Locating pleistocene refugia: comparing phylogeographic and ecological niche model predictions

Ecological niche models (ENMs) provide a means of characterizing the spatial distribution of suitable conditions for species, and have recently been applied to the challenge of locating potential distributional areas at the Last Glacial Maximum (LGM) when unfavorable climate conditions led to range contractions and fragmentation. Here, we compare and contrast ENM-based reconstructions of LGM refugial locations with those resulting from the more traditional molecular genetic and phylogeographic predictions. We examined 20 North American terrestrial vertebrate species from different regions and with different range sizes for which refugia have been identified based on phylogeographic analyses, using ENM tools to make parallel predictions. We then assessed the correspondence between the two approaches based on spatial overlap and areal extent of the predicted refugia. In 14 of the 20 species, the predictions from ENM and predictions based on phylogeographic studies were significantly spatially correlated, suggesting that the two approaches to development of refugial maps are converging on a similar result. Our results confirm that ENM scenario exploration can provide a useful complement to molecular studies, offering a less subjective, spatially explicit hypothesis of past geographic patterns of distribution.     

Tools for integrating range change,extinction risk and climate change information into conservation management

Ecological niche models (ENMs) are the primary tool used to describe and forecast the potential influence of climate change on biodiversity. However, ENMs do not directly account for important biological and landscape processes likely to affect range dynamics at a variety of spatial scales. Recent advances to link ENMs with population models have focused on the fundamental step of integrating dispersal and metapopulation dynamics into forecasts of species geographic ranges. Here we use a combination of novel analyses and a synthesis of findings from published plant and animal case studies to highlight three seldom recognised, yet important, advantages of linking ENMs with demographic modelling approaches: 1) they provide direct measures of extinction risk in addition to measures of vulnerability based on change in the potential range area or total habitat suitability. 2) They capture life‐history traits that permit population density to vary in different ways in response to key spatial drivers, conditioned by the processes of global change. 3) They can be used to explore and rank the cost effectiveness of regional conservation alternatives and demographically oriented management interventions. Given these advantages, we argue that coupled methods should be used preferentially where data permits and when conservation management decisions require intervention, prioritization, or direct estimates of extinction risk.     

Ectomycorrhizal fungi: exploring the mycelial frontier

Ectomycorrhizal (ECM) fungi form mutualistic symbioses with many tree species and are regarded as key organisms in nutrient and carbon cycles in forest ecosystems. Our appreciation of their roles in these processes is hampered by a lack of understanding of their soil-borne mycelial systems. These mycelia represent the vegetative thalli of ECM fungi that link carbon-yielding tree roots with soil nutrients, yet we remain largely ignorant of their distribution, dynamics and activities in forest soils. In this review we consider information derived from investigations of fruiting bodies, ECM root tips and laboratory-based microcosm studies, and conclude that these provide only limited insights into soil-borne ECM mycelial communities. Recent advances in understanding soil-borne mycelia of ECM fungi have arisen from the combined use of molecular technologies and novel field experimentation. These approaches have the potential to provide unprecedented insights into the functioning of ECM mycelia at the ecosystem level, particularly in the context of land-use changes and global climate change.     

Profound climatic effects on two East Asian black-throated tits (Ave: Aegithalidae), revealed by ecological niche models and phylogeographic analysis

Although a number of studies have assessed the effects of geological and climatic changes on species distributions in East Asian, we still have limited knowledge of how these changes have impacted avian species in south-western and southern China. Here, we aim to study paleo-climatic effects on an East Asian bird, two subspecies of black-throated tit (A. c. talifuensis-concinnus) with the combined analysis of phylogeography and Ecological Niche Models (ENMs). We sequenced three mitochondrial DNA markers from 32 populations (203 individuals) and used phylogenetic inferences to reconstruct the intra-specific relationships among haplotypes. Population genetic analyses were undertaken to gain insight into the demographic history of these populations. We used ENMs to predict the distribution of target species during three periods; last inter-glacial (LIG), last glacial maximum (LGM) and present. We found three highly supported, monophyletic MtDNA lineages and different historical demography among lineages in A. c. talifuensis-concinnus. These lineages formed a narrowly circumscribed intra-specific contact zone. The estimated times of lineage divergences were about 2.4 Ma and 0.32 Ma respectively. ENMs predictions were similar between present and LGM but substantially reduced during LIG. ENMs reconstructions and molecular dating suggest that Pleistocene climate changes had triggered and shaped the genetic structure of black-throated tit. Interestingly, in contrast to profound impacts of other glacial cycles, ENMs and phylogeographic analysis suggest that LGM had limited effect on these two subspecies. ENMs also suggest that Pleistocene climatic oscillations enabled the formation of the contact zone and thus support the refuge theory.     

Paleoecology and coalescence: phylogeographic analysis of hypotheses from the fossil record

The application of principles from coalescence theory to genealogical relationships within species can provide insights into the process of diversification and the influence of biogeography on distributional patterns. There are several features that make some organisms more suitable for detailed studies of historical processes; in particular, limited dispersal, which serves to conserve the patterns of genetic variation that developed during colonization. We describe the potential benefits of studies that integrate analyses of genetic variation with information from the fossil pollen record and present recent examples of the application of quantitative methods of phylogeographic analysis.     

Meta‐analyzing the likely cross‐species responses to climate change

Ecological Niche Models (ENMs) have different performances in predicting potential geographic distributions. Here we meta‐analyzed the likely effects of climate change on the potential geographic distribution of 1,205 bird species from the Neotropical region, modeled using eight ENMs and three Atmosphere‐Ocean General Circulation Models (AOGCM). We considered the variability in ENMs performance to estimate a weighted mean difference between potential geographic distributions for baseline and future climates. On average, potential future ranges were projected to be from 25.7% to 44.5% smaller than current potential ranges across species. However, we found that 0.2% to 18.3% of the total variance in range shifts occurred “within species” (i.e., owing to the use of different modeling techniques and climate models) and 81.7% to 99.8% remained between species (i.e., it could be explained by ecological correlates). Using meta‐analytical techniques akin to regression, we also showed that potential range shifts are barely predicted by bird biological traits. We demonstrated that one can combine and reduce species‐specific effects with high uncertainty in ENMs and also explore potential causes of climate change effect on species using meta‐analytical tools. We also highlight that the search for powerful correlates of climate change‐induced range shifts can be a promising line of investigation.     

Conservation of phylogeographic lineages under climate change

Aim We address the unexplored question of whether the lack of information on intra‐specific diversity inherent in species‐level niche modelling might bias evaluation of the conservation requirements of species and phylogeographic lineages under changing climates. We test for directional biases that might arise due to these methodological differences in ways of assessing risks from climate change. Location The African continent. Methods We identified from peer‐reviewed studies that used both nuclear and plastid markers the distribution of deep phylogeographic divisions within nine species of African mammals and their phylogeographic lineages. We fitted ecological niche models to describe currently suitable, occupied climates and to project the shift of suitable climate to two future time slices. We applied gap analysis to reveal potential changes in the protection of phylogeographic diversity owing to climatic shifts. Results We found that, within species, most phylogeographic lineages differ in the climates they experience and have substantial geographic separation. Models that do not distinguish these subspecific units often fail to identify potential risks of climate change to lineages. Modelled potential effects of climate change on the geographic extent of suitable climate vary in both direction and magnitude. Predictions of the persistence of suitable climate in current protected areas for the resident lineages differ on average by factor of 2 between species and lineage models. Main conclusions Our study develops an original synthetic approach by combining niche modelling, projected climate change, phylogeographic information and gap analysis. We clearly identify the potential benefits of using the new approach to evaluate risks to the conservation of intra‐specific genetic diversity that are posed by climate change. Our results suggest that prudent conservation strategies need to incorporate potential differences in climate tolerance among lineages when planning conservation measures for species confronted with environmental change.     

Postglacial recolonization of North America by spadefoot toads: integrating niche and corridor modeling to study species’ range dynamics over geologic time

Understanding the factors that shape species’ distributions is a key topic in biogeography. As climates change, species can either cope with these changes through evolution, plasticity or by shifting their ranges to track the optimal climatic conditions. Ecological niche modeling (ENM) is a widespread technique in biogeography that estimates the niche of the organism by using occurrences and environmental data to estimate species’ potential distributions. ENMs are often criticized for failing to take species’ dispersal abilities into consideration. Here, we attempt to fill this gap by combining ENMs with dispersal and corridor modeling to study the range dynamics of North American spadefoot toads (Scaphiopodidae) over the Holocene. We first estimated the current and past distributions of spadefoot toads and then estimated their past distributions from the Last Glacial Maximum (LGM) to the present day. Then, we estimated how each taxon recolonized North American by using dispersal and corridor modeling. By combining these two modeling approaches we were able to 1) estimate the LGM refugia used by the North American spadefoot toads, 2) further refine these projections by estimating which of the putative LGM refugia contributed to the recolonization of North America via dispersal, and 3) estimate the relative influence of each LGM refugium to the current species’ distributions. The models were tested using previously published phylogeographic data, revealing a high degree of congruence between our models and the genetic data. These results suggest that combining ENMs and dispersal modeling over time is a promising approach to investigate both historical and future species’ range dynamics.     

Phylogeographic insights into cryptic glacial refugia

The glacial episodes of the Quaternary (2.6 million years ago-present) were a major factor in shaping the present-day distributions of extant flora and fauna, with expansions and contractions of the ice sheets rendering large areas uninhabitable for most species. Fossil records suggest that many species survived glacial maxima by retreating to refugia, usually at lower latitudes. Recently, phylogeographic studies have given support to the existence of previously unknown, or cryptic, refugia. Here we summarise many of these insights into the glacial histories of species in cryptic refugia gained through phylogeographic approaches. Understanding such refugia might be important as the Earth heads into another period of climate change, in terms of predicting the effects on species distribution and survival.     

Northern glacial refugia for the pygmy shrew Sorex minutus in Europe revealed by phylogeographic analyses and species distribution modelling

The southern European peninsulas (Iberian, Italian and Balkan) are traditionally recognized as glacial refugia from where many species colonized central and northern Europe after the Last Glacial Maximum (LGM). However, evidence that some species had more northerly refugia is accumulating from phylogeographic, palaeontological and palynological studies, and more recently from species distribution modelling (SDM), but further studies are needed to test the idea of northern refugia in Europe. Here, we take a rarely implemented multidisciplinary approach to assess if the pygmy shrew Sorex minutus, a widespread Eurasian mammal species, had northern refugia during the LGM, and if these influenced its postglacial geographic distribution. First, we evaluated the phylogeographic and population expansion patterns using mtDNA sequence data from 123 pygmy shrews. Then, we used SDM to predict present and past (LGM) potential distributions using two different training data sets, two different algorithms (Maxent and GARP) and climate reconstructions for the LGM with two different general circulation models. An LGM distribution in the southern peninsulas was predicted by the SDM approaches, in line with the occurrence of lineages of S. minutus in these areas. The phylogeographic analyses also indicated a widespread and strictly northern‐central European lineage, not derived from southern peninsulas, and with a postglacial population expansion signature. This was consistent with the SDM predictions of suitable LGM conditions for S. minutus occurring across central and eastern Europe, from unglaciated parts of the British Isles to much of the eastern European Plain. Hence, S. minutus likely persisted in parts of central and eastern Europe during the LGM, from where it colonized other northern areas during the late‐glacial and postglacial periods. Our results provide new insights into the glacial and postglacial colonization history of the European mammal fauna, notably supporting glacial refugia further north than traditionally recognized.     

Modelle für den Wandel der Natur

Ecological Modelling – Scenarios of future environmental changes Climate change affects ecosystems at different levels. Changes in species phenology, distribution and interactions are today well described phenomena documenting species responses to increasing temperatures. Environmental niche models (ENMs) have developed as powerful tools to address various questions in macroecology. Aiming at a species environmental niche, statistical modelling can be employed to predict a species' potential occurrence by projecting environmental information recorded at locality records over space and time. In climate change biology, ENMs are used to identify individual species' fates as range expansions or retractions as well as features that affect the structure of species assemblages and species interactions within and across different taxonomic groups. ENMs help to promote the persistence of species by identifying spatial patterns of species richness or endangerment to target conservation priorities. Moreover, they are an essential part of risk assessments to set up preventive measures against non‐native species most likely to adversely impact native ecosystems.     

Integrating expert knowledge and ecological niche models to estimate Mexican primates’ distribution

Ecological niche modeling is used to estimate species distributions based on occurrence records and environmental variables, but it seldom includes explicit biotic or historical factors that are important in determining the distribution of species. Expert knowledge can provide additional valuable information regarding ecological or historical attributes of species, but the influence of integrating this information in the modeling process has been poorly explored. Here, we integrated expert knowledge in different stages of the niche modeling process to improve the representation of the actual geographic distributions of Mexican primates (Ateles geoffroyi, Alouatta pigra, and A. palliata mexicana). We designed an elicitation process to acquire information from experts and such information was integrated by an iterative process that consisted of reviews of input data by experts, production of ecological niche models (ENMs), and evaluation of model outputs to provide feedback. We built ENMs using the maximum entropy algorithm along with a dataset of occurrence records gathered from a public source and records provided by the experts. Models without expert knowledge were also built for comparison, and both models, with and without expert knowledge, were evaluated using four validation metrics that provide a measure of accuracy for presence-absence predictions (specificity, sensitivity, kappa, true skill statistic). Integrating expert knowledge to build ENMs produced better results for potential distributions than models without expert knowledge, but a much greater improvement in the transition from potential to realized geographic distributions by reducing overprediction, resulting in better representations of the actual geographic distributions of species. Furthermore, with the combination of niche models and expert knowledge we were able to identify an area of sympatry between A. palliata mexicana and A. pigra. We argue that the inclusion of expert knowledge at different stages in the construction of niche models in an explicit and systematic fashion is a recommended practice as it produces overall positive results for representing realized species distributions.     

Revealing cryptic diversity using molecular phylogenetics and phylogeography in frogs of the Scinax ruber and Rhinella margaritifera species groups

Few studies to date have examined genetic variability of widespread tropical amphibian species over their distributional range using different kinds of molecular markers. Here, we use genetic data in an attempt to delimit evolutionary entities within two groups of Neotropical frogs, the Scinax ruber species group and the Rhinella margaritifera species group. We combined mitochondrial and nuclear markers for a phylogenetic (a total of approximately 2500 bp) and phylogeographic study (approximately 1300 bp) to test the reliability of the currently accepted taxonomic assignments and to explore the geographic structure of their genetic variation, mainly based upon samples from the French Guianan region. Phylogenetic analyses demonstrated the polyphyly of Scinax ruber and Rhinella margaritifera. S. ruber consists of six lineages that may all merit species status. Conflicting signals of mitochondrial and nuclear markers indicated, among some Scinax lineages and species, the possibility of ongoing hybridization processes. R. margaritifera consisted of 11 lineages which might represent distinct species as well. Phylogeographic analyses added further information in support of the specific status of these lineages. Lineages of low divergence were found in sympatry and were reciprocally monophyletic for mitochondrial as well as nuclear genes, indicating the existence of young lineages that should be awarded species status. Our results highlight the utility of combining phylogenetic and phylogeographic methods, as well as the use of both mitochondrial and nuclear markers within one study. This approach helped to better understand the evolutionary history of taxonomically complex groups of species. The assessment of the geographic distribution of genetic diversity in tropical amphibian communities can lead to conclusions that differ strongly from prior analyses based on the occurrence of currently recognized species alone. Such studies, therefore, hold the potential to contribute to a more objective assessment of amphibian conservation priorities in tropical areas.     

Ecological niche models of invasive seaweeds

Ecological niche models (ENMs) are commonly used to calculate habitat suitability from species’ occurrence and macroecological data. In invasive species biology, ENMs can be applied to anticipate whether invasive species are likely to establish in an area, to identify critical routes and arrival points, to build risk maps and to predict the extent of potential spread following an introduction. Most studies using ENMs focus on terrestrial organisms and applications in the marine realm are still relatively rare. Here, we review some common methods to build ENMs and their application in seaweed invasion biology. We summarize methods and concepts involved in the development of niche models, show examples of how they have been applied in studies on algae and discuss the application of ENMs in invasive algae research and to predict effects of climate change on seaweed distributions.     

Faunal histories from Holocene ancient DNA

Recent studies using ancient DNA have been instrumental in advancing understanding of the impact of Holocene climate change on biodiversity. Ancient DNA has been used to track demography, migration and diversity, and is providing new insights into the long-term dynamics of species and population distributions. The Holocene is key to understanding how the past has impacted on the present, as it bridges the gap between contemporary phylogeographic studies and those with inference on Pleistocene patterns, based on ancient DNA studies. Here, we examine the major patterns of Holocene faunal population dynamics and connectivity; highlighting the dynamic nature of species and population responses to Holocene climatic change, thereby providing an 'analogue' for understanding potential impacts of future change.     

Phylogeography and demographic history of the neotropical otter (Lontra longicaudis)

The Neotropical otter (Lontra longicaudis) is a medium-sized semiaquatic carnivore with a broad distribution in the Neotropical region. Despite being apparently common in many areas, it is one of the least known otters, and genetic studies on this species are scarce. Here, we have investigated its genetic diversity, population structure, and demographic history across a large portion of its geographic range by analyzing 1471 base pairs (bp) of mitochondrial DNA from 52 individuals. Our results indicate that L. longicaudis presents high levels of genetic diversity and a consistent phylogeographic pattern, suggesting the existence of at least 4 distinct evolutionary lineages in South America. The observed phylogeographic partitions are partially congruent with the subspecies classification previously proposed for this species. Coalescence-based analyses indicate that Neotropical otter mitochondrial DNA lineages have shared a rather recent common ancestor, approximately 0.5 Ma, and have subsequently diversified into the observed phylogroups. A consistent scenario of recent population expansion was identified in Eastern South America based on several complementary analyses of historical demography. The results obtained here provide novel insights on the evolutionary history of this largely unknown Neotropical mustelid and should be useful to design conservation and management policies on behalf of this species and its habitats.     

应用生态位模型研究外来入侵物种生态位漂移

由于基础生态位和实际生态位的改变,外来入侵物种在入侵地成功定殖、扩散后常会发生生态位漂移,而物种生态位漂移往往很难直接证明。生态位模型在假设入侵物种的生态位需求保守的前提下,以物种在其原产地的生态位需求为基础,预测其在入侵地的潜在分布,通过比较预测分布与实际分布的差异可以从一定程度上得到外来入侵物种的生态位是否发生漂移的间接证据。以我国入侵杂草胜红蓟在原产地的生态位需求为基础,应用生态位模型预测其在其他地区的潜在分布。研究结果表明,生态位模型可以很好地预测胜红蓟在亚太平洋地区和非洲地区的分布,但在我国,其预测分布与实际分布存在较大差别。胜红蓟在我国预测分布主要为云南、海南、台湾部分地区,而胜红蓟入侵我国后现已广泛分布于长江以南地区,其实际分布比预测分布广泛得多,由此推测胜红蓟在入侵我国后其生态位已经产生了漂移。     

Historical biogeography of Drosophila simulans based on Y-chromosomal sequences

Y-chromosomal sequences have been used for phylogeographic studies in humans and other mammals, but so far have been ignored as a source of historical information in Drosophila and other insects with X/Y sex determination. Here, we present the first phylogeographic study of Drosophila simulans based on the Y chromosome. Geographic distribution of Y-chromosomal haplotypes suggests a high degree of population subdivision within Africa, as well as between the African and cosmopolitan groups of populations. Consistent with earlier studies based on autosomal and X-linked loci, our results suggest that D. simulans originated in Madagascar or East Africa, and that the South and West African populations of this species are derived.     

CpDNA-based species identification and phylogeography: application to African tropical tree species

Despite the importance of the African tropical rainforests as a hotspot of biodiversity, their history and the processes that have structured their biodiversity are understood poorly. With respect to past demographic processes, new insights can be gained through characterizing the distribution of genetic diversity. However, few studies of this type have been conducted in Central Africa, where the identification of species in the field can be difficult. We examine here the distribution of chloroplast DNA (cpDNA) diversity in Lower Guinea in two tree species that are difficult to distinguish, Erythrophleum ivorense and Erythrophleum suaveolens (Fabaceae). By using a blind-sampling approach and comparing molecular and morphological markers, we first identified retrospectively all sampled individuals and determined the limits of the distribution of each species. We then performed a phylogeographic study using the same genetic data set. The two species displayed essentially parapatric distributions that were correlated well with the rainfall gradient, which indicated different ecological requirements. In addition, a phylogeographic structure was found for E. suaveolens and, for both species, substantially higher levels of diversity and allelic endemism were observed in the south (Gabon) than in the north (Cameroon) of the Lower Guinea region. This finding indicated different histories of population demographics for the two species, which might reflect different responses to Quaternary climate changes. We suggest that a recent period of forest perturbation, which might have been caused by humans, favoured the spread of these two species and that their poor recruitment at present results from natural succession in their forest formations.     

Comparative multilocus phylogeography of two Palaearctic spruce bark beetles: influence of contrasting ecological strategies on genetic variation

While phylogeographic patterns of organisms are often interpreted through past environmental disturbances, mediated by climate changes, and geographic barriers, they may also be strongly influenced by species‐specific traits. To investigate the impact of such traits, we focused on two Eurasian spruce bark beetles that share a similar geographic distribution, but differ in their ecology and reproduction. Ips typographus is an aggressive tree‐killing species characterized by strong dispersal, whereas Dendroctonus micans is a discrete inbreeding species (sib mating is the rule), parasite of living trees and a poor disperser. We compared genetic variation between the two species over both beetles’ entire range in Eurasia with five independent gene fragments, to evaluate whether their intrinsic differences could have an influence over their phylogeographic patterns. We highlighted widely divergent patterns of genetic variation for the two species and argue that the difference is indeed largely compatible with their contrasting dispersal strategies and modes of reproduction. In addition, genetic structure in I. typographus divides European populations in a northern and a southern group, as was previously observed for its host plant, and suggests past allopatric divergence. A long divergence time was estimated between East Asian and other populations of both species, indicating their long‐standing presence in Eurasia, prior to the last glacial maximum. Finally, the strong population structure observed in D. micans for the mitochondrial locus provides insights into the recent colonization history of this species, from its native European range to regions where it was recently introduced.