Linking traits to energetics and population dynamics to predict lizard ranges in changing environments

I present a dynamic bioenergetic model that couples individual energetics and population dynamics to predict current lizard ranges and those following climate warming. The model predictions are uniquely based on first principles of morphology, life history, and thermal physiology. I apply the model to five populations of a widespread North American lizard, Sceloporus undulatus, to examine how geographic variation in traits and life histories influences ranges. This geographic variation reflects the potential for species to adapt to environmental change. I then consider the range dynamics of the closely related Sceloporus graciosus. Comparing predicted ranges and actual current ranges reveals how dispersal limitations, species interactions, and habitat requirements influence the occupied portions of thermally suitable ranges. The dynamic model predicts individualistic responses to a uniform 3 degrees C warming but a northward shift in the northern range boundary for all populations and species. In contrast to standard correlative climate envelope models, the extent of the predicted northward shift depends on organism traits and life histories. The results highlight the limitations of correlative models and the need for more dynamic models of species' ranges.     

Allee effects, invasion pinning, and species' borders

All species' ranges are the result of successful past invasions. Thus, models of species' invasions and their failure can provide insight into the formation of a species' geographic range. Here, we study the properties of invasion models when a species cannot persist below a critical population density known as an "Allee threshold." In both spatially continuous reaction-diffusion models and spatially discrete coupled ordinary-differential-equation models, the Allee effect can cause an invasion to fail. In patchy landscapes (with dynamics described by the spatially discrete model), range limits caused by propagation failure (pinning) are stable over a wide range of parameters, whereas, in an uninterrupted habitat (with dynamics described by a spatially continuous model), the zero velocity solution is structurally unstable and thus unlikely to persist in nature. We derive conditions under which invasion waves are pinned in the discrete space model and discuss their implications for spatially complex dynamics, including critical phenomena, in ecological landscapes. Our results suggest caution when interpreting abrupt range limits as stemming either from competition between species or a hard environmental limit that cannot be crossed: under a wide range of plausible ecological conditions, species' ranges may be limited by an Allee effect. Several example systems appear to fit our general model.     

Distribution-wide effects of climate on population densities of a declining migratory landbird

1. Increases in global temperatures have created concern about effects of climatic variability on populations, and climate has been shown to affect population dynamics in an increasing number of species. Testing for effects of climate on population densities across a species' distribution allows for elucidation of effects of climate that would not be apparent at smaller spatial scales. 2. Using autoregressive population models, we tested for effects of the North Atlantic Oscillation (NAO) and the El Ni?o Southern Oscillation (ENSO) on annual population densities of a North American migratory landbird, the yellow-billed cuckoo Coccyzus americanus, across the species' breeding distribution over a 37-year period (1966-2002). 3. Our results indicate that both the NAO and ENSO have affected population densities of C. americanus across much of the species' breeding range, with the strongest effects of climate in regions in which these climate systems have the strongest effects on local temperatures. Analyses also indicate that the strength of the effect of local temperatures on C. americanus populations was predictive of long-term population decline, with populations that were more negatively affected by warm temperatures experiencing steeper declines. 4. Results of this study highlight the importance of distribution-wide analyses of climatic effects and demonstrate that increases in global temperatures have the potential to lead to additional population declines.     

A method for simulating patterns of habitat availability at static and dynamic range margins

The dynamics of populations inhabiting range margins are likely to be critically important in determining the response of species to climate change. Despite this, there is a lack of both empirical and theoretical work that examines the behaviour of these populations. Populations living on the edge of a species' range frequently inhabit a more patchily distributed habitat than those that live closer to the centre of the range. This difference is likely to play an important role in determining the dynamics of range margin populations, both when the range is static and when it is dynamic, for example shifting in response to climate change. Here, we present a simple method that simulates the distribution of suitable habitat sites at the edge of a range. Habitat availability is determined as a function of both latitudinal and local environmental variability, and the relative importance of the two can be adjusted. The method is readily extended to describe shifting habitat availability during a period of climate change. We suggest that there is a need for a greater effort to examine the ecology of range margin populations, and believe that the method presented here could be of considerable use in future theoretical studies.     

Conclusions about niche expansion in introduced Impatiens walleriana populations depend on method of analysis

Determining the degree to which climate niches are conserved across plant species' native and introduced ranges is valuable to developing successful strategies to limit the introduction and spread of invasive plants, and also has important ecological and evolutionary implications. Here, we test whether climate niches differ between native and introduced populations of Impatiens walleriana, globally one of the most popular horticultural species. We use approaches based on both raw climate data associated with occurrence points and ecological niche models (ENMs) developed with Maxent. We include comparisons of climate niche breadth in both geographic and environmental spaces, taking into account differences in available habitats between the distributional areas. We find significant differences in climate envelopes between native and introduced populations when comparing raw climate variables, with introduced populations appearing to expand into wetter and cooler climates. However, analyses controlling for differences in available habitat in each region do not indicate expansion of climate niches. We therefore cannot reject the hypothesis that observed differences in climate envelopes reflect only the limited environments available within the species' native range in East Africa. Our results suggest that models built from only native range occurrence data will not provide an accurate prediction of the potential for invasiveness if applied to areas containing a greater range of environmental combinations, and that tests of niche expansion may overestimate shifts in climate niches if they do not control carefully for environmental differences between distributional areas.     

Variation in abundance across a species' range predicts climate change responses in the range interior will exceed those at the edge: a case study with North American beaver

The absence of information about how abundance varies across species' ranges restricts most modeling and monitoring of climate change responses to the range edge. We examine spatial variation in abundance across the northeastern range of North American beaver ( Castor canadensis ), evaluate the extent to which climate and nonclimate variables explain this variation, and use a species–climate envelope model that includes spatial variation in abundance to predict beaver abundance responses to projected climate change. The density of beaver colonies across Québec follows a roughly logistic pattern, with high but variable density across the southern portion of the province, a sharp decline in density at about 49°N, and a long tail of low density extending as far as 58°N. Several climate and nonclimate variables were strong predictors of variation in beaver density, but 97% of the variation explained by nonclimate variables could be accounted for by climate variables. Because of the peak and tail density pattern, beaver climate sensitivity (change in density per unit change in climate) was greatest in the interior and lowest at the edge of the range. Combining our best density–climate models with projections from general circulation models (GCM) predicts a relatively modest expansion of the species' northern range limit by 2055, but density increases in the range interior that far exceed those at the range edge. Thus, some of the most dramatic responses to climate change may be occurring in the core of species' ranges, far away from the edge-of-the-range focus of most current modeling and monitoring efforts.     

Species' borders: a unifying theme in ecology

Biologists have long been fascinated by species' borders, and with good reason. Understanding the ecological and evolutionary dynamics of species' borders may prove to be the key that unlocks new understanding across a wide range of biological phenomena. After all, geographic range limits are a point of entry into understanding the ecological niche and threshold responses to environmental change. Elucidating patterns of gene flow to, and returning from, peripheral populations can provide important insights into the nature of adaptation, speciation and coevolution. Species' borders form natural laboratories for the study of the spatial structure of species interactions. Comparative studies from the center to the margin of species' ranges allow us to explore species' demographic responses along gradients of increasing environmental stress. Range dynamics further permit investigation into invasion dynamics and represent bellwethers for a changing climate. This set of papers explores ecological and evolutionary dynamics of species' borders from diverse empirical and theoretical perspectives.     

How complex should models be? Comparing correlative and mechanistic range dynamics models

Criticism has been levelled at climate‐change‐induced forecasts of species range shifts that do not account explicitly for complex population dynamics. The relative importance of such dynamics under climate change is, however, undetermined because direct tests comparing the performance of demographic models vs. simpler ecological niche models are still lacking owing to difficulties in evaluating forecasts using real‐world data. We provide the first comparison of the skill of coupled ecological‐niche‐population models and ecological niche models in predicting documented shifts in the ranges of 20 British breeding bird species across a 40‐year period. Forecasts from models calibrated with data centred on 1970 were evaluated using data centred on 2010. We found that more complex coupled ecological‐niche‐population models (that account for dispersal and metapopulation dynamics) tend to have higher predictive accuracy in forecasting species range shifts than structurally simpler models that only account for variation in climate. However, these better forecasts are achieved only if ecological responses to climate change are simulated without static snapshots of historic land use, taken at a single point in time. In contrast, including both static land use and dynamic climate variables in simpler ecological niche models improve forecasts of observed range shifts. Despite being less skilful at predicting range changes at the grid‐cell level, ecological niche models do as well, or better, than more complex models at predicting the magnitude of relative change in range size. Therefore, ecological niche models can provide a reasonable first approximation of the magnitude of species' potential range shifts, especially when more detailed data are lacking on dispersal dynamics, demographic processes underpinning population performance, and change in land cover.     

Inclusion of local environmental conditions alters high-latitude vegetation change predictions based on bioclimatic models

Current predictions of how species will respond to climate change are typically based on coarse-grained climate surfaces utilizing bioclimate envelope modelling. However, the suitability of environmental conditions for a given species might result from a variety of factors including some unrelated to climate. To address this issue, we investigated whether the inclusion of topographical and soil information in bioclimatic envelope models would significantly alter predictions of climate change—induced fine-scale tree and shrub species range size changes at the tree-limit in subarctic Europe. Using generalized additive models and data on current climate and species distributions and three different climate scenarios for the period 2040–2069, we developed predictions of the currently suitable area and potential range size changes of seven tree and shrub species in an area of 1,100 km² at a resolution of 1-ha. The inclusion of topography and soil information increased the predictive accuracy of climate-only models for all studied species. The predicted changes in species distribution volumes were contradictory, and the predicted occurrences varied greatly depending on the model used. Our results therefore support the arguments that vegetation responses to climate change can be influenced by local environmental conditions and that attention should be paid to the combined effects of these factors. We conclude that disregarding local topography and soil conditions in bioclimatic models may result in biased projections of range expansions and the associated colonization, extinction and turnover assessments.     

Pleistocene climate, phylogeny, and climate envelope models: an integrative approach to better understand species' response to climate change

Mean annual temperature reported by the Intergovernmental Panel on Climate Change increases at least 1.1°C to 6.4°C over the next 90 years. In context, a change in climate of 6°C is approximately the difference between the mean annual temperature of the Last Glacial Maximum (LGM) and our current warm interglacial. Species have been responding to changing climate throughout Earth's history and their previous biological responses can inform our expectations for future climate change. Here we synthesize geological evidence in the form of stable oxygen isotopes, general circulation paleoclimate models, species' evolutionary relatedness, and species' geographic distributions. We use the stable oxygen isotope record to develop a series of temporally high-resolution paleoclimate reconstructions spanning the Middle Pleistocene to Recent, which we use to map ancestral climatic envelope reconstructions for North American rattlesnakes. A simple linear interpolation between current climate and a general circulation paleoclimate model of the LGM using stable oxygen isotope ratios provides good estimates of paleoclimate at other time periods. We use geologically informed rates of change derived from these reconstructions to predict magnitudes and rates of change in species' suitable habitat over the next century. Our approach to modeling the past suitable habitat of species is general and can be adopted by others. We use multiple lines of evidence of past climate (isotopes and climate models), phylogenetic topology (to correct the models for long-term changes in the suitable habitat of a species), and the fossil record, however sparse, to cross check the models. Our models indicate the annual rate of displacement in a clade of rattlesnakes over the next century will be 2 to 3 orders of magnitude greater (430-2,420 m/yr) than it has been on average for the past 320 ky (2.3 m/yr).     

Spatial-temporal population dynamics across species range: from centre to margin

Understanding the boundaries of species' ranges and the variations in population dynamics from the centre to margin of a species' range is critical. This study simulated spatial-temporal patterns of birth and death rates and migration across a species' range in different seasons. Our results demonstrated the importance of dispersal and migration in altering birth and death rates, balancing source and sink habitats, and governing expansion or contraction of species' ranges in changing environments. We also showed that the multiple equilibria of metapopulations across a species' range could be easily broken following climatic changes or physical disturbances either local or regional. Although we refer to our models as describing the population dynamics across whole species' range, they should also apply to small-scale habitats (metapopulations) in which species abundance follows a humped pattern or to any ecosystem or landscape where strong central-marginal (C-M) environmental gradients exist. Conservation of both central and marginal populations would therefore be equally important considerations in making management decisions.     

The response of two butterfly species to climatic variation at the edge of their range and the implications for poleward range shifts

To predict changes in species' distributions due to climate change we must understand populations at the poleward edge of species' ranges. Ecologists generally expect range shifts under climate change caused by the expansion of edge populations as peripheral conditions increasingly resemble the range core. We tested whether peripheral populations of two contrasting butterflies, a small-bodied specialist (Erynnis propertius) and a large-bodied generalist (Papilio zelicaon), respond favorably to warmer conditions. Performance of populations related to climate was evaluated in seven peripheral populations spanning 1.2 degrees latitude (160 km) using: (1) population density surveys, an indirect measure of site suitability; and (2) organismal fitness in translocation experiments. There was evidence that population density increased with temperature for P. zelicaon whose population density declined with latitude in 1 of 3 sample years. On the other hand, E. propertius showed a positive relationship of population density with latitude, apparently unrelated to climate or measured habitat variables. Translocation experiments showed increased larval production at increased temperatures for both species, and in P. zelicaon, larval production also increased under drier conditions. These findings suggest that both species may increase at their range edge with warming but the preference for core-like conditions may be stronger in P. zelicaon. Further, populations of E. propertius at the range boundary may be large enough to act as sources of colonists for range expansions, but range expansion in this species may be prevented by a lack of available host plants further north. In total, the species appear to respond differently to climate and other factors that vary latitudinally, factors that will likely affect poleward expansion.     

Individualistic vs community modelling of species distributions under climate change

Studies investigating the consequences of future climate changes on species distributions usually start with the assumption that species respond to climate changes in an individualistic fashion. This assumption has led researchers to use bioclimate envelope models that use present climate-range relationships to characterize species' limits of tolerance to climate, and then apply climate-change scenarios to enable projections of altered species distributions. However, there are techniques that combine climate variables together with information on the composition of assemblages to enable projections that are expected to mimic community dynamics. Here, we compare, for the first time, the performance of GLM (generalized linear model) and CQO (canonical quadratic ordination; a type of community-based GLM) for projecting distributions of species under climate change scenarios. We found that projections from these two methods varied both in terms of accuracy (GLM providing generally more accurate projections than CQO) and in the broad diversity patterns yielded (higher species richness values projected with CQO). Model outputs were also affected by species-specific traits, such as species range size and species geographical positions, supporting the view that methods are sensitive to different degrees of equilibrium of species distributions with climate. This study reveals differences in projections between individual- and community-based approaches that require further scrutiny, but it does not find support for unsupervised use community-based models for investigating climate change impacts on species distributions. Reasons for this lack of support are discussed.     

Performance of climate envelope models in retrodicting recent changes in bird population size from observed climatic change

Twenty-five-year population trends of 42 bird species rare as breeders in the UK were examined in relation to changes in climatic suitability simulated using climatic envelope models. The effects of a series of potential 'nuisance' variables were also assessed. A statistically significant positive correlation was found across species between population trend and climate suitability trend. The demonstration that climate envelope models are able to retrodict species' population trends provides a valuable validation of their use in studies of the potential impacts of future climatic changes.     

Time as an ecological constraint

Conventional approaches to population biology emphasise the roles of climatic conditions, nutrient flow and predation as constraints on population dynamics. We argue here that this focus has obscured the role of time as a crucial constraint on species' abilities to survive in some habitats. Time constraints may be particularly intrusive both for species that live in intensely bonded groups (where the need to devote time to social interaction may ultimately limit the size of group that a species can maintain in a particular habitat) and for taxa that face constraints on the length of the active day. We use a linear programming approach that allows us to specify both how time allocations to different activities are influenced by local environmental and climatic variables and how these in turn limit group size and population density. The linear programming approach identifies the realizable niche space within which a species can maintain coherent groups that are larger than the minimum viable group size (or density). This approach thus allow us to understand better why a given taxon can survive in some habitats but not others, as well as the demographic stress that a population may face. In addition, they also allow us to evaluate the implications of both past and future climate change for a taxon's ability to cope with particular habitats.     

Genetic differentiation across a latitudinal gradient in two co-occurring butterfly species: revealing population differences in a context of climate change

Genetic differentiation within a species' range is determined by natural selection, genetic drift, and gene flow. Selection and drift enhance genetic differences if populations are sufficiently isolated, while gene flow precludes differentiation and local adaptation. Over large geographical areas, these processes can create a variety of scenarios, ranging from admixture to a high degree of population differentiation. Genetic differences among populations may signal functional differences within a species' range, potentially leading to population or ecotype-specific responses to global change. We investigated differentiation within the geographical range of two butterfly species along a broad latitudinal gradient. This gradient is the primary axis of climatic variation, and many ecologists expect populations at the poleward edge of this gradient to expand under climate change. Our study species inhabit a shared ecosystem and differ in body size and resource specialization; both also find their poleward range limit on an island. We find evidence for divergence of peripheral populations from the core in both taxa, suggesting the potential for genetic distinctiveness at the leading edge of climate change. We also find differences between the species in the extent of peripheral differentiation with the smaller and more specialized species showing greater population divergence (microsatellites and mtDNA) and reduced gene flow (mtDNA). Finally, gene flow estimates in both species differed strongly between two marker types. These findings suggest caution in assuming that populations are invariant across latitude and thus will respond as a single ecotype to climatic change.     

The frequency of fitness peak shifts is increased at expanding range margins due to mutation surfing

Dynamic species' ranges, those that are either invasive or shifting in response to environmental change, are the focus of much recent interest in ecology, evolution, and genetics. Understanding how range expansions can shape evolutionary trajectories requires the consideration of nonneutral variability and genetic architecture, yet the majority of empirical and theoretical work to date has explored patterns of neutral variability. Here we use forward computer simulations of population growth, dispersal, and mutation to explore how range-shifting dynamics can influence evolution on rugged fitness landscapes. We employ a two-locus model, incorporating sign epistasis, and find that there is an increased likelihood of fitness peak shifts during a period of range expansion. Maladapted valley genotypes can accumulate at an expanding range front through a phenomenon called mutation surfing, which increases the likelihood that a mutation leading to a higher peak will occur. Our results indicate that most peak shifts occur close to the expanding front. We also demonstrate that periods of range shifting are especially important for peak shifting in species with narrow geographic distributions. Our results imply that trajectories on rugged fitness landscapes can be modified substantially when ranges are dynamic.     

Demographic consequences of climate change and land cover help explain a history of extirpations and range contraction in a declining snake species

Developing conservation strategies for threatened species increasingly requires understanding vulnerabilities to climate change, in terms of both demographic sensitivities to climatic and other environmental factors, and exposure to variability in those factors over time and space. We conducted a range‐wide, spatially explicit climate change vulnerability assessment for Eastern Massasauga (Sistrurus catenatus), a declining endemic species in a region showing strong environmental change. Using active season and winter adult survival estimates derived from 17 data sets throughout the species' range, we identified demographic sensitivities to winter drought, maximum precipitation during the summer, and the proportion of the surrounding landscape dominated by agricultural and urban land cover. Each of these factors was negatively associated with active season adult survival rates in binomial generalized linear models. We then used these relationships to back‐cast adult survival with dynamic climate variables from 1950 to 2008 using spatially explicit demographic models. Demographic models for 189 population locations predicted known extant and extirpated populations well (AUC = 0.75), and models based on climate and land cover variables were superior to models incorporating either of those effects independently. These results suggest that increasing frequencies and severities of extreme events, including drought and flooding, have been important drivers of the long‐term spatiotemporal variation in a demographic rate. We provide evidence that this variation reflects nonadaptive sensitivity to climatic stressors, which are contributing to long‐term demographic decline and range contraction for a species of high‐conservation concern. Range‐wide demographic modeling facilitated an understanding of spatial shifts in climatic suitability and exposure, allowing the identification of important climate refugia for a dispersal‐limited species. Climate change vulnerability assessment provides a framework for linking demographic and distributional dynamics to environmental change, and can thereby provide unique information for conservation planning and management.     

A changing climate is eroding the geographical range of the Namib Desert tree Aloe through population declines and dispersal lags

While poleward species migration in response to recent climatic warming is widely documented, few studies have examined entire range responses of broadly distributed sessile organisms, including changes on both the trailing (equatorward) and the leading (poleward) range edges. From a detailed population census throughout the entire geographical range of Aloe dichotoma Masson, a long-lived Namib Desert tree, together with data from repeat photographs, we present strong evidence that a developing range shift in this species is a 'fingerprint' of anthropogenic climate change. This is explained at a high level of statistical significance by population level impacts of observed regional warming and resulting water balance constraints. Generalized linear models suggest that greater mortalities and population declines in equatorward populations are virtually certainly the result, due to anthropogenic climate change, of the progressive exceedance of critical climate thresholds that are relatively closer to the species' tolerance limits in equatorward sites. Equatorward population declines are also broadly consistent with bioclimatically modelled projections under anticipated anthropogenic climate change but, as yet, there is no evidence of poleward range expansion into the area predicted to become suitable in future, despite good evidence for positive population growth trends in poleward populations. This study is among the first to show a marked lag between trailing edge population extinction and leading edge range expansion in a species experiencing anthropogenic climate change impacts, a pattern likely to apply to most sessile and poorly dispersed organisms. This provides support for conservative assumptions of species' migration rates when modelling climate change impacts for such species. Aloe dichotoma 's response to climate change suggests that desert ecosystems may be more sensitive to climate change than previously suspected.     

Climate and the range dynamics of species with imperfect detection

Reliable predictions for species range changes require a mechanistic understanding of range dynamics in relation to environmental variation. One obstacle is that most current models are static and confound occurrence with the probability of detecting a species if it occurs at a site. Here we draw attention to recently developed occupancy models, which can be used to examine colonization and local extinction or changes in occupancy over time. These models further account for detection probabilities, which are likely to vary spatially and temporally in many datasets. Occupancy models require repeated presence/absence surveys, for example checklists used in bird atlas projects. As an example, we examine the recent range expansion of hadeda ibises (Bostrychia hagedash) in South African protected areas. Colonization exceeded local extinction in most biomes, and the probability of occurrence was related to local climate. Extensions of the basic occupancy models can estimate abundance or species richness. Occupancy models are an appealing additional tool for studying species' responses to global change.