Transient LTRE analysis reveals the demographic and trait-mediated processes that buffer population growth.

Temporal variation in environmental conditions affects population growth directly via its impact on vital rates, and indirectly through induced variation in demographic structure and phenotypic trait distributions. We currently know very little about how these processes jointly mediate population responses to their environment. To address this gap, we develop a general transient life table response experiment (LTRE) which partitions the contributions to population growth arising from variation in (1) survival and reproduction, (2) demographic structure, (3) trait values and (4) climatic drivers. We apply the LTRE to a population of yellow-bellied marmots (Marmota flaviventer) to demonstrate the impact of demographic and trait-mediated processes. Our analysis provides a new perspective on demographic buffering, which may be a more subtle phenomena than is currently assumed. The new LTRE framework presents opportunities to improve our understanding of how trait variation influences population dynamics and adaptation in stochastic environments

Evolution of size-dependent flowering in a variable environment: construction and analysis of a stochastic integral projection model

Understanding why individuals delay reproduction is a classic problem in evolutionary biology. In plants, the study of reproductive delays is complicated because growth and survival can be size and age dependent, individuals of the same size can grow by different amounts and there is temporal variation in the environment. We extend the recently developed integral projection approach to include size- and age-dependent demography and temporal variation. The technique is then applied to a long-term individually structured dataset for Carlina vulgaris, a monocarpic thistle. The parameterized model has excellent descriptive properties in terms of both the population size and the distributions of sizes within each age class. In Carlina, the probability of flowering depends on both plant size and age. We use the parameterized model to predict this relationship, using the evolutionarily stable strategy approach. Considering each year separately, we show that both the direction and the magnitude of selection on the flowering strategy vary from year to year. Provided the flowering strategy is constrained, so it cannot be a step function, the model accurately predicts the average size at flowering. Elasticity analysis is used to partition the size- and age-specific contributions to the stochastic growth rate, λs. We use λs to construct fitness landscapes and show how different forms of stochasticity influence its topography. We prove the existence of a unique stochastic growth rate, λs, which is independent of the initial population vector, and show that Tuljapurkar's perturbation analysis for log(λs) can be used to calculate elasticities

Cumulative weather effects can impact across the whole life cycle

Predicting how species will be affected by future climatic change requires the underlying environmental drivers to be identified. As vital rates vary over the lifecycle, structured population models derived from statistical environment-demography relationships are often used to inform such predictions. Environmental drivers are typically identified independently for different vital rates and demographic classes. However, these rates often exhibit positive temporal covariance, suggesting the vital rates respond to common environmental drivers. Additionally, models often only incorporate average weather conditions during a single, a priori chosen time window (e.g. monthly means). Mismatches between these windows and the period when the vital rates are sensitive to variation in climate decrease the predictive performance of such approaches. We used a demographic structural equation model (SEM) to demonstrate that a single axis of environmental variation drives the majority of the (co)variation in survival, reproduction, and twinning across six age-sex classes in a Soay sheep population. This axis provides a simple target for the complex task of identifying the drivers of vital rate variation. We used functional linear models (FLMs) to determine the critical windows of three local climatic drivers, allowing the magnitude and direction of the climate effects to differ over time. Previously unidentified lagged climatic effects were detected in this well-studied population. The FLMs had a better predictive performance than selecting a critical window a priori, but not than a large-scale climate index. Positive covariance amongst vital rates and temporal variation in the effects of environmental drivers are common, suggesting our SEM-FLM approach is a widely applicable tool for exploring the joint responses of vital rates to environmental change

Funder Restrictions on Application Numbers Lead to Chaos

Restricting application rates is an attractive way for funders to reduce time and money wasted evaluating uncompetitive applications. However, mathematical models show that this could induce chaotic cycles in total application numbers, increasing uncertainty in the funding process. One emergent property is that smaller institutions spend disproportionally more time unfunded

Exploring population responses to environmental change when there is never enough data: a factor analytic approach

© 2018 The Authors. Methods in Ecology and Evolution published by John Wiley & Sons Ltd on behalf of British Ecological Society Temporal variability in the environment drives variation in vital rates, with consequences for population dynamics and life-history evolution. Integral projection models (IPMs) are data-driven structured population models widely used to study population dynamics and life-history evolution in temporally variable environments. However, many datasets have insufficient temporal replication for the environmental drivers of vital rates to be identified with confidence, limiting their use for evaluating population level responses to environmental change. Parameter selection, where the kernel is constructed at each time step by randomly selecting the time-varying parameters from their joint probability distribution, is one approach to including stochasticity in IPMs. We consider a factor analytic (FA) approach for modelling the covariance matrix of time-varying parameters, whereby latent variable(s) describe the covariance among vital rate parameters. This decreases the number of parameters to estimate and, where the covariance is positive, the latent variable can be interpreted as a measure of environmental quality. We demonstrate this using simulation studies and two case studies. The simulation studies suggest the FA approach provides similarly accurate estimates of stochastic population growth rate to estimating an unstructured covariance matrix. We demonstrate how the latent parameter can be perturbed to show how selection on reproductive delays in the monocarp Carduus nutans changes under different environmental conditions. We develop a demographic model of the fire dependent herb Eryngium cuneifolium to show how a putative driver of the variation in environmental quality can be incorporated with the addition of a single parameter. Using perturbation analyses we determine optimal management strategies for this species. This approach estimates fewer parameters than previous approaches and allows novel eco-evolutionary insights. Predictions on population dynamics and life-history evolution under different environmental conditions can be made without necessarily identifying causal factors. Putative environmental drivers can be incorporated with relatively few parameters, allowing for predictions on how populations will respond to changes in the environment

Changes in age-structure over four decades were a key determinant of population growth rate in a long-lived mammal

A changing environment directly influences birth and mortality rates, and thus population growth rates. However, population growth rates in the short term are also influenced by population age-structure. Despite its importance, the contribution of age-structure to population growth rates has rarely been explored empirically in wildlife populations with long-term demographic data. Here we assessed how changes in age-structure influenced short-term population dynamics in a semi-captive population of Asian elephantsElephas maximus. We addressed this question using a demographic dataset of female Asian elephants from timber camps in Myanmar spanning 45 years (1970-2014). First, we explored temporal variation in age-structure. Then, using annual matrix population models, we used a retrospective approach to assess the contributions of age-structure and vital rates to short-term population growth rates with respect to the average environment. Age-structure was highly variable over the study period, with large proportions of juveniles in the years 1970 and 1985, and made a substantial contribution to annual population growth rate deviations. High adult birth rates between 1970 and 1980 would have resulted in large positive population growth rates, but these were prevented by a low proportion of reproductive-aged females. We highlight that an understanding of both age-specific vital rates and age-structure is needed to assess short-term population dynamics. Furthermore, this example from a human-managed system suggests that the importance of age-structure may be accentuated in populations experiencing human disturbance where age-structure is unstable, such as those in captivity or for endangered species. Ultimately, changes to the environment drive population dynamics by influencing birth and mortality rates, but understanding demographic structure is crucial for assessing population growth

The implications of seasonal climatic effects for managing disturbance dependent populations under a changing climate

The frequency of ecological disturbances, such as fires, is changing due to changing land use and climatic conditions. Disturbance-adapted species may thus require the manipulation of disturbance regimes to persist. However, the effects of changes in other abiotic factors, such as climatic conditions, are frequently disregarded in studies of such systems. Where climatic effects are included, relatively simple approaches that disregard seasonal variation in the effects are typically used. We compare predictions of population persistence using different fire return intervals (FRIs) under recent and predicted future climatic conditions for the rare fire-dependent herb Eryngium cuneifolium. We used functional linear models (FLMs) to estimate the cumulative effect of climatic variables across the annual cycle, allowing the strength and direction of the climatic impacts to differ over the year. We then estimated extinction probabilities and minimum population sizes under past and forecasted future climatic conditions and a range of FRIs. Under forecasted climate change, E. cuneifolium is predicted to persist under a much broader range of FRIs, because increasing temperatures are associated with faster individual growth. Climatic impacts on fecundity do not result in a temporal trend in this vital rate due to antagonistic seasonal effects operating through winter and summer temperatures. These antagonistic seasonal climatic effects highlight the importance of capturing the seasonal dependence of climatic effects when forecasting their future fate. Synthesis. Awareness of the potential effects of climate change on disturbance-adapted species is necessary for developing suitable management strategies for future environmental conditions. However, our results suggest that widely used simple methods for modelling climate impacts, that disregard seasonality in such effects, may produce misleading inferences

The epidemiology underlying age-related avian malaria infection in a long-lived host: the mute swan Cygnus olor

Quantifying the factors that predict parasite outbreak and persistence is a major challenge for both applied and fundamental biology. Key to understanding parasite prevalence and disease outbreaks is determining at what age individuals show signs of infection, and whether or not they recover. Age-dependent patterns of the infection of a host population by parasites can indicate among-individual heterogeneities in their susceptibility to, or rate of recovery from, parasite infections. Here, we present a cross-sectional study of avian malaria in a long-lived bird species, the mute swan Cygnus olor, examining age-related patterns of parasite prevalence and modelling patterns of infection and recovery. One-hundred and fifteen swans, ranging from one to nineteen years old, were screened for infection with Plasmodium, Haemoproteus and Leucocytozoon parasites. Infections with three cytochrome-b lineages of Haemoproteus were found (pooled prevalence 67%), namely WW1 (26%), which is common in passerine birds, and two new lineages closely related to WW1: MUTSW1 (25%) and MUTSW2 (16%). We found evidence for age-related infection in one lineage, MUTSW1. Catalytic models examining patterns of infection and recovery in the population suggested that infections in this population were not life-long – recovery of individuals was included in the best fitting models. These findings support the results of recent studies that suggest hosts can clear infections, although patterns of infection-related mortality in older birds remain to be studied in more detail

Fine-scale spatial variation in fitness is comparable to disturbance-induced fluctuations in a fire-adapted species

The spatial scale at which demographic performance (e.g., net reproductive output) varies can profoundly influence landscape-level population growth and persistence, and many demographically pertinent processes such as species interactions and resource acquisition vary at fine scales. We compared the magnitude of demographic variation associated with fine-scale heterogeneity (1 ha) fluctuations associated with fire disturbance. We used a spatially explicit model within an IPM modeling framework to evaluate the demographic importance of fine-scale variation. We used a measure of expected lifetime fruit production, EF, that is assumed to be proportional to lifetime fitness. Demographic differences and their effects on EF were assessed in a population of the herbaceous perennial Hypericum cumulicola (~2,600 individuals), within a patch of Florida rosemary scrub (400 × 80 m). We compared demographic variation over fine spatial scales to demographic variation between years across 6 yr after a fire. Values of EF changed by orders of magnitude over <10 m. This variation in fitness over fine spatial scales (<10 m) is commensurate to postfire changes in fitness for this fire-adapted perennial. A life table response experiment indicated that fine-scale spatial variation in vital rates, especially survival, explains as much change in EF as demographic changes caused by time-since-fire, a key driver in this system. Our findings show that environmental changes over a few tens of meters can have ecologically meaningful implications for population growth and extinction