Range and niche shifts in response to past climate change in the desert horned lizard Phrynosoma platyrhinos

During climate change, species are often assumed to shift their geographic distributions (geographic ranges) in order to track environmental conditions – niches – to which they are adapted. Recent work, however, suggests that the niches do not always remain conserved during climate change but shift instead, allowing populations to persist in place or expand into new areas. We assessed the extent of range and niche shifts in response to the warming climate after the Last Glacial Maximum (LGM) in the desert horned lizard Phrynosoma platyrhinos, a species occupying the western deserts of North America. We used a phylogeographic approach with mitochondrial DNA sequences to approximate the species range during the LGM by identifying populations that exhibit a genetic signal of population stability versus those that exhibit a signal of a recent (likely post‐LGM) geographic expansion. We then compared the climatic niche that the species occupies today with the niche it occupied during the LGM using two models of simulated LGM climate. The genetic analyses indicated that P. platyrhinos persisted within the southern Mojave and Sonoran deserts throughout the latest glacial period and expanded from these deserts northwards, into the western and eastern Great Basin, after the LGM. The climatic niche comparisons revealed that P. platyrhinos expanded its climatic niche after the LGM towards novel, warmer and drier climates that allowed it to persist within the southern deserts. Simultaneously, the species shifted its climatic niche towards greater temperature and precipitation fluctuations after the LGM. We concluded that climatic changes at the end of the LGM promoted both range and niche shifts in this lizard. The mechanism that allowed the species to shift its niche remains unknown, but phenotypic plasticity likely contributes to the species ability to adjust to climate change.     

Effect of geographic background and equilibrium state on niche model transferability: predicting areas of invasion of Leptoglossus occidentalis

Niche conservatism providing support for using ecological niche modeling in biological invasions has been widely noticed, however, the equilibrium state and geographic background effect on niche model transferability has received scant attention. The western conifer seed bug, Leptoglossus occidentalis, native to western North America, has expanded its range eastward and has become an invasive pest in Europe and Asia. Niche models calibrated on the ranges of a small native population and two large expanding populations were compared. We found that the climate niche of L. occidentalis is conserved during its steady expansion in North America and rapid spread in Europe. Models based on the small western native range successfully captured the eastern expanding and introduced European populations, whereas the large area-based models varied with the presumed state of equilibrium. The equilibrium state based model succeeded but the non-equilibrium based model failed to predict the range in Europe. Our study estimates global invasion risk zones for L. occidentalis and suggests that, based on niche conservatism, modeling based on a reasonable geographic distribution at a climatic equilibrium of a species could guarantee the transferability of niche model prediction. Caution is warranted in interpreting low niche model transferability with niche differentiation and forwarding message for management strategy.     

Ecological niches as stable distributional constraints on mammal species, with implications for Pleistocene extinctions and climate change projections for biodiversity

Aim Theoretical work suggests that species’ ecological niches should remain relatively constant over long‐term ecological time periods, but empirical tests are few. We present longitudinal studies of 23 extant mammal species, modelling ecological niches and predicting geographical distributions reciprocally between the Last Glacial Maximum and present to test this evolutionary conservatism. Location This study covered distributional shifts in mammal species across the lower 48 states of the United States. Methods We used a machine‐learning tool for modelling species’ ecological niches, based on known occurrences and electronic maps summarizing ecological dimensions, to assess the ability of ecological niches as modelled in one time period to predict the geographical distribution of the species in another period, and vice versa. Results High intertemporal predictivity between niche models and species’ occurrences indicate that niche conservatism is widespread among the taxa studied, particularly when statistical power is considered as a reason for failure of reciprocal predictions. Niche projections to the present for 8 mammal taxa that became extinct at the end of the Pleistocene generally increased in area, and thus do not support the hypothesis of niche collapse as a major driving force in their extinction. Main conclusions Ecological niches represent long‐term stable constraints on the distributional potential of species; indeed, this study suggests that mammal species have tracked consistent climate profiles throughout the drastic climate change events that marked the end of the Pleistocene glaciations. Many current modelling efforts focusing on anticipating climate change effects on species’ potential geographical distributions will be bolstered by this result — in essence, the first longitudinal demonstration of niche conservatism.     

Phylogeography and ecological niche modelling of the New Zealand stick insect Clitarchus hookeri (White) support survival in multiple coastal refugia

Aim Increasing our understanding of the effects of the Last Glacial Maximum (LGM) and determining the location of refugia requires studies on widely distributed species with dense sampling of populations. We have reconstructed the biogeographic history of Clitarchus hookeri (White), a widespread species of New Zealand stick insect that exhibits geographic parthenogenesis, using phylogeographic analysis and ecological niche modelling. Location New Zealand. Methods We used DNA sequence data from the mitochondrial cytochrome c oxidase subunit I gene to reconstruct phylogenetic relationships among haplotypes from C. hookeri and two undescribed Clitarchus species. We also used distribution data from our own field surveys and museum records to reconstruct the geographic distribution of C. hookeri during the present and the LGM, using ecological niche modelling. Results The ecological niche models showed that the geographic distribution of C. hookeri has expanded dramatically since the LGM. Our model predicted large areas of suitable LGM habitat in upper North Island, and small patches along the east coast of South Island. The phylogeographic analysis shows that populations in the northern half of North Island contain much higher levels of genetic variation than those from southern North Island and South Island, and is congruent with the ecological niche model. The distribution of bisexual populations is also non-random, with males completely absent from South Island and very rare in southern North Island. Main conclusions During the LGM C. hookeri was most likely restricted to several refugia in upper North Island and one or more smaller refugia along the east coast of South Island. The unisexual populations predominate in post-glacial landscapes and are clearly favoured in the recolonization of such areas. Our study exemplifies the utility of integrating ecological niche modelling and phylogeographic analysis.     

Wild Felid Range Shift Due to Climatic Constraints in the Americas: a Bottleneck Explanation for Extinct Felids?

Theoretical and empirical evidence suggests that the ecological niche of species tends to be conservative over evolutionary time in many taxonomic groups, thus representing long-term stable constraints on species geographic distributions. Using an ecological niche modeling approach, we assessed the impact of climatic change on wild felid species potential range shifts over the last 130 K years in the Americas and the potential of such shifts as an extinction driver. We found a significant range shift for most species (both living and extinct) across their distributions driven by large-scale environmental changes. Proportionally, the most drastic range increase for all species occurred in the Last Glacial Maximum (LGM: 18 K years)–Current transition, while for the Last Inter-Glacial (LIG: 130 K years)–LGM transition an important range reduction occurred, which was larger for extinct North American species. In conclusion, the reduction of climatically suitable areas for many species in the transition LIG–LGM may have produced population reductions, which, in turn, may have played an important role in species’ extinction throughout the continent.     

Ecological niche conservatism in North American freshwater fishes

There are many hypotheses of relationships, and also of speciation processes, in North American freshwater fishes, although, to date, there have been no direct tests of whether there is evidence of ecological niche conservatism. In the present study, ecological niche modeling is used to look for evidence of ecological niche conservatism in six clades of freshwater fishes: the starheaded topminnows, sand darters, black basses, Notropis rubellus species group, Notropis longirostris species group, and the Hybopsis amblops species group. This is achieved by evaluating the reciprocal predictivity of distributional predictions based on ecological niche models developed for each individual taxon in a clade under the assumption that high reciprical predictivity between sister species can be taken as evidence of niche conservatism. Omission percentages, total and average commission, and the area under the curve in a receiver operating characteristic analysis, where calculated, are used to evaluate predictive ability. Occurrence data for each species were subset into a training and independent validation data set where possible. Across all clades and species, models predicted the validation data for a given species well. Ecological niche conservatism was found generally across the starheaded topminnows, the sand darters, and the N. longirostris species group. There was some inter-predictivity within the N. rubellus group, but almost no inter-predictivity within the black basses, indicating a lack of conservatism. These results demonstrate that ecological niches generally act as stable constraints on freshwater fish distributions in North America.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 282–295.     

Uncommon paleodistribution patterns of Chrysolophus pheasants in east Asia: explanations and implications

Some modeling studies indicated that the past distributions of species in east Asia during the Last Interglacial (LIG) and Last Glacial Maximum (LGM) periods differ from those of European and North American species and the deviant Asian distribution pattern is known under the term ‘pre‐LGM expansion’. It represents the unusually similar distribution patterns between the current and the LGM scenario. However, there is still no satisfying explanation for this phenomenon so far. Therefore, we took the two recently separated pheasant species of genus Chrysolophus in east Asia as an example to test the pattern by performing ecological niche models. The main findings of this study include: 1) the paleodistributions of these two pheasants also corresponded to the ‘pre‐LGM expansion’ pattern; 2) climatic similarity results from mobility‐oriented parity analysis also revealed similar pattern for both species; 3) climate regimes of east Asia showed patterns different from those in Europe and North America in a climate shift towards drier conditions and stronger seasonality and to more extreme temperatures of the coldest months particularly during the LIG; 4) the two Chrysolophus species occupied significantly different ecological niches according to current distribution. We suggest that ecological segregation established in allopatric glacial refugia should be the main determinants for the separation of two Chrysolophus species until they came into extant post‐Pleistocene contact.     

Using ecological niche modelling to evaluate niche stability in deep time

Aim To investigate relative niche stability in species responses to various types of environmental pressure (biotic and abiotic) on geological time‐scales using the fossil record. Location The case study focuses on Late Ordovician articulate brachiopods of the Cincinnati Arch in eastern North America. Methods Species niches were modelled for a suite of fossil brachiopod species based on five environmental variables inferred from sedimentary parameters using GARP and Maxent . Niche stability was assessed by comparison of (1) the degree of overlap of species distribution models developed for a time‐slice and those generated by projecting niche models of the previous time‐slice onto environmental layers of a second time‐slice using GARP and Maxent , (2) Schoener’s D statistic, and (3) the similarity of the contribution of each environmental parameter within Maxent niche models between adjacent time‐slices. Results Late Ordovician brachiopod species conserved their niches with high fidelity during intervals of gradual environmental change but responded to inter‐basinal species invasions through niche evolution. Both native and invasive species exhibited similar levels of niche evolution in the invasion and post‐invasion intervals. Niche evolution was related mostly to decreased variance within the former ecological niche parameters rather than to shifts to new ecospace. Main conclusions Although the species examined exhibited morphological stasis during the study interval, high levels of niche conservatism were observed only during intervals of gradual environmental change. Rapid environmental change, notably inter‐basinal species invasions, resulted in high levels of niche evolution among the focal taxa. Both native and invasive species responded with similar levels of niche evolution during the invasion interval and subsequent environmental reorganization. The assumption of complete niche conservatism frequently employed in ecological niche modelling (ENM) analyses to forecast or hindcast species geographical distributions is more likely to be accurate for climate change studies than for invasive species analyses over geological time‐scales.     

A comparative analysis reveals little evidence for niche conservatism in aquatic macrophytes among four areas on two continents

One of the most intriguing questions in current ecology is the extent to which the ecological niches of species are conserved in space and time. Niche conservatism has mostly been studied using coarse‐scale data of species' distributions, although it is at the local habitat scales where species' responses to ecological variables primarily take place. We investigated the extent to which niches of aquatic macrophytes are conserved among four study regions (i.e. Finland, Sweden and the US states of Minnesota and Wisconsin) on two continents (i.e. Europe and North America) using data for 11 species common to all the four study areas. We studied how ecological variables (i.e. local, climate and spatial variables) explain variation in the distributions of these common species in the four areas using species distribution modelling. In addition, we examined whether species' niche parameters vary among the study regions. Our results revealed large variation in both species' responses to the studied ecological variables and in species' niche parameters among the areas. We found little evidence for niche conservatism in aquatic macrophytes, though local environmental conditions among the studied areas were largely similar. This suggests that niche shifts, rather than different environmental conditions, were responsible for variable responses of aquatic macrophytes to local ecological variables. Local habitat niches of aquatic macrophytes are mainly driven by variations in local environmental conditions, whereas their climate niches are more or less conserved among regions. This highlights the need to study niche conservatism using local‐scale data to better understand whether species' niches are conserved, because different niches (e.g. local versus climate) operating at various scales may show different degrees of conservatism. The extent to which species' niches are truly conserved has wide practical implications, including for instance, predicting changes in species' distributions in response to global change.     

Inferring the past to predict the future: climate modelling predictions and phylogeography for the freshwater gastropod Radix balthica (Pulmonata, Basommatophora)

Understanding the impact of past climatic events on species may facilitate predictions of how species will respond to future climate change. To this end, we sampled populations of the common pond snail Radix balthica over the entire species range (northwestern Europe). Using a recently developed analytical framework that employs ecological niche modelling to obtain hypotheses that are subsequently tested with statistical phylogeography, we inferred the range dynamics of R. balthica over time. A Maxent modelling for present-day conditions was performed to infer the climate envelope for the species, and the modelled niche was used to hindcast climatically suitable range at the last glacial maximum (LGM) c . 21 000 years ago. Ecological niche modelling predicted two suitable areas at the LGM within the present species range. Phylogeographic model selection on a COI mitochondrial DNA data set confirmed that R. balthica most likely spread from these two disjunct refuges after the LGM. The match observed between the potential range of the species at the LGM given its present climatic requirements and the phylogeographically inferred refugial areas was a clear argument in favour of niche conservatism in R. balthica , thus allowing to predict the future range. The subsequent projection of the potential range under a global change scenario predicts a moderate pole-ward shift of the northern range limits, but a dramatic loss of areas currently occupied in France, western Great Britain and southern Germany.     

Ecological differentiation among key plant mutualists from a cryptic ant guild

As key dispersers of herbaceous seeds, Aphaenogaster ants strongly influence the distribution of woodland plants in eastern North America. Ants within this genus are difficult to distinguish and often are identified by subgroup, but emerging research suggests they occupy species-specific ecological niches. As such, distinct climatic requirements among Aphaenogaster spp. might result in transient plant interactions with climate change. We examine whether there are ecological and distributional differences among Aphaenogaster species that coincide with current taxonomic differentiations. We use occurrence records for six Aphaenogaster spp. that occur in deciduous forests in eastern North America. We associate the geographic patterning of species occurrence with temperature and precipitation data, and we examine whether unique climatic niches characterize each species. We then predict habitat suitability throughout eastern North America using species distribution models. For verification, we test how well the predicted ranges fit observed occurrences using novel data sets for each species. We find that Aphaenogaster species within this cryptic genus demonstrate unique ecological and geographic signatures. Each species within the subgroup generally responds differently to temperature, and somewhat differently to precipitation and seasonal variance, suggesting unique ecological niches for each species. Our results indicate that each ant species may respond uniquely to changes in climate. Such shifts could disrupt current community associations and biotic interactions with ant-dispersed plants.     

Climatic niche conservatism and the evolutionary dynamics in species range boundaries: global congruence across mammals and amphibians

Aim Comparative evidence for phylogenetic niche conservatism – the tendency for lineages to retain their ancestral niches over long time scales – has so far been mixed, depending on spatial and taxonomic scale. We quantify and compare conservatism in the climatic factors defining range boundaries in extant continental mammals and amphibians in order to identify those factors that are most evolutionarily conserved, and thus hypothesized to have played a major role in determining the geographic distributions of many species. We also test whether amphibians show stronger signals of climatic niche conservatism, as expected from their greater physiological sensitivity and lower dispersal abilities. Location Global; continental land masses excluding Antarctica. Methods We used nearly complete global distributional databases to estimate the climatic niche conservatism in extant continental mammals and amphibians. We characterized the climatic niche of each species by using a suite of variables and separately investigate conservatism in each variable using both taxonomic and phylogenetic approaches. Finally, we explored the spatial, taxonomic and phylogenetic patterns in recent climatic niche evolution. Results Amphibians and mammals showed congruent patterns of conservatism in cold tolerance, with assemblages of escapee species (i.e. those escaping most from the climatic constraints of their ancestors) aggregated in the North Temperate Zone. Main conclusions The relative strength of climatic niche conservatism varies across the variables tested, but is strongest for cold tolerance in both mammals and amphibians. Despite the apparent conservatism in this variable, there is also a strong signal of recent evolutionary shifts in cold tolerance in assemblages inhabiting the North Temperate Zone. Our results thus indicate that distribution patterns of both taxa are influenced by both niche conservatism and niche evolution.     

Diversification in North American arid lands: Niche conservatism, divergence and expansion of habitat explain speciation in the genus Ephedra

A lineage of 12 arid land shrubby species in the gymnosperm genus Ephedra (Gnetales) from North America is used to evaluate the influence of climate on speciation. With a long evolutionary history, and a well documented fossil record this lineage is an ideal model for understanding the process of speciation under a niche conservatism scenario. Using seven DNA molecular markers, Bayesian inference is carried out to uncover sister species and to estimate time of divergence of the lineages. Ecological niche models are generated for four parapatric and sympatric sister species and two analyses of niche evolution are performed, one based on ecological niche models and another using raw data and multivariate analysis. As previous analyses suggest, the diversification of North America Ephedra species may be the result of a recent secondary radiation. Both parapatric and sympatric species diverged mostly in a scenario of climatic niche conservatism. However, we also found strong evidence for niche divergence for one of the sister species pairs (E. californica-E. trifurca). Moreover, the multivariate analysis found environmental differences for some variables between sister species. The estimated divergence time of three pairs of sister species distributed in southwestern North America (E. cutleri-E. aspera, E. californica-E. trifurca and E. torreyana-E. viridis) is inferred to have occurred in the Late Miocene to Pliocene and for the sister species pair E. antisyphilitica-E. coryi distributed in the southern United States and northeastern Mexico, it was inferred from the Pliocene to Pleistocene. The orogenetic and climatic changes documented for these regions related to expansion of arid lands, may have contributed to the diversification in North American Ephedra, rather than adaptations to new climatic conditions.     

Continental-scale tree population response to rapid climate change, competition and disturbance

Aim  Using a new approach to analyse fossil pollen data, we investigate temporal and spatial patterns in Populus ( poplar, cottonwood, aspen) from the Late Glacial to the present at regional to continental scales.
Location  North America.
Methods  We extracted data on the timing and magnitude of the maximum value of Populus pollen from each pollen diagram in the North American Pollen Database (NAPD). The information was plotted in histograms of 150-year bins to identify times when Populus was abundant on the landscape. We also mapped the maximum values to identify spatial patterns and their causes.
Results  Our analyses show that there have been several periods since the Late Glacial when Populus was abundant on the landscape: (1) from 12.35 to 12.65 kyr  bp , in eastern North America, largely in response to the opening of the forest following the onset of the Younger Dryas; (2) from 10.85 to 11.75 kyr  bp , following the termination of the Younger Dryas; and (3) during the last 150 years, as land was cleared for agricultural use, especially in the midwestern United States.
Main conclusion  Since the Late Glacial, changes in the abundance of Populus were caused more by the effects of abrupt climate change on its major competitors, rather than the direct effects of climate on Populus itself.     

Niche shift versus niche conservatism? Climatic characteristics of the native and invasive ranges of the Mediterranean house gecko (Hemidactylus turcicus)

Aim The use of ecological niche models (ENMs) to predict potential distributions of species is steadily increasing. A necessary assumption is that climatic niches are conservative, but recent findings of niche shifts during biological invasion indicate that this assumption is not always valid. Selection of predictor variables may be one reason for the observed shifts. In this paper we assess differences in climatic niches in the native and invaded ranges of the Mediterranean house gecko (Hemidactylus turcicus) in terms of commonly applied climate variables in ENMs. We analyse which variables are more conserved versus relaxed (i.e. subject to niche shift). Furthermore, we study the predictive power of different sets of climate variables. Location The Mediterranean region and North America. Methods We developed models using Maxent and various subsets of variables out of 19 bioclimatic layers including: (1) two subsets comprising almost all variables excluding only highly collinear ones; (2) two subsets with minimalistic variable sets of water availability and energy measures; (3) two subsets focused on temperature‐related parameters; (4) two subsets with precipitation‐related parameters; and (5) one subset comprising variables combining temperature and precipitation characteristics. Occurrence data from the native Mediterranean range were used to predict the potential introduced range in North America and vice versa. Degrees of niche similarity and conservatism were assessed using both Schoener's index and Hellinger distances. The significance of the results was tested using null models. Results The degree of niche similarity and conservatism varied greatly among the predictors and variable sets applied. Shifts observed in some variables could be attributed to active habitat selection while others apparently reflected background effects. Main conclusions The study was based on comprehensive occurrence data from all regions where Hemidactylus turcicus is present in Europe and North America, providing a robust foundation. Our results clearly indicate that the degree of conservatism of niches in H. turcicus largely varies among predictors and variable sets applied. Therefore, the extent of niche conservatism of variables applied should always be tested in ENMs. This has an important impact on studies of biological invasion, impacts of climate change and niche evolution.     

Niche shifting in response to warming climate after the last glacial maximum: inference from genetic data and niche assessments in the chisel‐toothed kangaroo rat (Dipodomys microps)

During Pleistocene glacial‐interglacial cycles, the geographic range is often assumed to have shifted as a species tracks its climatic niche. Alternatively, the geographic range would not necessarily shift if a species can adapt in situ to a changing environment. The potential for a species to persist in place might increase with the diversity of habitat types that a species exploits. We evaluate evidence for either range shift or range stability between the last glacial maximum (LGM) and present time in the chisel‐toothed kangaroo rat (Dipodomys microps), an endemic of the Great Basin and Mojave deserts. We modeled how the species’ range would have changed if the climatic niche of the species remained conserved between the LGM and present time. The climatic models imply that if D. microps inhabited the same climatic niche during the LGM as it does today, the species would have persisted primarily within the warm Mojave Desert and expanded northwards into the cold Great Basin only after the LGM. Contrary to the climatic models, the mitochondrial DNA assessment revealed signals of population persistence within the current distribution of the species throughout at least the latest glacial‐interglacial cycle. We concluded that D. microps did not track its climatic niche during late Pleistocene oscillations, but rather met the challenge of a changing environment by shifting its niche and retaining large portions of its distribution. We speculate that this kind of response to fluctuating climate was possible because of ‘niche drifting’, an alteration of the species’ realized niche due to plasticity in various biological characters. Our study provides an example of an approach to reconstruct species’ responses to past climatic changes that can be used to evaluate whether and to what extent taxa have capacity to shift their niches in response to the changing environment – information becoming increasingly important to predicting biotic responses to future environmental changes.     

Pollen–plant–climate relationships in sub-Saharan Africa

Aim  To demonstrate that incorporating the bioclimatic range of possible contributor plants leads to improved accuracy in interpreting the palaeoclimatic record of taxonomically complex pollen types.
Location  North Tropical Africa.
Methods  The geographical ranges of selected African plants were extracted from the literature and geo-referenced. These plant ranges were compared with the pollen percentages obtained from a network of surface sediments. Climate-response surfaces were graphed for each pollen taxon and each corresponding plant species.
Results  Several patterns can be identified, including taxa for which the pollen and plant distributions coincide, and others where the range limits diverge. Some pollen types display a reduced climate range compared with that of the corresponding plant species, due to low pollen production and/or dispersal. For other taxa, corresponding to high pollen producers such as pioneer taxa, pollen types display a larger climatic envelope than that of the corresponding plants. The number of species contained in a pollen taxon is an important factor, as the botanical species included in a taxon may have different geographical and climate distributions.
Main conclusions  The comparison between pollen and plant distributions is an essential step towards more precise vegetation and climate reconstructions in Africa, as it identifies taxa that have a high correspondence between pollen and plant distribution patterns. Our method is a useful tool to reassess biome reconstructions in Africa and to characterize accurately the vegetation and climate conditions at a regional scale, from pollen data.     

Evaluating niche changes during invasion with seasonal models in Capsella bursa-pastoris

Premise

Researchers often use ecological niche models to predict where species might establish and persist under future or novel climate conditions. However, these predictive methods assume species have stable niches across time and space. Furthermore, ignoring the time of occurrence data can obscure important information about species reproduction and ultimately fitness. Here, we assess compare ecological niche models generated from full-year averages to seasonal models.

Methods

In this study, we generate full-year and monthly ecological niche models for Capsella bursa-pastoris in Europe and North America to see if we can detect changes in the seasonal niche of the species after long-distance dispersal.

Results

We find full-year ecological niche models have low transferability across continents and there are continental differences in the climate conditions that influence the distribution of C. bursa-pastoris. Monthly models have greater predictive accuracy than full-year models in cooler seasons, but no monthly models can predict North American summer occurrences very well.

Conclusions

The relative predictive ability of European monthly models compared to North American monthly models suggests a change in the seasonal timing between the native range to the non-native range. These results highlight the utility of ecological niche models at finer temporal scales in predicting species distributions and unmasking subtle patterns of evolution.     

DOES NICHE CONSERVATISM PROMOTE SPECIATION? A CASE STUDY IN NORTH AMERICAN SALAMANDERS

Recent speciation research has generally focused on how lineages that originate in allopatry evolve intrinsic reproductive isolation, or how ecological divergence promotes nonallopatric speciation. However, the ecological basis of allopatric isolation, which underlies the most common geographic mode of speciation, remains poorly understood and largely unstudied. Here, we explore the ecological and evolutionary factors that promote speciation in Desmognathus and Plethodon salamanders from temperate eastern North America. Based on published molecular phylogenetic estimates and the degree of geographic range overlap among extant species, we find strong evidence for a role for geographic isolation in speciation. We then examine the relationship between climatic variation and speciation in 16 sister-taxon pairs using geographic information system maps of climatic variables, new methods for modeling species' potential geographic distributions, and data on geographic patterns of genetic variation. In contrast to recent studies in tropical montane regions, we found no evidence for parapatric speciation along climatic gradients. Instead, many montane sister taxa in the Appalachian Highlands inhabit similar climatic niches and seemingly are allopatric because they are unable to tolerate the climatic conditions in the intervening lowlands. This temporal and spatial-ecological pattern suggests that niche conservatism, rather than niche divergence, plays the primary role in promoting allopatric speciation and montane endemism in this species-rich group of vertebrates. Our results demonstrate that even the relatively subtle climatic differences between montane and lowland habitats in eastern North America may play a key role in the origin of new species.     

Measuring ecological niche overlap from occurrence and spatial environmental data

Aim Concerns over how global change will influence species distributions, in conjunction with increased emphasis on understanding niche dynamics in evolutionary and community contexts, highlight the growing need for robust methods to quantify niche differences between or within taxa. We propose a statistical framework to describe and compare environmental niches from occurrence and spatial environmental data. Location Europe, North America and South America. Methods The framework applies kernel smoothers to densities of species occurrence in gridded environmental space to calculate metrics of niche overlap and test hypotheses regarding niche conservatism. We use this framework and simulated species with pre‐defined distributions and amounts of niche overlap to evaluate several ordination and species distribution modelling techniques for quantifying niche overlap. We illustrate the approach with data on two well‐studied invasive species. Results We show that niche overlap can be accurately detected with the framework when variables driving the distributions are known. The method is robust to known and previously undocumented biases related to the dependence of species occurrences on the frequency of environmental conditions that occur across geographical space. The use of a kernel smoother makes the process of moving from geographical space to multivariate environmental space independent of both sampling effort and arbitrary choice of resolution in environmental space. However, the use of ordination and species distribution model techniques for selecting, combining and weighting variables on which niche overlap is calculated provide contrasting results. Main conclusions The framework meets the increasing need for robust methods to quantify niche differences. It is appropriate for studying niche differences between species, subspecies or intra‐specific lineages that differ in their geographical distributions. Alternatively, it can be used to measure the degree to which the environmental niche of a species or intra‐specific lineage has changed over time.