of climate-change effects on bee-foraging periods during blueberry bloom. Osmia appeared to be more stable over a 22- year period in their species richness and relative abundances in Lowbush Blueberry fields when compared to Megachile over a similar 17-year period. The native bee community in a single location in Winterport was observed to fluctuate in abundance 2 to 3 times annually. Modeling of the total bee community and taxa-specific group abundances (Bombus, Megachilids, Andrenids, and Halictids and other bees) suggest that while stochastic density-independent processes such as weather can play a role in determining their annual oscillations, density-dependent lags of 1 and 2 years appear to be the main driving forces. Estimation of fruit set over the same 29-year period, based upon native bee abundance, suggests that pollination is more buffered than community bee abundance, resulting in a lesser degree of fluctuation over time. We speculate that this finding is due to redundancy in floral preferences, multiple floral visitations, and differing pollination efficiencies by the highly diverse native bee community associated with Lowbush Blueberry. Effects of climate change in Maine Lowbush Blueberry fields during May bloom was investigated using a historic weather database. Since the early 1990s, precipitation has, to a large degree, reduced the number of optimal bee foraging days during bloom, with implications for pollination and bee species abundances. This new information reinforces the need for provision of pollinator gardens to support native pollinators of Lowbush Blueberry.

Europe-wide consistency in density-dependence of red deer (Cervus elaphus) fertility

Population dynamics of large herbivores are affected by density dependent and density independent processes. Changing population size causes variation in age- and sex-specific demographic parameters such as reproduction, survival and dispersal. With our study, based on meta-analyses of literature data, we aimed at (1) describing age-specific fertility rates of red deer (Cervus elaphus) throughout Europe, and (2) tracking negative density feedback in reproduction of female red deer. Across the species range, in most European populations (84%) females began breeding as yearlings. We observed age-specific variation in female fertility, which was the lowest and most diverse in yearling females. Increasing population density had a negative effect on female fertility in all age groups. There was no difference in the strength of the relationship between different age classes of females. In 1.5-year old females, at low and moderate densities, fertility exhibited a high variation, which could indicate strong susceptibility of youngest females to detrimental external factors. Our study showed that meta-analysis of Europe-wide data was able to detect biologically relevant relationships which may be difficult to observe when applying locally-based and time-scale approach.

Interactions among habitat, management, and demography for an invasive annual grass

Effective long-term management of noxious plant species depends on our ability to identify and manage ecological processes-driving invasion and to use site-specific information to design realistic management strategies and goals. However, there is a surprising lack of research that investigates relationships among habitat characteristics, weed demography, and management efforts. We investigated the interactions among these factors using the invasive annual grass medusahead (Elymus caput-medusae). In Yuba County, California, we seeded medusahead in densities from 0 to 50,000 seeds/m2 in open grassland and oak woodland habitats. We also exposed plots to defoliation none, once, or twice in a season and captured how medusahead germination, establishment, and persistence responded to experimental treatments. We found that medusahead establishment was higher in the grassland, compared to the woodland habitat, likely due, in part, to the presence of litter under oak canopies. After a single defoliation event, medusahead persistence was higher in the oak, but not the grassland plots. We also found that defoliation once or twice reduced medusahead density. However, a single clipping treatment actually resulted in increased seed production. This was likely a compensatory response by a younger cohort, and provides an explanation for why single control efforts do not generally result in successful long-term outcomes. This work highlights the complex relationships between density-dependent and density-independent processes that may influence invasion dynamics.

Extended density-dependent mortality in mature conifer forests: causes and implications for ecosystem management.

Understanding processes driving mortality in forests is important for comprehension of natural stand dynamics and for informing natural disturbance-based ecosystem management. There has been considerable study of mortality in forests during the self-thinning phase but we know much less about processes driving mortality in stands at later successional stages. We addressed this through study of five 1-ha spatially explicit permanent plots in mature (111-186yr old in 2012) Pinus contorta stands in the Canadian Rocky Mountains using data from repeated measurements over a 45-yr period, dendrochronological information, and point pattern analysis. We tested the hypothesis that these stands had completed the self-thinning/density-dependent mortality stage of succession. Contrary to our expectations, the self-thinning phase can persist for more than 140yr following stand establishment. Our findings suggest this was attributable to prolonged post-fire establishment periods due to surface fires in three of the plots while in the other two plots moist conditions and slow growth most likely delayed the onset of competition. Several pieces of evidence indicated the importance of density-dependent mortality in these stands over the study period: (1) The diameter distribution of individuals changed from initially right-skewed toward normality as a result of mortality of smaller-diameter stems. (2) Individuals of lower canopy positions were proportionally more affected by mortality. (3) When compared to the pre-mortality pattern, surviving stems in all stands had an increasingly uniform spatial distribution. In two of the plots, recent windthrow and/or ingrowth initially hindered our ability to detect density-dependent mortality but our dendrochronological sampling and permanent plot data allowed us to untangle the different processes at play; in doing so we demonstrate for the first time how density-independent processes can mask underlying density-dependent mortality processes in older stands. Mortality of larger dominant canopy trees increased over the study period and mortality of dominant stems was a random process in all stands suggesting these stands were approaching the end of the self-thinning stage and that density-independent processes might soon become more important. Our results provide an improved understanding of mortality processes that can be applied to natural disturbance-based ecosystem management.

Testing for effects of climate change on competitive relationships and coexistence between two bird species.

Climate change is expected to have profound ecological effects, yet shifts in competitive abilities among species are rarely studied in this context. Blue tits (Cyanistes caeruleus) and great tits (Parus major) compete for food and roosting sites, yet coexist across much of their range. Climate change might thus change the competitive relationships and coexistence between these two species. Analysing four of the highest-quality, long-term datasets available on these species across Europe, we extend the textbook example of coexistence between competing species to include the dynamic effects of long-term climate variation. Using threshold time-series statistical modelling, we demonstrate that long-term climate variation affects species demography through different influences on density-dependent and density-independent processes. The competitive interaction between blue tits and great tits has shifted in one of the studied sites, creating conditions that alter the relative equilibrium densities between the two species, potentially disrupting long-term coexistence. Our analyses show that long-term climate change can, but does not always, generate local differences in the equilibrium conditions of spatially structured species assemblages. We demonstrate how long-term data can be used to better understand whether (and how), for instance, climate change might change the relationships between coexisting species. However, the studied populations are rather robust against competitive exclusion.

Effects of spatial structure of population size on the population dynamics of barnacles across their elevational range.

Explanations for why population dynamics vary across the range of a species reflect two contrasting hypotheses: (i) temporal variability of populations is larger in the centre of the range compared to the margins because overcompensatory density dependence destabilizes population dynamics and (ii) population variability is larger near the margins, where populations are more susceptible to environmental fluctuations. In both of these hypotheses, positions within the range are assumed to affect population variability. In contrast, the fact that population variability is often related to mean population size implies that the spatial structure of the population size within the range of a species may also be a useful predictor of the spatial variation in temporal variability of population size over the range of the species. To explore how population temporal variability varies spatially and the underlying processes responsible for the spatial variation, we focused on the intertidal barnacle Chthamalus dalli and examined differences in its population dynamics along the tidal levels it inhabits. Changes in coverage of barnacle populations were monitored for 10.5 years at 25 plots spanning the elevational range of this species. Data were analysed by fitting a population dynamics model to estimate the effects of density-dependent and density-independent processes on population growth. We also examined the temporal mean-variance relationship of population size with parameters estimated from the population dynamics model. We found that the relative variability of populations tended to increase from the centre of the elevational range towards the margins because of an increase in the magnitude of stochastic fluctuations of growth rates. Thus, our results supported hypothesis (2). We also found that spatial variations in temporal population variability were well characterized by Taylor's power law, the relative population variability being inversely related to the mean population size. Results suggest that understanding the population dynamics of a species over its range may be facilitated by taking the spatial structure of population size into account as well as by considering changes in population processes as a function of position within the range of the species.

Dispersal‐mediated effect of microhabitat availability and density dependence determine population dynamics of a forest floor web spider

Landscapes in nature can be viewed as a continuum of small total habitable area with high fragmentation to widely spreading habitats. The dispersal-mediated rescue effect predominates in the former landscapes, while classical density-dependent processes generally prevail in widely spread habitats. A similar principle should be applied to populations of organisms utilizing microhabitats in limited supply. To test this hypothesis, we examined the population dynamics of a web spider, Neriene brongersmai, in 16 populations with varying degrees of microhabitat availability, and we explored whether: (i) high microhabitat availability improves survival rate during density-independent movement, while the resultant high density reduces survival rate in a density-dependent manner; and (ii) temporal population stability increases with microhabitat availability at the population level. Furthermore, we conducted two types of field experiments to verify whether high microhabitat availability actually reduces mortality associated with web-site movement. Field observations revealed that demographic change in N.brongersmai populations was affected by three factors at different stages, namely the microhabitat limitation from the early to late juvenile stages, the density dependence from the late juvenile to adult stages and the food limitation from the adult to the next early juvenile stages. In addition, there was a tendency for a positive association between population stability and microhabitat availability at the population level. A small-scale experiment, where the frequency of spider web relocation was equalized artificially, revealed that high microhabitat availability elevated the survival rate during a movement event between web-sites. The larger spatiotemporal scale experiment also revealed an improved spider survival rate following treatment with high microhabitat availability, even though spider density was kept at a relatively low level. The population dynamics of N.brongersmai can be determined primarily by density-independent processes based on web-site fragmentation and density-dependent processes driven by interference competition. We conclude that depending on the amount of habitat resources, the relative importance of the two contrasting paradigms-equilibrium and non-equilibrium-appears to vary, even within a particular system.

Density-Dependent Regulation of Brook Trout Population Dynamics along a Core-Periphery Distribution Gradient in a Central Appalachian Watershed

Spatial population models predict strong density-dependence and relatively stable population dynamics near the core of a species' distribution with increasing variance and importance of density-independent processes operating towards the population periphery. Using a 10-year data set and an information-theoretic approach, we tested a series of candidate models considering density-dependent and density-independent controls on brook trout population dynamics across a core-periphery distribution gradient within a central Appalachian watershed. We sampled seven sub-populations with study sites ranging in drainage area from 1.3–60 km2 and long-term average densities ranging from 0.335–0.006 trout/m. Modeled response variables included per capita population growth rate of young-of-the-year, adult, and total brook trout. We also quantified a stock-recruitment relationship for the headwater population and coefficients of variability in mean trout density for all sub-populations over time. Density-dependent regulation was prevalent throughout the study area regardless of stream size. However, density-independent temperature models carried substantial weight and likely reflect the effect of year-to-year variability in water temperature on trout dispersal between cold tributaries and warm main stems. Estimated adult carrying capacities decreased exponentially with increasing stream size from 0.24 trout/m in headwaters to 0.005 trout/m in the main stem. Finally, temporal variance in brook trout population size was lowest in the high-density headwater population, tended to peak in mid-sized streams and declined slightly in the largest streams with the lowest densities. Our results provide support for the hypothesis that local density-dependent processes have a strong control on brook trout dynamics across the entire distribution gradient. However, the mechanisms of regulation likely shift from competition for limited food and space in headwater streams to competition for thermal refugia in larger main stems. It also is likely that source-sink dynamics and dispersal from small headwater habitats may partially influence brook trout population dynamics in the main stem.

What makes a species common? No evidence of density-dependent recruitment or mortality of the sea urchin Diadema antillarum after the 1983–1984 mass mortality

A potential consequence of individuals compensating for density-dependent processes is that rare or infrequent events can produce profound and long-term shifts in species abundance. In 1983-1984 a mass mortality event reduced the numbers of the abundant sea urchin Diadema antillarum by 95-99% throughout the Caribbean and western Atlantic. Following this event, the abundance of macroalgae increased and the few surviving D. antillarum responded by increasing in body size and fecundity. These initial observations suggested that populations of D. antillarum could recover rapidly following release from food limitation. In contrast, published studies of field manipulations indicate that this species had traits making it resistant to density-dependent effects on offspring production and adult mortality; this evidence raises the possibility that density-independent processes might keep populations at a diminished level. Decadal-scale (1983-2011) monitoring of recruitment, mortality, population density and size structure of D. antillarum from St John, US Virgin Islands, indicates that population density has remained relatively stable and more than an order of magnitude lower than that before the mortality event of 1983-1984. We detected no evidence of density-dependent mortality or recruitment since this mortality event. In this location, model estimates of equilibrium population density, assuming density-independent processes and based on parameters generated over the first decade following the mortality event, accurately predict the low population density 20 years later (2011). We find no evidence to support the notion that this historically dominant species will rebound from this temporally brief, but spatially widespread, perturbation.

Climate, fishing, and fluctuations of sardine and anchovy in the California Current

Since the days of Elton, population cycles have challenged ecologists and resource managers. Although the underlying mechanisms remain debated, theory holds that both density-dependent and density-independent processes shape the dynamics. One striking example is the large-scale fluctuations of sardine and anchovy observed across the major upwelling areas of the world. Despite a long history of research, the causes of these fluctuations remain unresolved and heavily debated, with significant implications for fisheries management. We here model the underlying causes of these fluctuations, using the California Current Ecosystem as a case study, and show that the dynamics, accurately reproduced since A.D. 1661 onward, are explained by interacting density-dependent processes (i.e., through species-specific life-history traits) and climate forcing. Furthermore, we demonstrate how fishing modifies the dynamics and show that the sardine collapse of the 1950s was largely unavoidable given poor recruitment conditions. Our approach provides unique insight into the origin of sardine-anchovy fluctuations and a knowledge base for sustainable fisheries management in the California Current Ecosystem and beyond.

Winter climate, age, and population density affect the timing of conception in female moose (Alces alces)

Phenological events such as conception or parturition dates may have profound impact on several key life-history traits of ungulates at the individual as well as the population level. However, relatively little is known about the causes of variation in the timing of reproduction. Based on a 17-year survey of reproductive tracts, we investigated the effect of climate, population density, and age on the conception date of female moose (Alces alces) harvested in Estonia. Ninety-five percent of studied moose cows were conceived within a period of 9 weeks (29 August–30 October), while more than 45 % of all moose cows were conceived from 19 September to 2 October. Conception date was negatively related to population density and nonlinearly to the regional measure of winter climate reflecting the maximal extent of ice on the Baltic Sea (MIE) in the previous winter. High air temperatures during rut (in September) delayed the conception date. The timing of conception also depended on female age. Yearlings conceived significantly later as compared to females of all other age groups. Our findings corroborate the importance of density-dependent as well density-independent processes on the timing of conception of this ungulate. We also propose that the effect of population density on conception date may be mediated by increasing ecological carrying capacity concurrent with increasing population abundance.