Diversification of Chionochloa (Poaceae) and biogeography of the New Zealand Southern Alps

Aim We test hypotheses regarding the origin of diversity and patterns of species richness in and around the New Zealand Southern Alps with 25 species of Chionochloa (Poaceae, Danthonioideae). Location New Zealand. Methods We inferred a well‐resolved and mostly robustly supported chloroplast phylogeny based on multiple DNA sequence markers (trnT–L–F, rpl16, trnD–psbM, atpB–rbcL, matK and ndhF), sampling 92% of the recognized species and 82% of the subspecific taxa. Nuclear ribosomal internal transcribed spacer sequences were also sampled, but proved uninformative. Biogeographic reconstruction and character optimization were done using both parsimony and likelihood approaches, and molecular dating used relaxed clock approaches. Results Most of the species diversity in Chionochloa stemmed from a common ancestor in the southern South Island with subsequent dispersal between areas. One clade of apparently cryptic taxa diversified within the central South Island ‘endemism gap’, persisting there throughout at least the latter half of the Pleistocene. Exclusively alpine and other habitat specialist species originated independently, the former relatively recently (between 7.6 Ma and the present). Main conclusions The phylogeny of Chionochloa and other published phylogenies of New Zealand plant groups demonstrate that the higher degree of endemism in the north and south of the New Zealand South Island relative to a central endemism gap cannot be explained by Alpine Fault displacement. Furthermore, our results suggest that if extinctions resulting from glaciations played a role in the origin of the central endemism gap, their impact was less than might be presumed on the basis of the distribution of taxa as they are currently defined. The diversification of Chionochloa and a number of New Zealand plant groups, such as Ranunculus, was contemporaneous with the initiation of the uplift of the Southern Alps. In contrast to patterns of diversifications within the alpine regions typical of the hyperdiverse Andes, exclusively alpine species in New Zealand arose independently from ancestors distributed in more lowland areas. Similarly, habitat specialists in Chionochloa arose independently from more generalist ancestors. Thus, although diversification in these groups may have been stimulated by mountain building and Pleistocene climatic oscillations, cladogenesis did not occur within the high alpine habitat itself.     

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.     

Glacial refugia in a maritime temperate climate: Cicada (Kikihia subalpina) mtDNA phylogeography in New Zealand

Understanding the biological significance of Pleistocene glaciations requires knowledge of the nature and extent of habitat refugia during glacial maxima. An opportunity to examine evidence of glacial forest refugia in a maritime, Southern Hemisphere setting is found in New Zealand, where the extent of Pleistocene forests remains controversial. We used the mitochondrial phylogeography of a forest-edge cicada ( Kikihia subalpina ) to test the hypothesis that populations of this species survived throughout South Island during the Last Glacial Maximum. We also compared mitochondrial DNA phylogeographic patterns with male song patterns that suggest allopatric divergence across Cook Strait. Cytochrome oxidase I and II sequences were analyzed using network analysis, maximum-likelihood phylogenetic estimation, Bayesian dating and Bayesian skyline plots. K. subalpina haplotypes from North Island and South Island form monophyletic clades that are concordant with song patterns. Song divergence corresponds to approximately 2% genetic divergence, and Bayesian dating suggests that the North Island and South Island population-lineages became isolated around 761 000 years bp . Almost all South Island genetic variation is found in the north of the island, consistent with refugia in Marlborough Sounds, central Nelson and northwest Nelson. All central and southern South Island and Stewart Island haplotypes are extremely similar to northern South Island haplotypes, a 'northern richness/southern purity' pattern that mirrors genetic patterns observed in many Northern Hemisphere taxa. Proposed southern South Island forest habitat fragments may have been too small to sustain populations of K. subalpina , and/or they may have harboured ecological communities with no modern-day analogues.     

Phylogeny, biogeography and adaptive radiation of Pachycladon (Brassicaceae) in the mountains of South Island, New Zealand

Aim To report analyses and propose hypotheses of adaptive radiation that explain distributional patterns of the alpine genus Pachycladon Hook.f. – a morphologically diverse genus from New Zealand closely related to Arabidopsis thaliana. Location South Island, New Zealand. Methods Morphological and nrDNA ITS sequence phylogenies were generated for Pachycladon. An analysis is presented of species distributional patterns and attributes. Results Phylogenetic analyses of morphological characters and nrDNA ITS sequence data were found to be congruent in supporting three New Zealand clades for Pachycladon. Monophyletic groups identified within the genus are geographically distinct and are associated with different geological parent materials. Distribution maps, latitude and altitude range, and data on geological parent material are presented for the nine named and one unnamed species of Pachycladon from New Zealand. Main conclusions (a) Panbiogeographic hypotheses accounting for the origin and present‐day distribution of Pachycladon in New Zealand are not supported.
(b) Species diversity and distributions of Pachycladon are explained by a Late Tertiary–Quaternary adaptive radiation associated with increasing specialization to geological substrates. Pachycladon cheesemanii Heenan & A.D.Mitch. is morphologically similar to the closest overseas relatives. It is a geological generalist and has wide latitudinal and altitudinal ranges, and we suggest it resembles the ancestral form of the genus in New Zealand. Pachycladon novae‐zelandiae (Hook.f.) Hook.f. and P. wallii (Carse) Heenan & A.D.Mitch. are a southern South Island group that predominantly occurs on Haast Schist, are polycarpic, have lobed leaves, and lateral inflorescences. Pachycladon enysii (Cheeseman) Heenan & A.D.Mitch., P. fastigiata (Hook.f.) Heenan & A.D.Mitch., and P. stellata (Allan) Heenan & A.D.Mitch. are restricted to greywacke in the eastern South Island, and are facultatively monocarpic, have serrate leaves, and stout terminal inflorescenes.
(c) Present distributions of Pachycladon species may relate to Pleistocene climate change. Pachycladon enysii reaches the highest altitude of New Zealand species of Pachycladon and is most common in the Southern Alps in Canterbury. We propose that this species survived on nunataks at the height of the last glaciation. In contrast, P. fastigiata grows at a lower altitude and is absent from the high mountains of the Southern Alps. We suggest it was extirpated from this area during the last glaciation.     

The integration of multiple independent data reveals an unusual response to Pleistocene climatic changes in the hard tick Ixodes ricinus

In the last few years, improved analytical tools and the integration of genetic data with multiple sources of information have shown that temperate species exhibited more complex responses to ice ages than previously thought. In this study, we investigated how Pleistocene climatic changes affected the current distribution and genetic diversity of European populations of the tick Ixodes ricinus, an ectoparasite with high ecological plasticity. We first used mitochondrial and nuclear genetic markers to investigate the phylogeographic structure of the species and its Pleistocene history using coalescent‐based methods; then we used species distribution modelling to infer the climatic niche of the species at last glacial maximum; finally, we reviewed the literature on the I. ricinus hosts to identify the locations of their glacial refugia. Our results support the scenario that during the last glacial phase, I. ricinus never experienced a prolonged allopatric divergence in separate glacial refugia, but persisted with interconnected populations across Southern and Central Europe. The generalist behaviour in host choice of I. ricinus would have played a major role in maintaining connections between its populations. Although most of the hosts persisted in separate refugia, from the point of view of I. ricinus, they represented a continuity of ‘bridges’ among populations. Our study highlights the importance of species‐specific ecology in affecting responses to Pleistocene glacial–interglacial cycles. Together with other cases in Europe and elsewhere, it contributes to setting new hypotheses on how species with wide ecological plasticity coped with Pleistocene climatic changes.     

Identifying glacial refugia in a geographic parthenogen using palaeoclimate modelling and phylogeography: the New Zealand stick insect Argosarchus horridus (White)

We have used phylogeographic analysis of mitochondrial DNA (COI and COII genes) and ecological niche modelling (ENM) to reconstruct the population history of Argosarchus horridus (White), a widespread species of New Zealand stick insect. These data were used to address outstanding questions on the role of glacial refugia in determining the distribution and genetic structure of New Zealand species. Phylogeographic analysis shows a general pattern of high diversity in upper North Island and reduced diversity in lower North Island and South Island. The ENM indicates that during the last glacial maximum, A. horridus was largely restricted to refugia around coastal areas of North Island. The ENM also suggests refugia on the northeast coast of South Island and southeast coast of North Island and this prediction is verified by phylogeographic analysis, which shows a clade restricted to this region. Argosarchus horridus is also most likely a geographic parthenogen where males are much rarer at higher latitudes. The higher levels of genetic variation in northern, bisexual populations suggest southern and largely unisexual populations originated from southwardly expanding parthenogenetic lineages. Bayesian skyline analysis also provides support for a recent population size increase consistent with a large increase in geographic distribution in the late Pleistocene. These results exemplify the utility of integrating ENM and phylogeographic analysis in testing hypotheses on the origin of geographic parthenogenesis and effects of Pleistocene environmental change on biodiversity.     

Phylogeography of Australian and New Zealand spray zone spiders (Anyphaenidae: Amaurobioides): Moa's Ark loses a few more passengers

Amaurobioides are restricted to the spray zone of southern continents, where they live in small, isolated populations and hunt from silk retreats built in rock crevices. A Star BEAST species tree based on ITS1 nuclear and ND1 mitochondrial genes did not support the hypothesis that this unusual niche linked the evolutionary history of these spiders to geological events reshaping Gondwana into present‐day Australia and New Zealand. Instead, it showed that Amaurobioides reached Australia approximately 4.5 Mya and dispersed twice to New Zealand. Approximately 2.37 Mya, spiders from Tasmania colonized the Deep South of the South Island and, approximately 0.38 Mya, those from South Australia colonized more northern regions. Thus, the present study further limits the scope of the Moa's Ark hypothesis of vicariant New Zealand biogeography.     

Phylogeography of <Emphasis Type="Italic">Leiopelma hochstetteri</Emphasis> reveals strong genetic structure and suggests new conservation priorities

Leiopelma hochstetteri belongs to a singular anuran lineage endemic to New Zealand that diverged from other frogs about 200 million years ago. The species now is reduced to a series of isolated populations in the northern half of the North Island and Great Barrier Island. We have used mitochondrial and nuclear sequence data to examine the genetic affinities of extant populations of L. hochstetteri. Phylogenetic reconstructions reveal that populations are highly structured. Each of the geographically isolated populations harbours independent mtDNA lineages as well as different degrees of nuDNA differentiation. Moreover, molecular dating reveals that this structure originated in the early Pleistocene. This pattern of genetic structure likely results from unfavourable climatic conditions during the Pleistocene combined with the low dispersal ability of the species. Isolated populations in forested refugia existed even in the southern part of the distribution of the species during glacial cycles. Previously published variation in chromosome numbers and isozyme data are consistent with the new evidence. We identified 13 evolutionary significant units (ESUs) that should serve as the focus for future management and conservation of this species.     

Identifying populations for management: fine-scale population structure in the New Zealand alpine rock wren (<Emphasis Type="Italic">Xenicus gilviventris</Emphasis>)

Examining the spatial genetic structure of cryptic species occupying challenging terrain can afford otherwise unattainable insights into ecological and evolutionary processes, such as population dynamics and dispersal patterns; information important for optimising conservation management. Using 13 microsatellite markers, we evaluated patterns of fine-scale gene flow and the spatial extent of genetic structuring of rock wren (Xenicus gilviventris), a threatened alpine passerine endemic to mountainous regions of the South Island, New Zealand. Through spatial autocorrelation analysis, we found that the ‘genetic patch size’, i.e. the distance over which individuals were not genetically independent, was surprisingly large (c. 70 km), given the rock wren’s limited flight ability. By estimating recent migration rates among sampling locations we also found asymmetries in gene flow indicative of source–sink dynamics. An area with intensive deer and predator control, in the Murchison Mountains, Fiordland, appears to be a particularly important source of migrants for other populations. These findings suggest that management to maintain connectivity is required across relatively large spatial scales and source populations may be those where introduced mammals are managed.     

Identifying climate refugia and its potential impact on Tibetan brown bear (Ursus arctos pruinosus) in Sanjiangyuan National Park,China

Climate change has direct impacts on wildlife and future biodiversity protection efforts. Vulnerability assessment and habitat connectivity analyses are necessary for drafting effective conservation strategies for threatened species such as the Tibetan brown bear (Ursus arctos pruinosus). We used the maximum entropy (MaxEnt) model to assess the current (1950–2000) and future (2041–2060) habitat suitability by combining bioclimatic and environmental variables, and identified potential climate refugia for Tibetan brown bears in Sanjiangyuan National Park, China. Next, we selected Circuit model to simulate potential migration paths based on current and future climatically suitable habitat. Results indicate a total area of potential suitable habitat under the current climate scenario of approximately 31,649.46 km², of which 28,778.29 km² would be unsuitable by the 2050s. Potentially suitable habitat under the future climate scenario was projected to cover an area of 23,738.6 km². Climate refugia occupied 2,871.17 km², primarily in the midwestern and northeastern regions of Yangtze River Zone, as well as the northern region of Yellow River Zone. The altitude of climate refugia ranged from 4,307 to 5,524 m, with 52.93% lying at altitudes between 4,300 and 4,600 m. Refugia were mainly distributed on bare rock, alpine steppe, and alpine meadow. Corridors linking areas of potentially suitable brown bear habitat and a substantial portion of paths with low‐resistance value were distributed in climate refugia. We recommend various actions to ameliorate the impact of climate change on brown bears, such as protecting climatically suitable habitat, establishing habitat corridors, restructuring conservation areas, and strengthening monitoring efforts.     

Surviving glacial ages within the Biotic Gap: phylogeography of the New Zealand cicada Maoricicada campbelli

Aim New Zealand is an ideal location in which to investigate the roles of landscape and climate change on speciation and biogeography. An earlier study of the widespread endemic cicada Maoricicada campbelli (Myers) found two phylogeographically distinguishable major clades – northern South Island plus North Island (northern‐SI + NI) and Otago. These two clades appeared to have diverged on either side of an area of the South Island known as the Biotic Gap. We sampled more intensively to test competing theories for this divergence. We aimed to discover if M. campbelli had survived within the Biotic Gap during recent glacial maxima, and if predicted areas of secondary contact between the two major clades existed. Location New Zealand. Methods We analysed mitochondrial DNA sequences (1520 bp; 212 individuals; 91 populations) using phylogenetic (maximum likelihood, Bayesian), population genetic (analysis of molecular variance) and molecular dating methods (Bayesian relaxed clock with improved priors). Results We found strong geographical structuring of genetic variation. Our dating analyses suggest that M. campbelli originated 1.83–2.58 Ma, and split into the two major clades 1.45–2.09 Ma. The main subclades in the northern‐SI + NI clade arose almost simultaneously at 0.69–1.03 Ma. Most subclades are supported by long internal branches and began to diversify 0.40–0.78 Ma. We found four narrow areas of secondary contact between the two major clades. We also found a difference between calling songs of the Otago vs. northern‐SI + NI clades. Main conclusions Phylogeographical patterns within M. campbelli indicate an early Pleistocene split into two major clades, followed by late Pleistocene range expansion and in situ population differentiation of subclades. The northern‐SI + NI clade diversified so rapidly that the main subclade relationships cannot be resolved, and we now have little evidence for a disjunction across the Biotic Gap. Structure within the main subclades indicates rapid divergence after a common bottlenecking event, perhaps attributable to an extremely cold glacial maximum at c. 0.43 Ma. Clade structure and dating analyses indicate that M. campbelli survived in many refugia during recent glacial maxima, including within the Biotic Gap. The narrow overlap between the two major clades is attributed to recent contact during the current interglacial and slow gene diffusion. The two major clades appear to be in the early stages of speciation based on genetic and behavioural differences.     

Tracking the ice: Subterranean harvestmen distribution matches ancient glacier margins

Biogeographic studies often underline the role of glacial dynamism during Pleistocene (1.806–0.011 Mya) in shaping the distribution of subterranean species. Accordingly, it is presumed that present‐day distribution of most specialized cold‐adapted (cryophilic) cave‐dwelling species should bear the signatures of past climatic events. To test this idea, we modelled the distribution of specialized cold‐adapted subterranean alpine harvestmen (Arachnida: Opiliones: Ischyropsalididae: Ischyropsalis). We found that the distance from the glacier margins during Last Glacial Maximum (LGM; about 22,000 years ago) was the most important predictor of their present‐day distribution. In particular, the peak in the probability of occurrence of alpine subterranean Ischyropsalis was found to be in close proximity to the LGM glacier, with a sharp drop at a distance of 30 km from the ice margin. In light of the role played by past climatic events in determining the species current range, we briefly discuss their biogeographic history and the role played by glacial refugia dynamics in determining the current distribution of these species. We argue that low dispersal harvestmen such as our model species can be used as biological indicators for tracking past glaciations and other similar biogeographic events.     

Late‐glacial recolonization and phylogeography of European red deer (Cervus elaphus L.)

The Pleistocene was an epoch of extreme climatic and environmental changes. How individual species responded to the repeated cycles of warm and cold stages is a major topic of debate. For the European fauna and flora, an expansion–contraction model has been suggested, whereby temperate species were restricted to southern refugia during glacial times and expanded northwards during interglacials, including the present interglacial (Holocene). Here, we test this model on the red deer (Cervus elaphus) a large and highly mobile herbivore, using both modern and ancient mitochondrial DNA from the entire European range of the species over the last c. 40 000 years. Our results indicate that this species was sensitive to the effects of climate change. Prior to the Last Glacial Maximum (LGM) haplogroups restricted today to South‐East Europe and Western Asia reached as far west as the UK. During the LGM, red deer was mainly restricted to southern refugia, in Iberia, the Balkans and possibly in Italy and South‐Western Asia. At the end of the LGM, red deer expanded from the Iberian refugium, to Central and Northern Europe, including the UK, Belgium, Scandinavia, Germany, Poland and Belarus. Ancient DNA data cannot rule out refugial survival of red deer in North‐West Europe through the LGM. Had such deer survived, though, they were replaced by deer migrating from Iberia at the end of the glacial. The Balkans served as a separate LGM refugium and were probably connected to Western Asia with genetic exchange between the two areas.     

Phylogeography and ecological niche modelling implicate coastal refugia and trans‐alpine dispersal of a New Zealand fungus beetle

The Last Glacial Maximum (LGM) severely restricted forest ecosystems on New Zealand’s South Island, but the extent of LGM distribution for forest species is still poorly understood. We used mitochondrial DNA phylogeography (COI) and ecological niche modelling (ENM) to identify LGM refugia for the mycophagous beetle Agyrtodes labralis (Leiodidae), a forest edge species widely distributed in the South Island. Both the phylogenetic analyses and the ENM indicate that A. labralis refuged in Kaikoura, Nelson, and along much of the South Island’s west coast. Phylogeography of this species indicates that recolonization of the largely deforested east and southeast South Island occurred in a west–east direction, with populations moving through the Southern Alps, and that the northern refugia participated little in interglacial population expansion. This contradicts published studies of other New Zealand species, in which recolonization occurs in a north–south fashion from many of the same refugia.     

Plio‐Pleistocene diversification and genetic population structure of an endangered lizard (the Blue Mountains water skink,Eulamprus leuraensis) in south‐eastern Australia

Aim Although climatic fluctuations occurred world‐wide during the Pleistocene, the severity of glacial and drought events – and hence their influence on animal and plant biogeography – differed among regions. Many Holarctic species were forced to warmer‐climate refugia during glacial periods, leaving the genetic signature of recent expansion and gene flow among modern‐day populations. Montane south‐eastern Australia experienced less extreme glaciation, but the effects of drier and colder climatic conditions over this period on biotic distributions, and hence on the present‐day genetic structure of animal and plant populations, are poorly known. Location South‐eastern Australia. Methods The endangered Blue Mountains water skink (Eulamprus leuraensis) is a viviparous lizard known from fewer than 40 isolated small swamps at 560–1060 m elevation in south‐eastern Australia. We conducted molecular phylogenetic, dating and population genetics analyses using the mitochondrial NADH dehydrogenase 4 (ND4) of 224 individuals of E. leuraensis sampled across the species’ distribution. Results Ancient divergences in haplotype groups between lizards from the Blue Mountains and the Newnes Plateau, and strong genetic differences, even between swamps separated by only a few kilometres, suggest that the species has persisted as a series of relatively isolated populations within its current distribution for about a million years. Presumably, habitat patches similar to current‐day swamps persisted throughout glacial–interglacial cycles in this region, allowing the development of high levels of genetic structuring within and among present‐day populations. Main conclusions Our results suggest that less extreme glacial conditions occurred in the Southern Hemisphere compared with the Northern Hemisphere, allowing cold‐adapted species (such as E. leuraensis) to persist in montane areas. However, additional studies are needed before we can assemble a comprehensive view of the impact of Pleistocene climatic variation on the phylogeography of Southern Hemisphere taxa.     

History and evolution of alpine plants endemic to the Qinghai-Tibetan Plateau: Aconitum gymnandrum (Ranunculaceae)

How Quaternary climatic oscillations affected range distributions and intraspecific divergence of alpine plants on the Qinghai‐Tibetan Plateau (QTP) remains largely unknown. Here, we report a survey of chloroplast DNA (cpDNA) and nuclear ribosomal internal transcribed spacer (ITS) DNA variation aimed at exploring the phylogeographical history of the QTP alpine endemic Aconitum gymnandrum. We sequenced three cpDNA fragments (rpl20–rps12 intergenic spacer, the trnV intron and psbA‐trnH spacer) and also the nuclear (ITS) region in 245 individuals from 23 populations sampled throughout the species’ range. Two distinct lineages, with eastern and western geographical distributions respectively, were identified from a phylogenetic analysis of ITS sequence variation. Based on a fast substitution rate, these were estimated to have diverged from each other in the early Pleistocene approximately 1.45 Ma. The analysis of cpDNA variation identified nine chlorotypes that clustered into two major clades that were broadly congruent in geographical distribution with the two ITS lineages. The east–west split of cpDNA divergence was supported by an amova which partitioned approximately half of the total variance between these two groups of populations. Analysis of the spatial distribution of chlorotypes showed that each clade was subdivided into two groups of populations such that a total of four population groups existed in the species. It is suggested that these different groups derive from four independent glacial refugia that existed during the Last Glacial Maximum (LGM), and that three of these refugia were located at high altitude on the QTP platform itself at that time. Coalescent simulation of chlorotype genealogies supported both an early Pleistocene origin of the two main cpDNA clades and also the ‘four‐refugia’ hypothesis during the LGM. Two previous phylogeographical studies of QTP alpine plants indicated that such plants retreated to refugia at the eastern/south‐eastern plateau edge during the LGM and/or previous glacial maxima. However, the results for A. gymnandrum suggest that at least some of these cold‐tolerant species may have also survived centrally on the QTP platform throughout the Quaternary.     

Phylogeography and historical demography of the Lusitanian snail <Emphasis Type="Italic">Elona quimperiana</Emphasis> reveal survival in unexpected separate glacial refugia

Background  

Present day distributions of Palearctic taxa in northern latitudes mainly result from populations having survived in local patches during the Late Pleistocene and/or from recolonizing populations from southern temperate refugia. If well-studied Mediterranean and eastern European refugia are widely accepted, some recent biogeographical assumptions still remain unclear, such as the occurrence of multiple glacial refugia in Iberia and cryptic refugia in northern Europe during the last glaciations. The Lusitanian snail Elona quimperiana has a remarkably disjunct distribution, limited to northwestern France (Brittany), northwestern Spain and the Basque Country. By describing the phylogeographical structure of this species across its entire range, the present study attempts to identify refugia and subsequent recolonization routes.     

An explicit test of Pleistocene survival in peripheral versus nunatak refugia in two high mountain plant species

Pleistocene climate fluctuations had profound influence on the biogeographical history of many biota. As large areas in high mountain ranges were covered by glaciers, biota were forced either to peripheral refugia (and possibly beyond to lowland refugia) or to interior refugia (nunataks). However, nunatak survival remains controversial as it relies solely on correlative genetic evidence. Here, we test hypotheses of glacial survival using two high alpine plant species (the insect‐pollinated Pedicularis asplenifolia and wind‐pollinated Carex fuliginosa) in the European Alps. Employing the iDDC (integrative Distributional, Demographic and Coalescent) approach, which couples species distribution modelling, spatial and temporal demographic simulation and Approximate Bayesian Computation, we explicitly test three hypotheses of glacial survival: (a) peripheral survival only, (b) nunatak survival only and (c) peripheral plus nunatak survival. In P. asplenifolia the peripheral plus nunatak survival hypothesis was supported by Bayes factors (BF> 100), whereas in C. fuliginosa the peripheral survival only hypothesis, although best supported, could not be unambiguously distinguished from the peripheral plus nunatak survival hypothesis (BF = 5.58). These results are consistent with current habitat preferences (P. asplenifolia extends to higher elevations) and the potential for genetic swamping (i.e., replacement of local genotypes via hybridization with immigrating genotypes [expected to be higher in the wind‐pollinated C. fuliginosa]). Although the persistence of plants on nunataks during glacial periods has been debated and studied over decades, this is one of the first studies to explicitly test the hypothesis instead of solely using correlative evidence.     

Use of ISSR profiles and ITS-sequences to study the biogeography of alpine cushion plants in the genus Raoulia (Asteraceae)

Nuclear ITS sequences and ISSR profiles provide evidence that Raoulia rubra is endemic to the Tararua Range in the southern part of the North Island of New Zealand. Populations in the South Island previously ascribed to R. rubra are better ascribed to R. eximia. Our findings suggest that glaciation of the central mountains of the South Island during the last ice age have had a major impact on the evolution of the South Island cushion Raoulia species in New Zealand. However, simple hypotheses accounting for the effect of Pleistocene climate change are insufficient to explain patterns of endemism in the group.     

Phylogeography and niche modelling of the relict plant Amborella trichopoda (Amborellaceae) reveal multiple Pleistocene refugia in New Caledonia

Amborella trichopoda Baill. (Amborellaceae, Amborellales), the sole living member of the sister group to all other extant angiosperms, is endemic to New Caledonia. We addressed the intraspecific phylogeography of Amborella by investigating whether its present population genetic structure could be related to its current and past habitats. We found moderate range‐wide genetic diversity based on nuclear microsatellite data and detected four well‐differentiated, geographically distinct genetic groups using Bayesian clustering analyses. We modelled the ecological niche of Amborella based on the current climatic and environmental conditions. The predictive ability of the model was very good throughout the Central East mainland zone, but Amborella was predicted in the northern part of the island where this plant has not been reported. Furthermore, no significant barrier was detected based on habitat suitability that could explain the genetic differentiation across the area. Conversely, we found that the main genetic clusters could be related to the distribution of the suitable habitat at the last glacial maximum (LGM, c. 21 000 years BP), when Amborella experienced a dramatic 96.5% reduction in suitable area. At least two lineages survived in distinct putative refugia located in the Massif des Lèvres and in the vicinity of Mount Aoupinié. Our findings finally confirmed the importance of LGM rainforest refugia in shaping the current intra‐ and interspecific diversity in New Caledonian plants and revealed the possibility of an as yet unreported refugium. The combination of niche modelling and population genetics thereby offered novel insight into the biogeographical history of an emblematic taxon.