Predicting changes in the distribution and abundance of species under environmental change

Environmental changes are expected to alter both the distribution and the abundance of organisms. A disproportionate amount of past work has focused on distribution only, either documenting historical range shifts or predicting future occurrence patterns. However, simultaneous predictions of abundance and distribution across landscapes would be far more useful. To critically assess which approaches represent advances towards the goal of joint predictions of abundance and distribution, we review recent work on changing distributions and on effects of environmental drivers on single populations. Several methods have been used to predict changing distributions. Some of these can be easily modified to also predict abundance, but others cannot. In parallel, demographers have developed a much better understanding of how changing abiotic and biotic drivers will influence growth rate and abundance in single populations. However, this demographic work has rarely taken a landscape perspective and has largely ignored the effects of intraspecific density. We advocate a synthetic approach in which population models accounting for both density dependence and effects of environmental drivers are used to make integrated predictions of equilibrium abundance and distribution across entire landscapes. Such predictions would constitute an important step forward in assessing the ecological consequences of environmental changes.     

Evolutionary tracking is determined by differential selection on demographic rates and density dependence

Recent ecological forecasts predict that ~25% of species worldwide will go extinct by 2050. However, these estimates are primarily based on environmental changes alone and fail to incorporate important biological mechanisms such as genetic adaptation via evolution. Thus, environmental change can affect population dynamics in ways that classical frameworks can neither describe nor predict. Furthermore, often due to a lack of data, forecasting models commonly describe changes in population demography by summarizing changes in fecundity and survival concurrently with the intrinsic growth rate (r). This has been shown to be an oversimplification as the environment may impose selective pressure on specific demographic rates (birth and death) rather than directly on r (the difference between the birth and death rates). This differential pressure may alter population response to density, in each demographic rate, further diluting the information combined to produce r. Thus, when we consider the potential for persistence via adaptive evolution, populations with the same r can have different abilities to persist amidst environmental change. Therefore, we cannot adequately forecast population response to climate change without accounting for demography and selection on density dependence. Using a continuous‐time Markov chain model to describe the stochastic dynamics of the logistic model of population growth and allow for trait evolution via mutations arising during birth events, we find persistence via evolutionary tracking more likely when environmental change alters birth rather than the death rate. Furthermore, species that evolve responses to changes in the strength of density dependence due to environmental change are less vulnerable to extinction than species that undergo selection independent of population density. By incorporating these key demographic considerations into our predictive models, we can better understand how species will respond to climate change.     

Predicting conditions for migration: effects of density dependence and habitat quality

Migration is widespread among animals, but the factors that influence the decision to migrate are poorly understood. Within a single species, populations may be completely migratory, completely sedentary or partially migratory. We use a population model to derive conditions for migration and demonstrate how migratory survival, habitat quality and density dependence on both the breeding and non-breeding grounds influence conditions for migration and the proportion of migrants within a population. Density dependence during the season in which migratory and sedentary individuals use separate sites is necessary for partial migration. High levels of density dependence at the non-shared sites widen the range of survival values within which we predict partial migration, whereas increasing the strength of density dependence at the shared sites narrows the range of survival values within which we predict partial migration. Our results have important implications for predicting how contemporary populations with variable migration strategies may respond to changes in the quality or quantity of habitat.     

Pattern of variation in avian population growth rates

A central question in population ecology is to understand why population growth rates differ over time. Here, we describe how the long-term growth of populations is not only influenced by parameters affecting the expected dynamics, for example form of density dependence and specific population growth rate, but is also affected by environmental and demographic stochasticity. Using long-term studies of fluctuations of bird populations, we show an interaction between the stochastic and the deterministic components of the population dynamics: high specific growth rates at small densities r(1) are typically positively correlated with the environmental variance sigma(e)(2). Furthermore, theta, a single parameter describing the form of the density regulation in the theta-logistic density-regulation model, is negatively correlated with r(1). These patterns are in turn correlated with interspecific differences in life-history characteristics. Higher specific growth rates, larger stochastic effects on the population dynamics and stronger density regulation at small densities are found in species with large clutch sizes or high adult mortality rates than in long-lived species. Unfortunately, large uncertainties in parameter estimates, as well as strong stochastic effects on the population dynamics, will often make even short-term population projections unreliable. We illustrate that the concept of population prediction interval can be useful in evaluating the consequences of these uncertainties in the population projections for the choice of management actions.     

Talkin' 'bout my generation: environmental variability and cohort effects

In variable environments, it is probable that environmental conditions in the past can influence demographic performance now. Cohort effects occur when these delayed life-history effects are synchronized among groups of individuals in a population. Here we show how plasticity in density-dependent demographic traits throughout the life cycle can lead to cohort effects and that there can be substantial population dynamic consequences of these effects. We show experimentally that density and food conditions early in development can influence subsequent juvenile life-history traits. We also show that conditions early in development can interact with conditions at maturity to shape future adult performance. In fact, conditions such as food availability and density at maturity, like conditions early in development, can generate cohort effects in mature stages. Based on these data, and on current theory about the effects of plasticity generated by historical environments, we make predictions about the consequences of such changes on density-dependent demography and on mite population dynamics. We use a stochastic cohort effects model to generate a range of population dynamics. In accordance with the theory, we find the predicted changes in the strength of density dependence and associated changes in population dynamics and population variability.     

Growth and development rates have different thermal responses

Growth and development rates are fundamental to all living organisms. In a warming world, it is important to determine how these rates will respond to increasing temperatures. It is often assumed that the thermal responses of physiological rates are coupled to metabolic rate and thus have the same temperature dependence. However, the existence of the temperature-size rule suggests that intraspecific growth and development are decoupled. Decoupling of these rates would have important consequences for individual species and ecosystems, yet this has not been tested systematically across a range of species. We conducted an analysis on growth and development rate data compiled from the literature for a well-studied group, marine pelagic copepods, and use an information-theoretic approach to test which equations best describe these rates. Growth and development rates were best characterized by models with significantly different parameters: development has stronger temperature dependence than does growth across all life stages. As such, it is incorrect to assume that these rates have the same temperature dependence. We used the best-fit models for these rates to predict changes in organism mass in response to temperature. These predictions follow a concave relationship, which complicates attempts to model the impacts of increasing global temperatures on species body size.     

Bird behaviour and environmental planning: approaches in the study of wader populations

Predicting the effect of changes in land-use on bird populations requires a degree of understanding of their population dynamics which is seldom available. Such knowledge is especially difficult to acquire if the birds occupy a variety of habitats of differing quality and they are also migratory. The effects of habitat loss at a particular time of year on the year-round population dynamics are then difficult to predict. The problem is discussed using wintering waders as examples. Unless a species can extend its present range, the initial effect of habitat loss is to increase bird density. Whether this affects population size will depend on whether bird density affects either winter survival or subsequent breeding success. Measuring such density-dependent relationships is in practice extremely difficult in migratory populations. However, behavioural studies help in testing the key hypothesis that birth and death rates are affected by bird density, and may even allow the form of any density-dependent functions to be deduced. Simulation modelling then allows the effects of habitat loss on the overall population dynamics to be explored. The general point is that behavioural studies play an important part in Environmental Impact Studies because behaviour is the main way in which birds respond to environmental changes and compete for limited resources, such as diminishing habitat. They are therefore likely to provide insights when making predictions about the responses of birds and populations to habitat loss.     

Single‐ or multistage regulation in complex life cycles: does it make a difference?

Data on the different stages of complex life cycles are often rather unbalanced, especially those concerning the effects of density. How does this affect our understanding of a species’ population dynamics? Two discrete three‐stage models with overlapping generations and delayed maturation are constructed to address this question. They assume that survival or emigration in any life stage and/or reproduction can be density dependent. A typical pond‐breeding amphibian species with a well‐studied larval stage serves as an example. Numerical results show that the population dynamics resulting from density dependence at a single (e.g. the larval) stage can be decisively and unpredictably modified by density dependence in additional stages. Superposition of density‐dependent processes could thus be one reason for the difficulties in identifying density dependence in the field. Moreover, in a simulated source‐refuge system with habitat‐specific density‐dependent dispersal of juveniles density dependence in multiple stages can stabilize or destabilize the dynamics and produce misleading age structures. From an applied perspective this model shows that excluding multistage regulation prematurely clearly affects our ability to predict consequences of human impacts.     

Toward an ecological synthesis: a case for habitat selection

Habitat selection, and its associated density and frequency-dependent evolution, has a profound influence on such vital phenomena as population regulation, species interactions, the assembly of ecological communities, and the origin and maintenance of biodiversity. Different strategies of habitat selection, and their importance in ecology and evolution, can often be revealed simply by plots of density in adjacent habitats. For individual species, the strategies are closely intertwined with mechanisms of population regulation, and with the persistence of populations through time. For interacting species, strategies of habitat selection are not only responsible for species coexistence, but provide one of the most convenient mechanisms for measuring competition, and the various community structures caused by competitive interactions. Other kinds of interactions, such as those between predators and prey, demonstrate that an understanding of the coevolution of habitat-selection strategies among strongly interacting species is essential to properly interpret their spatial and temporal dynamics. At the evolutionary scale, the frequency dependence associated with habitat selection may often allow populations to diverge and diversify into separate species. Habitat selection thereby demonstrates how we can map microevolutionary strategies in behavior onto their population and community consequences, and from there, onto macroevolutionary patterns of speciation and adaptive radiation. We can anticipate that future studies of habitat selection will not only help us complete those maps, but that they will also continue to enrich the panoply of ideas that shape evolutionary ecology.     

Individual‐based ecology of coastal birds

Conservation objectives for non‐breeding coastal birds (shorebirds and wildfowl) are determined from their population size at coastal sites. To advise coastal managers, models must predict quantitatively the effects of environmental change on population size or the demographic rates (mortality and reproduction) that determine it. As habitat association models and depletion models are not able to do this, we developed an approach that has produced such predictions thereby enabling policy makers to make evidence‐based decisions. Our conceptual framework is individual‐based ecology, in which populations are viewed as having properties (e.g. size) that arise from the traits (e.g. behaviour, physiology) and interactions of their constituent individuals. The link between individuals and populations is made through individual‐based models (IBMs) that follow the fitness‐maximising decisions of individuals and predict population‐level consequences (e.g. mortality rate) from the fates of these individuals. Our first IBM was for oystercatchers Haematopus ostralegus and accurately predicted their density‐dependent mortality. Subsequently, IBMs were developed for several shorebird and wildfowl species at several European sites, and were shown to predict accurately overwinter mortality, and the foraging behaviour from which predictions are derived. They have been used to predict the effect on survival in coastal birds of sea level rise, habitat loss, wind farm development, shellfishing and human disturbance. This review emphasises the wider applicability of the approach, and identifies other systems to which it could be applied. We view the IBM approach as a very useful contribution to the general problem of how to advance ecology to the point where we can routinely make meaningful predictions of how populations respond to environmental change.     

Environmental variation and population responses to global change

Species' responses to environmental changes such as global warming are affected not only by trends in mean conditions, but also by natural and human‐induced environmental fluctuations. Methods are needed to predict how such environmental variation affects ecological and evolutionary processes, in order to design effective strategies to conserve biodiversity under global change. Here, we review recent theoretical and empirical studies to assess: (1) how populations respond to changes in environmental variance, and (2) how environmental variance affects population responses to changes in mean conditions. Contrary to frequent claims, empirical studies show that increases in environmental variance can increase as well as decrease long‐term population growth rates. Moreover, environmental variance can alter and even reverse the effects of changes in the mean environment, such that even if environmental variance remains constant, omitting it from population models compromises their ability to predict species' responses to changes in mean conditions. Drawing on theory relating these effects of environmental variance to the curvatures of population growth responses to the environment, we outline how species' traits such as phylogenetic history and body mass could be used to predict their responses to global change under future environmental variability.     

Variation in plant quality and the population dynamics of herbivores: there is nothing average about aphids

In the attempt to use results from small-scale studies to make large-scale predictions, it is critical that we take into account the greater spatial heterogeneity encountered at larger spatial scales. An important component of this heterogeneity is variation in plant quality, which can have a profound influence on herbivore population dynamics. This influence is particularly relevant when we consider that the strength of density dependence can vary among host plants and that the strength of density dependence determines the difference between exponential and density- dependent growth. Here, we present some simple models and analyses designed to examine the impact of variable plant quality on the dynamics of insect herbivore populations, and specifically the consequences of variation in the strength of density dependence among host plants. We show that average values of herbivore population growth parameters, calculated from plants that vary in quality, do not predict overall population growth. Furthermore, we illustrate that the quality of a few individual plants within a larger plant population can dominate herbivore population growth. Our results demonstrate that ignoring spatial heterogeneity that exists in herbivore population growth on plants that differ in quality can lead to a misunderstanding of the mechanisms that underlie population dynamics.     

Predicting when climate‐driven phenotypic change affects population dynamics

Species' responses to climate change are variable and diverse, yet our understanding of how different responses (e.g. physiological, behavioural, demographic) relate and how they affect the parameters most relevant for conservation (e.g. population persistence) is lacking. Despite this, studies that observe changes in one type of response typically assume that effects on population dynamics will occur, perhaps fallaciously. We use a hierarchical framework to explain and test when impacts of climate on traits (e.g. phenology) affect demographic rates (e.g. reproduction) and in turn population dynamics. Using this conceptual framework, we distinguish four mechanisms that can prevent lower‐level responses from impacting population dynamics. Testable hypotheses were identified from the literature that suggest life‐history and ecological characteristics which could predict when these mechanisms are likely to be important. A quantitative example on birds illustrates how, even with limited data and without fully‐parameterized population models, new insights can be gained; differences among species in the impacts of climate‐driven phenological changes on population growth were not explained by the number of broods or density dependence. Our approach helps to predict the types of species in which climate sensitivities of phenotypic traits have strong demographic and population consequences, which is crucial for conservation prioritization of data‐deficient species.     

Increased outbreak frequency associated with changes in the dynamic behaviour of populations of two aphid species

Sudden changes in the variability of natural populations can result in increased likelihood of extinction or in greater frequency and intensity of pest outbreaks. These changes could be associated with changes in some relevant population parameters such as the equilibrium density or the maximum population growth rate. However, changes in these parameters have very different consequences. An increase in equilibrium density results in a higher variance in population fluctuations according to the relationship between mean and variance described by Taylor's power law, but does not modify the stability properties of the system. On the other hand, changes in the maximum growth rate induce changes in the dynamic regimes and stability properties of the population. In this study, using statistical and mathematical methods borrowed from econometrics and engineering, we identify structural changes to the variance in the population dynamics of the sycamore aphid Drepanosiphum platanoidis and the green spruce aphid Elatobium abietinum in the UK. Some localities showed strong changes in their population parameters, such that their dynamic regime changed completely. These changes in the population dynamic regimes increase the expected frequency of outbreaks, which has enormous economic and ecological consequences. Through this study we show the application of methods that could be helpful to pest and wildlife managers in the task of evaluating changes in the risk of outbreaks or extinction of animal populations under changing global environmental scenarios.     

Predicting the effect of disturbance on coastal birds

Assessments of whether disturbance is having a deleterious effect on populations have often measured behavioural responses to disturbance and assumed that populations with a larger behavioural response are more susceptible to disturbance. However, there is no guarantee that the behavioural response to disturbance is related to the population consequence, measured in terms of decreased reproduction or increased mortality. Individual-based models, consisting of fitness-maximizing individuals, are one means of linking the behavioural responses to disturbance to population consequences. This paper reviews how individual-based models have been used to predict the effect of disturbance on populations of shorebirds and wildfowl at several European sites, and shows how these models could be improved in the future by incorporating a range of alternative responses to disturbance.     

The evolution of dispersal – the importance of information about population density and habitat characteristics

The evolution of mobility patterns and dispersal strategies depend on different population, habitat and life history characteristics. The ability to perceive and make use of information about the surrounding environment for dispersal decisions will also differ between organisms. To investigate the evolutionary consequences of such differences, we have used a simulation model with nearest-neighbour dispersal in a metapopulation to study how variation in the ability to obtain and make use of information about habitat quality and conspecific density affects the evolution of dispersal strategies. We found a rather strong influence of variation in information on the overall rate of dispersal in a metapopulation. The highest emigration rate evolved in organisms with no information about either density or habitat quality and the lowest rate was found in organisms with information about both the natal and the neighbouring patches. For organisms that can make use of information about conspecific density, positively density-dependent dispersal evolved in the majority of cases, with the strongest density dependence occurring when an individual only has information about density in the natal patch. However, we also identified situations, involving strong local population fluctuations and frequent local extinctions, where negatively density-dependent dispersal evolved.     

The role of temperature variability on insect performance and population dynamics in a warming world

Despite the amount of research on the consequences of global warming on ecological systems, most studies examine the impact of increases in average temperature. However, there are few studies concerning the role of thermal variability on ecological processes. Based on insect thermal and population ecology, we propose a theoretical framework for organizing the study of the role that thermal mean and variability plays in individual performance, and how it may affect population dynamics. Starting with three predictions of global warming scenarios, we develop null models of the expected changes in individual physiological performance and population dynamics. Ecological consequences in each scenario may range from simple changes in performance to drastic changes in population fluctuations and geographic ranges. In particular, our null models show that potential changes in the intrinsic population growth rate (R_m) will depend on the interaction of mean temperature and thermal variability, and that the net effect of the interaction could be synergistic or antagonistic. To evaluate these null models, we fit performance curves to compiled data from the literature on measurements of R_m at several constant and fluctuating temperatures. The fitted models showed that several of the qualitative characteristics predicted by the null model may be found in the fitted curves. We expect that this framework will be useful as a guide to study the influence of thermal changes on the dynamics of natural populations. Synthesis Despite the common assertion that global warming impacts depend on not only the mean temperatures but also on thermal variability, theoretical approaches to explain how the interaction of thermal mean and variability determines fitness are lacking. Here we propose a framework for studying the role of thermal mean and variability on individual performance and population dynamics. We developed null models that show how changes in the intrinsic population growth rate (R_m) will depend on the interaction of mean temperature and thermal variability, and that the net effect could be synergistic or antagonistic. We expect that this framework will be useful to study the influence of thermal changes on natural populations.     

Patterns and processes of population dynamics with fluctuating habitat size: a case study of a marine copepod inhabiting tide pools

The logistic model is a fundamental population model often used as the basis for analyzing wildlife population dynamics. In the classic logistic model, however, population dynamics may be difficult to characterize if habitat size is temporally variable because population density can vary at a constant abundance, which results in variable strength of density‐dependent feedback for a given population size. To incorporate habitat size variability, we developed a general population model in which changes in population abundance, density, and habitat size are taken into account. From this model, we deduced several predictions for patterns and processes of population dynamics: 1) patterns of fluctuation in population abundance and density can diverge, with respect of their correlation and relative variability; and 2) along with density dependence, habitat size fluctuation can affect population growth with a time lag because changes in habitat size result in changes in population density. In order to test these predictions, we applied our model to population dynamics data of 36 populations of Tigriopus japonicus, a marine copepod inhabiting tide pools of variable sizes caused by weather processes. As expected, we found a significant difference in the fluctuation patterns of population abundance and density of T. japonicus populations with respect to the correlation between abundance and density and their relative variability, which correlates positively with the variability of habitat size. In addition, we found direct and lagged‐indirect effects of weather processes on population growth, which were associated with density dependence and impose regulatory forces on local and regional population dynamics. These results illustrate how changes in habitat size can have an impact on patterns and processes of wildlife population dynamics. We suggest that without knowledge of habitat size fluctuation, measures of population size and its variability as well as inferences about the processes of population dynamics may be misleading.     

Changes in dispersal during species' range expansions

Explanations for rapid species' range expansions have typically been purely ecological, with little attention given to evolutionary processes. We tested predictions for the evolution of dispersal during range expansion using four species of wing-dimorphic bush cricket (Conocephalus discolor, Conocephalus dorsalis, Metrioptera roeselii, and Metrioptera brachyptera). We observed distinct changes in dispersal in the two species with expanding ranges. Recently colonized populations at the range margin showed increased frequencies of dispersive, long-winged (macropterous) individuals, compared with longer-established populations in the range core. This increase in dispersal appeared to be short-lived because 5-10 years after colonization populations showed similar incidences of macroptery to populations in the range core. These changes are consistent with evolutionary change; field patterns persisted when nymphs were reared under controlled environmental conditions, and range margin individuals reared in the laboratory flew farther than range core individuals in a wind tunnel. There was also a reproductive trade-off with dispersal in both females and males, which could explain the rapid reversion to lower rates of dispersal once populations become established. The effect of population density on wing morphology differed between populations from the range core (no significant effect of density) and expanding range margins (negative density dependence), which we propose is part of the mechanism of the changes in dispersal. Transient changes in dispersal are likely to be common in many species undergoing range expansion and can have major population and biogeographic consequences.     

Social consequences of rapid environmental change

While direct influences of the environment on population growth and resilience are well studied, indirect routes linking environmental changes to population consequences are less explored. We suggest that social behavior is key for understanding how anthropogenic environmental changes affect the resilience of animal populations. Social structures of animal groups are evolved and emergent phenotypes that often have demographic consequences for group members. Importantly, environmental drivers may directly influence the consequences of social structure or indirectly influence them through modifications to social interactions, group composition, or group size. We have developed a framework to study these demographic consequences. Estimating the strength of direct and indirect pathways will give us tools to understand, and potentially manage, the effect of human-induced rapid environmental changes.