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Chance pervades life. In turn, life histories are described by probabilities (e.g. survival and breeding) and averages across individuals (e.g. mean growth rate and age at maturity). In this study, we explored patterns of luck in lifetime outcomes by analysing structured population models for a wide array of plant and animal species. We calculated four response variables: variance and skewness in both lifespan and lifetime reproductive output (LRO), and partitioned them into contributions from different forms of luck. We examined relationships among response variables and a variety of life history traits. We found that variance in lifespan and variance in LRO were positively correlated across taxa, but that variance and skewness were negatively correlated for both lifespan and LRO. The most important life history trait was longevity, which shaped variance and skew in LRO through its effects on variance in lifespan. We found that luck in survival, growth, and fecundity all contributed to variance in LRO, but skew in LRO was overwhelmingly due to survival luck. Rapidly growing populations have larger variances in LRO and lifespan than shrinking populations. Our results indicate that luck-induced genetic drift may be most severe in recovering populations of species with long mature lifespan and high iteroparity.

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Variation in life history traits in animals and plants can often be structured along major axes of life history strategies. The position of a species along these axes can inform on their sensitivity to environmental change. For example, species with slow life histories are found to be less sensitive in their long-term population responses to environmental change than species with fast life histories. This provides a tantalizing link between sets of traits and population responses to change, contained in a highly generalizable theoretical framework. Life history strategies are assumed to reflect the outcome of life history tradeoffs that, by their very nature, act at the individual level. Examples include the tradeoff between current and future reproductive success, and allocating energy into growth versus reproduction. But the importance of such tradeoffs in structuring population-level responses to environmental change remains understudied. We aim to increase our understanding of the link between individual-level life history tradeoffs and the structuring of life history strategies across species, as well as the underlying links to population responses to environmental change. We find that the classical association between lifehistory strategies and population responses to environmental change breaks down when accounting for individual-level tradeoffs and energy allocation. Therefore, projecting population responses to environmental change should not be inferred based only on a limited set of species traits. We summarize our perspective and a way forward in a conceptual framework.

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AbstractDispersal emerges as an outcome of organismal traits and external forcings. However, it remains unclear how the emergent dispersal kernel evolves as a by-product of selection on the underlying traits. This question is particularly compelling in coastal marine systems, where dispersal is tied to development and reproduction and where directional currents bias larval dispersal downstream, causing selection for retention. We modeled the dynamics of a metapopulation along a finite coastline using an integral projection model and adaptive dynamics to understand how asymmetric coastal currents influence the evolution of larval (pelagic larval duration) and adult (spawning frequency) life history traits, which indirectly shape the evolution of marine dispersal kernels. Selection induced by alongshore currents favors the release of larvae over multiple time periods, allowing long pelagic larval durations and long-distance dispersal to be maintained in marine life cycles in situations where they were previously predicted to be selected against. Two evolutionarily stable strategies emerged: one with a long pelagic larval duration and many spawning events, resulting in a dispersal kernel with a larger mean and variance, and another with a short pelagic larval duration and few spawning events, resulting in a dispersal kernel with a smaller mean and variance. Our theory shows how coastal ocean flows are important agents of selection that can generate multiple, often co-occurring evolutionary outcomes for marine life history traits that affect dispersal.

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Climate is assumed to strongly influence species distribution and abundance. Although the performance of many organisms is influenced by the climate in their immediate proximity, the climate data used to model their distributions often have a coarse spatial resolution. This is problematic because the local climate experienced by individuals might deviate substantially from the regional average. This problem is likely to be particularly important for sessile organisms like plants and in environments where small-scale variation in climate is large. To quantify the effect of local temperature on vital rates and population growth rates, we used temperature values measured at the local scale (in situ logger measures) and integral projection models with demographic data from 37 populations of the forest herb Lathyrus vernus across a wide latitudinal gradient in Sweden. To assess how the spatial resolution of temperature data influences assessments of climate effects, we compared effects from models using local data with models using regionally aggregated temperature data at several spatial resolutions (≥1 km). Using local temperature data, we found that spring frost reduced the asymptotic population growth rate in the first of two annual transitions and influenced survival in both transitions. Only one of the four regional estimates showed a similar negative effect of spring frost on population growth rate. Our results for a perennial forest herb show that analyses using regionally aggregated data often fail to identify the effects of climate on population dynamics. This emphasizes the importance of using organism-relevant estimates of climate when examining effects on individual performance and population dynamics, as well as when modeling species distributions. For sessile organisms that experience the environment over small spatial scales, this will require climate data at high spatial resolutions.

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Body size shifts in ectotherms are mostly attributed to the Temperature Size Rule (TSR) stating that warming speeds up initial growth rate but leads to smaller size when food does not limit growth. Investigating the links between temperature, growth, and life history traits is key to understand the adaptive value of TSR, which might be context dependent. In particular, global warming can affect food quantity or quality which is another major driver of growth, fecundity, and survival. However, we have limited information on how temperature and food jointly influence life history traits in vertebrate predators and how changes in different life history traits combine to influence fitness and population demography. We investigate (1) whether TSR is maintained under different food conditions, (2) if food exacerbates or dampens the effects of temperature on growth and life history traits and (3) if food influences the adaptive value of TSR. We combine experiments on the medaka with Integral Projection Models to scale from life history traits to fitness consequences. Our results confirm that warming triggers a higher initial growth rate and a lower adult size, reduces generation time and increases mean fitness. A lower level of food exacerbates the effects of warming on growth trajectories. Although lower feeding frequency increased survival and decreased fecundity, it did not influence the effects of warming on fish development rates, fecundity, and survival. In contrast, feeding frequency influenced the adaptive value of TSR, as, under intermittent feeding, generation time decreased faster with warming and the increase in growth rate with warming was weaker compared to continuously fed fish. These results are of importance in the context of global warming as resources are expected to change with increasing temperatures but, surprisingly, our results suggest that feeding frequency have a lower impact on fitness at high temperature.

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The persistent exposure of coral assemblages to more variable abiotic regimes is assumed to augment their resilience to future climatic variability. Yet, while the determinants of coral population resilience across species remain unknown, we are unable to predict the winners and losers across reef ecosystems exposed to increasingly variable conditions. Using annual surveys of 3171 coral individuals across Australia and Japan (2016-2019), we explore spatial variation across the short- and long-term dynamics of competitive, stress-tolerant, and weedy assemblages to evaluate how abiotic variability mediates the structural composition of coral assemblages. We illustrate how, by promoting short-term potential over long-term performance, coral assemblages can reduce their vulnerability to stochastic environments. However, compared to stress-tolerant, and weedy assemblages, competitive coral taxa display a reduced capacity for elevating their short-term potential. Accordingly, future climatic shifts threaten the structural complexity of coral assemblages in variable environments, emulating the degradation expected across global tropical reefs.

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Identifying and accounting for unobserved individual heterogeneity in vital rates in demographic models is important for estimating population-level vital rates and identifying diverse life-history strategies, but much less is known about how this individual heterogeneity influences population dynamics. We aimed to understand how the distribution of individual heterogeneity in reproductive and survival rates influenced population dynamics using vital rates from a Weddell seal population by altering the distribution of individual heterogeneity in reproduction, which also altered the distribution of individual survival rates through the incorporation of our estimate of the correlation between the two rates and assessing resulting changes in population growth. We constructed an integral projection model (IPM) structured by age and reproductive state using estimates of vital rates for a long-lived mammal that has recently been shown to exhibit large individual heterogeneity in reproduction. Using output from the IPM, we evaluated how population dynamics changed with different underlying distributions of unobserved individual heterogeneity in reproduction. Results indicate that the changes to the underlying distribution of individual heterogeneity in reproduction cause very small changes in the population growth rate and other population metrics. The largest difference in the estimated population growth rate resulting from changes to the underlying distribution of individual heterogeneity was less than 1%. Our work highlights the differing importance of individual heterogeneity at the population level compared to the individual level. Although individual heterogeneity in reproduction may result in large differences in the lifetime fitness of individuals, changing the proportion of above- or below-average breeders in the population results in much smaller differences in annual population growth rate. For a long-lived mammal with stable and high adult-survival that gives birth to a single offspring, individual heterogeneity in reproduction has a limited effect on population dynamics. We posit that the limited effect of individual heterogeneity on population dynamics may be due to canalization of life-history traits.

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Fed aquaculture is one of the fastest-growing and most valuable food production industries in the world. The efficiency with which farmed fish convert feed into biomass influences both environmental impact and economic revenue. Salmonid species, such as king salmon (Oncorhynchus tshawytscha), exhibit high levels of plasticity in vital rates such as feed intake and growth rates. Accurate estimations of individual variability in vital rates are important for production management. The use of mean trait values to evaluate feeding and growth performance can mask individual-level differences that potentially contribute to inefficiencies. Here, the authors apply a cohort integral projection model (IPM) framework to investigate individual variation in growth performance of 1625 individually tagged king salmon fed one of three distinct rations of 60%, 80%, and 100% satiation and tracked over a duration of 276 days. To capture the observed sigmoidal growth of individuals, they compared a nonlinear mixed-effects (logistic) model to a linear model used within the IPM framework. Ration significantly influenced several aspects of growth, both at the individual and at the cohort level. Mean final body mass and mean growth rate increased with ration; however, variance in body mass and feed intake also increased significantly over time. Trends in mean body mass and individual body mass variation were captured by both logistic and linear models, suggesting the linear model to be suitable for use in the IPM. The authors also observed that higher rations resulted in a decreasing proportion of individuals reaching the cohort's mean body mass or larger by the end of the experiment. This suggests that, in the present experiment, feeding to satiation did not produce the desired effects of efficient, fast, and uniform growth in juvenile king salmon. Although monitoring individuals through time is challenging in commercial aquaculture settings, recent technological advances combined with an IPM approach could provide new scope for tracking growth performance in experimental and farmed populations. Using the IPM framework might allow the exploration of other size-dependent processes affecting vital rate functions, such as competition and mortality.

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The center-periphery hypothesis predicts a decline in population performance toward the periphery of a species' range, reflecting an alteration of environmental conditions at range periphery. However, the rare demographic tests of this hypothesis failed to disentangle the role of geography from that of ecological niche and are biased toward temperate regions. We hypothesized that, because species are expected to experience optimal abiotic conditions at their climatic niche center, (1) central populations will have better demographic growth, survival, and fertility than peripheral populations. As a result, (2) central populations are expected to have higher growth rates than peripheral populations. Peripheral populations are expected to decline, thus limiting species range expansion beyond these boundaries. Because peripheral populations are expected to be in harsh environmental conditions, (3) population growth rate will be more sensitive to perturbation of survival-growth rather than fertility in peripheral populations. Finally, we hypothesized that (4) soils properties will drive the variations in population growth rates for narrowly distributed species for which small scale ecological factors could outweigh landscape level drivers. To test these hypotheses, we studied the demography of Thunbergia atacorensis (Acanthaceae), a range-limited herb in West Africa. We collected three years of demographic data to parameterize an integral projection model (IPM) and estimated population level demographic statistics. Demographic vital rates and population growth rates did not change significantly with distance from geographic or climatic center, contrary to predictions. However, populations at the center of the geographic range were demographically more resilient to perturbation than those at the periphery. Soil nitrogen was the main driver of population growth rate variation. The relative influence of survival-growth on population growth rates exceeded that of fertility at the geographic range center while we observed the opposite pattern for climatic niche. Our study highlights the importance of local scale processes in shaping the dynamics and distribution of range-limited species. Our findings also suggest that the distinction between geographic distribution and climatic niche is important for a robust demographic test of the center-periphery hypothesis.

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Competition is among the most important factors regulating plant population and community dynamics, but we know little about how different vital rates respond to competition and jointly determine population growth and species coexistence. We conducted a field experiment and parameterised integral projection models to model the population growth of 14 herbaceous plant species in the absence and presence of neighbours across an elevation gradient (284 interspecific pairs). We found that suppressed individual growth and seedling establishment contributed the most to competition-induced declines in population growth, although vital rate contributions varied greatly between species and with elevation. In contrast, size-specific survival and flowering probability and seed production were frequently enhanced under competition. These compensatory vital rate responses were nearly ubiquitous (occurred in 92% of species pairs) and significantly reduced niche overlap and stabilised coexistence. Our study highlights the importance of demographic processes for regulating population and community dynamics, which has often been neglected by classic coexistence theories.

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Acute environmental stressors such as short-term exposure to pollutants can have lasting effects on organisms, potentially impacting future generations. Parental exposure to toxicants can result in changes to the epigenome (e.g., DNA methylation) that are passed down to subsequent, unexposed generations. However, it is difficult to gauge the cumulative population-scale impacts of epigenetic effects from laboratory experiments alone. Here, we developed a size- and age-structured delay-coordinate population model to evaluate the long-term consequences of epigenetic modifications on population sustainability. The model emulated changes in growth, mortality, and fecundity in the F0, F1, and F2 generations observed in experiments in which larval Menidia beryllina were exposed to environmentally relevant concentrations of bifenthrin (Bif), ethinylestradiol (EE2), levonorgestrel (LV), or trenbolone (TB) in the parent generation (F0) and reared in clean water up to the F2 generation. Our analysis suggests potentially dramatic population-level effects of repeated, chronic exposures of early-life stage fish that are not captured by models not accounting for those effects. Simulated exposures led to substantial declines in population abundance (LV and Bif) or near-extinction (EE2 and TB) with the exact trajectory and timeline of population decline dependent on the combination of F0, F1, and F2 effects produced by each compound. Even acute one-time exposures of each compound led to declines and recovery over multiple years due to lagged epigenetic effects. These results demonstrate the potential for environmentally relevant concentrations of commonly used compounds to impact the population dynamics and sustainability of an ecologically relevant species and model organism.

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Quantifying relationships between genetic variation and population viability is important from both basic biological and applied conservation perspectives, yet few populations have been monitored with both long-term demographic and population genetics approaches. To empirically test whether and how genetic variation and population dynamics are related, we present one such paired approach. First, we use eight years of historical demographic data from five populations of Boechera fecunda (Brassicaceae), a rare, self-compatible perennial plant endemic to Montana, USA, and use integral projection models to estimate the stochastic population growth rate (λ S) and extinction risk of each population. We then combine these demographic estimates with previously published metrics of genetic variation in the same populations to test whether genetic diversity within populations is linked to demographic performance. Our results show that in this predominantly inbred species, standing genetic variation and demography are weakly positively correlated. However, the inbreeding coefficient was not strongly correlated with demographic performance, suggesting that more inbred populations are not necessarily less viable or at higher extinction risk than less inbred populations. A contemporary re-census of these populations revealed that neither genetic nor demographic parameters were consistently strong predictors of current population density, although populations showing lower probabilities of extinction in demographic models had larger population sizes at present. In the absence of evidence for inbreeding depression decreasing population viability in this species, we recommend conservation of distinct, potentially locally adapted populations of B. fecunda rather than alternatives such as translocations or reintroductions.

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AbstractExtinctions are predicted to rise by an order of magnitude over the next century. Although contemporary documented extinctions are uncommon, local extirpations likely provide hints about global extinction risks. Comparing responses to global change of locally extinct versus extant species pairs in a phylogenetic framework could highlight why certain species are more vulnerable to extinction than others and which anthropogenic changes are most relevant to their decline. As anthropogenic changes likely interact to affect population declines, demographic studies partitioning the effects of multifactorial stressors are needed but remain rare. I examine demographic responses to nitrogen addition and deer herbivory, two major drivers of species losses in grasslands, in experimental reintroductions of 14 locally extinct and extant confamilial native plants from Michigan prairies. Nitrogen consistently reduces survival, especially in locally extinct species, and growth of locally extinct species benefits less from nitrogen than growth of extant species. Nitrogen reduces population growth rates, largely via reductions in survival. Deer herbivory, meanwhile, had inconsistent effects on vital rates among species and did not affect population growth. Nitrogen and herbivory rarely interacted to affect vital rates. These results link community-level patterns of species loss under nitrogen addition to the population-level processes underlying those losses.

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Demographic compensation-the opposing responses of vital rates along environmental gradients-potentially delays anticipated species' range contraction under climate change, but no consensus exists on its actual contribution. We calculated population growth rate (λ) and demographic compensation across the distributional ranges of 81 North American tree species and examined their responses to simulated warming and tree competition. We found that 43% of species showed stable population size at both northern and southern edges. Demographic compensation was detected in 25 species, yet 15 of them still showed a potential retraction from southern edges, indicating that compensation alone cannot maintain range stability. Simulated climatic warming caused larger decreases in λ for most species and weakened the effectiveness of demographic compensation in stabilising ranges. These findings suggest that climate stress may surpass the limited capacity of demographic compensation and pose a threat to the viability of North American tree populations.

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Climate change alters many aspects of weed performance and may also alter the effectiveness of management practices to control pests. Despite this concern, entire categories of widely used management practices, such as physical control, remain understudied in this context. We conducted a field experiment growing the invasive pest musk thistle (Carduus nutans) at ambient and experimentally elevated temperatures. We tested mowing management strategies that varied in the timing of a single mowing event relative to thistles' stem elongation phenology and compared these with an unmowed control. Results from this experiment informed demographic models to project population growth rates for different warming/mowing scenarios. Compared to plants grown under ambient conditions, warmed thistles were more likely to survive the same mowing treatment, flowered earlier in the season, grew to taller heights, and produced more flowering capitula. Proportional reductions in plant height and capitulum production caused by mowing were smaller under warming. Warming did not change the relative ranking of mowing treatments; mowing late in the growing season (2 weeks after individuals first reached a height of 40 cm) was most effective at ambient temperatures and under warming. Warming caused significant increases in projected local population growth rate for all mowing treatments. For invasive musk thistle, warmed individuals outperformed individuals grown at ambient temperatures across all the mowing treatments we considered. Our results suggest that to achieve outcomes comparable to those attainable at today's temperatures, farmers will need to apply supplemental management, possibly including additional mowing effort or alternative practices such as chemical control. We recommend that scientists test management practices under experimental warming, where possible, and that managers monitor ongoing management to identify changes in effectiveness. Information about changes in managed weeds' mortality, fecundity, and phenology can then be used to make informed decisions in future climates.

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The effectiveness of hedgerows as functional corridors in the face of climate warming has been little researched. Here we investigated the effects of warming temperatures on plant performance and population growth of Geum urbanum in forests versus hedgerows in two European temperate regions. Adult individuals were transplanted in three forest-hedgerow pairs in each of two different latitudes, and an experimental warming treatment using open-top chambers was used in a full factorial design. Plant performance was analysed using mixed models and population performance was analysed using Integral Projection Models and elasticity analyses. Temperature increases due to open-top chamber installation were higher in forests than in hedgerows. In forests, the warming treatment had a significant negative effect on the population growth rate of G. urbanum. In contrast, no significant effect of the warming treatment on population dynamics was detected in hedgerows. Overall, the highest population growth rates were found in the forest control sites, which was driven by a higher fecundity rather than a higher survival probability. Effects of warming treatments on G. urbanum population growth rates differed between forests and hedgerows. In forests, warming treatments negatively affected population growth, but not in hedgerows. This could be a consequence of the overall lower warming achieved in hedgerows. We conclude that maintenance of cooler forest microclimates coul, at least temporarily, moderate the species response to climate warming.

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In many ecosystems, especially aquatic ecosystems, size plays a critical role in the factors that determine an individual's ability to survive and reproduce. In aquatic ecotoxicology, size informs both realized and potential acute and chronic effects of chemical exposure. This paper demonstrates how chemical and nonchemical effects on growth, survival, and reproduction can be linked to population-level dynamics using size-structured integral projection models (IPM). The modeling approach was developed with the goals and constraints of ecological risk assessors in mind, who are tasked with estimating the effects of chemical exposures to wildlife populations in a data-limited environment. The included case study is a collection of daily time-step IPMs parameterized for the life history and annual cycle of fathead minnows (Pimephales promelas), which motivated the development of modeling techniques for seasonal, iteroparous reproduction, density dependent growth effects, and size-dependent over-winter survival. The effects of a time-variable annual chemical exposure were interpreted using a toxicokinetic-toxicodynamic model for acute survival and sub-lethal growth effects model for chronic effects and incorporated into the IPMs. This paper presents a first application of integral projection models to ecotoxicology. Our research demonstrates that size-structured IPMs provide a promising, flexible, framework for synthesizing ecotoxicologically relevant data and theory to explore the effects of chemical and nonchemical stressors and the resulting impacts on exposed populations.

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Estimates of the percentage of species "committed to extinction" by climate change range from 15% to 37%. The question is whether factors other than climate need to be included in models predicting species' range change. We created demographic range models that include climate vs. climate-plus-competition, evaluating their influence on the geographic distribution of Pinus edulis, a pine endemic to the semiarid southwestern U.S. Analyses of data on 23,426 trees in 1941 forest inventory plots support the inclusion of competition in range models. However, climate and competition together only partially explain this species' distribution. Instead, the evidence suggests that climate affects other range-limiting processes, including landscape-scale, spatial processes such as disturbances and antagonistic biotic interactions. Complex effects of climate on species distributions-through indirect effects, interactions, and feedbacks-are likely to cause sudden changes in abundance and distribution that are not predictable from a climate-only perspective.

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Climate change is expected to alter the distribution and abundance of tree species, impacting ecosystem structure and function. Yet, anticipating where this will occur is often hampered by a lack of understanding of how demographic rates, most notably recruitment, vary in response to climate and competition across a species range. Using large-scale monitoring data on two dry woodland tree species (Pinus edulis and Juniperus osteosperma), we develop an approach to infer recruitment, survival, and growth of both species across their range. In doing so, we account for ecological and statistical dependencies inherent in large-scale monitoring data. We find that drying and warming conditions generally lead to declines in recruitment and survival, but the strength of responses varied between species. These climate conditions point to geographic regions of high vulnerability for particular species, such as Pinus edulis in northern Arizona, where both survival and recruitment are low. Our approach provides a path forward for leveraging emerging large-scale monitoring and remotely sensed data to anticipate the impacts of global change on species distributions.

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Increases in the frequency and intensity of acute and chronic disturbances are causing declines of coral reefs world-wide. Although quantifying the responses of corals to acute disturbances is well documented, detecting subtle responses of coral populations to chronic disturbances is less common, but can also result in altered population and community structures. We investigated the population dynamics of two key reef-building Merulinid coral species, Dipsastraea favus and Platygyra lamellina, with similar life-history traits, in the Gulf of Eilat and Aqaba, Red Sea from 2015 to 2018, to assess potential differences in their population trajectories. Demographic processes, which included rates of survival, growth, reproduction and recruitment were used to parametrize integral projection models and estimate population growth rates and the likely population trajectories of both coral species. The survival and reproduction rates of both D. favus and P. lamellina were positively related to coral colony size, and elasticity analyses showed that large colonies most influenced population dynamics. Although both species have similar life-history traits and growth morphologies and are generally regarded as 'stress-tolerant', the populations showed contrasting trajectories-D. favus appears to be increasing whereas P. lamellina appears to be decreasing. As many corals have long-life expectancies, the process of local and regional decline might be subtle and slow. Ecological assessments based on total living coral coverage, morphological groups or functional traits might overlook subtle, species-specific trends. However, demographic approaches capable of detecting subtle species-specific population changes can augment ecological studies and provide valuable early warning signs of decline before major coral loss becomes evident.

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