Glacial refugia of mammals in Europe: evidence from fossil records

• 1 Glacial refugia were core areas for the survival of temperate species during unfavourable environmental conditions and were the sources of postglacial recolonizations. Unfortunately, the locations of glacial refugia of animals and plants are usually described by models, without reference to facts about real geographical ranges at that time.
• 2 Careful consideration of the faunal assemblages of archaeological sites from the Younger Palaeolithic, which are precisely dated to the Last Glacial Maximum (LGM), gives indications about the distribution of species during the LGM (23 000–16 000 bp ) and provides evidence for the locations of glacial refugia for mammalian species in Europe.
• 3 In Europe, 47 LGM sites, dating from 23 000 to 16 000 bp and containing typical temperate mammal species, have been described. The geographical range of these archaeological sites clearly shows a distribution which differs from the hypothesized traditional refuge areas of the temperate fauna. A considerable number of sites situated in the Dordogne in south‐western France and the Carpathian region contain records of red deer Cervus elaphus, roe deer Capreolus capreolus, wild boar Sus scrofa and red fox Vulpes vulpes.
• 4 The faunal composition of the majority of the evaluated Palaeolithic sites in the southern European peninsulas (with the exception of Greece), as well as France and the Carpathian region, indicates the co‐occurrence of these temperate species with cold‐adapted faunal elements such as mammoth Mammuthus primigenius and/or reindeer Rangifer tarandus.
• 5 The survival of species in Central European refugia would have significant consequences for phylogeography and would be revealed by the dominant distribution of haplotypes, originating from this region. A Carpathian refuge could also be the reason for the very early records of small mammals or mustelids from the Late‐Glacial or Interstadials before the LGM in regions like southern Germany.

     

Postglacial colonization of Europe by the barbastelle bat: agreement between molecular data and past predictive modelling

The barbastelle (Barbastella barbastellus) is a rare forest bat with a wide distribution in Europe. Here, we combine results from the analysis of two mtDNA fragments with species distribution modelling to determine glacial refugia and postglacial colonization routes. We also investigated whether niche conservatism occurs in this species. Glacial refugia were identified in the three southern European peninsulas: Iberia, Italy and the Balkans. These latter two refugia played a major role in the postglacial colonization process, with their populations expanding to England and central Europe, respectively. Palaeo‐distribution models predicted that suitable climatic conditions existed in the inferred refugia during the last glacial maximum (LGM). Nevertheless, the overlap between the current and the LGM distributions was almost inexistent in Italy and in the Balkans, meaning that B. barbastellus populations were forced to shift range between glacial and interglacial periods, a process that probably caused some local extinctions. In contrast, Iberian populations showed a ‘refugia within refugium’ pattern, with two unconnected areas containing stable populations (populations that subsisted during both glacial and interglacial phases). Moreover, the match between LGM models and the refugial areas determined by molecular analysis supported the hypothesis of niche conservatism in B. barbastellus. We argue that geographic patterns of genetic structuring, altogether with the modelling results, indicate the existence of four management units for conservation: Morocco, Iberia, Italy and UK, and Balkans and central Europe. In addition, all countries sampled possessed unique gene pools, thus stressing the need for the conservation of local populations.     

Modelling invasion for a habitat generalist and a specialist plant species

Predicting suitable habitat and the potential distribution of invasive species is a high priority for resource managers and systems ecologists. Most models are designed to identify habitat characteristics that define the ecological niche of a species with little consideration to individual species' traits. We tested five commonly used modelling methods on two invasive plant species, the habitat generalist Bromus tectorum and habitat specialist Tamarix chinensis , to compare model performances, evaluate predictability, and relate results to distribution traits associated with each species. Most of the tested models performed similarly for each species; however, the generalist species proved to be more difficult to predict than the specialist species. The highest area under the receiver-operating characteristic curve values with independent validation data sets of B. tectorum and T. chinensis was 0.503 and 0.885, respectively. Similarly, a confusion matrix for B. tectorum had the highest overall accuracy of 55%, while the overall accuracy for T. chinensis was 85%. Models for the generalist species had varying performances, poor evaluations, and inconsistent results. This may be a result of a generalist's capability to persist in a wide range of environmental conditions that are not easily defined by the data, independent variables or model design. Models for the specialist species had consistently strong performances, high evaluations, and similar results among different model applications. This is likely a consequence of the specialist's requirement for explicit environmental resources and ecological barriers that are easily defined by predictive models. Although defining new invaders as generalist or specialist species can be challenging, model performances and evaluations may provide valuable information on a species' potential invasiveness.     

Improving species distribution models for climate change studies: variable selection and scale

Statistical species distribution models (SDMs) are widely used to predict the potential changes in species distributions under climate change scenarios. We suggest that we need to revisit the conceptual framework and ecological assumptions on which the relationship between species distributions and environment is based. We present a simple conceptual framework to examine the selection of environmental predictors and data resolution scales. These vary widely in recent papers, with light inconsistently included in the models. Focusing on light as a necessary component of plant SDMs, we briefly review its dependence on aspect and slope and existing knowledge of its influence on plant distribution. Differences in light regimes between north‐ and south‐facing aspects in temperate latitudes can produce differences in temperature equivalent to moves 200 km polewards. Local topography may create refugia that are not recognized in many climate change SDMs using coarse‐scale data. We argue that current assumptions about the selection of predictors and data resolution need further testing. Application of these ideas can clarify many issues of scale, extent and choice of predictors, and potentially improve the use of SDMs for climate change modelling of biodiversity.     

Distribution of cryptic blue oat mite species in Australia: current and future climate conditions

• 1 Invertebrate pests, such as blue oat mites Penthaleus spp., cause significant economic damage to agricultural crops in Australia. Climate is a major driver of invertebrate species distributions and climate change is expected to shift pest assemblages and pest prevalence across Australia. At this stage, little is known of how individual species will respond to climate change.
• 2 We have mapped the current distribution for each of the three pest Penthaleus spp. in Australia and built ecological niche models for each species using the correlative modelling software, maxent . Predictor variables useful for describing the climate space of each species were determined and the models were projected into a range of future climate change scenarios to assess how climate change may alter species‐specific distribution patterns in Australia.
• 3 The distributions of the three cryptic Penthaleus spp. are best described with different sets of climatic variables. Suitable climate space for all species decreases under the climate change scenarios investigated in the present study. The models also indicate that the assemblage of Penthaleus spp. is likely to change across Australia, particularly in Western Australia, South Australia and Victoria.
• 4 These results show the distributions of the three Penthaleus spp. are correlated with different climatic variables, and that regional control of mite pests is likely to change in the future. A further understanding of ecological and physiological processes that may influence the distribution and pest status of mites is required.

     

Potential source and sink locations for climate‐driven species range shifts in Europe since the Last Glacial Maximum

Aim To identify potential source and sink locations for climate‐driven species range shifts in Europe since the Last Glacial Maximum (LGM). Location Europe. Methods We developed a new approach combining past‐climate simulations with the concept of analogous climate space. Our index gives a continuous measure of the potential of a location to have acted as a source or a sink for species that have shifted their ranges since the LGM. High glacial source potential is indicated by LGM climatic conditions that are widespread now; high post‐glacial sink potential is indicated by current climatic conditions that were widespread at the LGM. The degree of isolation of source and sink areas was calculated as the median distance to areas with analogous climate conditions. Results We identified areas of high glacial source potential in the previously recognized refugial areas in the southern European peninsulas, but also in large areas in central‐western Europe. The most climatically isolated source areas were located in northern Spain, in north‐western Europe and in eastern Turkey. From here species would have had to cover substantial distances to find current climate conditions analogous to LGM conditions of these areas. Areas with high post‐glacial sink potential were mainly located in Fennoscandia and in central and south‐eastern Europe. Some of the most isolated sink areas were located in the Spanish highlands and around the Baltic Sea. Main conclusions Our species‐independent approach successfully identified previously recognized glacial refugial areas with high source potential for species range shifts in southern Europe and in addition highlighted other potential source areas in central Europe. This study offers new insights into how the distribution of past and current climatic conditions may have influenced past species range shifts and current large‐scale biodiversity patterns.     

Impact of landscape predictors on climate change modelling of species distributions: a case study with Eucalyptus fastigata in southern New South Wales,Australia

Aim We consider three questions. (1) How different are the predicted distribution maps when climate‐only and climate‐plus‐terrain models are developed from high‐resolution data? (2) What are the implications of differences between the models when predicting future distributions under climate change scenarios, particularly for climate‐only models at coarse resolution? (3) Does the use of high‐resolution data and climate‐plus‐terrain models predict an increase in the number of local refugia? Location South‐eastern New South Wales, Australia. Methods We developed two species distribution models for Eucalyptus fastigata under current climate conditions using generalized additive modelling. One used only climate variables as predictors (mean annual temperature, mean annual rainfall, mean summer rainfall); the other used both climate and landscape (June daily radiation, topographic position, lithology, nutrients) variables as predictors. Predictions of the distribution under current climate and climate change were then made for both models at a pixel resolution of 100 m. Results The model using climate and landscape variables as predictors explained a significantly greater proportion of the deviance than the climate‐only model. Inclusion of landscape variables resulted in the prediction of much larger areas of existing optimal habitat. An overlay of predicted future climate on the current climate space indicated that extrapolation of the statistical models was not occurring and models were therefore more robust. Under climate change, landscape‐defined refugia persisted in areas where the climate‐only model predicted major declines. In areas where expansion was predicted, the increase in optimal habitat was always greater with landscape predictors. Recognition of extensive optimal habitat conditions and potential refugia was dependent on the use of high‐resolution landscape data. Main conclusions Using only climate variables as predictors for assessing species responses to climate change ignores the accepted conceptual model of plant species distribution. Explicit statements justifying the selection of predictors based on ecological principles are needed. Models using only climate variables overestimate range reduction under climate change and fail to predict potential refugia. Fine‐scale‐resolution data are required to capture important climate/landscape interactions. Extrapolation of statistical models to regions in climate space outside the region where they were fitted is risky.     

Saxifraga rivularis L. on Beinn Eighe

Summary

A new locality for Saxifraga rivularis is described from Beinn Eighe, Wester Ross. Species lists from the site and from two nearby sites are compared in relation to soil analyses.     

Conservatism of ecological niche characteristics in North American plant species over the Pleistocene-to-Recent transition

Aim  To provide a test of the conservatism of a species' niche over the last 20,000 years by tracking the distribution of eight pollen taxa relative to climate type as they migrated across eastern North America following the Last Glacial Maximum (LGM).
Location  North America.
Methods  We drew taxon occurrence data from the North American pollen records in the Global Pollen Database, representing eight pollen types – all taxa for which ≥5 distinct geographic occurrences were available in both the present day and at the LGM (21,000 years ago ± 3000 years). These data were incorporated into ecological niche models based on present-day and LGM climatological summaries available from the Palaeoclimate Modelling Intercomparison Project to produce predicted potential geographic distributions for each species at present and at the LGM. The output for each time period was projected onto the 'other' time period, and tested using independent known occurrence information from that period.
Results  The result of our analyses was that all species tested showed general conservatism in ecological characteristics over the climate changes associated with the Pleistocene-to-Recent transition.
Main conclusions  This analysis constitutes a further demonstration of general and pervasive conservatism in ecological niche characteristics over moderate periods of time despite profound changes in climate and environmental conditions. As such, our results reinforce the application of ecological niche modelling techniques to the reconstruction of Pleistocene biodiversity distribution patterns, and to project the future potential distribution range of species in the face of global-scale climatic changes.     

Testing the efficiency of nested barriers to dispersal in the Mediterranean high mountain plant Edraianthus graminifolius (Campanulaceae)

Due to strong spatial heterogeneity and limited Pleistocene glaciation, the Balkan Peninsula is a major European biodiversity hot spot. Surprisingly little, however, is known about patterns and processes of intraspecific diversification of its biota in general and of high‐altitude species in particular. A well‐suited system to test hypotheses with respect to various isolating factors acting at different geographic scales and to explore full‐range phylogeographic patterns on the Balkan Peninsula is Edraianthus graminifolius (Campanulaceae), distributed in the western Balkan mountain systems, the southwestern Carpathians and the Apennine Peninsula. To this end, we used a dense population sampling and employed amplified fragment length polymorphism (AFLP) markers and plastid DNA sequences supplemented by ecological niche modelling. The strongest splits were inferred to separate southern and northern Balkan populations from the central ones, from where range extension occurred to the Carpathians and, in more recent times, once or twice to the Apennine Peninsula. The three genetic groups in the western Balkan Peninsula were remarkably congruent among molecular markers, suggesting that the barriers to gene flow acted over long time periods facilitating allopatric differentiation. Each main group of Balkan populations contained genetically and geographically distinct subgroups, which likely are the result of local refugia during warmer periods. Evidently, the topographically highly complex and during the Last Glacial Maximum only locally glaciated Balkan Peninsula is a hot spot of species richness and endemism as well as a sanctuary of intraspecific genetic diversity, even if the underlying causes remain insufficiently understood.     

Global patterns in the shape of species geographical ranges reveal range determinants

Aim Do species range shapes follow general patterns? If so, what mechanisms underlie those patterns? We show for 11,582 species from a variety of taxa across the world that most species have similar latitudinal and longitudinal ranges. We then seek to disentangle the roles of climate, extrinsic dispersal limitation (e.g. barriers) and intrinsic dispersal limitation (reflecting a species’ ability to disperse) as constraints of species range shape. We also assess the relationship between range size and shape. Location Global. Methods Range shape patterns were measured as the slope of the regression of latitudinal species ranges against longitudinal ranges for each taxon and continent, and as the coefficient of determination measuring the degree of scattering of species ranges from the 1:1 line (i.e. latitudinal range = longitudinal range). Two major competing hypotheses explaining species distributions (i.e. dispersal or climatic determinism) were explored. To this end, we compared the observed slopes and coefficients of determination with those predicted by a climatic null model that estimates the potential range shapes in the absence of dispersal limitation. The predictions compared were that species distribution shapes are determined purely by (1) intrinsic dispersal limitation, (2) extrinsic dispersal limitations such as topographic barriers, and (3) climate. Results  Using this methodology, we show for a wide variety of taxa across the globe that species generally have very similar latitudinal and longitudinal ranges. However, neither neutral models assuming random but spatially constrained dispersal, nor models assuming climatic control of species distributions describe range shapes adequately. The empirical relationship between the latitudinal and longitudinal ranges of species falls between the predictions of these competing models. Main conclusions We propose that this pattern arises from the combined effect of macroclimate and intrinsic dispersal limitation, the latter being the major determinant among restricted‐range species. Hence, accurately projecting the impact of climate change onto species ranges will require a solid understanding of how climate and dispersal jointly control species ranges.     

Postglacial expansion and not human influence best explains the population structure in the endangered kea (Nestor notabilis)

Inferring past demography is a central question in evolutionary and conservation biology. It is, however, sometimes challenging to infer the processes that shaped the current patterns of genetic variation in endangered species. Population substructuring can occur as a result of survival in several isolated refugia and subsequent recolonization processes or via genetic drift following a population decline. The kea (Nestor notabilis) is an endemic parrot widely distributed in the mountains of the South Island of New Zealand that has gone through a major human‐induced population decline during the 1860s–1970s. The aims of this study were to understand the glacial and postglacial history of kea and to determine whether the recent population decline played a role in the shaping of the current genetic variation. We examined the distribution of genetic variation, differentiation and admixture in kea using 17 microsatellites and the mitochondrial control region. Mitochondrial data showed a shallow phylogeny and a genetic distinction between the North and South of the range consistent with the three genetic clusters identified with microsatellite data. Both marker types indicated an increase in genetic isolation by geographic distance. Approximate Bayesian Computation supported a scenario of postglacial divergence from a single ancestral glacial refugium, suggesting that the contemporary genetic structure has resulted from recolonization processes rather than from a recent population decline. The recent evolutionary origin of this genetic structure suggests that each genetic cluster does not need to be considered as independent conservation units.     

Nunatak survival vs. tabula rasa in the Central Pyrenees: a study on the endemic plant species Borderea pyrenaica (Dioscoreaceae)

Aim  Borderea pyrenaica (Dioscoreaceae) is a Tertiary relict plant endemic to the Central Pyrenees. Because of its narrow distribution in a small geographical area and the fact that it is restricted to high alpine habitats, it constitutes an ideal model species for inferring the historical dynamics of population survival and migration during and after Quaternary glaciations in the Pyrenees.
Location  Central Pyrenees and pre-Pyrenees, Spain–France.
Methods  Eleven primer pairs were used to amplify 18 microsatellite loci in this allotetraploid species in a sample of 804 individuals from 15 populations, revealing a total of 77 alleles. Genotypic data of individuals and populations were analysed using clustering and Bayesian methods of analysis of population structure.
Results  A higher number of private alleles and a significantly higher allelic richness ( A *) were found in the southern area (21, A * = 2.295) than in the northern area (5, A * = 1.791). Furthermore, the allelic composition of the northern area represented a subset of that from the southern area.
Main conclusions  The hypothesis of in situ survival in northern Pyrenean nunataks was rejected, while peripheral refugia were considered to be restricted to the southern Pyrenees and pre-Pyrenees, where historical geographical fragmentation probably caused the divergence among southern Pyrenean populations. Molecular evidence indicates that these refugial populations probably colonized the northern area after sheet-ice retreat. Borderea pyrenaica lineages followed two migratory pathways in their northward colonization, suggesting several founder events for the populations that eventually reached the territory of the Gavarnie cirque.     

The geography and ecology of plant speciation: range overlap and niche divergence in sister species

A goal of evolutionary biology is to understand the roles of geography and ecology in speciation. The recent shared ancestry of sister species can leave a major imprint on their geographical and ecological attributes, possibly revealing processes involved in speciation. We examined how ecological similarity, range overlap and range asymmetry are related to time since divergence of 71 sister species pairs in the California Floristic Province (CFP). We found that plants exhibit strikingly different age-range correlation patterns from those found for animals; the latter broadly support allopatric speciation as the primary mode of speciation. By contrast, plant sisters in the CFP were sympatric in 80% of cases and range sizes of sisters differed by a mean of 10-fold. Range overlap and range asymmetry were greatest in younger sisters. These results suggest that speciation mechanisms broadly grouped under ‘budding’ speciation, in which a larger ranged progenitor gives rise to a smaller ranged derivative species, are probably common. The ecological and reproductive similarity of sisters was significantly greater than that of sister–non-sister congeners for every trait assessed. However, shifts in at least one trait were present in 93% of the sister pairs; habitat and soil shifts were especially common. Ecological divergence did not increase with range overlap contrary to expectations under character displacement in sympatry. Our results suggest that vicariant speciation is more ubiquitous in animals than plants, perhaps owing to the sensitivity of plants to fine-scale environmental heterogeneity. Despite high levels of range overlap, ecological shifts in the process of budding speciation may result in low rates of fine-scale spatial co-occurrence. These results have implications for ecological studies of trait evolution and community assembly; despite high levels of sympatry, sister taxa and potentially other close relatives, may be missing from local communities.     

Niche shift in four non‐native estrildid finches and implications for species distribution models

Non‐native species can have severe impacts on ecosystems. Therefore, predictions of potentially suitable areas that are at risk of the establishment of non‐native populations are desirable. In recent years, species distribution models (SDMs) have been widely applied for this purpose. However, the appropriate selection of species records, whether from the native area alone or also from the introduced range, is still a matter of debate. We combined analyses of native and non‐native realized climate niches to understand differences between models based on all locations, as well as on locations from the native range only. Our approach was applied to four estrildid finch species that have been introduced to many regions around the world. Our results showed that SDMs based on location data from native areas alone may underestimate the potential distribution of a given species. The climatic niches of species in their native ranges differed from those of their non‐native ranges. Niche comparisons resulted in low overlap values, indicating considerable niche shifts, at least in the realized niches of these species. All four species have high potential to spread over many tropical and subtropical areas. However, transferring these results to temperate areas has a high degree of uncertainty, and we urge caution when assessing the potential spread of tropical species that have been introduced to higher latitudes.