Molecular phylogeny of Cissus L. of Vitaceae (the grape family) and evolution of its pantropical intercontinental disjunctions

Pantropical intercontinental disjunct distribution is a major biogeographic pattern in plants, and has been explained mainly by boreotropical migration via the North Atlantic land bridges (NALB) and transoceanic long-distance dispersal (LDD), and sometimes by vicariance. However, well-resolved phylogenies of pantropical clades are still relatively few. Cissus is the largest genus of the grape family Vitaceae and shows a pantropical intercontinental disjunction with its 300 species distributed in all major tropical regions. This study constructed the phylogenetic relationships and biogeographic diversification history of Cissus, employing five plastid markers (rps16, trnL-F, atpB-rbcL, trnH-psbA and trnC-petN). The results confirmed that Cissus polyphyletic, consisting of three main clades: the core Cissus, the Cissus striata complex, and the Australian–Neotropical disjunct Cissus antarctica – C. trianae clade. The latter two clades need to be removed from Cissus to maintain the monophyly of the genus. The core Cissus is inferred to have originated in Africa and is estimated to have diverged from its relatives in Vitaceae in the late Cretaceous. It diversified in Africa into several main lineages in the late Paleocene to the early Eocene, colonized Asia at least three times in the Miocene, and the Neotropics in the middle Eocene. The NALB seems the most plausible route for the core Cissus migration from Africa to the Neotropics in the middle Eocene. Three African–Asian and two Neotropical–Australian disjunctions in Cissus s.l. are estimated to have originated in the Miocene and may be best explained by LDD.     

Plastid and nuclear phylogenomic incongruences and biogeographic implications of Magnolia s.l. (Magnoliaceae)

Magnoliaceae, an assemblage of early diverged angiosperms, comprises two subfamilies, speciose Magnolioideae with approximately 300 species in varying numbers of genera and monogeneric Liriodendroideae with two species in Liriodendron L. This family occupies a pivotal phylogenetic position with important insights into the diversification of early angiosperms, and shows intercontinentally disjunct distribution patterns between eastern Asia and the Americas. Widespread morphological homogeneity and slow substitution rates in Magnolia L. s.l. resulted in poorly supported phylogenetic relationships based on morphology or molecular evidence, which hampers our understanding of the genus’ temporal and spacial evolution. Here, based on the newly generated genome skimming data for 48 Magnolia s.l. species, we produced robust Magnolia phylogenies using genome-wide markers from both plastid genomes and single nucleotide polymorphism data. Contrasting the plastid and nuclear phylogenies revealed extensive cytonuclear conflicts in both shallow and deep relationships. ABBA-BABA and PhyloNet analyses suggested hybridization occurred within sect. Yulania, and sect. Magnolia, which is in concordance with the ploidy level of the species in these two sections. Divergence time estimates and biogeographic reconstruction indicated that the timing of the three tropical Magnolia disjunctions coincided with the mid-Eocene cooling climate and/or late Eocene climate deterioration, and two temperate disjunctions occurred much later, possibly during the warm periods of the Miocene, hence supporting the boreotropical flora concept of Magnolia s.l.     

The diversification of the northern temperate woody flora – A case study of the Elm family (Ulmaceae) based on phylogenomic and paleobotanical evidence

Ulmaceae is a woody family widespread in northern temperate forests. Despite the ecological importance of this family, its phylogeny and biogeographic history are poorly understood. In this study, we reconstruct phylogenetic relationships within the family and infer spatio-temporal diversification patterns based on chloroplast genome (complete cpDNA) and nuclear ribosomal DNA sequences (nrDNA). The seven Ulmaceae genera are resolved in two main clades (temperate vs. tropical) by both cpDNA and nrDNA sequences. The temperate clade includes four genera, Hemiptelea, Zelkova, Planera, and Ulmus. The relationships among Planera and other genera are controversial because of inconsistent topologies between plastid and nuclear data. The tropical clade includes three genera ((Ampelocera, Phyllostylon), Holoptelea). Molecular dating and diversification analyses show that Ulmaceae originated in the Early Cretaceous (ca. 110–125 Ma) with the main lineages establishing from the Late Cretaceous to the early Eocene. The diversification rate slowed during the middle to the late Paleogene (ca. 23–45 Ma), followed by a rapid diversification of the East Asian temperate group in the Neogene, congruent with a global cooling event. The ancestral state optimization analysis suggests an East Asian origin of the temperate Ulmaceae clade during the Paleocene, which is consistent with the fossil record. Both phylogenomic and fossil evidence support East Asia as a center of origin and diversification for the temperate woody lineages.     

根据叶绿体ndhF和核核糖体DNA序列推断梓属(紫葳科)的系统发育

Phylogenetics of Chilopsis and Catalpa (Bignoniaceae) was elucidated based on sequences of chloroplast ndhF and the nrDNA ITS region. In Bignoniaceae, Chilopsis and Catalpa are most closely related as sister genera. Our data supported section Macrocatalpa of the West Indies and section Catalpa of eastern Asian and North American continents. Within section Catalpa, Catalpa ovata of eastern Asia form a clade with North American species, C. speciosa and C. bignonioides, while the other eastern Asian species comprise a clade where C. duclouxii is sister to the clade of C. bungei and C. fargesii. The Caribbean species of Catalpa diverged early from the continental species. More studies are needed to test whether the phylogenetic pattern is common in eastern Asian-North American disjunct genera with species in the West Indies.     

Extent and timing of floristic exchange between Australian and Asian rain forests

Aim We tested an entrenched concept – that the Australian rain forest flora is essentially a Gondwanan relict. We also assessed the role of regional‐level source–sink dynamics in the assembly of this flora. Location Eastern Australia. Methods To avoid potential biases inherent in selective studies undertaken to date, we used an analytical, whole‐of‐flora approach integrated with the fossil record. We identified disjunctions between woody Australian rain forest plant taxa and relatives on other land masses. To test the strength of the fossil evidence for the regional antiquity of this flora, we evaluated the proportion of these disjunct clades represented in the Australian fossil record, and to minimize the effects of biases in this record, we compared late Quaternary (i.e. late Pleistocene and Holocene, 126–0 ka), Pliocene and late Oligocene–early Miocene Australian pollen records interpreted as tropical rain forest. Using within‐species disjunctions as a proxy, we assessed the role of recent immigration from Asia into Australia. To assess the role of source–sink dynamics, we performed comparative analyses of disjunctions in major rain forest categories representing a north–south/climatic gradient. Results Southern Australian, cool temperate (microthermal) rain forests contain many floristic disjunctions with Gondwanan fragments and most of these clades have Gondwanan fossils. Disjunct clades in Australian mesothermal rain forest mostly occur in Asia/Malesia and a low proportion of these clades show pre‐Neogene records. Many clades in lowland tropical and ‘dry’ rain forest show disjunctions with Asia/Malesia and few have Australian fossil records. Rates of recent immigration from Asia/Malesia are high in these northern forests, and outweigh rates of recent emigration approximately nine‐fold. The late Quaternary fossil record has many more rain forest angiosperms than Oligocene–Miocene and Pliocene floras, consistent with extensive late Cenozoic immigration. Main conclusions The microthermal rain forests are largely Gondwanan relicts, but there is progressively greater, and more recent contribution from Asia/Malesia into more northern, and more lowland tropical rain forests. This variation reflects a strong gradient in geographic and ecological proximity between these forests and source floras in Asia/Malesia, and is consistent with a source–sink size model of immigration driven by late Cenozoic contractions and expansions of Australian rain forest.     

Geographical affinities of the Cape flora, South Africa

Aim The flora characteristic of the Cape Floristic Region (CFR) is dominated by a relatively small number of clades that have been proposed as ‘Cape clades’. These clades have variously been suggested to have African or Austral affinities. Here we evaluate the support for these conflicting hypotheses. In addition, we test the hypothesis that these clades share a common time of differentiation from their geographical neighbours. Location The Cape Floristic Region, South Africa Methods We use both published and unpublished phylogenetic information to investigate the geographical sister areas of the Cape clades as well as the timing and the direction of biogeographical disjunctions. Results Almost half of the Cape clades for which unambiguous sister areas could be established show a trans‐Indian Ocean disjunction. The earliest trans‐Indian Ocean disjunction dates from 80 Ma. Other disjunctions date from various times in the Cenozoic, and we suggest that the process of recruiting lineages into the Cape flora might be ongoing. Relatively few Cape clades show a sister relationship with South America and tropical Africa, despite their relative geographical proximity. Numerous Cape clades contain species also found on tropical African mountains; in all cases tested, these species are shown to be embedded within the Cape clades. While many Cape clades show a relationship with the Eurasian temperate flora, this is complicated by their presence in tropical Africa. The single case study addressing this to date suggests that the Cape clade is nested within a European grade. Main conclusions Although many Cape clades show Austral rather than African relationships, there are numerous other patterns suggestive of a cosmopolitan flora. This spatial variation is echoed in the temporal data, from which, although there is wide variance around the dates of disjunctions, it is clear the Cape flora has been assembled over a long time period. There is no simple hypothesis that can account for the geographical sources of the currently distinctive Cape flora. The phylogenetic positions of Afromontane members of Cape clades suggest a history of dispersal from the CFR, rather than the reverse.     

Phylogenetic and biogeographic complexity of Magnoliaceae in the Northern Hemisphere inferred from three nuclear data sets

This study employs three nuclear genes (PHYA, LFY, and GAI1) to reconstruct the phylogenetic and biogeographic history of Magnoliaceae. A total of 104 samples representing 86 taxa from all sections and most subsections were sequenced. Twelve major groups are well supported to be monophyletic within Magnoliaceae and these groups are largely consistent with the recent taxonomic revision at the sectional and subsectional levels. However, relationships at deeper nodes of the subfamily Magnolioideae remain not well resolved. A relaxed clock relying on uncorrelated rates suggests that the complicated divergent evolution of Magnolioideae began around the early Eocene (54.57mya), concordant with paleoclimatic and fossil evidence. Intercontinental disjunctions of Magnoliaceae in the Northern Hemisphere appear to have originated during at least two geologic periods. Some occurred after the middle Miocene, represented by two well-recognized temperate lineages disjunct between eastern Asia and eastern North America. The others may have occurred no later than the Oligocene, with ancient separations between or within tropical and temperate lineages.     

Phylogenetic analysis of Toxicodendron （Anacardiaceae） and its biogeographic implications on the evolution of north temperate and tropical intercontinental disjunctions

Toxicodendron is a genus in the Rhus complex of Anacardiaceae with a disjunct distribution between eastern Asia and North America, extending to southeastern Asia and the neotropics. Nuclear （internal transcribed spacer, external transcribed spacer, and NIA-i3） and chloroplast （ndhF and trnL-F） sequences were used to construct phylogenetic relationships of Toxicodendron. Phylogenetic analysis of these data strongly support Toxieodendron as a monophyletic group distinct from other genera of the Rhus complex, and the phylogeny does not fully corroborate classification at the sectional level. Two temperate disjunct lineages were detected, one from section Toxicodendron and the other between the eastern North American Toxicodendron vernix and the eastern Asian Toxicodendron vernieifluum. Their divergence times were estimated to be 13.46 （7.95-19.42） and 7.53 （2.76-12.86） mya, respectively. The disjunction between section Griffithii （taxa from warm temperate to tropical Asia） and Toxieodendron striatum （from the neotropics） was supported and their divergence time was estimated to be 20.84 （11.1 6-30.52） mya in the early Miocene. Our biogeographic results and the paleontological data support the Bering land bridge as the most likely route to explain the temperate disjunctions, yet the tropical disjunction in Toxicodendron seems to be best explained by the North Atlantic land bridge hypothesis.     

Evolution and biogeography of the slipper orchids: Eocene vicariance of the conduplicate genera in the Old and New World Tropics

Intercontinental disjunctions between tropical regions, which harbor two-thirds of the flowering plants, have drawn great interest from biologists and biogeographers. Most previous studies on these distribution patterns focused on woody plants, and paid little attention to herbs. The Orchidaceae is one of the largest families of angiosperms, with a herbaceous habit and a high species diversity in the Tropics. Here we investigate the evolutionary and biogeographical history of the slipper orchids, which represents a monophyletic subfamily (Cypripedioideae) of the orchid family and comprises five genera that are disjunctly distributed in tropical to temperate regions. A relatively well-resolved and highly supported phylogeny of slipper orchids was reconstructed based on sequence analyses of six maternally inherited chloroplast and two low-copy nuclear genes (LFY and ACO). We found that the genus Cypripedium with a wide distribution in the northern temperate and subtropical zones diverged first, followed by Selenipedium endemic to South America, and finally conduplicate-leaved genera in the Tropics. Mexipedium and Phragmipedium from the neotropics are most closely related, and form a clade sister to Paphiopedilum from tropical Asia. According to molecular clock estimates, the genus Selenipedium originated in Palaeocene, while the most recent common ancestor of conduplicate-leaved slipper orchids could be dated back to the Eocene. Ancestral area reconstruction indicates that vicariance is responsible for the disjunct distribution of conduplicate slipper orchids in palaeotropical and neotropical regions. Our study sheds some light on mechanisms underlying generic and species diversification in the orchid family and tropical disjunctions of herbaceous plant groups. In addition, we suggest that the biogeographical study should sample both regional endemics and their widespread relatives.     

Early evolutionary history of the flowering plant family Annonaceae: steady diversification and boreotropical geodispersal

Aim Rain forest‐restricted plant families show disjunct distributions between the three major tropical regions: South America, Africa and Asia. Explaining these disjunctions has become an important challenge in biogeography. The pantropical plant family Annonaceae is used to test hypotheses that might explain diversification and distribution patterns in tropical biota: the museum hypothesis (low extinction leading to steady accumulation of species); and dispersal between Africa and Asia via Indian rafting versus boreotropical geodispersal. Location Tropics and boreotropics. Methods Molecular age estimates were calculated using a Bayesian approach based on 83% generic sampling representing all major lineages within the family, seven chloroplast markers and two fossil calibrations. An analysis of diversification was carried out, which included lineage‐through‐time (LTT) plots and the calculation of diversification rates for genera and major clades. Ancestral areas were reconstructed using a maximum likelihood approach that implements the dispersal–extinction–cladogenesis model. Results The LTT plots indicated a constant overall rate of diversification with low extinction rates for the family during the first 80 Ma of its existence. The highest diversification rates were inferred for several young genera such as Desmopsis, Uvariopsis and Unonopsis. A boreotropical migration route was supported over Indian rafting as the best fitting hypothesis to explain present‐day distribution patterns within the family. Main conclusions Early diversification within Annonaceae fits the hypothesis of a museum model of tropical diversification, with an overall steady increase in lineages possibly due to low extinction rates. The present‐day distribution of species within the two largest clades of Annonaceae is the result of two contrasting biogeographic histories. The ‘long‐branch clade’ has been diversifying since the beginning of the Cenozoic and underwent numerous geodispersals via the boreotropics and several more recent long‐distance dispersal events. In contrast, the ‘short‐branch clade’ dispersed once into Asia via the boreotropics during the Early Miocene and further dispersal was limited.     

Phylogeny and biogeography of Rhododendron subsection Pontica, a group with a tertiary relict distribution

Rhododendron subgenus Hymenanthes subsection Pontica is exceptional among Tertiary relict groups in having a high proportion of species (4 of 11) native to SW Eurasia. A phylogeny based on cpDNA matK and trnL-F indicated that multiple Pontica lineages colonised each of SW Eurasia, SE North America, and NE Asia, with little or no speciation within regions thereafter. Therefore, multiple (3-4) Pontica lineages survived the Quaternary in SW Eurasia, in contrast to other Tertiary relict genera. Pontica comprises two major clades, one of which is wholly Eurasian, and paraphyletic with respect to at least some of the remaining 200 species of subgenus Hymenanthes, which are all distributed in SE Asia. The other clade has species from W and SE North America, SW Eurasia, and NE Asia. According to synonymous matK substitution data, the two clades diverged 9-6 million years ago (mya), whereas most divergence within them happened 5-3 mya. Although the phylogeny indicates probable trans-Atlantic migration for one of two America-Eurasia disjunctions in Pontica, the timing supports migration via Beringia for both.     

New Eocene fossil fruits and leaves of Menispermaceae from the central Tibetan Plateau and their biogeographic implications

Menispermaceae are a pantropical and temperate family with an extensive fossil record during the Paleogene period, especially in North America and Europe, but with much less evidence from Asia. The latest fossil evidence indicates a succession of tropical to subtropical flora on the central Tibetan Plateau during the Paleogene. However, the biogeographic histories of these floras are still unresolved. Here, we report on endocarps and leaves of Menispermaceae from the Middle Eocene of Jianglang village, Bangor County, central Tibetan Plateau. The endocarps belong to two genera: Stephania, which is characterized by a horseshoe-shaped endocarp and with one lateral crest ornamented by spiny to rectangular ribs, and a condyle area; and Cissampelos (s.l.), which has two characteristic lateral ridges and a conspicuous external condyle. Associated leaves belong to the genus Menispermites, and are characterized by actinodromous primary venation, brochidodromous secondary veins, entire margins, and the presence of marginal secondary veins. The biogeographic history of Menispermaceae is complex, but evidence from these new fossils indicates an early diversification of the group in Asia, probably in response to the warming climate during the Eocene. The Jianglang flora appears to be part of a boreotropical flora, connecting Asia with North American and European floras during the Middle Eocene. The modern distribution of menispermaceous taxa found in Jianglang, as well as other families represented in the Jianglang flora, show that a tropical to subtropical climate occurred during the Eocene in central Tibet.     

Molecular phylogeny and biogeographic diversification of Parthenocissus (Vitaceae) disjunct between Asia and North America

? Premise of the study: Parthenocissus is a genus of the grape family Vitaceae and has a disjunct distribution in Asia and North America with members in both tropical and temperate regions. The monophyly of Parthenocissus has not yet been tested, and the species relationships and the evolution of its intercontinental disjunction have not been investigated with extensive sampling and molecular phylogenetic methods. ? Methods: Plastid (trnL-F, rps16, and atpB-rbcL) and nuclear GAI1 sequences of 56 accessions representing all 12 Parthenocissus species were analyzed with parsimony, likelihood, and Bayesian inference. Divergence times of disjunct lineages were estimated with relaxed Bayesian dating. Evolution of the leaflet number was assessed by tracing this character onto Bayesian trees using the Trace Character Over Trees option in the program Mesquite. ? Key results: Parthenocissus is monophyletic and sister to the newly described segregate genus Yua. Two major clades within Parthenocissus are recognizable corresponding to their distribution in Asia and North America. The disjunction between the two continents is estimated to be at 21.64 (95% higher posterior densities 10.23-34.89) million years ago. ? Conclusions: Parthenocissus is likely to have derived from the Eocene boreotropical element. Its current Asian-North American disjunction is dated to the early Miocene, congruent with fossil and paleoclimatic evidence. The tropical species is nested within the temperate clade and is inferred to have dispersed from the adjacent temperate regions. Parthenocissus and Yua are best treated as distinct genera. Leaflet number in this genus has a complex history and cannot be used as a character for infrageneric classification.     

Phylogeny,niche conservatism and the latitudinal diversity gradient in mammals

Biologists have long searched for mechanisms responsible for the increase in species richness with decreasing latitude. The strong correlation between species richness and climate is frequently interpreted as reflecting a causal link via processes linked to energy or evolutionary rates. Here, we investigate how the aggregation of clades, as dictated by phylogeny, can give rise to significant climate–richness gradients without gradients in diversification or environmental carrying capacity. The relationship between climate and species richness varies considerably between clades, regions and time periods in a global-scale phylogenetically informed analysis of all terrestrial mammal species. Many young clades show negative richness–temperature slopes (more species at cooler temperatures), with the ages of these clades coinciding with the expansion of temperate climate zones in the late Eocene. In carnivores, we find steeply positive richness–temperature slopes in clades with restricted distributions and tropical origins (e.g. cat clade), whereas widespread, temperate clades exhibit shallow, negative slopes (e.g. dog–bear clade). We show that the slope of the global climate–richness gradient in mammals is driven by aggregating Chiroptera (bats) with their Eutherian sister group. Our findings indicate that the evolutionary history should be accounted for as part of any search for causal links between environment and species richness.     

Phylogeny and biogeography of Altingiaceae: evidence from combined analysis of five non-coding chloroplast regions

The Altingiaceae consist of approximately 15 species that are disjunctly distributed in Asia and North America. The genus Liquidambar has been employed as a biogeographic model for studying the Northern Hemisphere intercontinental disjunctions. Parsimony and Bayesian analyses based on five non-coding chloroplast regions support that (1) Liquidambar is paraphyletic; (2) the temperate Liquidambar acalycina and Liquidambar formosana are nested within a large tropical to subtropical Asian clade; (3) Semiliquidambar is scattered in the eastern Asian clade and is of hybrid origin involving at least two maternal species: L. formosana and L. acalycina; and (4) the eastern North American Liquidambar styraciflua groups with the western Asian Liquidambar orientalis, but is highly distinct from other lineages. Biogeographically, our results demonstrate the complexity of biogeographic migrations throughout the history of Altingiaceae since the Cretaceous, with migration across both the Bering and the North Atlantic land bridges.     

Phylogeny of Acridocarpus-Brachylophon (Malpighiaceae): implications for tertiary tropical floras and Afroasian biogeography

Abstract.— A major tenet of African Tertiary biogeography posits that lowland rainforest dominated much of Africa in the late Cretaceous and was replaced by xeric vegetation as a response to continental uplift and consequent widespread aridification beginning in the late Paleogene. The aridification of Africa is thought to have been a major factor in the extinction of many African humid-tropical lineages, and in the present-day disparity of species diversity between Africa and other tropical regions. This primarily geologically based model can be tested with independent phylogenetic evidence from widespread African plant groups containing both humid- and xeric-adapted species. We estimated the phylogeny and lineage divergence times within one such angiosperm group, the acridocarpoid clade (Malpighiaceae), with combined ITS, ndhF , and trnL-F data from 15 species that encompass the range of morphological and geographic variation within the group. Dispersal-vicariance analysis and divergence-time estimates suggest that the basal acridocarpoid divergence occurred between African and Southeast Asian lineages approximately 50 million years ago (mya), perhaps after a southward ancestral retreat from high-latitude tropical forests in response to intermittent Eocene cooling. Dispersion of Acridocarpus from Africa to Madagascar is inferred between approximately 50 and 35 mya, when lowland humid tropical forest was nearly continuous between these landmasses. A single dispersal event within Acridocarpus is inferred from western Africa to eastern Africa between approximately 23 and 17 mya, coincident with the widespread replacement of humid forests by savannas in eastern Africa. Although the spread of xeric environments resulted in the extinction of many African plant groups, our data suggest that for others it provided an opportunity for further diversification.     

Molecular phylogeny and historical biogeography of the Holarctic wetland leaf beetle of the genus Plateumaris

Leaf beetles of the genus Plateumaris inhabit wetlands across the temperate zone of the Holarctic region. To explore the phylogeographic relationships among North American, East Asian, and European members of this genus and the origin of the species endemic to Japan, we studied the molecular phylogeny of 20 of the 27 species in this genus using partial sequences of mitochondrial cytochrome oxidase subunit I (COI) and the 16S and nuclear 28S rRNA genes. The molecular phylogeny revealed that three species endemic to Europe are monophyletic and sister to the remaining 11 North American and six Asian species. Within the latter clade, North American and Asian species did not show reciprocal monophyly. Dispersal-vicariance analysis and divergence time estimation revealed that the European and North America-Asian lineages diverged during the Eocene. Moreover, subsequent differentiation occurred repeatedly between North American and Asian species, which was facilitated by three dispersal events from North America to Asia and one in the opposite direction during the late Eocene through the late Miocene. Two Japanese endemics originated from different divergence events; one differentiated from the mainland lineage after differentiation from the North American lineage, whereas the other showed a deep coalescence from the North American lineage with no present-day sister species on the East Asian mainland. This study of extant insects provides molecular phylogenetic evidence for ancient vicariance between Europe and East Asia-North America, and for more recent (but pre-Pleistocene) faunal exchanges between East Asia and North America.     

Intercontinental disjunctions between eastern Asia and western North America in vascular plants highlight the biogeographic importance of the Bering land bridge from late Cretaceous to Neogene

This review shows a close biogeographic connection between eastern Asia and western North America from the late Cretaceous to the late Neogene in major lineages of vascular plants (flowering plants, gymnosperms, ferns and lycophytes). Of the eastern Asian–North American disjuncts, conifers exhibit a high proportion of disjuncts between eastern Asia and western North America. Several lineages of ferns also show a recent disjunct pattern in the two areas. In flowering plants, the pattern is commonly shown in temperate elements between northeastern Asia and northwestern North America, as well as elements of the relict boreotropical and Neogene mesophytic and coniferous floras. The many cases of intercontinental biogeographic disjunctions between eastern Asia and western North America in plants supported by recent phylogenetic analyses highlight the importance of the Bering land bridge and/or the plant migrations across the Beringian region from the late Cretaceous to the late Neogene, especially during the Miocene. The Beringian region has permitted the filtering and migration of certain plant taxa since the Pliocene after the opening of the Bering Strait, as many conspecific taxa or closely related species occur on both sides of Beringia.     

Biogeographical Divergence of the Flora of Yunnan,Southwestern China Initiated by the Uplift of Himalaya and Extrusion of Indochina Block

The floral composition of Yunnan is conspicuously linked to the biogeographical history of this extremely species-rich province in southwestern China. The floristic compositions of three representative regions in Yunnan were compared to reveal their variation with geography. From southern Yunnan, 4150 native species (including subspecies and varieties) from 1240 genera and 183 families of seed plants were recognized. From central Yunnan 3389 native species from 1095 genera and 167 families of seed plants were recognized. From northwestern Yunnan 6807 native species from 1296 genera and 166 families of seed plants were recognized. Although these three floras across Yunnan are similar in familial composition, similarities between the floras of southern and northwestern Yunnan are low at the generic and specific levels. The flora of northwestern Yunnan is dominated by families and genera with cosmopolitan and north temperate distributions, while the flora of southern Yunnan is dominated by tropical families and genera. Northwestern Yunnan is composed largely of temperate genera, of which the highest proportion has a north temperate distribution. In contrast, southern Yunnan has mainly tropical genera, of which most have a tropical Asian distribution. The flora of central Yunnan is a combination of southern and northwestern Yunnan. These three floras might be derived from a common Tertiary tropical or subtropical East Asian flora, but the geological history of each region has influenced its flora, and they have remained divergent since the late Tertiary. The flora of northwestern Yunnan has evolved with the uplift of the Himalayas and by gradual proliferation of mainly cosmopolitan and north temperate floristic elements, while the flora of southern Yunnan has evolved with extrusion of the Indochina block and the influence of mainly tropical Asian elements.     

A fossil-calibrated relaxed clock for Ephedra indicates an Oligocene age for the divergence of Asian and New World clades and Miocene dispersal into South America

Ephedra comprises approximately 50 species, which are roughly equally distributed between the Old and New World deserts, but not in the intervening regions （amphitropical range）. Great heterogeneity in the substitution rates of Gnetales （Ephedra, Gnetum, and Welwitschia） has made it difficult to infer the ages of the major divergence events in Ephedra, such as the timing of the Beringian disjunction in the genus and the entry into South America. Here, we use data from as many Gnetales species and genes as available from GenBank and from a recent study to investigate the timing of the major divergence events. Because of the tradeoff between the amount of missing data and taxon/gene sampling, we reduced the initial matrix of 265 accessions and 12 loci to 95 accessions and 10 loci, and further to 42 species （and 7736 aligned nucleotides） to achieve stationary distributions in the Bayesian molecular clock runs. Results from a relaxed clock with an uncorrelated rates model and fossil-based calibration reveal that New World species are monophyletic and diverged from their mostly Asian sister clade some 30 mya, fitting with many other Beringian disjunctions. The split between the single North American and the single South American clade occurred approximately 25 mya, well before the closure of the Panamanian Isthmus. Overall, the biogeographic history of Ephedra appears dominated by long-distance dispersal, but finer-scale studies are needed to test this hypothesis.