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.     

Predation and the evolutionary dynamics of species ranges

Gene flow that hampers local adaptation can constrain species distributions and slow invasions. Predation as an ecological factor mainly limits prey species ranges, but a richer array of possibilities arises once one accounts for how predation alters the interplay of gene flow and selection. We extend previous single-species theory on the interplay of demography, gene flow, and selection by investigating how predation modifies the coupled demographic-evolutionary dynamics of the range and habitat use of prey. We consider a model for two discrete patches and a complementary model for species along continuous environmental gradients. We show that predation can strongly influence the evolutionary stability of prey habitat specialization and range limits. Predators can permit prey to expand in habitat or geographical range or, conversely, cause range collapses. Transient increases in predation can induce shifts in prey ranges that persist even if the predator itself later becomes extinct. Whether a predator tightens or loosens evolutionary constraints on the invasion speed and ultimate size of a prey range depends on the predator effectiveness, its mobility relative to its prey, and the prey's intraspecific density dependence, as well as the magnitude of environmental heterogeneity. Our results potentially provide a novel explanation for lags and reversals in invasions.     

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.     

Theoretical models of species' borders: single species approaches

The range of potential mechanisms limiting species' distributions in space is nearly as varied and complex as the diversity of life itself. Yet viewed abstractly, a species' border is a geographic manifestation of a species' demographic responses to a spatially and temporally varying world. Population dynamic models provide insight into the different routes by which range limits can arise owing to gradients in demographic rates. In a metapopulation context, for example, range limits may be caused by gradients in extinction rates, colonization rates or habitat availability. We have consider invasion models in uniform and heterogeneous environments as a framework for understanding non-equilibrium range limits, and explore conditions under which invasions may cease to spread leaving behind a stationary range limit. We conclude that non-equilibrial range dynamics need further theoretical and empirical attention.     

Interspecific Competition, Environmental Gradients, Gene Flow, and the Coevolution of Species' Borders

Darwin viewed species range limits as chiefly determined by an interplay between the abiotic environment and interspecific interactions. Haldane argued that species' ranges could be set intraspecifically when gene flow from a species' populous center overwhelms local adaptation at the periphery. Recently, Kirkpatrick and Barton have modeled Haldane's process with a quantitative genetic model that combines density-dependent local population growth with dispersal and gene flow across a linear environmental gradient in optimum phenotype. To address Darwin's ideas, we have extended the Kirkpatrick and Barton model to include interspecific competition and the frequency-dependent selection that it generates, as well as stabilizing selection on a quantitative character. Our model includes local population growth, movements over space, natural selection, and gene flow. It simultaneously addresses the evolution of character displacement and species borders. It reproduces the Kirkpatrick and Barton single-species result that limited ranges can be produced with sufficiently steep environmental gradients and strong dispersal. Further, in the absence of environmental gradients or barriers to dispersal, interspecific competition will not limit species ranges at evolutionary equilibrium. However, interspecific competition can interact with environmental gradients and gene flow to generate limited ranges with much less extreme gradient and dispersal parameters than in the single-species case. Species display character displacement in sympatry, yet the reduction in competition that results from this displacement does not necessarily allow the two species to become sympatric everywhere. When species meet, competition reduces population densities in the region of overlap, which, in turn, intensifies the asymmetry in gene flow from center to margin. This reduces the ability of each species to adapt to local physical conditions at their range limits. If environmental gradients are monotonic but not linear, the transition zone between species at coevolutionary equilibrium occurs where the environmental gradient is steepest. If productivity gradients are also introduced into the model, then patterns similar to Rapoport's rule emerge. Interacting species respond to climate change, as it affects the optimal phenotype over space, by a combination of range shifts and local evolution in mean phenotype, while solitary species respond solely by range shifts. Finally, we compare empirical estimates for intrinsic growth rates and diffusion coefficients for several species to those needed by the single-species model to produce a stable limited range. These empirical values are generally insufficient to produce limited ranges in the model suggesting a role for interspecific interactions.     

Spatial structure and dynamics of breeding bird populations at a distribution margin,southern California

Aim Local‐scale processes at species distribution margins can affect larger‐scale distribution dynamics, but are rarely studied. The objective of this research was to elucidate the nature of distribution limits by studying the comparative structure, dynamics and environmental associations of breeding bird populations at their distribution margin. We hypothesized that climate is principally responsible for setting distribution limits, whereas biotic habitat features are more strongly associated with distribution patterns within the range. Location Southern California, USA. Methods During 2005–2007 we studied the distribution patterns of breeding birds in three study areas, each spanning a low‐elevation (200–1800 m) desert scrub‐to‐chaparral gradient. We used logistic regression with hierarchical partitioning to assess the independent effects of environmental variables (e.g. climate versus habitat) on distributions. We tested for shifts in the relative importance of these environmental variables in determining distribution limits versus within‐range patterns, and we also compared higher‐ and lower‐elevation groups of species. Results Distribution patterns were highly variable among species, but were remarkably static over the three study areas and 3‐year study period. Across species, habitat floristic variables performed relatively well at explaining distribution patterns. For higher‐elevation species (chaparral birds), climate was relatively important in setting their lower distribution limits, and there was a shift to a greater importance of biotic habitat (mainly habitat structural variables) for determining within‐range patterns. Relationships were more mixed for lower‐elevation species (desert scrub birds), but with respect to distribution limits, biotic habitat variables tended to be more important relative to climate than we observed for chaparral birds. Main conclusions Along this warm, arid elevational gradient, higher‐elevation chaparral birds are more limited by climate at their lower margin than are lower‐elevation desert birds at their upper margin, suggesting that climate plays a strong role (relative to other values) in excluding non‐desert birds from desert. However, given the strong differences among species, predictive distribution models will need to be individually tailored, and for most species biotic habitat variables were of greater importance than climate in determining limits. This research highlights the usefulness of studying environmental relationships at distribution margins and the importance of considering biotic relationships in forecasting distribution shifts under changing climates.     

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.     

Topography, energy and the global distribution of bird species richness

A major goal of ecology is to determine the causes of the latitudinal gradient in global distribution of species richness. Current evidence points to either energy availability or habitat heterogeneity as the most likely environmental drivers in terrestrial systems, but their relative importance is controversial in the absence of analyses of global (rather than continental or regional) extent. Here we use data on the global distribution of extant continental and continental island bird species to test the explanatory power of energy availability and habitat heterogeneity while simultaneously addressing issues of spatial resolution, spatial autocorrelation, geometric constraints upon species' range dynamics, and the impact of human populations and historical glacial ice-cover. At the finest resolution (1 degree), topographical variability and temperature are identified as the most important global predictors of avian species richness in multi-predictor models. Topographical variability is most important in single-predictor models, followed by productive energy. Adjusting for null expectations based on geometric constraints on species richness improves overall model fit but has negligible impact on tests of environmental predictors. Conclusions concerning the relative importance of environmental predictors of species richness cannot be extrapolated from one biogeographic realm to others or the globe. Rather a global perspective confirms the primary importance of mountain ranges in high-energy areas.     

Alternative causes for range limits: a metapopulation perspective

All species have limited distributions at broad geographical scales. At local scales, the distribution of many species is influenced by the interplay of the three factors of habitat availability, local extinctions and colonization dynamics. We use the standard Levins metapopulation model to illustrate how gradients in these three factors can generate species' range limits. We suggest that the three routes to range limits have radically different evolutionary implications. Because the Levins model makes simplifying assumptions about the spatial coupling of local populations, we present numerical studies of spatially explicit metapopulation models that complement the analytical model. The three routes to range limits give rise to distinct spatiotemporal patterns. Range limits in one species can also arise because of environmental gradients impinging upon other species. We briefly discuss a predator–prey example, which illustrates indirect routes to range limits in a metacommunity context.     

Mechanistic niche modelling: combining physiological and spatial data to predict species' ranges

Species distribution models (SDMs) use spatial environmental data to make inferences on species' range limits and habitat suitability. Conceptually, these models aim to determine and map components of a species' ecological niche through space and time, and they have become important tools in pure and applied ecology and evolutionary biology. Most approaches are correlative in that they statistically link spatial data to species distribution records. An alternative strategy is to explicitly incorporate the mechanistic links between the functional traits of organisms and their environments into SDMs. Here, we review how the principles of biophysical ecology can be used to link spatial data to the physiological responses and constraints of organisms. This provides a mechanistic view of the fundamental niche which can then be mapped to the landscape to infer range constraints. We show how physiologically based SDMs can be developed for different organisms in different environmental contexts. Mechanistic SDMs have different strengths and weaknesses to correlative approaches, and there are many exciting and unexplored prospects for integrating the two approaches. As physiological knowledge becomes better integrated into SDMs, we will make more robust predictions of range shifts in novel or non-equilibrium contexts such as invasions, translocations, climate change and evolutionary shifts.     

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).     

Species' borders and dispersal barriers

Range limits of species are determined by combined effects of physical, historical, ecological, and evolutionary forces. We consider a subset of these factors by using spatial models of competition, hybridization, and local adaptation to examine the effects of partial dispersal barriers on the locations of borders between similar species. Prompted by results from population genetic models and biogeographic observations, we investigate the conditions under which species' borders are attracted to regions of reduced dispersal. For borders maintained by competition or hybridization, we find that dispersal barriers can attract borders whose positions would otherwise be either neutrally stable or moving across space. Borders affected strongly by local adaptation and gene flow, however, are repelled from dispersal barriers. These models illustrate how particular biotic and abiotic factors may combine to limit species' ranges, and they help to elucidate mechanisms by which range limits of many species may coincide.     

Testing abundance-range size relationships in European carabid beetles (Coleoptera, Carabidae)

Four of the eight hypotheses proposed in the literature for explaining the relationship between abundance and range size (the sampling artifact, phylogenetic non-independence, range position and resource breadth hypotheses) were tested by using atlas data for carabid beetles (Coleoptera, Carabidae) from Belgium, Denmark and the Netherlands. A positive relationship between abundance and partial range size was found in all three countries, and the variation in abundance was lower for widespread species. Analysis of the data did not support three of the proposed hypotheses, but did support the resource breadth hypothesis (species having broader environmental tolerances and being able to use a wider range or resources will have higher local densities and be more widely distributed than more specialised species). Examination of species' characteristics revealed that widespread species are generally large bodied, generalists (species with wide niche breadths occurring in a variety of habitat types) and are little influenced by human-altered landscapes, while species with restricted distributions are smaller bodied, specialists (species with small niche breadths occurring in only one or two habitat types), and favour natural habitat. Landscape alteration may be an important factor influencing carabid abundance and range size in these three countries with a long history of human-induced environmental changes.     

Population genetics and the evolution of geographic range limits in an annual plant

Abstract Theoretical models of species' geographic range limits have identified both demographic and evolutionary mechanisms that prevent range expansion. Stable range limits have been paradoxical for evolutionary biologists because they represent locations where populations chronically fail to respond to selection. Distinguishing among the proposed causes of species' range limits requires insight into both current and historical population dynamics. The tools of molecular population genetics provide a window into the stability of range limits, historical demography, and rates of gene flow. Here we evaluate alternative range limit models using a multilocus data set based on DNA sequences and microsatellites along with field demographic data from the annual plant Clarkia xantiana ssp. xantiana. Our data suggest that central and peripheral populations have very large historical and current effective population sizes and that there is little evidence for population size changes or bottlenecks associated with colonization in peripheral populations. Whereas range limit populations appear to have been stable, central populations exhibit a signature of population expansion and have contributed asymmetrically to the genetic diversity of peripheral populations via migration. Overall, our results discount strictly demographic models of range limits and more strongly support evolutionary genetic models of range limits, where adaptation is prevented by a lack of genetic variation or maladaptive gene flow.     

Bioclimatic Analysis to Enhance Reintroduction Biology of the Endangered Helmeted Honeyeater (Lichenostomus melanops cassidix) in Southeastern Australia

Reintroduction programs are a high-risk conservation strategy for restoring populations of endangered species. The success of these programs often depends on the ability to identify suitable habitat within the species' former range. Bioclimatic analysis offers an empirical, explicit, robust, and repeatable method to analyze large areas rapidly using a small number of locality records, and in turn predicting (and/or reconstructing) its potential distribution limits. This approach therefore can estimate the broad limits of the distribution of a taxon, using data that may be inadequate for standard forms of statistical analysis. We illustrate the potential value of bioclimatic modeling for reintroduction biology using a case study of the highly endangered Helmeted Honeyeater ( Lichenostomus melanops cassidix ) from Victoria, southeastern Australia. The results of our analyses assisted us to both predict the former range limits of the Helmeted Honeyeater and determine the broad limits of those areas that may contain potentially suitable sites for future reintroduction programs for the subspecies. The analysis predicted that the range of the Helmeted Honeyeater extends from the Yarra River district east of Melbourne, south to the Western Port Bay and east as far as the Morwell area of Victoria. The climatic characteristics of habitat occupied by the extant population of the Helmeted Honeyeater were found to be unique within its predicted range. We recommend that reintroduction efforts therefore be concentrated within this small area, as has occurred to date.     

Climate structures genetic variation across a species' elevation range: a test of range limits hypotheses

Gene flow may influence the formation of species range limits, and yet little is known about the patterns of gene flow with respect to environmental gradients or proximity to range limits. With rapid environmental change, it is especially important to understand patterns of gene flow to inform conservation efforts. Here we investigate the species range of the selfing, annual plant, Mimulus laciniatus, in the California Sierra Nevada. We assessed genetic variation, gene flow, and population abundance across the entire elevation‐based climate range. Contrary to expectations, within‐population plant density increased towards both climate limits. Mean genetic diversity of edge populations was equivalent to central populations; however, all edge populations exhibited less genetic diversity than neighbouring interior populations. Genetic differentiation was fairly consistent and moderate among all populations, and no directional signals of contemporary gene flow were detected between central and peripheral elevations. Elevation‐driven gene flow (isolation by environment), but not isolation by distance, was found across the species range. These findings were the same towards high‐ and low‐elevation range limits and were inconsistent with two common centre‐edge hypotheses invoked for the formation of species range limits: (i) decreasing habitat quality and population size; (ii) swamping gene flow from large, central populations. This pattern demonstrates that climate, but not centre‐edge dynamics, is an important range‐wide factor structuring M. laciniatus populations. To our knowledge, this is the first empirical study to relate environmental patterns of gene flow to range limits hypotheses. Similar investigations across a wide variety of taxa and life histories are needed.     

里海西北部Caspian地区草原扩大与啮齿类群落结构的变化

于1997-1999年检测了俄联邦卡尔梅克(Kalmyki)共和国ChernieZemly半干旱草原由10个物种组成的啮齿类群落。该区域的植被特点是家畜大量减少后形成的恢复植被。我们的任务之一就是了解不同啮齿动物的生境需求,以预测不同物种对环境变化的特异性反应。我们也检测了群落空间结构、生态位参数和物种多样性。逐步回归分析表明,各种啮齿动物的空间分布仅部分地决定于已知环境因子(2-6个变量)。虽然方程式具有强显著性,但决定系数R2很低,不超过18%。对于大多数种类,这种结果可由最近草原扩大而来的生境异质性降低解释。物种生境选择条件和资源的明显低水平可能是由物种对环境变化的惯性反应造成的。判别函数分析和主分量分析结果表明,群落结构以及单个物种的生态位参数在不同年间并不稳定。空间分层结构在种间中等和低重叠的高物种多样性年度表现明显。多样性、生态位组成和生态位宽度分析结果表明,不同物种对环境条件和资源变化的反应具有不同的个性化方式。同时,物种对环境动态的明显个性化反应成为物种多样性定向变化的原因。限定空间和相对单一条件下的结果表明,啮齿类α多样性与生境结构复杂性呈正相关,而与生境生产力特征呈负相关。观察表明,卡尔梅克国草原扩大伴随着初级生产力的升高和生境异质性的降低。随着时间的推移会导致大多数草原和半干旱区域啮齿类多样性的降低,并形成动物区系核心.     

Human land use, agriculture, pesticides and losses of imperiled species

Aim  Anthropogenic habitat loss is usually cited as the most important cause of recent species' extinctions. We ask whether species losses are in fact more closely related to habitat loss than to any other aspect of human activity such as use of agricultural pesticides, or human population density (which reflects urbanization).
Location  Canada.
Methods  We statistically compared areas in Canada where imperiled species currently occur, versus areas where they have been lost. Using multiple regressions, we relate the numbers of species that had suffered range reductions in an ecoregion to variables that represent present habitat loss, pesticide use and human population density.
Results  We find high losses of imperiled species in regions with high proportions of agricultural land cover. However, losses of imperiled species are significantly more strongly related to the proportion of the region treated with agricultural pesticides. The relationship between species losses and area treated with pesticides remains significant after controlling for area in agriculture.
Main conclusions  Our results are consistent with the hypothesis that agricultural pesticide use, or something strongly collinear with it (perhaps intensive agriculture more generally), has contributed significantly to the decline of imperiled species in Canada. Habitat conversion per se may be a less important cause of species declines than how that converted habitat is used.     

Developmental stability across the breeding distribution of the scissor-tailed flycatcher (Tyrannus forficatus)

A central question in ecology is how individual fitness interacts with the spatial variation in population density and habitat characteristics across species' ranges. We used fluctuating asymmetry (FA) as a measure of developmental stability (DS) in individuals of Tyrannus forficatus to estimate the suitability of sites of varying abundance and position within the species' range. FA in the inner-tail feathers of males and females is not spatially correlated across the species' range. FA in males increases towards the centre of the range and is not correlated with abundance. FA in females is not correlated with position in the range or abundance. Our results suggest that optimal sites are found throughout the range of the species, whereas suboptimal sites are mainly found towards the centre of the range. Additionally, our results suggest that abundance may not reflect the suitability of sites across species' ranges.