Assembly of Alaska‐Yukon boreal steppe communities: Testing biogeographic hypotheses via modern ecological distributions

Beringia (eastern Asia, Alaska, northwest Canada) has been a land‐bridge dispersal route between Asia and North America intermittently since the Mesozoic Era. The Quaternary, the most recent period of exchange, is characterized by large, geologically rapid climate fluctuations and sea‐level changes that alternately expose and inundate the land‐bridge region. Insights into how Quaternary land‐bridge geography has controlled species exchange and assembly of the North American flora comes from focusing on a restricted community with narrow ecological tolerances: species that are today restricted to isolated steppe habitats (dry grasslands) in the Subarctic. We evaluated (i) potential controls over current spatial distributions of steppe plants and their pollinators in Alaska and Yukon and (ii) their ecological distributions in relation to potential biogeographic histories. Taxa present in North America that are disjunct from Asia tended to have larger altitudinal ranges (tolerating colder temperatures) than taxa disjunct from farther south in North America, which were largely restricted to the warmest, lowest‐elevation sites. Ecological findings support the following biogeographic scenarios. Migration from Asia via the land‐bridge occurred during Quaternary glacial periods when conditions were colder and drier than today. While a corridor for migration of cold‐tolerant species of cold steppe and tundra, the land bridge acted as a filter that excluded warmth‐demanding species. Migration from North America occurred under warm, dry interglacial conditions; thermophilous North American disjuncts taking this route may have long histories in Beringia, or they may have migrated recently during the relatively warm and dry early Holocene, when forest cover was incomplete.     

The phylogeographical history of the Iberian steppe plant Ferula loscosii (Apiaceae): a test of the abundant-centre hypothesis

The geology and climate of the western Mediterranean area were strongly modified during the Late Tertiary and the Quaternary. These geological and climatic events are thought to have induced changes in the population histories of plants in the Iberian Peninsula. However, fine-scale genetic spatial architecture across western Mediterranean steppe plant refugia has rarely been investigated. A population genetic analysis of amplified fragment length polymorphism variation was conducted on present-day, relict populations of Ferula loscosii (Apiaceae). This species exhibits high individual/population numbers in the middle Ebro river valley and, according to the hypothesis of an abundant-centre distribution, these northern populations might represent a long-standing/ancestral distribution centre. However, our results suggest that the decimated southern and central Iberian populations are more variable and structured than the northeastern ones, representing the likely vestiges of an ancestral distribution centre of the species. Phylogeographical analysis suggests that F. loscosii likely originated in southern Spain and then migrated towards the central and northeastern ranges, further supporting a Late Miocene southern-bound Mediterranean migratory way for its oriental steppe ancestors. In addition, different glacial-induced conditions affected the southern and northern steppe Iberian refugia during the Quaternary. The contrasting genetic homogeneity of the Ebro valley range populations compared to the southern Iberian ones possibly reflects more severe bottlenecks and subsequent genetic drift experienced by populations of the northern Iberia refugium during the Pleistocene, followed by successful postglacial expansion from only a few founder plants.     

Phylogenetics and evolution of the eastern Asian-eastern North American disjunct aphid tribe,Hormaphidini (Hemiptera: Aphididae)

A conspicuous biogeographic pattern of the Northern Hemisphere is the disjunct occurrence of related taxa on different continents. Perhaps best studied in plants, this pattern includes disjunct distributions of genera in eastern Asia and eastern North America. Such continental disjunctions are thought to be the remnants of a mostly continuously distributed, mixed mesophytic forest dating to the Miocene, which subsequently became fragmented by geological and climatic changes. Some highly host-specific insects, namely aphids, live on descendants of the mixed mesophytic forest taxa and exhibit the same disjunct distributions as that of their host plants. We estimated the phylogeny of Hormaphidini aphids, which host-alternate between witch-hazel (Hamamelis; an eastern Asian-eastern North American disjunct genus) and birch (Betula). Based on partial nuclear elongation factor 1alpha and mitochondrial tRNA leucine/cytochrome oxidase II sequences, trees inferred from maximum-parsimony and maximum-likelihood showed strong support for two monophyletic genera (Hamamelistes and Hormaphis), each containing a clade of Japanese and a clade of North American species. The estimated divergence dates of Asian and North American clades in both genera was 20-30 million years ago, consistent with the idea that aphids may have experienced the same vicariance events as those of their host plants.     

Pleistocene range expansions might explain striking disjunctions between eastern Brazil,Andes and Mesoamerica in Leandra s.str. (Melastomataceae)

Leandra s.str. clade has around 200 species nearly restricted to eastern Brazil. Most species in this group are narrow endemics, but a few present striking disjunct distributions between eastern Brazil and Andes or Mesoamerica. Given the predominantly “montane” distribution observed in most Leandra s.str., we hypothesized that cyclical range expansions during colder Pleistocene periods, followed by local extinctions during warmer interglacial periods, could have shaped the distribution of the disjunct species in this clade. In order to gather support for this biogeographical scenario in a phylogenetic framework, the species that occur outside eastern Brazil were identified, ages of the dispersal events estimated, climatic niche models for the disjuncts were generated, and the climatic envelope of these species compared. Our results place all dispersal events from eastern Brazil to Andes or Mesoamerica during the Pleistocene. Climatic niche modeling indicates a potential range expansion during the Pleistocene colder times for the disjunct species. Although the surpassing of the “dry diagonal” could have been facilitated during glacial periods, this open corridor is an effective barrier for Leandra, given the reduced number of species that dispersed beyond an eastern Brazilian origin. Additionally, the disjunct species do not present significant differences in their climatic envelopes to the non‐disjunct species. Our results provide support to a short‐dispersion/stepping‐stone migration scenario to account for the observed disjunctions in this clade. Range expansions during Pleistocene colder periods followed by local extinctions during interglacial periods could have shaped the distribution of Leandra s.str.     

The lizard assemblage from Seasonally Dry Tropical Forest enclaves in the Cerrado biome, Brazil, and its association with the Pleistocenic Arc

Aim  To determine if the distributions of lizard species from Seasonally Dry Tropical Forest (SDTF) enclaves within the Cerrado biome in central Brazil are associated with the Tropical Seasonal Forests Region, a recently proposed phytogeographic unit of South America, corroborating the existence of a Pleistocenic Arc of SDTFs.
Location  SDTF remnants in the Paranã River valley, municipality of São Domingos, Goiás, Brazil.
Methods  Lizards were extensively sampled using haphazard sampling, funnel traps, and pitfall traps with drift fences during four expeditions. The composition of the SDTF lizard assemblage was compared with those from other South American phytogeographic regions (Caatinga, Cerrado, Chaco, Llanos, and the dry forests of Colombia and Bolivia), based on the literature and our own unpublished data.
Results  The SDTF lizard assemblage contained 20 species, including 11 species with extensive distributions among the regions considered, seven species shared exclusively with Cerrado localities, a single species shared exclusively with other SDTFs, and one endemic species. The presence of Lygodactylus klugei (Smith, Martin & Swain, 1977), presumably endemic to the Pleistocenic Arc formed by the Tropical Seasonal Forests Region, considerably extends the known distribution of this species, suggesting historical connections between Caatinga and Cerrado SDTF enclaves.
Main conclusions  The composition of the lizard assemblage in Cerrado SDTF enclaves seems to corroborate the recent proposal that the SDTF should be recognized as a phytogeographic unit (or dominium). The presence of disjunct populations and endemic species highlights the urgency of considering the uniqueness of the Paranã River valley SDTFs and the importance of its conservation.     

Disjunct distributions during glacial and interglacial periods in mountain butterflies: Erebia epiphron as an example

Over several decades, the distribution patterns and evolution of alpine disjunct species has become an increasingly discussed subject. Large scale genetic analysis has allowed the resolution of the past range changes and intraspecific evolution of many species, in Europe especially of Mediterranean origin. However, the phylogeographic structures of species with arctic-alpine disjunct distribution patterns are relatively poorly studied. The existing phylogeographic analysis (mostly of alpine plant species) supports disjunct distributions during glacial as well as post-glacial periods for a number of species. However, several questions still remain unresolved and we therefore analysed the Mountain Ringlet Erebia epiphron as a model for such alpine disjunct species. We found strong differentiation into five different lineages supporting five differentiation centres: (i) the eastern Pyrenees, (ii) the mountain ranges between the central Pyrenees and south-western Alps, (iii, iv) two areas along the southern Alps margin and (v) the northern Alps margin. We propose that these patterns evolved due to the humidity requirements of this species, which did not allow survival in the dry glacial steppes, but along the margins of the wetter glaciated high mountain ranges.     

Genetic diversity analysis and conservation implications for the Iberian threatened populations of the irano-turanian relict Krascheninnikovia ceratoides (Chenopodiaceae)

Krascheninnikovia ceratoides (L.) Gueldenst. (Chenopodiaceae) is an irano‐turanian steppe plant with a striking geographical vicariance between the western Mediterranean and the central Asian regions. Its westernmost populations are located in the north‐east part of the Iberian Peninsula, in the narrow middle Ebro and Alfambra valleys, where they are threatened and have been catalogued as ‘vulnerable’. Genetic diversity and structure was studied in 150 individuals from five populations of these two valleys using inter‐simple sequence repeats (ISSR). A total of 121 informative bands allowed the identification of 150 genotypes. Nei's genetic diversity found in all natural populations (h = 0.448) was higher than that of other geographical restricted species. Analysis of molecular variance and spatial correlations analyses showed a strong genetic differentiation among populations (35.88%) and among valleys (22.33%). The unweighted pair‐group method of arithmetical averages and Neighbour‐joining clusterings further demonstrated a substructure of two population cores within the Ebro valley. Our genetic data suggest that the high levels of genetic diversity and the strong genetic structure found among the northern Iberian populations and ranges of K. ceratoides might result from the summed effects of its tetraploidy, its outbreeding nature, and its success in colonizing newly altered areas. However, the ISSR data might also reflect the existence of an ancient wider distribution range of the plant in the Iberian Peninsula coupled with a more recent history of habitat fragmentation. Different management conservation guidelines are recommended for this plant after our genetic analysis: a microreserve of approximately 2 km² is proposed for the highly threatened population of Osera (Ebro valley), whereas seed collection strategies and occasional population reinforcements are suggested for the remaining populations. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 419–429.     

Conservation genetics of the endangered Iberian steppe plant Ferula loscosii (Apiaceae)

Ferula loscosii (Lange) Willk (Apiaceae) is a threatened endemic species native to the Iberian Peninsula. The plant has a narrow and disjunct distribution in three regions, NE, C and SE Spain. Genetic variability within and among 11 populations from its natural distribution was assessed using allozymes. Intermediate levels of genetic diversity were detected in F. loscosii (P(99%) = 36.83; H(E) = 0.125; H(T) = 0.152). However, the highest genetic diversity (58%) corresponded to the threatened populations from SE and C Spain (H(T) = 0.169) rather than the more abundant and larger populations from NE Spain (Ebro valley) (H(T) = 0.122). Low to moderate levels of genetic structure were found among regional ranges (G(ST) = 0.134), and several statistical spatial correlation analyses corroborated substantial genetic differentiation among the three main regional ranges. However, no significant genetic differentiation was found among the NE Spain populations, except for a northernmost population that is geographically isolated. Outcrossing mating and other biological traits of the species could account for the maintenance of the present values of genetic diversity within populations. The existence of an ancestral late Tertiary wider distribution of the species in SE and C Spain, followed by the maintenance of different Quaternary refugia in these warmer areas, together with a more recent and rapid post-glacial expansion towards NE Spain, are arguments that could explain the low genetic variability and structure found in the Ebro valley and the higher levels of diversity in the southern Iberian populations.     

The coastal-disjunct mesic flora in the inland Pacific Northwest of USA and Canada: refugia, dispersal and disequilibrium

Aim Understanding the history of the mesic‐adapted plant species of eastern British Columbia and northern Idaho, disjunct from their main coastal distribution, may suggest how biotas reorganize in the face of climate change and dispersal barriers. For different species, current evidence supports establishment of the disjunction via an inland glacial refugium, via recent dispersal from the coast, or via a combination of both. In this study, the modern distributions of the coastal‐disjunct vascular plants are analysed with respect to modern climate to examine how refugia and/or dispersal limitation control regional patterns in species richness. Location North‐west North America. Methods The distributions of nine tree and 58 understorey species with a coastal‐disjunct pattern were compiled on a 50‐km grid. The relationship between species richness and an estimate of available moisture was calculated separately for formerly glaciated and unglaciated portions of the coastal and inland regions. Growth habit and dispersal mode were assessed as possible explanatory variables for species distributions. Results Species richness shows a strong relationship to climate in coastal‐unglaciated areas but no relationship to climate in inland‐glaciated areas. In inland‐glaciated areas, richness is c. 70% lower than that expected from climate. Species with animal‐dispersed seeds occupy a larger portion of coastal and inland regions than species with less dispersal potential. Main conclusions Modern patterns of diversity are consistent with both refugia and dispersal processes in establishing the coastal‐disjunct pattern. The inland glacial refugium is marked by locally high diversity and several co‐distributed endemics. In the inland‐glaciated area, dispersal limitation has constrained diversity despite the nearby refugia. Onset of mesic climate within only the last 3000 years and the low dispersal capacity of many species in the refugium may explain this pattern. This study suggests that vascular plant species will face significant challenges responding to climate change on fragmented landscapes.     

几种石耳科地衣生物地理学关系的分子评估(英文)

对具有北美东部和亚洲东部近缘生物地理学关系的大型地衣种类进行系统发育检测较少。笔者获得了石耳科中8个种10个样本nrDNA的ITS完整序列,并对本科中在分布上具有重要意义的4个种的生物地理学关系进行了分析和评估。通过对细胞核核糖体DNA变异性较强的ITS段碱基序列的比较分析,支持放射盘石耳(Umbilicaria muehlenbergii)、宾州疱脐衣(Lasallia pennsylvanica)为同种型间断分布,美味石耳(U.escu-lenta)和大叶石耳(U.mammulata)间已经达到了相当高的分化程度,为异种型间断分布。地理学分析显示该两种石耳的分化可能是由于太平洋的扩展和隔离所造成的结果。同种型间断分布种类的不同地区的样本之间的差异较小。推测了不同类型间断分布种类的分化时间,美味石耳与大叶石耳的分化发生在中新世(23～5.33 Mya)中期,白令海峡形成期间。     

Phylogeographic signal variation in mitochondrial DNA among geographically isolated grassland butterflies

Aim We surveyed mitochondrial DNA (mtDNA) sequence variation among regionally isolated populations of 10 grassland‐associated butterfly species to determine: (1) the utility of phylogeographic comparisons among multiple species for assessing recent evolutionary patterns, and (2) the respective roles of isolation attributable to range disjunction versus isolation attributable solely to geographic distance in establishing divergence patterns. Location The Peace River grasslands of northern Alberta and British Columbia, Canada, which are isolated by 300+ km from similar communities to the south. Methods We sequenced mtDNA (1420 bp of cytochrome c oxidase subunit I) from five grassland‐restricted butterfly species that have geographically disjunct populations and from five ecologically broader species that have more continuous distributions across the same regions. Using analysis of molecular variance (AMOVA), Mantel and partial Mantel tests, and haplotype networks, we compared population structure within and between species in order to assess the validity of single‐species phylogeographic characterizations. We then contrasted variance components between disjunct and continuously distributed species to assess whether divergences were correlated more with disjunction or with geographic distance. Results Single‐species analyses varied substantially within both the disjunct and the continuous groups. One species in each of these groups had mtDNA with unusually deep intraspecific mitochondrial lineage divergences. On the whole, however, the five species with disjunct ranges exhibited greater divergence between geographically distant populations than did the five species with continuous distributions. Comparison of variance components between disjunct and continuous species indicated that isolation attributable only to geographic distance was responsible for up to half of the total sequence variation between disjunct populations of grassland butterflies. Main conclusions Our findings show that single‐species phylogeographic analyses of post‐Pleistocene butterfly distributions are inadequate for characterizing regional biogeographic divergence histories. However, comparison of mtDNA sequence divergences between groups of disjunct and continuously distributed species can allow isolation attributable to range interruption to be quantitatively distinguished from isolation attributable solely to gene flow attenuation over the same geographic area.     

Southern isolation and northern long‐distance dispersal shaped the phylogeography of the widespread,but highly disjunct,European high mountain plant Artemisia eriantha (Asteraceae)

We investigated the range dynamics of Artemisia eriantha, a widespread, but rare, mountain plant with a highly disjunct distribution in the European Alpine System. We focused on testing the roles of vicariance and long‐distance dispersal in shaping the current distribution of the species. To this end, we collected AFLP and plastid DNA sequence data for 17 populations covering the entire distributional range of the species. Strong phylogeographical structure was found in both datasets. AFLP data suggested that almost all populations were genetically strongly differentiated, with 58% of the overall genetic variation partitioned among populations. Bayesian clustering identified five groups of populations: Balkans, Pyrenees, Central Apennines, one southwestern Alpine population and a Widespread cluster (eastern Pyrenees, Alps, Carpathians). Major groups were supported by neighbor‐joining and NeighbourNet analyses. Fourteen plastid haplotypes were found constituting five strongly distinct lineages: Alps plus Pyrenees, Apennines, Balkans, southern Carpathians, and a Widespread group (eastern Pyrenees, northern Carpathians, Mt. Olympus). Plastid DNA data suggested that A. eriantha colonized the European Alpine System in a westward direction. Although, in southern Europe, vicariant differentiation among the Iberian, Italian and Balkan Peninsulas predominated, thus highlighting their importance as glacial refugia for alpine species, in temperate mountain ranges, long‐distance dispersal prevailed. This study emphasizes that currently highly disjunct distributions can be shaped by both vicariance and long‐distance dispersal, although their relative importance may be geographically structured along, for instance, latitude, as in A. eriantha. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 214–226.     

Investigating the cause of the disjunct distribution of <Emphasis Type="Italic">Amietophrynus pantherinus</Emphasis>, the Endangered South African western leopard toad

More amphibians are threatened through loss of habitat than any other single factor. Conservation measures to restore habitat are dependent on historical data indicating the original extent of a species. When historical data is absent, disjunct distributions create a special problem for conservationists who need to determine whether they have an anthropogenic cause. The Endangered western leopard toad (Amietophrynus pantherinus) has a disjunct distribution in the south-western tip of South Africa. We use mitochondrial sequences from 153 individuals to show that the disjunct distribution is unlikely to have an anthropogenic origin. Two distinct populations are separated by 100 km, with highest probability for the most recent common ancestor arising some 5 Kya. The causes of this disjunct distribution appear to be too recent to be attributable to changes in sea level; instead we suggest that there was a range retraction associated with a distinct drying period in the area during the Holocene, possibly indicating that this species will be susceptible to future climate change. Further, we find that the eastern population is less genetically diverse and appears to be undergoing a serious reduction in range, despite its occurrence in the least urbanized habitat. Conservation measures suggested include surveys for breeding sites in the eastern population, custodian agreements for existing breeding sites and foraging areas and the need for a Biodiversity Management Plan to be drawn up and implemented.     

Phylogenomics,biogeography, and evolution of the blue‐ or white‐fruited dogwoods (Cornus)—Insights into morphological and ecological niche divergence following intercontinental geographic isolation

The eastern Asian (EA)–eastern North American (ENA) floristic disjunction represents a major pattern of phytogeography of the Northern Hemisphere. Despite 20 years of studies dedicated to identification of taxa that display this disjunct pattern, its origin and evolution remain an open question, especially regarding post‐isolation evolution. The blue‐ or white‐fruited dogwoods (BW) are the most species‐rich among the four major clades of Cornus L., consisting of ~35 species divided into three subgenera (subg. Yinquania, subg. Mesomora, and subg. Kraniopsis). The BW group provides an excellent example of the EA–ENA floristic disjunction for biogeographic study due to its diversity distribution centered in eastern Asia and eastern North America, yet its species relationships and delineation have remained poorly understood. In this study, we combined genome‐wide markers from RAD‐seq, morphology, fossils, and climate data to understand species relationships, biogeographic history, and ecological niche and morphological evolution. Our phylogenomic analyses with RAxML and MrBayes recovered a strongly supported and well‐resolved phylogeny of the BW group with three intercontinental disjunct clades in EA and ENA or Eurasia and North America, of which two are newly identified within subg. Kraniopsis. These analyses also recovered a potential new species but failed to resolve relationships within the C. hemsleyi–C. schindleri complex. In an effort to develop an approach to reduce computation time, analysis of different nodal age settings in treePL suggests setting a node's minimum age constraint to the lower bound of a fossil's age range to obtain similar ages to that of BEAST. Divergence time analyses with BEAST and treePL dated the BW stem back to the very Late Cretaceous and the divergence of the three subgenera in the Paleogene. By integrating fossil ages and morphology, a total evidence‐based dating approach was used in conjunction with time‐slice probabilities of dispersal under a DEC model to resolve ancestral ranges of each disjunct in the Miocene: Eurasia and ENA (disjunct 1), EA and western North America (disjunct 2), and EA (disjunct 3). The dated biogeographic history supports dispersal via the North Atlantic Land Bridge in the late Paleogene in disjunct 1 and dispersal via the Bering Land Bridge in the Miocene for disjuncts 2 and 3. Character mapping with a stochastic model in phytools and comparison of ecological niche, morphospace, and rate of evolution indicated differential divergence patterns in morphology, ecological niche, and molecules between disjunct sisters. Although morphological stasis was observed in most of the characters, evolutionary changes in growth habit and some features of leaf, flower, and fruit morphology occurred in one or both sister clades. A significant differentiation of ecological habitats in temperature, precipitation, and elevation between disjunct sisters was observed, suggesting a role of niche divergence in morphological evolution post‐isolation. The patterns of evolutionary rate between morphology and molecules varied among disjunct clades and were not always congruent between morphology and molecules, suggesting cases of non‐neutral morphological evolution driven by ecological selection. Our phylogenetic evidence and comparisons of evolutionary rate among disjunct lineages lend new insights into the formation of the diversity anomaly between EA and ENA, with particular support of an early diversification in EA. These findings, in conjunction with previous studies, again suggest that the EA–ENA disjunct floras are an assembly of lineages descended from the Mesophytic Forests that evolved from the early Paleogene “boreotropical flora” through varied evolutionary pathways across lineages.     

Strong genetic impoverishment from the centre of distribution in southern Europe to peripheral Baltic and isolated Scandinavian populations of the pearly heath butterfly

Aim Climatic changes and fluctuations in the past have strongly influenced the distribution of animal and plant species. Such fluctuations are also reflected in the patterns of genetic diversity on both local and global scales. The genetic pattern of the pearly heath butterfly, Coenonympha arcania, was used to evaluate the genetic differentiation of isolated (in north‐western Europe), peripheral (in north‐eastern Europe) and central (in southern Europe) populations in the context of post‐glacial distributional changes of the species. Location Europe (Sweden, Germany, the Baltic states, Italy, Slovenia, Hungary, Romania, Bulgaria). Thus, samples were collected from large parts of the species’ distribution representing the three categories mentioned above. Methods We analysed 18 loci of 569 individuals from 28 populations by allozyme electrophoresis. We used both individual‐based and population‐based analyses, including F‐statistics, various clustering methods and Markov chain Monte Carlo simulations. Results All loci, except Fum, were polymorphic. The mean F_ST for all samples was 0.18. The mean genetic distance among populations was 0.046. Two major genetic lineages were distinguished. Populations from the centre of the distributional range in southern Europe and the northern periphery of the distributional range differed significantly in their level of genetic variability. The central populations of south‐eastern Europe showed high levels of genetic diversity and no differentiation among populations. Main conclusions Most probably the two major genetic lineages evolved during glacial isolation in two disjunct Mediterranean refugia. The lack of genetic differentiation across south‐eastern Europe implies a continuous Würm ice age distribution in this area, thus supporting the functional existence of steppe forests throughout this region. The peripheral‐isolated populations in Sweden seem to have suffered from one or more severe bottlenecks, resulting in substantial genetic impoverishment. The peripheral‐connected eastern Baltic populations, on the other hand, are affected by post‐glacial and possibly recurrent gene flow from more central parts of the distribution.     

Genetic Differentiation,Niche Divergence,and the Origin and Maintenance of the Disjunct Distribution in the Blossomcrown Anthocephala floriceps (Trochilidae)

Studies of the origin and maintenance of disjunct distributions are of special interest in biogeography. Disjunct distributions can arise following extinction of intermediate populations of a formerly continuous range and later maintained by climatic specialization. We tested hypotheses about how the currently disjunct distribution of the Blossomcrown (Anthocephala floriceps), a hummingbird species endemic to Colombia, arose and how is it maintained. By combining molecular data and models of potential historical distributions we evaluated: (1) the timing of separation between the two populations of the species, (2) whether the disjunct distribution could have arisen as a result of fragmentation of a formerly widespread range due to climatic changes, and (3) if the disjunct distribution might be currently maintained by specialization of each population to different climatic conditions. We found that the two populations are reciprocally monophyletic for mitochondrial and nuclear loci, and that their divergence occurred ca. 1.4 million years before present (95% credibility interval 0.7–2.1 mybp). Distribution models based on environmental data show that climate has likely not been suitable for a fully continuous range over the past 130,000 years, but the potential distribution 6,000 ybp was considerably larger than at present. Tests of climatic divergence suggest that significant niche divergence between populations is a likely explanation for the maintenance of their disjunct ranges. However, based on climate the current range of A. floriceps could potentially be much larger than it currently is, suggesting other ecological or historical factors have influenced it. Our results showing that the distribution of A. floriceps has been discontinous for a long period of time and that populations exhibit different climatic niches have taxonomic and conservation implications.     

Gene frequencies, genetic background and environment in Pyrenean populations of Cepaea nemoralis (L.)

Microgeographical genetic differentiation of populations of the polymorphic land snail Cepaea nemoralis (L.) has been the subject of intensive investigation. The importance of environmental selection acting at the loci which control the shell polymorphism in determining the distribution of genes in this species remains a matter of considerable controversy. Some populations of C. nemoralis in the central Pyrenees show striking local associations of morph frequency (and particularly of the frequency of unhanded shells) with topography which have been ascribed to the action of natural selection.
In the Ter valley (eastern Spanish Pyrenees) populations described here, however, the pattern of association of the frequencies of the shell morphs with topography is quite different from that found in the central Pyrenees. In particular, the distribution of unhanded shells is almost the inverse of that found in some previously studied populations. The general picture of variation in relation to topography in Pyrenean C. nemoralis populations is now seen to be one of great local consistency within valley systems, but of considerable differences between valley systems. This variation in population structure may be due either to cryptic and undetected environmental differences between valley systems or to evolutionary divergence between their C. nemoralis populations. The difficulty of deciding between these two conflicting hypotheses emphasizes the limitations of the technique of simply examining the distribution of individual genes in natural populations as a method of analysing their evolutionary history.     

What is natural? The scale of cryptogenesis in the North Atlantic Ocean

Aim Cryptogenic species may include those taxa that were historically introduced and are now falsely viewed as native. Investigated here is the scale of cryptogenesis in the North Atlantic Ocean by examining disjunct distributions, defined as temperate species occurring only on both sides of the North Atlantic. Disjunct distributions can be explained by four scenarios: glacial relicts, taxonomic artefacts, natural trans‐oceanic dispersal and human‐mediated introduction. Location North Atlantic Ocean. Methods Model taxa included ascidians, bivalves and hydrozoans. Biogeographic status (native, introduced or cryptogenic) was assigned to all species exhibiting a disjunct distribution, based upon multiple criteria. Results Of 1030 species, 60 have a strictly disjunct distribution. Of these disjunct species, for five species there is no reason to doubt their native status, and 55 species are cryptogenic or introduced. Groups with high relative dispersal capacities do not have disjunct distributions more often. Infaunal bivalves have the lowest relative number of disjunct species; none are cryptogenic or naturally disjunct. This supports the concept that glaciations are unlikely to cause disjunct distributions: there are no studies that provide conclusive evidence for the glacial relict model. Hydrozoa have the highest relative number of disjunct species, which, while historically explained by undocumented rafting, may more likely be the result of dispersal by ships, which travel relatively fast, are independent of currents and provide greater surface area. Main conclusions This reanalysis of the historical biogeography of the North Atlantic marine biota reveals that far more species may have been introduced than previously recognized, potentially significantly altering our fundamental understanding of community evolution and ecology. Species that have been present for centuries and can be important ecological engineers who have shaped contemporary communities are possibly falsely viewed as native: they may in fact be the unrecognized introductions of historical times.     

中国西南地区与台湾种子植物间断分布现象

中国西南地区和台湾植物的间断分布是东亚生物地理学上一个十分有趣的研究课题。该文对种子植物进行了统计,发现有50属具有该分布式样的种对或种（包括变种和亚种）;30属呈台-琼（至粤南）-西南间断分布,而其中大多数类群的分布区在粤南-桂南-滇南,即中国热带线以南。统计结果显示,有41属被子植物在台湾和西南之间呈连续分布;有35属从台湾分布至海南（和/或华南）或至福建（和华南）。中国台湾和大陆之间类群的连续分布,以及两者植物区系的巨大相似性表明,台湾作为大陆性岛屿,与中国大陆曾经属于一个统一的植物区系。新近纪以来台湾中央山脉和中国西南至东喜马拉雅地区均经历了海拔升高的过程,形成相似的低至中、高山生境;在第四纪冰期,一些类群在这两个地区之间形成连续的分布区。后来全球气温升高,这些植物类群从低海拔向高海拔山区迁移,由于华东至华中和华南缺少高山生境,使得原来连续分布的某些类群或其祖先类群在中国西南和台湾两地之间灭绝,从而形成间断分布。同时,台湾与大陆失去陆地连接后造成分类群在台湾和西南地区的隔离分化,以及中国南端热带生境的不连续性导致热带植物属种分布区的破碎也是形成台湾和西南间断分布的重要因素。最后将西南和台湾间断分布类群归纳为3种类型：孑遗型、分化型和热带型。     

Phylogeographical study of the alpine plant Cassiope lycopodioides (Ericaceae) suggests a range connection between the Japanese archipelago and Beringia during the Pleistocene

Given that East Asia is located south‐west of Beringia and was less glaciated during the Pleistocene, species at higher latitudes were able to expand their range in this region during climate cooling. Although southward migration is an inevitable colonization process, the biogeographical history of the disjunct ranges of higher‐latitude species in East Asia has been investigated less extensively. Here, we assess whether their disjunct distributions in the Japanese archipelago connected sufficiently with Beringia or persisted in isolation following their establishment. Sequences of nine nuclear loci were determined for Cassiope lycopodioides (Ericaceae) from the Japanese archipelago as well as its surrounding areas, Kamchatka and Alaska. According to the geographical pattern of genetic diversity, the northern populations from Kamchatka to the northern part of the Japanese archipelago were similar genetically and were differentiated from populations in central Japan. Our study suggested that the distribution of C. lycopodioides was connected between the northern part of the Japanese archipelago and south‐western Beringia due to Pleistocene climate cooling. Conversely, central Japan harboured a disjunct range after its establishment. These inferences suggest that widespread range expansion in northern East Asia was plausible for species distributed in Beringia. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 497–509.