Quaternary Climatic Oscillations Research Articles

Phylogeography of the Japanese common sedge, Carex conica complex (Cyperaceae), based on chloroplast DNA sequence data and chromosomal variation

Genetic and chromosomal variations in plants are often reflected in the geographical distribution patterns. Therefore, identifying such phylogeographical patterns on population is important for understanding the process of plant diversification and speciation, and analyzing both molecular and cytological aspects is necessary. • We investigated the phylogeographic pattern and genetic diversity of the widespread Japanese sedge, Carex conica complex, based on chloroplast DNA haplotypes and chromosomal variations. • Sixteen distinct haplotypes were detected from 258 individuals in the C. conica complex inferred from the cpDNA sequences of intergenic spacer (IGS) between atpB and rbcL, IGS between trnT and trnL, trnL intron, and IGS between trnL and trnF. Most haplotypes showed distinct geographical structures. Phylogenetic analyses revealed two major clades, clades I and II, among 16 haplotypes of the complex. The haplotypes of clade I were mainly found in eastern Japan, while most haplotypes of clade II were found in western Japan. Four intraspecific aneuploids of 2n = 32, 34, 36, and 38, with forming normal bivalents in meiotic divisions, were also found in the complex. The geographical distribution of these intraspecific aneuploids corresponded well with those of the haplotypes. • Our results suggest that the genetic diversity and chromosomal variations in the C. conica complex may have originated from contractions and expansions of geographical ranges affected by Quaternary climatic oscillations.

Multiple Pleistocene refugia and Holocene range expansion of an abundant southwestern American desert plant species (Melampodium leucanthum, Asteraceae)

Pleistocene climatic fluctuations had major impacts on desert biota in southwestern North America. During cooler and wetter periods, drought-adapted species were isolated into refugia, in contrast to expansion of their ranges during the massive aridification in the Holocene. Here, we use Melampodium leucanthum (Asteraceae), a species of the North American desert and semi-desert regions, to investigate the impact of major aridification in southwestern North America on phylogeography and evolution in a widespread and abundant drought-adapted plant species. The evidence for three separate Pleistocene refugia at different time levels suggests that this species responded to the Quaternary climatic oscillations in a cyclic manner. In the Holocene, once differentiated lineages came into secondary contact and intermixed, but these range expansions did not follow the eastwardly progressing aridification, but instead occurred independently out of separate Pleistocene refugia. As found in other desert biota, the Continental Divide has acted as a major migration barrier for M. leucanthum since the Pleistocene. Despite being geographically restricted to the eastern part of the species' distribution, autotetraploids in M. leucanthum originated multiple times and do not form a genetically cohesive group.

Disjunct distribution of chloroplast DNA haplotypes in the understory perennial Veratrum album ssp. oxysepalum (Melanthiaceae) in Japan as a result of ancient introgression

• The Quaternary climatic changes resulted in range shifts of species, providing chances for hybridization. However, the genetic signatures of such ancient introgression have rarely been reported. To investigate such signatures, we performed a phylogeographical study on the perennial plant Veratrum album ssp. oxysepalum, which may have hybridized long ago with another congeneric species, V. stamineum. • Sequence variations in chloroplast DNA (cpDNA) were examined in 43 populations in Japan and adjacent areas. Phylogenetic analyses of different cpDNA haplotypes were conducted on the basis of cpDNA and nuclear ribosomal internal transcribed spacer (nrITS) variations. • In the Japanese archipelago, two major groups of haplotypes were detected, one of which was distributed in a disjunct pattern. The major haplotype, occupying the central part of the species' distribution, formed a monophyletic group with V. stamineum in phylogenetic trees on the basis of cpDNA variation, although the two species did not form a monophyletic group in phylogenetic trees on the basis of nrITS variation. • Historical hybridization between V. album ssp. oxysepalum and V. stamineum in refugia during the Quaternary climatic oscillations, and the resulting chloroplast capture of V. stamineum by V. album ssp. oxysepalum, are most probably responsible for the disjunct distribution of cpDNA in V. album ssp. oxysepalum.

Phylogeography of Japanese horse chestnut (Aesculus turbinata) in the Japanese Archipelago based on chloroplast DNA haplotypes

Japanese horse chestnut (Aesculus turbinata: Hippocastanaceae) is one of the typical woody plants that grow in temperate riparian forests in the Japanese Archipelago. To analyze the phylogeography of this plant in the Japanese Archipelago, we determined cpDNA haplotypes for 337 samples from 55 populations covering the entire distribution range. Based on 1,313bp of two spacers, we determined ten haplotypes that are distinguished from adjacent haplotypes by one or two steps. Most of the populations had a single haplotype, suggesting low diversity. Spatial analysis of molecular variance suggested three obvious phylogeographic structures in western Japan, where Japanese horse chestnut is scattered and isolated in mountainous areas. Conversely, no clear phylogeographic structure was observed from the northern to the southern limit of this species, including eastern Japan, where this plant is more common. Rare and private haplotypes were also found in southwestern Japan, where Japanese horse chestnuts are distributed sparsely. These findings imply that western Japan might have maintained a relatively large habitat for A. turbinata during the Quaternary climatic oscillations, while northerly regions could not.

Early Wisconsinan (MIS 4) Arctic ground squirrel middens and a squirrel-eye-view of the mammoth-steppe

Fossil arctic ground squirrel ( Spermophilus parryii) middens were recovered from ice-rich loess sediments in association with Sheep Creek-Klondike and Dominion Creek tephras (ca 80 ka) exposed in west-central Yukon. These middens provide plant and insect macrofossil evidence for a steppe-tundra ecosystem during the Early Wisconsinan (MIS 4) glacial interval. Midden plant and insect macrofossil data are compared with those previously published for Late Wisconsinan middens dating to ∼25–29 14C ka BP (MIS 3/2) from the region. Although multivariate statistical comparisons suggest differences between the relative abundances of plant macrofossils, the co-occurrence of steppe-tundra plants and insects (e.g., Elymus trachycaulus, Kobresia myosuroides, Artemisia frigida, Phlox hoodii, Connatichela artemisiae) provides evidence for successive reestablishment of the zonal steppe-tundra habitats during cold stages of the Late Pleistocene. Arctic ground squirrels were well adapted to the cold, arid climates, steppe-tundra vegetation and well-drained loessal soils that characterize cold stages of Late Pleistocene Beringia. These glacial conditions enabled arctic ground squirrel populations to expand their range to the interior regions of Alaska and Yukon, including the Klondike, where they are absent today. Arctic ground squirrels have endured numerous Quaternary climate oscillations by retracting populations to disjunct “interglacial refugia” during warm interglacial periods (e.g., south-facing steppe slopes, well-drained arctic and alpine tundra areas) and expanding their distribution across the mammoth-steppe biome during cold, arid glacial intervals.

Plant recolonization in the Himalaya from the southeastern Qinghai-Tibetan Plateau: Geographical isolation contributed to high population differentiation

The Himalaya–Hengduan Mountains region (HHM) in the southern and southeastern Qinghai-Tibetan Plateau (QTP) is considered an important reservoir and a differentiation center for temperate and alpine plants in the Cenozoic. To reveal how plants responded to the Quaternary climatic oscillations in the QTP, the phylogeographical histories of a few subalpine and alpine plants have been investigated, but nearly all studies used only uniparentally inherited cytoplasmic DNA markers, and only a couple of them included sampling from the Himalaya. In this study, range-wide genetic variation of the Himalayan hemlock ( Tsuga dumosa), an important forest species in the HHM, was surveyed using DNA markers from three genomes. All markers revealed genetic depauperation in the Himalaya and richness in the Hengduan Mountains populations. Surprisingly, population differentiation of this wind-pollinated conifer is very high in all three genomes, with few common and many private nuclear gene alleles. These results, together with fossil evidence, clearly indicate that T. dumosa recolonized the Himalaya from the Hengduan Mountains before the Last Glacial Maximum (LGM), accompanied with strong founder effects, and the influence of the earlier glaciations on demographic histories of the QTP plants could be much stronger than that of the LGM. The strong population differentiation in T. dumosa could be attributed to restricted gene flow caused by the complicated topography in the HHM that formed during the uplift of the QTP, and thus sheds lights on the importance of geographical isolation in the development of high plant species diversity in this biodiversity hotspot.

Relations between climatic changes and prehistoric human migrations during Holocene between Gissar Range, Pamir, Hindu Kush and Kashmir: The archaeological and ecological data

In the beginning of the Holocene, hunter–gatherer populations settled in the Pamir plateau, especially in the tectonic depression of Markansu near the Kara Kul Lake (Osh Kona site, 4100 m, 9580 BP and 7145 BP). These populations were well adapted to high altitude life conditions in summer. They were using stone tools related to a cobble tool tradition that Ranov considered as the local continuation of an Asian Middle Palaeolithic (Ferghana, Darwaz or Punjab). However, they disappeared during the second half of the Holocene. As well, hunters–gatherers characterised by a tradition close to Markansu Culture, namely Hissar Culture, appeared during the Holocene in the northern mountains of the Tajik–Afghan depression (Amu Darya upper basin) occupying the middle altitude valleys (Gissar Range) when Neolithic populations began to develop at lower altitudes. For Ranov, origin and destiny of both Markansu and Hissar cultures were the most important questions to solve in this region. The Department of Prehistory, National Museum of Natural History (Paris) conducted field work in collaboration with the University of Peshawar between 1996 and 1998 in the northernmost valley of the Indus basin giving access to the Wakhan corridor. The upper Yarkhun valley (Chitral district, Pakistan), accessible only by foot, was surveyed from 2500 to 4000 m altitude. This allowed the discovery of 6 sites yielding lithic artefacts, among which half are cobble tools. They attest to human activity later than 5500 BP, suggesting population movements between the Amu Darya and Indus basins. The consequences of the Quaternary climate oscillations between South Asian monsoon and Northern hemisphere influences on the one hand, and the impact on biotopes of the increasing pastoral activity on the other hand, are discussed as probable causes for the disappearance of the mountainous nomadic hunters of Central Asia.

Multiple autopolyploidizations and range expansion ofAllium przewalskianumRegel. (Alliaceae) in the Qinghai-Tibetan Plateau

We used the Allium przewalskianum diploid-tetraploid complex on the Qinghai-Tibetan Plateau (QTP) as a model to examine how this complex responded to the Quaternary climatic oscillations, and whether multiple autopolyploidizations have occurred. We sequenced five chloroplast DNA (cpDNA) fragments (accD-psaI, trnH-psbA, trnL-trnF, trnS-trnG and rpl16-intron) in 306 individuals (all of known ploidy level) from 48 populations across the distribution of this species complex. We identified a total of 32 haplotypes-11 in diploids only, 13 in tetraploids only, and 8 found in both cytotypes. This, plus network analyses, indicated that tetraploids have arisen independently from diploids at least eight times. Most populations in the eastern QTP contained multiple haplotypes, but only a single haplotype was found for 17 tetraploid populations on the western QTP, suggesting a recent colonization of the western QTP. We further found that this species complex underwent an earlier range expansion around 5-150 thousand years ago (kya), after the largest glacial period (800-170 kya) in the QTP. In addition, the high frequencies of tetraploids in the QTP suggested that the tetraploid A. przewalskianum cytotype has evolutionary advantages over diploids in colonizing and/or surviving the arid habitats of the QTP.

Phylogeography of the Middle Eastern tree frogs ( Hyla, Hylidae, Amphibia) as inferred from nuclear and mitochondrial DNA variation, with a description of a new species

Evolutionary relationships of the tree frogs from the Middle East and the demographic histories of their populations were studied using a combination of mitochondrial and nuclear genes. Hyla savignyi and neighboring populations of H. orientalis (former eastern populations of H. arborea) were the main focus taxa. Within H. savignyi, a deep phylogenetic divergence dated about 8.4 Ma was discovered. Southern populations from Yemen, Jordan, southern Syria and extreme north-eastern Israel are hereby described as a new species, H. felixarabica sp. nov. Our study points to a biogeographic connection of the south-western Arabian Peninsula and southern Levant and to the importance of the Dead Sea Rift as a historical barrier geographically separating the new species from H. savignyi. Major genetic breaks revealed within species ( H. felixarabica: Yemen vs. Jordan–Syria; H. savignyi sensu stricto: Levant vs. Turkey–Iran) are probably connected to climate changes during the Plio–Pleistocene boundary, while the finer phylogeographic structuring probably resulted from the Quaternary climate oscillations. The Cypriote population of H. savignyi originated from southern Anatolia relatively recently. Hyla orientalis from the southern Black Sea region seems to be genetically quite uniform, although two phylogeographic units with western Turkish and Caucasus–Caspian affinities might be detected. Hyla savignyi and H. orientalis carry signals of population expansions dated to the middle to late Pleistocene, while populations of H. felixarabica seem to have rather been constant in size, which might indicate more stable climatic conditions in the southern regions during the Quaternary.

Multiple northern refugia for Asian sacred lotus, an aquatic plant with characteristics of ice-age endurance

The effect of palaeoclimate on the distribution of aquatic plants is little known, although these plants typically have much broader geographical distributions than their terrestrial counterparts. We investigated the structure of genetic variation of chloroplast DNA in the Asian sacred lotus (Nelumbo nucifera Gaertn (Nelumbonaceae)) in 15 wild populations across China as well as in four populations from India, Japan and Thailand, to infer the refugia of this ancient plant during Quaternary climatic oscillations. We obtained 37 cpDNA haplotypes in 417 individuals from 19 populations. A moderate to high level of chloroplast genetic differentiation (GST = 0.547, NST = 0.691) and significant phylogeographic structure (NST > GST) were observed, suggesting a low level of recurrent seed-mediated gene flow among the populations. The results of AMOVA analysis also showed that more variation was partitioned among (71%) than within populations (29%). The phylogenetic relationships for the recovered haplotypes showed that haplotypes of wild lotus in north-eastern China significantly diverged from those distributed in central and southern regions of China. A high level of haplotype diversity, rather than reduced genetic diversity, in north-eastern China indicated multiple refugia in northern China during the Quaternary glaciations. Lotus plants have the following two characteristics that facilitated survival through Quaternary glaciations: seeds buried in mud remain viable for thousands of years, and self-heating flowers can reproduce in cold temperatures.

The east-west-north colonization history of the Mediterranean and Europe by the coastal plant Carex extensa (Cyperaceae)

Coastal plants are ideal models for studying the colonization routes of species because of the simple linear distributions of these species. Carex extensa occurs mainly in salt marshes along the Mediterranean and European coasts. Variation in cpDNA sequences, amplified fragment length polymorphisms (AFLPs) and simple sequence repeats (SSRs) of 24 populations were analysed to reconstruct its colonization history. Phylogenetic relationships indicate that C. extensa together with the South American Carex vixdentata and the southern African Carex ecklonii form a monophyletic group of halophilic species. Analyses of divergence times suggest that early lineage diversification may have occurred between the late Miocene and the late Pliocene (Messinian crisis). Phylogenetic and network analyses of cpDNA variation revealed the monophyly of the species and an ancestral haplotype contained in populations of the eastern Mediterranean. The AFLP and SSR analyses support a pattern of variation compatible with these two lineages. These analyses also show higher levels of genetic diversity and differentiation in the eastern population group, which underwent an east-to-west Mediterranean colonization. Quaternary climatic oscillations appear to have been responsible for the split between these two lineages. Secondary contacts may have taken place in areas near the Ligurian Sea in agreement with the gene flow detected in Corsican populations. The AFLP and SSR data accord with the 'tabula rasa' hypothesis in which a recent and rapid colonization of northern Europe took place from the western Mediterranean after the Last Glacial Maximum. The unbalanced west-east vs. west-north colonization may be as a result of 'high density blocking' effect.

Phylogeographic analyses suggest that a deciduous species (Ostryopsis davidiana Decne., Betulaceae) survived in northern China during the Last Glacial Maximum

AbstractAim Palaeontologial data suggest that all temperate forest species in northern China migrated southwards during the Last Glacial Maximum (LGM) and recolonized post‐glacially within the Holocene. We tested this assumption using phylogeographical studies of a temperate deciduous shrub species (Ostryopsis davidiana Decne., Betulaceae), which has a wide distribution in northern China.Location Northern China.Methods We sequenced two chloroplast DNA (cpDNA) fragments (trnL–trnF and psbA–trnH, together about 1300 bp in length) of 294 plants from 21 populations across the total distribution range of this species. We used maximum parsimony and haplotype network methods to construct phylogenetic relationships among haplotypes.Results The analysis of cpDNA variation identified eight haplotypes. A single haplotype was fixed in all populations except for one population that was polymorphic, having two haplotypes. The population subdivisions were extremely high (GST = 0.972 and NST = 0.974), suggesting very low gene flow between populations. Haplotypes clustered into two tentative clades, both of which occur in the southern region of the species’ range but only one of which occurs in the northern region. Across the sampled populations, the haplotype distributions were differentiated geographically.Main conclusions Our analyses suggest that multiple refugia were maintained across the range of O. davidiana in both northern (north of the Qing Mountains) and southern (south of the Qing Mountains) regions during the LGM rather than that the species survived only in the south and subsequently colonized northwards. The extremely low within‐population diversity of this species suggests strong bottleneck or founder effects within each fragmented region during the Quaternary climatic oscillations. These findings provide important clues for understanding range shifts and changes in within‐ and/or between‐population genetic diversity of temperate forests in response to past climatic oscillations in northern China.

Molecular inference of a Late Pleistocene diversification shift inNigellas. lat. (Ranunculaceae) resulting from increased speciation in the Aegean archipelago

AbstractAim To infer the temporal course and geographical mode of speciation in Mediterranean/Southwest AsianNigellas. lat.Location Mediterranean Basin, Aegean archipelago.Methods Phylogenies forNigellaL. andGaridellaL. (= Nigellas. lat.) were obtained from maximum‐likelihood analyses of internal transcribed spacer (ITS) sequences. Diversification through time was analysed by log lineages‐through‐time (LTT) plots and survival analyses. Relative node age estimates were regressed against the degree of sympatry between sister clades to infer the predominant mode of geographical speciation inNigellas. lat.Results The Late Pleistocene radiation of theNigella arvensiscomplex in the Aegean region caused a significant departure from a stochastic speciation/extinction process of diversification during the evolution ofNigellas. lat., a lineage of (at least) Late Miocene origin. Speciation withinNigellas. lat. predominantly took place in allopatry.Main conclusions No significant effect on diversification rate was found regarding the establishment of a Mediterranean‐type climate, or the onset of the Quaternary climatic oscillations. Rather, the accelerated rate of speciation in theN. arvensiscomplex is plausibly related toincreasedopportunities for allopatric speciation afforded by the (palaeo)geographical complexity of the Aegean archipelago combined with Late Pleistocene changes in climate and sea level. The evolution of self‐pollination and associated changes in habitat preference and flowering time further augmented speciation and niche differentiation within the complex, but these changes did not act as the primary promoters of the radiation process.

Species‐wide phylogeography of North American mule deer (Odocoileus hemionus): cryptic glacial refugia and postglacial recolonization

Quaternary climatic oscillations greatly influenced the present-day population genetic structure of animals and plants. For species with high dispersal and reproductive potential, phylogeographic patterns resulting from historical processes can be cryptic, overshadowed by contemporary processes. Here we report a study of the phylogeography of Odocoileus hemionus, a large, vagile ungulate common throughout western North America. We examined sequence variation of mitochondrial DNA (control region and cytochrome b) within and among 70 natural populations across the entire range of the species. Among the 1766 individual animals surveyed, we recovered 496 haplotypes. Although fine-scale phylogenetic structure was weakly resolved using phylogenetic methods, network analysis clearly revealed the presence of 12 distinct haplogroups. The spatial distribution of haplogroups showed a strong genetic discontinuity between the two morphological types of O. hemionus, mule deer and black-tailed deer, east and west of the Cascade Mountains in the Pacific Northwest. Within the mule deer lineage, we identified several haplogroups that expanded before or during the Last Glacial Maximum, suggesting that mule deer persisted in multiple refugia south of the ice sheets. Patterns of genetic diversity within the black-tailed deer lineage suggest a single refugium along the Pacific Northwest coast, and refute the hypothesis that black-tailed deer persisted in one or more northern refugia. Our data suggest that black-tailed deer recolonized areas in accordance with the pattern of glacial retreat, with initial recolonization northward along a coastal route and secondary recolonization inland.

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.

Quaternary history and contemporary patterns in a currently expanding species

BackgroundQuaternary climatic oscillations had dramatic effects on species evolution. In northern latitudes, populations had to survive the coldest periods in refugial areas and recurrently colonized northern regions during interglacials. Such a history usually results in a loss of genetic diversity. Populations that did not experience glaciations, in contrast, probably maintained most of their ancestral genetic diversity. These characteristics dramatically affected the present-day distribution of genetic diversity and may influence the ability of species to cope with the current global changes. We conducted a range-wide study of mitochondrial genetic diversity in the pine processionary moth (Thaumetopoea pityocampa/T. wilkinsoni complex, Notodontidae), a forest pest occurring around the Mediterranean Basin and in southern Europe. This species is responding to the current climate change by rapid natural range expansion and can also be accidentally transported by humans. Our aim was to assess if Quaternary climatic oscillations had a different effect across the species' range and to determine if genetic footprints of contemporary processes can be identified in areas of recent introduction.ResultsWe identified three main clades that were spatially structured. In most of Europe, the genetic diversity pattern was typical for species that experienced marked glaciation cycles. Except in refugia, European populations were characterized by the occurrence of one main haplotype and by a strong reduction in genetic diversity, which is expected in regions that were rapidly re-colonized when climatic conditions improved. In contrast, all other sub-clades around the Mediterranean Basin occurred in limited parts of the range and were strongly structured in space, as is expected in regions in which the impact of glaciations was limited. In such places, genetic diversity was retained in most populations, and almost all haplotypes were endemic. This pattern was extreme on remote Mediterranean islands (Crete, Cyprus, Corsica) where highly differentiated, endemic haplotypes were found. Recent introductions were typified by the existence of closely-related haplotypes in geographically distant populations, which is difficult to detect in most of Europe because of a lack of overall genetic structure.ConclusionIn regions that were not prone to marked glaciations, recent moth introductions/expansions could be detected due to the existence of a strong spatial genetic structure. In contrast, in regions that experienced the most intense Quaternary climatic oscillations, the natural populations are not genetically structured, and contemporary patterns of population expansion remain undetected.

Divergence and diversification in North American Psoraleeae (Fabaceae) due to climate change

BackgroundPast studies in the legume family (Fabaceae) have uncovered several evolutionary trends including differential mutation and diversification rates across varying taxonomic levels. The legume tribe Psoraleeae is shown herein to exemplify these trends at the generic and species levels. This group includes a sizable diversification within North America dated at approximately 6.3 million years ago with skewed species distribution to the most recently derived genus, Pediomelum, suggesting a diversification rate shift. We estimate divergence dates of North American (NAm) Psoraleeae using Bayesian MCMC sampling in BEAST based on eight DNA regions (ITS, waxy, matK, trnD-trnT, trnL-trnF, trnK, trnS-trnG, and rpoB-trnC). We also test the hypothesis of a diversification rate shift within NAm Psoraleeae using topological and temporal methods. We investigate the impact of climate change on diversification in this group by (1) testing the hypothesis that a shift from mesic to xeric habitats acted as a key innovation and (2) investigating diversification rate shifts along geologic time, discussing the impact of Quaternary climate oscillations on diversification.ResultsNAm Psoraleeae represents a recent, rapid radiation with several genera originating during the Pleistocene, 1 to 2 million years ago. A shift in diversification rate is supported by both methods with a 2.67-fold increase suggested around 2 million years ago followed by a 8.73-fold decrease 440,000 years ago. The hypothesis that a climate regime shift from mesic to xeric habitats drove increased diversification in affected taxa was not supported. Timing of the diversification rate increase supports the hypothesis that glaciation-induced climate changes during the Quaternary influenced diversification of the group. Nonrandom spatial diversification also exists, with greater species richness in the American Southwest.ConclusionThis study outlines NAm Psoraleeae as a model example of a recent, rapid radiation. Diversification rate shifts in NAm Psoraleeae are not due to current climate regimes as represented by habitat, but instead to past global climate change resulting from Quaternary glaciations. NAm Psoraleeae diversification is a good example of how earthly dynamics including global climate change and topography work together to shape biodiversity.

Extensive population expansion of Pedicularis longiflora (Orobanchaceae) on the Qinghai‐Tibetan Plateau and its correlation with the Quaternary climate change

The Qinghai-Tibetan Plateau (QTP) is thought to be more strongly affected by the Quaternary glaciations than most other regions of the same latitude. It would be of great interest to investigate the population genetic structure of organisms distributed on the platform and its correlation with the Quaternary climatic oscillations. Here we used the chloroplast (cp)DNA trnT-trnF sequence to study genetic variation and phylogeography of Pedicularis longiflora, an alpine herb with extensive distribution on the QTP. Based on a range-wide sampling comprising 41 populations and 910 individuals, we detected 30 cpDNA haplotypes that were divided into five clades by phylogenetic and network analyses and a strong phylogeographical structure. All haplotypes but one in the three basal clades occur exclusively in the southeast QTP, whereas haplotypes in the young clade V occupy almost the whole species range. In particular, the young haplotype H18 occurs in 420 individuals, even at a frequency of 100% in some QTP platform populations and the Altai population. The haplotype distribution pattern, together with molecular clock estimation and mismatch distribution analysis, suggests that the southeast QTP was either a refuge for P. longiflora during the Quaternary climatic change or is the place of origin of the species. The present wide distribution of the species on the QTP platform has resulted from recent population expansions which could be dated back to 120,000-17,000 years ago, a period mostly before the last glacial maximum. The possible relationships among geographic genetic structure, climatic change and species diversification in Pedicularis are also discussed.

Phylogeography of Pinus tabulaeformis Carr. (Pinaceae), a dominant species of coniferous forest in northern China

How coniferous trees in northern China changed their distribution ranges in response to Quaternary climatic oscillations remains largely unknown. Here we report a study of the phylogeography of Pinus tabulaeformis, an endemic and dominant species of coniferous forest in northern China. We examined sequence variation of maternally inherited, seed-dispersed mitochondrial DNA (mtDNA) (nad5 intron 1 and nad4/3-4) and paternally inherited, pollen- and seed-dispersed chloroplast DNA (cpDNA) (rpl16 and trnS-trnG) within and among 30 natural populations across the entire range of the species. Six mitotypes and five chlorotypes were recovered among 291 trees surveyed. Population divergence was high for mtDNA variation (G(ST) = 0.738, N(ST) = 0.771) indicating low levels of seed-based gene flow and significant phylogeographical structure (N(ST) > G(ST), P < 0.05). The spatial distribution of mitotypes suggests that five distinct population groups exist in the species: one in the west comprising seven populations, a second with a north-central distribution comprising 15 populations, a third with a southern and easterly distribution comprising five populations, a fourth comprising one central and one western population, and a fifth comprising a single population located in the north-central part of the species' range. Each group apart from the fourth group is characterized by a distinct mitotype, with other mitotypes, if present, occurring at low frequency. It is suggested, therefore, that most members of each group apart from Group 4 are derived from ancestors that occupied different isolated refugia in a previous period of range fragmentation of the species, possibly at the time of the Last Glacial Maximum. Possible locations for these refugia are suggested. A comparison of mitotype diversity between northern and southern subgroups within the north-central group of populations (Group 2) showed much greater uniformity in the northern part of the range both within and between populations. This could indicate a northward migration of the species from a southern refugium in this region during the postglacial period, although alternative explanations cannot be ruled out. Two chlorotypes were distributed across the geographical range of the species, resulting in lower levels of among-population chlorotype variation. The geographical pattern of variation for all five chlorotypes provided some indication of the species surviving past glaciations in more than one refugium, although differentiation was much less marked, presumably due to the greater dispersal of cpDNA via pollen.

Distinct geographical structure across species units evidenced by chloroplast DNA haplotypes and nuclear ribosomal ITS genotypes ofCorylopsis(Hamamelidaceae) in the Japanese islands

The geographical distribution of chloroplast DNA (cpDNA) haplotypes and nuclear ribosomal internal transcribed spacer (nrITS) genotypes of Japanese Corylopsis (Hamamelidaceae), which consists of four species, was investigated. Two hundred and five individuals belonging to four species from 30 populations, covering the entire geographical range, were studied. Based on approximately 1108 bp of the three non-coding regions of cpDNA, nine haplotypes were detected, and each was distinguished from adjacent haplotypes by one substitution. Based on approximately 507-bp nrITS sequences, 47 genotypes were detected, for which three clades were identified in the phylogenetic analysis. There was inconsistency between the cpDNA haplotypes, nrITS genotypes, and classification of Corylopsis taxa, possibly because of incomplete lineage sorting or introgressive hybridization. The distribution of the haplotypes was highly structured geographically, and NST (0.893) was significantly greater than GST (0.819), implying that the current distribution of Corylopsis species was structured phylogeographically during Quaternary climatic oscillations. The haplotype composition and results of analysis of molecular variance showed that the populations in Hokuriku were highly divergent, suggesting that they are long-term persistent populations arising from refugia during the Quaternary climatic oscillations. Refugial populations in Chugoku and Shikoku may have lost genetic diversity because of a bottleneck resulting from a small population size, followed by post-glacial range expansion. Pre-existing refugia may have been so small that the subsequent range expansion replaced the pre-existing genetic structure. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 501–518.