The phytogeographical studies of Thermopsis (Fabaceae)

The present article is the first comprehensive treatment of phytogeography of Thermopsis (Fabaceae) in the world. Thermopsis is one of the few genera within Fabaceae with the distribution pattern of the East Asia-North American disjunction. The distribution patterns of 5 recognized sections (including a new one) covering 21 species in Thermopsis are analyzed, and the results show four centres of frequency of the genus: the Eastern Asiatic Region (9 spp. / 3 sects., including 4 endemic species), the Irano-Turanian Region (7 spp./3 sects., including 3 endemic species), the Rocky Mountain Region (7 spp./2 sects., all endemic), and the Atlantic North American Region (3 spp. / 1 sect., all endemic). In the light of the fact that most species and sections, a number of phylogenetic series of the genus, and the most primitive sections and most advanced sections in Thermopsis occur in the East Asia, the Eastern Asiatic Region might be the centre of diversity of the genus. As the Irano-Turanian Region and the Rocky Mountain Region were just second to that of Eastern Asiatic Region in number of sections and species, and many polyploids appeared in these regions, they were considered as the secondary centres of distribution and speciation of the genus. The speciation looks to be frequent and complex in these regions, and many new taxa have been described from there while many new reduced or incorporated taxa have happened over there. However, recent molecular data has shown that two reduced taxa of Thermopsis are distinct in these regions. Based on the modern distribution patterns and evolutionary trends in morphological characters of the genus, and available fossil record of the genus and the historical geology, we speculate that Thermopsis had already existed on Eurasia and North America before the Late Miocene, and probably originated from an ancestral form of Sophora-like taxa with lupine alkaloids somewhere in the Laurasia in the Early Tertiary or Late Cretaceous. After the separation of the two continents, species on different continents developed distinctly under influences of different evolutionary factors. In Asia, the late Tertiary orogeny, disappearing of the Tethys and aridity and freezing caused by the Quaternary glaciation were the main forces to promote the speciation and evolutionary processes, whereas in North America it was the Quaternary glaciation and the orogeny of partial area to promote evolution of the genus. According to the evolutionary trends in Thermopsis and the distribution pattern of the primitive taxa, Sino-Japanese Subregion of Eastern Asiatic Region may be considered asthe centre of primitive forms of Thermopsis.     

The Origin,Dispersal and Formation of the Distribution Pattern of Swertia L. (Gentianaceae)

The genus Swertia is one of the large genera in Gentianaceae, including 154 species, 16 series and 11 sections. It is disjunctly distributed in Europe, Asia, Africa and N. America, but entirely absent from Oceania and S. America. According to Takhtajan’s (1978) regionalization of the world flora, Swertia is found in 14 regions. Eastern Asiatic region with 86 species, of which 58 are local endemics, 13 series and 9 sections, ranks the first among all the regions. The highest concentration of the taxa and endemics in Eastern Asiatic region occurs in SW China-Himalayan area (Sikang-Yunnan P. , W. Sichuan, W. Yunnan-Guichou Plateau of China and NE. Burma, N. Burmense P. , E. Himalayan P. and Khasi-Manipur P. ). In this area there are 74 species (48 endemics), 12 series, and 9 sections; thus about half species of the world total, three quarters of series and 82% of sections occur in this small area. Besides, the taxa at different evolutionary stages in Swertia also survive here. It is an indication that SW. China-Himalayan area is a major distribution centre of the genus Swertia. In addition, Sudan-Zambezian Region in Africa, with 22 species, 4 series and 2 sections, is a second distribution centre. The primitive type of the genus Swertia is Sect. Rugosa which consists of 2 series and 23 species. It is highly centred in the mountains of SW. China (Yunnan, Sichuan, Guizhou and SE. Xizang) where 2 series and 16 species occur. Among them 15 species of Ser. Rugosae were considered as the most primitive groups in this genus. From our study, the outgroup of Swertia is the genus Latouchea Frahch. , which is distributed in Yunnan, Sichuan, Guizhou, Hunan, Guangdong, Guangxi and Fujian. The two groups overlap in distribution in SW. China. According to the principle of common origin, the ancestor of two genera ap peared most probably in this overlapping area. It was inferred that SW. China Was the birth-place of the genus Swertia. Four sections of Swertia have different disjunct distribution patterns: Sect. Ophelia is of Tropic Asia, Africa and Madagascar disjunct distribution; sect. Swertia is of north temperate distribution; sect. Spinosisemina is in Tropical Asia (Trop. India to S. China and Philipines); sect. Platynema also is in Tropical Asia (Java, Sumatra, Himalayas to SW. China). These disjunct patterns indicate that the Swertia floras between the continents or between continent and islands have a connection with each other. From paleogeographical analysis, Swertia plants dispersed to Madagascar before the Late Cretaceous, to SE. Asian Islands in the Pleistocene, to North America in the Miocene. The distribution of Swertia in Madagascar might be later than that in Asia. Therefore the origin time of the genus Swertia was at least not later than the Late Cretaceous, and might be back to the Mid-Cretaceous. The genus Swertia first fully developed and differentiated, forming some taxa at different evolutionary stages (Rugosa, Swertia, Poephila, Ophelia and Platynema etc. ) in the original area, and these taxa quickly dispersed in certain directions during the Late Cretaceous-Middle Tertiary when the global climate was warm and no much change. There seem to be three main dispersal routes from the origin area to different continents; (1) The westward route i. e. from SW. China, along the Himalayas area to Kashmir, Pakistan, Afghanistan and Iran, and then southwestwards into Africa throuth Arabia. Four sections (Poephila, Macranthos, Kingdon-Wardia and Ophelia) took this dispersal route. Most species of sect. Ophelia dispersed along this route, but a few along southern route and north ern route. Sect. Ophelia greatly differentiated in Africa and the African endemic sectionSect. Montana was derived from it. The two sections form there a second distribution center of Swertia. (2) The southward route, i. e. towards S. India through the Himalayas, and towards SE. Asian islands through C. and S. China, Indo-China. Along this dispersal route sect. Platynema, Sect. Spinosisemina and a few species of Sect. Ophelia dispersed; (3) The northward rout, i. e. northwards across N. China, C. Asia to a high latitude of Euasia, and also through E. Asia into N. America. The following groups took this route: sect. Rugosa, sect. Swertia, sect. Frasera, sect. Heteranthos and sect. Ophelia ser. Dichotomae. Therefore, it seems that the genus Swertia originated in SW. China and then dispersed from there to N. and S. Asia, Africa, Europe and North America and formed the moderndistribution pattern of this genus.     

木兰科（Magnoliaceae）的起源、进化和地理分布

木兰科为亚洲－美洲间断分布科,全世界有１５属,２４６种,主要分布于亚洲东南部的热带、亚热带地区,从喜马拉雅至日本,向南达新几内亚及新不列颠;少数种类分布于北美东南部、中美至南美巴西．中国有１１属,约９９种．木兰科的现代分布中心在东亚－东南亚地区．根据木兰科的化石记录、系统发育和现代分布,推测其起源时间为早白垩纪,甚至更早．起源地可能在中国的西南地区,并由此向外辐射,向东经日本、俄罗斯远东地区经白令陆桥进入北美;向西经西亚、欧洲,通过格陵兰进入北美,然后到达南美;向南经印度支那、马来西亚,直至新几内亚．东亚－北美间断分布的形成是受第四纪冰期的影响;南美的木兰科是从北美迁移而来．     

木兰科（Magnoliaceae）的起源、进化和地理分布

木兰科为亚洲－美洲间断分布科,全世界有１５属,２４６种,主要分布于亚洲东南部的热带、亚热带地区,从喜马拉雅至日本,向南达新几内亚及新不列颠;少数种类分布于北美东南部、中美至南美巴西．中国有１１属,约９９种．木兰科的现代分布中心在东亚－东南亚地区．根据木兰科的化石记录、系统发育和现代分布,推测其起源时间为早白垩纪,甚至更早．起源地可能在中国的西南地区,并由此向外辐射,向东经日本、俄罗斯远东地区经白令陆桥进入北美;向西经西亚、欧洲,通过格陵兰进入北美,然后到达南美;向南经印度支那、马来西亚,直至新几内亚．东亚－北美间断分布的形成是受第四纪冰期的影响;南美的木兰科是从北美迁移而来．     

报春花科植物的地理分布

根据Ｔａｋｈｔａｊａｎ世界植物区系分区对报春花科２２属在世界各地以及在中国各省区的分布作了较详细的统计,在此基础上,将报春花科各属归纳为１０个分布型,认为中国西部横断山区和东西马拉雅为报春花科的现代分布中心和多样化中心;高加索—阿尔卑斯山脉为第二分布中心;中国云南、贵州南部,广西西部至越南、泰国北部和缅甸西北部山地是报春花科植物最可能的起源中心;报春花科的起源时间应在早第三纪或晚白垩纪．     

The Evolution and Distribution of Subfam. Spiraeoideae (Rosaceae) of China,with Special Reference to Distribution of the Subfamily in the World

The subfam. Spiraeoideae, consisting of 22 genera and more than 260 species in the world,is the most primitive subfamily of Rosaceae. It has developed into two groups,i.e. evergreen and deciduous ones, of which eight genera and 100 species in China are totally deciduous. In the present paper, the origin,evolution and distribution of the Chinese genera is discussed mainly, and the distribution of the whole subfamily in the floristic regions of the world is also mentioned. Based on evolutionary trends of morphological characters, Spiraea L. is considered as the most primitive genus in the deciduous group of subfam. Spiraeoideae, from which some genera are been derived, the systematic position and evolutionary relationships between different genera are elucidated in this paper. Through the analysis on the geographical distribution of the genera in China, the areal types may be divided as follows: (1) North Temperate Type: Spiraea, Physocarpus, Aruncus. (2) East Asian and North American Disjunct Type: Sorbaria. (3) Mediterranean, West Asian (or Central Asia) and East Asian Type: Sibiraea. (4) Temperate Asian Type: Exochorda.(5) East Asian Type: (a) Sino Himalayan Distribution: Neillia; (b) Sino Japan Distribution: Stephanandra. After analysis of the distribution of subfam. Spiraeoideae in the world, it is shown that the Eastern Asiatic Region, being the richest in genera, species and endemic species of the world,is not only the center of distribution and differentiation,but also an important region for occurrence and development of some deciduous genera of this subfamily, while in North America, the Madrean Region and Rocky Mountain Region, genera, species and endemic species are abundant, which indicates that the western part of North America is also the distribution center of this subfamily at the present, but it may be the secondary center of distribution. It can be seen that the relatively primitive and evergreen g enera, i.e. Quillaja and Kageneckia, are now confined to South America. The fact implies that the South America may be the region for early differentiation and development of the evergreen genera in Subfam. Spiraeoideae. The analysis of Chinese plants has shown that China has the most members of the subfamily in Eastern Asiatic Region, with eight genera, 82 species and 62 endemic species and that the maximum concentration is in western Sichuan, northwestern Yunnan and their adjacent areas. It is very obvious that the center of distribution and diversity of Subfam. Spiraeoideae in China lies in the Hengduan Mountain Region of Sino Himalayan Forest Subkingdom and the western part of Sino Japan Forest Subkingdom, where may be the birthplace of some genera in China. It may be considered that the deciduous genera of Subfam. Spiraeoideae might have originated in Laurasia.According to the fossil records, the time of origin of Subfam.Spiraeoideae dates back to the Lower Cretaceous.     

叶蜂总科昆虫生物地理研究II叶蜂总科广布属的地理分布特性(膜翅目)

本文系统分析了叶蜂总科广布属的地理分布特性。叶蜂总科广布属被分为１２个主要的分布类型,其中全北界分布型６９属,可再分为６种次类型。在各分布型下列举了全部具有该类分布特征的叶蜂总科属名,并提出了一些有关起源与扩散的设想和推论。在广布型属的地理分布研究基础上,对各大生物地理界之间的关系也提出了一些看法。     

中国蔷薇科绣线菊亚科的演化、分布——兼述世界绣线菊亚科植物的分布

绣线菊亚科是蔷薇科最原始的亚科,共有22属260余种, 包括常绿和落叶两大类群,前者是 原始类型。我国有8属100种,全都为落叶性。本文着重讨论中国各属的起源、演化和分布等 ,同时也概述全亚科植物在世界各植物区的分布等问题。绣线菊属Spiraea是该亚科落叶类群中最原始的属,它在早期发生趋异进化,衍生出形态各异而亲缘关系密切 的不同属,本文阐明了中国各属的系统位置和属间的亲缘关系。通过对我国各属地理分布的 分析对比,属的分布区可归纳为5个类型。对全球绣线菊亚科植物在世界各植物区中的属、种数统计表明,东亚区有8属105种,其中有96个特有种,是该亚科植物分布最多而又最集中 地区,具有在系统发育上处于各主要演化阶段的落叶类型,因此,东亚区是全球绣线菊亚科植 物的现代分布和分化中心,也是落叶类群发生和发展的关键地区。在北美洲,从马德雷区至落基山区一带分布着11属46种,均为特有种,显然北美洲西部也是该亚科植物的现代分布中心,但可能是第二分布中心。南美洲至今保存2个较古老的常绿属,即Quillaja和K ageneckia,基于此,南美洲可能是绣线菊亚科某些常绿属早期分化和发展的关键地区 。中国绣线菊亚科植物在东亚区占绝对优势,有8属82种,其中有62个特有种,分别占该区属 、种和 特有种数的100%、82%、和65%, 这些类群分布最密集地区是在中国喜马拉雅森林植物亚区 中的横断山脉地区和中国日本森林植物亚区的西部,这一带是中国绣线菊亚科的现代分布和多样性中心,很可能是某些属的发源地。由此看来,绣线菊亚科的落叶属可能起源于劳亚古陆。据化石记载,该亚科植物的起源时间可以追溯到白垩纪早白垩世。     

Origin,Differentiation, and Geographic Distribution of the Chenopodiaceae

Numbers of species and genera,endemic genera,extant primitive genera,relationship and distribution patterns of presently living Chenopodiaceae(two subfamilies,12 tribes,and 118 genera)are analyzed and compared for eight distributional areas,namely central Asia,Europe,the Mediterranean region,Africa,North America,South America, Australia and East Asia． The Central Asia,where the number of genera and diversity of taxa are greater than in other areas,appears to be the center of distribution of extant Chenopodiaceae．North America and Australia are two secondary centers of distribution． Eurasia has 11 tribes out of the 12,a total of 70 genera of extant chenopodiaceous plants,and it contains the most primitive genera of every tribe． Archiatriplex of Atripliceae,Hablitzia of Hablitzeae,Corispermum of Corispermeae,Camphorosma of Camphorosmaea,Kalidium of Salicornieae,Polecnemum of Polycnemeae,Alexandra of Suaedeae,and Nanophyton of Salsoleae,are all found in Eurasia,The Beteae is an Eurasian endemic tribe,demonstrating the antiquity of the Chenopodiaceae flora of Eurasia．Hence,Eurasia is likely the place of origin of chenopodiaceous plants． The presence of chenopodiaceous plants is correlated with an arid climate．During the Cretaceous Period,most places of the continent of Eurasia were occupied by the ancient precursor to the Mediterranean,the Tethys Sea．At that time the area of the Tethys Sea had a dry and warm climate．Therefore,primitive Chenopodiaceae were likely present on the beaches of this ancient land．This arid climatic condition resulted in differentiation of the tribes Chenopodieae,Atripliceae,Comphorosmeae,Salicornieae,etc．,the main primitive tribes of the subfamily Cyclolobeae. Then following continental drift and the Laurasian and Gondwanan disintegration, the Chenopodiaceae were brought to every continent to propagate and develop, and experience the vicissitudes of climates, forming the main characteristics and distribution patterns of recent continental floras. The tribes Atripliceae, Chenopodieae, Camphorosmeae, and Salicornieae of recent Chenopodiaceae in Eurasia, North America, South America, southern Africa, and Australia all became strongly differentiated. However, Australia and South America, have no genera of Spirolobeae except for a few maritime Suaeda species. The Salsoleae and Suaedeae have not arrived in Australia and South America, which indicates that the subfamily Spirolobeae developed in Eurasia after Australia separated from the ancient South America-Africa continent, and South America had left Africa. The endemic tribe of North America, the tribe Sarcobateae, has a origin different from the tribes Salsoleae and Suaedeae of the subfamily Spirolobeae. Sarcobateae flowers diverged into unisexuality and absence of bractlets. Clearly they originated in North America after North America had left the Eurasian continent. North America and southern Africa have a few species of Salsola, but none of them have become very much differentiated or developed, so they must have arrived through overland migration across ancient continental connections. India has no southern African Chenopodiaceae floristic components except for a few maritime taxa, which shows that when the Indian subcontinent left Africa in the Triassic period, the Chenopodiaceae had not yet developed in Africa. Therefore, the early Cretaceous Period about 120 million years ago, when the ancient Gondwanan and Laurasian continents disintegrated, could have been the time of origin of Chenopodiaceae plants.The Chinese flora of Chenopodiaceae is a part of Chenopodiaceae flora of central Asia. Cornulaca alaschnica was discovered from Gansu, China, showing that the Chinese Chenopodiaceae flora certainly has contact with the Mediterranean Chenopodiaceae flora. The contact of southeastern China with the Australia Chenopodiaceae flora, however, is very weak.     

棕榈科植物的地理分布

棕榈科是一个泛热带分布的科,共有１９８属,约２６７０种,下分６亚科,１４族。贝叶棕族是最原始的族,低地榈族则最进化。本科植物在世界上的分布可划分为１３个区,其中以印度－马来西区和新热带区的属、种最多。中国只有１６属和８５种,没有特有属。这些种大部分属热带亚洲分布,与热带亚洲植物区系关系非常密切。关于棕榈科起源地问题,有西冈瓦纳起源和劳亚起源之说。根据化石记录和形态特征的分析,棕榈科很可能于早白垩纪     

Construction of the phylogenetic model for the genera of the tribe Arctiini (Lepidoptera,Arctiidae) with the SYNAP method

A new scheme of the phylogeny of the tribe Arctiini is proposed. The Western Mediterranean genus Atlantarctia is considered the most primitive one in the tribe; the rest of genera form two large clades Arctia-Pericallia and Gonerda-Platyprepia. The first clade is supposed to have been subjected to radiation in western Eurasia, and the second clade, in Asia and North America in the Palaeogene when the eastern part of Asia was isolated from western Eurasia. Subsequently, most probably in the Neogene-Pleistocene, representatives of both clades spread over the whole Eurasia and North America. The Arctiini fauna of the tundra zone, which includes the genera Acerbia and Pararctia, was formed in Asia and North America, whereas the subboreal fauna (both steppe and nemoral) originated in western Eurasia. The boreal genus Borearctia has most likely also originated in Asia.     

藜科植物的起源、分化和地理分布

全球藜科植物共约130属1500余种,广泛分布于欧亚大陆、南北美洲、非洲和大洋洲的半干旱及盐碱地区。它基本上是一个温带科,对亚热带和寒温带也有一定的适应性。本文分析了该科包含的1l族的系统位置和分布式样,以及各个属的分布区,提出中亚区是现存藜科植物的分布中心,原始的藜科植物在古地中海的东岸即华夏陆台(或中国的西南部)发生,然后向干旱的古地中海沿岸迁移、分化,产生了环胚亚科主要族的原始类群;起源的时间可能在白垩纪初,冈瓦纳古陆和劳亚古陆进一步解体的时期。文章对其迁移途径及现代分布式样形成的原因进行了讨论。     

The genus Artemisia (Asteraceae: Anthemideae) at a continental crossroads: molecular insights into migrations, disjunctions, and reticulations among Old and New World species from a Beringian perspective

Artemisia is the largest genus (ca. 350-500+ spp.) in the tribe Anthemideae and is composed of ecologically, morphologically, and chemically diverse species that are found primarily throughout the Northern Hemisphere. Two major centers of diversity for the genus are located in Eurasia and western North America, but phytogeographic links connecting these two regions are observed all across the North Pacific Rim and adjacent areas in the Arctic, including many islands and archipelagos. Previous phylogenetic studies have helped to clarify major lineages and identify likely sister relationships, but many questions remain unanswered regarding the relationships and migration history of New and Old World species. Here we investigate the phylogenetics of Artemisia within a biogeographic context centered in the Beringian Region and offer new hypotheses concerning species relationships, migration history, and the likely role of reticulate evolution in the genus. Our sampling included many new taxa and emphasized multiple accessions of widespread species, species from proposed refugia, and species with disjunct/vicariant distributions. The ITS phylogeny contained 173 accessions (94 new and 79 from GenBank) and indicated that Artemisia is paraphyletic by the exclusion of several small Asian genera and the North American genus Sphaeromeria. Following a survey of thirteen chloroplast loci, phylogenies based on two plastid markers (psbA-trnH and rpl32-trnL spacers) were constructed with a reduced data set, and though largely consistent with the ITS topology, revealed several cases of possible introgression among New World and Beringian species. Our analysis reveals that North American Artemisia species have multiple origins, and that western North America has served as a source for some colonizing elements in eastern Asia and South America.     

粉条儿菜属(肺筋草科)的地理分布及其物种多样性和分化中心

粉条儿菜属(AletrisL.)隶属于肺筋草科,全世界有23种1变种,东亚有18种1变种,北美东南部有5种,为典型的东亚-北美间断分布的属.本文在种(变种)的水平上,研究了粉条儿菜属的地理分布及其分布中心和多样化中心,并对其起源和分化以及现代洲际间断分布格局的成因进行了分析.结果表明,(1)中国共分布有粉条儿菜属植物15种1变种,而广义的横断山地区集中分布有13种1变种,是东亚粉条儿菜属植物分布最为集中的地区,而且包含该属植物各个进化阶段的代表.因此,广义的横断山地区是粉条儿菜属在东亚的分布中心和多样化中心.(2)根据粉条儿菜属及其近缘属的分布格局推测,该属可能在不晚于第三纪早期,起源于古北大陆.东亚和北美的粉条儿菜属植物形态区别明显,应该是隔离分化的结果.(3)该属植物可能曾经广布于北半球,后来地质、气候以及冰川等因素的变化,导致该属在一些地区灭绝,而仅存于东亚和北美东南部.(4)尽管横断山及其周边地区是东亚粉条儿菜属的多样化中心,但该地区很可能并不是粉条儿菜属最早的分化中心,因横断山地区周边的一些特有种可能是在晚近的时期形成的新特有种;另外,东亚粉条儿菜属一些原始的种类主要分布于我国中东部到日本一带.所以,中国中东部到日本一带可能是粉条儿菜属早期的分化中心.     

Phylogeny and Phytogeography of Eupatorium (Eupatorieae, Asteraceae): Insights from Sequence Data of the nrDNA ITS Regions and cpDNA RFLP

Eupatorium were examined by sequencing the internal transcribed spacers (ITS) of nuclear ribosomal DNA and restriction site analysis of chloroplast DNA. Molecular data provided strong evidence that (1) this genus originated in North America, (2) the genus diverged into three morphological species groups, Eutrochium, Traganthes and Uncasia in North America, and (3) one of the North American Uncasia lineages migrated into temperate Europe and eastern Asia over the Bering land bridge. The estimated divergence times support a late Miocene to early Pliocene migration from North America to Eurasia via the Bering land bridge. A European species was sister to all of the eastern Asian species examined. The disjunct distribution pattern of the genus Eupatorium is incongruent with the classical Arcto-Tertiary geoflora concept. Received 13 September 1999/ Accepted in revised form 4 January 2000     

Phylogeny and biogeography of the flowering plant genus Styrax (Styracaceae) based on chloroplast DNA restriction sites and DNA sequences of the internal transcribed spacer region

Phylogenetic relationships within the flowering plant genus Styrax were investigated with DNA sequence data from the internal transcribed spacer (ITS) region of nuclear ribosomal DNA (nrDNA) and with chloroplast DNA restriction site data from the genes trnK, rpoC1, and rpoC2. The data sets from each genome were analyzed separately and in combination with parsimony methods. The results strongly support the monophyly of each of the four series of the genus but provide little phylogenetic resolution among them. Reticulate evolution may at least partly explain discordance between the molecular phylogenetic estimates and a prior morphological estimate within series Cyrta. The historical biogeography of the genus was inferred with unweighted parsimony character optimization of trees recovered from a combined ITS and morphological data set, after a series of combinability tests for data set congruence was conducted. The results are consistent with the fossil record in supporting a Eurasian origin of Styrax. The nested phylogenetic position of the South American members of the genus within those from southern North America and Eurasia suggests that the boreotropics hypothesis best explains the amphi-Pacific tropical disjunct distribution occurring within section Valvatae. The pattern of relationship recovered among the species of section Styrax ((western North America + western Eurasia) (eastern North America + eastern Eurasia)) is rare among north-temperate Tertiary forest relicts. The monophyly of the group of species from western North America and western Eurasia provides qualified support for the Madrean-Tethyan hypothesis, which posits a Tertiary floristic connection among the semiarid regions in which these taxa occur. A single vicariance event between eastern Asia and eastern North America accounts for the pattern of relationship among intercontinental disjuncts in series Cyrta.     

A Preliminary Study on the Taxonomy of the Family Magnoliaceae

A new system of classification of Magnoliaceae proposed. This paper deals mainly with taxonomy and phytogeography of the family Magnoliaceae on the basis of external morphology, wood anatomy and palynology. Different authors have had different ideas about the delimitation of genera of this family, their controversy being carried on through more than one hundred years (Table I). Since I have been engaged in the work of the Flora Reipublicae Popularis Sinicae, I have accumulated a considerable amount of information and material and have investigated the living plants at their natural localities, which enable me to find out the evolutionary tendencies and primitive morphological characters of various genera of the family. According to the evolutionary tendencies of the characters and the geographical distribution of this family I propose a new system by dividing it into two subfamilies, Magnolioideae and Liriodendroideae Law (1979), two tribes, Magnolieae and Michelieae Law, four subtribes, Manglietiinae Law, Magnoliinae, Elmerrilliinae Law and Micheliinae, and fifteen genera (Fig. 1 ), a system which is different from those by J. D. Dandy (1964-1974) and the other authors. The recent distribution and possible survival centre of Magnoliaceae. The members of Magnoliaceae are distributed chiefly in temperate and tropical zones of the Northern Hemisphere, ——Southeast Asia and southeast North America, but a few genera and species also occur in the Malay Archipelago and Brazil of the Southern Hemisphere. Forty species of 4 genera occur in America, among which one genus (Dugendiodendron) is endemic to the continent, while about 200 species of 14 genera occur in Southeast Asia, of which 12 genera are endemic. In China there are about 110 species of 11 genera which mostly occur in Guangxi, Guangdong and Yunnan; 58 species and more than 9 genera occur in the mountainous districts of Yunnan. Moreover, one genus (Manglietiastrum Law, 1979) and 19 species are endemic to this region. The family in discussion is much limited to or interruptedly distributed in the mountainous regions of Guangxi, Guangdong and Yunnan. The regions are found to have a great abundance of species, and the members of the relatively primitive taxa are also much more there than in the other regions of the world. The major genera, Manglietia, Magnolia and Michelia, possess 160 out of a total of 240 species in the whole family. Talauma has 40 species, while the other eleven genera each contain only 2 to 7 species, even with one monotypic genus. These three major genera are sufficient for indicating the evolutionary tendency and geographical distribution of Magnoliaceae. It is worthwhile discussing their morphological characters and distributional patterns as follows: The members of Manglietia are all evergreen trees, with flowers terminal, anthers dehiscing introrsely, filaments very short and flat, ovules 4 or more per carpel. This is considered as the most primitive genus in subtribe Manglietiinae. Eighteen out of a total of 35 species of the genus are distributed in the western, southwest to southeast Yunnan. Very primitive species, such as Manglietia hookeri, M. insignis and M. megaphylla, M. grandis, also occur in this region. They are distributed from Yunnan eastwards to Zhejiang and Fujian through central China, south China, with only one species (Manglietia microtricha) of the genus westwards to Xizang. There are several species distributing southwards from northeast India to the Malay Archipelago (Fig. 7). The members of Magnolia are evergreen and deciduous trees or shrubs, with flowers terminal, anthers dehiscing introrsely or laterally, ovules 2 per carpel, stipule adnate to the petiole. The genus Magnolia is the most primitive in the subtribe Magnoliinae and is the largest genus of the family Magnoliaceae. Its deciduous species are distributed from Yunnan north-eastwards to Korea and Japan (Kurile N. 46’) through Central China, North China and westwards to Burma, the eastern Himalayas and northeast India. The evergreen species are distributed from northeast Yunnan (China) to the Malay Archipelago. In China there are 23 species, of which 15 seem to be very primitive, e.g. Magnolia henryi, M. delavayi, M. officinalis and M. rostrata, which occur in Guangxi, Guangdong and Yunnan. The members of Michelia are evergreen trees or shrubs, with flowers axillary, anthers dehiscing laterally or sublaterally, gynoecium stipitate, carpels numerous or few. Michelia is considered to be the most primitive in the subtribe Micheliinae, and is to the second largest genus of the family. About 23 out of a total of 50 species of this genus are very primitive, e.g. Michelia sphaerantha, M. lacei, M. champaca, and M. flavidiflora, which occur in Guangdong, Guangxi and Yunnan (the distributional center of the family under discussion) and extend eastwards to Taiwan of China, southern Japan through central China, southwards to the Malay Archipelago through Indo-China. westwards to Xizang of China, and south-westwards to India and Sri Lanka (Fig. 7). The members of Magnoliaceae are concentrated in Guangxi, Guangdong and Yunnan and radiate from there. The farther away from the centre, the less members we are able to find, but the more advanced they are in morphology. In this old geographical centre there are more primitive species, more endemics and more monotypic genera. Thus it is reasonable to assume that the region of Guangxi, Guangdong and Yunnan, China, is not only the centre of recent distribution, but also the chief survival centreof Magnoliaceae in the world.     

Integrated Fossil and Molecular Data Reveal the Biogeographic Diversification of the Eastern Asian-Eastern North American Disjunct Hickory Genus (Carya Nutt.)

The hickory genus (Carya) contains ca. 17 species distributed in subtropical and tropical regions of eastern Asia and subtropical to temperate regions of eastern North America. Previously, the phylogenetic relationships between eastern Asian and eastern North American species of Carya were not fully confirmed even with an extensive sampling, biogeographic and diversification patterns had thus never been investigated in a phylogenetic context. We sampled 17 species of Carya and 15 species representing all other genera of the Juglandaceae as outgroups, with eight nuclear and plastid loci to reconstruct the phylogeny of Carya. The phylogenetic positions of seven extinct genera of the Juglandaceae were inferred using morphological characters and the molecular phylogeny as a backbone constraint. Divergence times within Carya were estimated with relaxed Bayesian dating. Biogeographic analyses were performed in DIVA and LAGRANGE. Diversification rates were inferred by LASER and APE packages. Our results support two major clades within Carya, corresponding to the lineages of eastern Asia and eastern North America. The split between the two disjunct clades is estimated to be 21.58 (95% HPD 11.07-35.51) Ma. Genus-level DIVA and LAGRANGE analyses incorporating both extant and extinct genera of the Juglandaceae suggested that Carya originated in North America, and migrated to Eurasia during the early Tertiary via the North Atlantic land bridge. Fragmentation of the distribution caused by global cooling in the late Tertiary resulted in the current disjunction. The diversification rate of hickories in eastern North America appeared to be higher than that in eastern Asia, which is ascribed to greater ecological opportunities, key morphological innovations, and polyploidy.     

On the Distribution and Origin of Salix in the World

1. The distribution of Salix species among the continents. There are about 526 species of Salix in the world, most of which are distributed in the Northern Hemisphere with only a few species in the Southern Hemisphere. In Asia, there are about 375 species, making up 71.29 percent of the total in the world, including 328 endemics; in Europe, about 114 species, 21.67 percent with 73 endemics; in North America, about 91 species, 17.3 percent with 71 endemics; in Africa, about 8 species, 1.5 percent, with 6 endemics. Only one species occurs in South America. Asia, Europe and North America have 8 species in common (excluding 4 cultivated species). There are 34 common species between Asia and Europe, 14 both between Europe and North America and between Asia and North America, 2 between Asia and Africa. Acording to the Continental Drift Theory, the natural circumstances which promoted speciation and protected newly originated and old species were created by the orogenic movement of the Himalayas in the middle and late Tertiary. Besides, the air temperature was a little higher in Asia than in Europe and North America (except its west part) and the dominant glaciers were mountainous in Asia during the glacial epoch in the Quaternary Period. Then willows of Europe moved southwards to Asia. During the interglacial period they moved in opposite direction. Such a to-and-fro willow migration between Asia and Europe and between and North America occurred so often that it resulted in the diversity of willow species in Asia. Those species of willows common among the continents belong to the Arctic flora. 2. The multistaminal willows are of the primitive group in Salix. Asia has 28 species of multistaminal willows, but Europe has only one which is also found in Asia. These 28 species are divided into two groups, “northern type” and “southern type”, according to morphology of the ovary. The boundary between the two forms in distribution is at 40°N. The multistaminal willows from south Asia, Africa and South America are very similar to each other and may have mutually communicated between these continents in the Middle or Late Cretaceous Period. The southern type willows in south Asia are similar to the North American multistaminal willows but a few species. The Asian southern type willows spreaded all over the continents of Europe, Asia and North America through the communication between them before the Quaternany Period. Nevertheless, it is possible that the willows growing in North America immigranted through the middle America from South America. The Asian northern type multistaminal willows may have originated during the ice period. The multistaminal willows are more closed to populars in features of sexual organs. They are more primitive than the willows with 1-3 stamens and the most primitive ones in the genus. 3. The center of origin and development of willows Based on the above discussion it is reasonable to say that the region between 20°-40°N in East Asia is the center of the origin and differentiation of multistaminal willows. It covers Southern and Southwestern China and northern Indo-China Pennisula.     

Species richness in plant genera disjunct between temperate eastern Asia and North America

The species richness of 109 amphi-Pacific disjunct genera was examined in eastern Asia and North America. Although the entire flora of eastern Asia contains approximately one-third more species than that of North America, the difference in species richness among disjunct taxa is less. When woody and herbaceous genera are considered separately, the former exhibit a strong diversity bias favouring eastern Asia whereas there is no significant difference in diversity between continents among herbaceous genera. This result is not due to habitat differences between woody and herbaceous genera, because the disjunct herbs inhabit primarily moist forests and woodlands. This result is also not related to relative phylogenetic advancement, even though older major lineages of plants tend to have a predominance of woody taxa. Woody genera are distributed in lower latitudes than herbaceous genera on both continents, and both woody and herbaceous genera are distributed in lower latitudes in eastern Asia than in North America. The North American temperate flora is primarily a relict of a flora form 7 more widespread throughout the Northern Hemisphere. Contemporary patterns of diversity suggest that the effects of climate changes in the late Tertiary were less severe in eastern Asia and promoted diversification, but were more severe in North America and may have caused widespread extinction. The difference in the effect of climate change on diversity in herbaceous and woody lineages reflects the different ecological relationships of species having these contrasting life forms. Clearly, the contemporary floras of eastern Asia and North America bear the imprint of history and emphasize the important interface between ecological relationships and evolutionary responses.