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.     

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.     

湖南省石松类和蕨类植物分布新记录

在2004年出版的《湖南植物志》中收录有标本考证的湖南省石松类和蕨类植物683种41变种10变型[1],截至2014年底,又陆续发表了70余个新记录[2-9]。2013年出版的《Flora of China》结合现代分子系统学和最新的研究成果,对大量植物物种进行归并和科属的重新划分,作为湖南省优势科属资源的一些种类数量也相应减少,如水龙骨科( Polypodiaceae)的鳞果星蕨属( Lepidomicrosorium Ching et K. H. Shing )和盾蕨属( Neolepisorus Ching ),鳞毛蕨科( Dryopteridaceae)的复叶耳蕨属( Arachniodes Blume)以及以武陵山区为中心分布的黔蕨属( Phanerophlebiopsis Ching)(归并到复叶耳蕨属Arachniodes Blume)等,书中仅记录湖南省分布的石松类和蕨类植物约434种(含变种、亚种和杂种)[10],数量相对较少,而湖南省分布的一些较珍稀的蕨类植物未有罗列,如川黔肠蕨也Diplaziopsis cavaleriana ( Christ) C. Chr.页、中华双扇蕨( Dipteris chinensis Christ )、全缘燕尾蕨也Cheiropleuria integrifolia ( D. C. Eaton ex Hook.) M. Kato页、睫毛蕨也Pleurosoriopsis makinoi ( Maxim. ex Makino) Fomin页[1]和心叶薄唇蕨也Leptochilus cantoniensis ( Baker) Ching页等[11]。作者根据近十年的相关文献[1-9,12],对湖南省分布的石松类和蕨类植物进行了梳理,统计出631种27变种2亚种和2杂种。自2013年开始,作者对湖南省东南部的莽山、西部的小溪以及西北部的八大公山等多个国家级自然保护区进行了野外考察,发现了湖南省石松类和蕨类植物新记录6种,分属于5科6属,它们的生境及形态特征见图1。凭证标本保存于上海辰山植物标本馆( CSH )。其中一些种类分布的海拔较低,对于湖南省蕨类植物区系特点、科属结构以及物种多样性演化等研究具有潜在价值。     

黄土高原石松类和蕨类植物的多样性与地理分布

物种编目是了解生物多样性的基础。本文收集已出版的专著和文献, 查阅标本馆的馆藏标本信息, 在此基础上, 对黄土高原的石松类和蕨类植物的多样性和地理分布数据进行统计和分析。科属排列采用Flora of China的分类系统, 科属界定依据分子系统学的最新研究成果。结果表明, 黄土高原共有石松类和蕨类植物18科42属165种2亚种5变种, 其中中国特有种35种, 占总种数的20.35%。种类数量最多的5个科依次为凤尾蕨科(29种, 含种下单位, 下同)、鳞毛蕨科(28种)、水龙骨科(19种)、蹄盖蕨科(16种)和铁角蕨科(16种); 种类数量最多的6个属依次为铁角蕨属(Asplenium, 16种)、鳞毛蕨属(Dryopteris, 15种)、耳蕨属(Polystichum, 12种)、岩蕨属(Woodsia, 11种)、卷柏属(Selaginella, 10种)和瓦韦属(Lepisorus, 10种)。黄土高原的石松类和蕨类植物中, 无珍稀濒危保护植物, 无中国特有属。依据生态类型可划分为4类, 即土生植物(112种)、石生植物(75种)、附生植物(18种)、水生植物(3种), 该区域石松类和蕨类植物土生或石生的种类占总种数的89.53%。在地理分布上, 黄土高原石松类和蕨类处于我国现生类群分布的边缘区域, 这与我国的石松类和蕨类分布规律基本一致。     

山东石松类和蕨类植物新记录

该研究报道了山东地区4种1变种和1变型石松类和蕨类植物新记录,分别是石松类的伏地卷柏( Sel-aginella nipponica Franch. et Sav.),蕨类的小戟叶耳蕨[Polystichum hancockii (Hance) Diels]、阔羽贯众(Cyrtomium yamamotoi Tagawa)、粗齿阔羽贯众[C. yamamotoi Tagawa var. intermedium (Diels) Ching et Shing ex Shing]、小羽贯众[C. fortunei J. Sm. f. polypterum (Diels) Ching]、远叶瓦韦[Lepisorus ussuriensis var. distans (Makino) Taga-wa]。同时采用扫描电镜对其孢子形态进行了观察,该研究结果为其分类鉴定提供了孢粉学依据。     

Evolution of the Eastern Asian and Eastern North American disjunct genus Symplocarpus (Araceae): insights from chloroplast DNA restriction site data

Chloroplast DNA (cpDNA) variation was surveyed with 20 restriction endonucleases for the eastern Asian and eastern North American disjunct genus Symplocarpus (Araceae). The cpDNA phylogeny reveals a sister group relationship between S. foetidus from eastern North America and S. renifolius from eastern Asia. The cpDNA divergence between the two intercontinental sister species is 0.61%, which suggests an estimated divergence time of 6.1 million years ago during the late Miocene. The Bering land bridge hypothesis is compatible with the estimated time of divergence for the migration of Symplocarpus between eastern Asia and North America. Furthermore, a single origin of the exothermic spadices in Symplocarpus is suggested by the phylogeny. The cpDNA data also provide independent support for the recognition of three species within the genus.     

石松类和蕨类植物质体基因组结构演化研究进展

近年来, 随着测序技术的发展, 石松类和蕨类植物的核基因组、质体基因组以及线粒体基因组研究发展迅速, 质体基因组研究工作更是呈爆发式增长。截至2019年3月1日, GenBank公布的石松类和蕨类植物的175个质体基因组中, 约3/4为最近两年新增。研究内容从早期对个别质体基因组结构和序列特征的简单报道, 逐渐发展到综合性的比较基因组学和系统发育基因组学研究。目前已发表的质体基因组覆盖了石松类和蕨类植物的所有目和大部分科, 这两大类群的质体基因组结构变异和系统发育的基本框架已逐渐清晰。这些研究为我们理解维管植物的早期演化提供了重要参考。本文对石松类和蕨类植物的质体基因组结构特征进行了系统梳理, 发现其结构变异主要包括大片段倒位、IR区边界变动、基因或内含子丢失等, 其中一些结构变异可作为较高分类阶元的共衍征。RNA编辑和长片段非编码序列插入普遍存在于石松类和蕨类植物的质体基因组中, 但其起源、演化机制和功能等仍不清楚。我们对质体基因组的应用、系统发育研究中质体和核基因组的优劣性, 以及系统发育基因组学的前景进行了评述。     

Morphological and molecular comparisons ofCampsis grandiflora andC. radicans (Bignoniaceae), an eastern Asian and eastern North American vicariad species pair

Morphological and molecular comparisons were made forCampsis grandiflora (Thunb.)K. Schumann (Bignoniaceae) from eastern Asia andC. radicans (L.)Seemann from eastern North America. Chloroplast DNA (cpDNA) variation was surveyed with 20 restriction endonucleases. The cpDNA divergence between the two vicariad species was 2.44%, which is the highest reported among North Temperate disjunct taxa and one of the highest reported for infrageneric taxa. Detailed morphological comparisons also suggest a high level of divergence. Cluster analyses based on 22 morphological characters and 39 OTUs revealed two distinct groups corresponding with the two species. The average taxonomic distance between the two species was 1.806. Analyses of variance (ANOVA) revealed that 12 of the 18 quantitative characters differed significantly ( 0.01) betweenC. grandiflora andC. radicans. Divergence time based on cpDNA data was estimated as 24.4 million years. The Bering land bridge hypothesis was favored over the North Atlantic land bridge hypothesis based on the estimated divergence time and the geological history of the North Temperate region. The high levels of morphological and cpDNA divergence are not consistent with morphological stasis, which has been proposed as a common mode of evolution for North Temperate disjunct taxa.     

Differential diversifications of South American and Eastern Asian disjunct genera Bocconia and Macleaya (Papaveraceae)

Bocconia (10 species) and Macleaya (2 species) are two disjunct genera between South America and eastern Asia (EAS) in the Papaveraceae offering an opportunity to compare its biogeographic history with that of the well‐known disjunction between EAS and eastern North American (ENA). Our phylogenetic analyses of the chloroplast matK and rbcL gene sequences of Ranunculales including two species of Macleaya and six species of Bocconia supported the monophyly of Bocconia, Macleaya, and Chelidonioideae to which Bocconia and Macleaya belong. Nucleotide sequences of matK, rbcL, and nrDNA ITS supported the sister relationship of Bocconia and Macleaya. Biogeographic analyses of Chelidonioideae using S‐DIVA (statistical dispersal vicariance analysis) and DEC (dispersal extinction cladogenesis) methods inferred Eurasia as the most likely ancestral area of Bocconia and Macleaya and suggested no extinction events in either Bocconia or Macleaya. This agrees with the “Out‐of‐Asia” pattern of the EAS‐ENA disjunction. Molecular dating of Ranunculales with fossil‐based calibrations showed that Bocconia and Macleaya diverged in the late Eocene and early Oligocene, which is much earlier than most EAS‐ENA disjunct taxa. The disjunction may have formed via long distance dispersal or boreotropical connections via the North Atlantic and Bering land bridges. Both Bocconia and Macleaya diversified in the late mid‐Miocene, but Bocconia has apparently experienced a greater diversification probably aided by the evolution of the bird dispersal syndrome in fruit and seed after migration to South America. The greater diversification of Bocconia is also evidenced by the diverse leaf morphology and growth habit in response to colonization in various local habitats in South America.     

Phylogeny and biogeography of Sassafras (Lauraceae) disjunct between eastern Asia and eastern North America

Sassafras (Lauraceae) consists of three species disjunct between eastern Asia (S. tzumu and S. randaiense) and eastern North America (S. albidum). Phylogenetic analysis based on sequences of nuclear ribosomal ITS and three chloroplast non-coding regions (rpl16, trnL-F, and psbA-trnH) showed that Sassafras is monophyletic and that the eastern North American S. albidum is sister to the clade of its two eastern Asian counterparts. Their intercontinental divergence was estimated to be 13.80 ± 2.29−16.69 ± 2.52 million years ago (mya) using the penalized likelihood method with the ITS and three chloroplast markers. Biogeographic analyses combined with fossil evidence suggest that Sassafras has a relict distribution in the Northern Hemisphere without a Gondwanan link. The divergence time of the two eastern Asian species (the continental Chinese Sassafras tzumu and S. randaiense endemic to Taiwan) is estimated to be 0.61 ± 0.75−2.23 ± 0.76 mya. Sassafras randaiense from Taiwan was most likely derived from an ancestor from continental China.     

Ribosomal DNA evidence and disjunctions of western American Portulacaceae

Phylogenetic analysis of ribosomal DNA internal transcribed spacer sequences from 35 members of western American Portulacaceae plus seven Portulacaceae outgroups generally supports morphologically based interpretations of multiple intercontinental disjunctions. The data neither support nor refute monophyly of the western American group but strongly support a group comprising the western American taxa plus Phemeranthus, the only strictly American genus of the morphology-based eastern American/African group of Portulacaceae, along with the Australian genus Parakeelya. Support is strong for the monophyly of Calandrinia, Montiopsis, Lewisia, Claytonia, and Montia, along with a sister relationship of the last two. The data neither strongly support nor refute the morphologically based diagnosis of Cistanthe, but they strongly support a clade including the North American Cistanthe section Calyptridium and the South American Cistanthe sections Amarantoideae and Philippiamra. The internal transcribed spacer data fail to resolve the phylogenetic relationships among most of the western American lineages, suggesting either rapid radiation or, alternatively, erratic evolution of the internal transcribed spacer. The internal transcribed spacer and morphological evidence together suggest that in this group there have been 8-13 dispersal and colonization events across >2000 km (1 for every 15-26 extant species in this group). The internal transcribed spacer data document complex molecular evolutionary patterns, including strong substitution biases, among-site rate heterogeneity, positional bias for deamination-type substitutions, nonstationarity, and variable rates of insertion/deletion. Our phylogenetic conclusions, however, do not appear to be sensitive to these patterns.     

The structure and development of haustorial placentas in leptosporangiate ferns provide a clear-cut distinction between euphyllophytes and lycophytes

This light and electron microscope study revealed that leptosporangiate ferns have highly distinctive gametophyte-sporophyte junctions characterized by sporophytic haustoria, the absence of intraplacental spaces and degenerating cells, and the early appearance of wall ingrowths in both generations. Other notable cytological features are highly pleomorphic plastids and mitochondrial aggregates in the gametophytic placental cells. Close similarities with the gametophyte-sporophyte junctions in Tmesipteris and major differences from those of homosporous lycophytes are in line with the placement of psilophytes and ferns in the same clade and distance both from lycophytes. A smooth interface between the two generations in Azolla suggests a clear-cut discontinuity between homosporous and heterosporous ferns, although this is the only heterosporous fern investigated to date. Similarities between the gametophyte-sporophyte junctions of leptosporangiate ferns and hornworts, when balanced against differences between them, are considered more likely the result of parallel evolution rather than homology.     

Phylogenetic relationships and biogeographic history of the unique Saxifraga sect. Irregulares (Saxifragaceae) from eastern Asia

Saxifraga L. is the largest genus in Saxifragaceae and a characteristic component of the herbaceous flora of the temperate and alpine mountains in the Northern Hemisphere. Section Irregulares is a small group of 15–20 species, representing one of the early‐diverged lineages in the genus characterized with unique zygomorphic flowers. We used both nuclear internal transcribed spacer and chloroplast DNA regions (psbA‐trnH, trnL‐F, and matK) to reconstruct its species relationships, estimate divergence times, and infer its historical biogeography. Our phylogenetic results corroborate the monophyly of sect. Irregulares and its sister relationship to sect. Heterisia from North America. The section was well resolved into two lineages corresponding to their morphological features and biogeographic distributions. One represents ser. Stoloniferae including S. stolonifera Curtis and S. nipponica Makino with long‐creeping stolons/rhizomes and small petals with spots and the other comprises the remaining taxa (ser. Rufescentes) which lack long‐creeping rhizomes. Spots on leaves (abaxially spotted vs. abaxially without spots) and spots on petals (without spots vs. with spots) are inferred to be phylogenetically informative within ser. Rufescentes. Divergence time estimates and ancestral area analysis suggested a western North American origin of sect. Irregulares with migration into East Asia by way of the Bering land bridge in the Middle Oligocene. The development of drying and desertification belt in the late Miocene could have played an important role in the subsequent restriction and separation of the north and south lineages within eastern Asia.