试论东亚被子植物区系的历史成分和第三纪源头——基于省沽油科、刺参科和忍冬科植物地理的研究

虽然在历史植物地理学文献中常见的有5种区系成分;地理成分、发生成分、迁移成分、历史成分和生态成分,但由于缺乏古地理、古气候和古植物资料,当前要全面分析一个区系中所有5种成分是不可能的。本文是基于省沽油科、刺参科和忍冬科植物研究的结果,企图分析东亚植物区系的历史成分,从一个侧面看到它的演变过程。除去本地产生的鬼吹箫属和六道木属之外,凡以上3科在东亚有分布的属,按其历史成分,可归为3类;北方第三纪成分——忍冬属、猬实属、锦带花属和莛子？属;提特斯第三纪成分-刺参属、刺续断属、蓟叶参属、野鸦椿属、毛核木属、糯米条属、北极花属、双盾木属和七子花属;北热带成分- 山香圆属、省沽油属和瘿椒树属。本文所指北方第三纪植物群和提特斯第三纪植物群是采用Takhtajan的概念,而北热带植物群是采用Wolfe的概念。这3种成分意味着东亚被子植物区系在演变过程中有3个第三纪源头。本文作者还认为中新世在东亚植物区系中似可作为区分被子植物“新特有属”和“古特有属”的一个分界期。大凡起源于中新世以前的属可视为“古特有属”,起源于中新世及其后的可归为“新特有属”。这个假设是基于下列理由：(1)东亚植物区系的主体部分形成于早中新世;(2)在中新世,植物长距离的传播基本停止,现在植物区系区划中的“区”(region),其迅速分化和丰富其多样性主要依靠本地产生的特有类群;(3)从中新世至上新世,喜马拉雅运动加剧,不仅改变了我国的地形,而且全面地影响了我国的自然地理环境。     

七子花的细胞学研究

七子花ＨｅｐｔａｃｏｄｉｕｍｍｉｃｏｎｉｏｉｄｅｓＲｅｈｄ .是我国特有单种属植物 ,属于忍冬科北极花族ＴｒｉｂｅＬｉｎｎａｅｅａｅＤｕｍｏｒｔｉｅｒ,主要分布于浙江、湖北两省 (徐炳声 ,1 988)。关于忍冬科的细胞学研究见诸报端的已有不少如 :六道木属的糯米条ＡｂｅｌｌｉａｃｈｉｎｅｎｓｉｓＲ .Ｂｒ.2ｎ =32 (Ｂｅｎｋｏ -Ｉｓｅｐｐｏｎ ,1 992 )、北极花属的北极花ＬｉｎｎａｅａｂｏｒｅａｌｉｓＬｉｎｎ .2ｎ =32 (Ｂｅｎｋｏ -Ｉｓｅｐｐｏｎ ,1 992 ;Ｓｅｍｅｒｅｎｋｏ ,1 990 )、忍冬属的Ｌｏｎｉｃｅｒａａｌｐｉｇ…     

六道木属六道木组种间关系的AFLP分析和黄花六道木中国分布新记录的认定

用荧光标记AFLP技术对东亚产六道木属六道木组作了系统关系分析。我们采用 5对AFLP引物 ,对六道木属 9个物种 ,锦带花属和猬实属各 1个物种共 16份样品进行分析 ,共获得 988个位点。不同引物所获得的结果具有很高的相容性 ,Mantel相关系数为 0 72 5～0 919,说明AFLP技术适用于六道木属的种间关系分析。用邻接法 (neighbor joining)对 16个样品 988个位点的分析生成的系统树获得很高的自展分析 (bootstrap)支持率。树系图表明 ,猬实属与六道木属管花六道木组有密切的关系。在六道啪组内 ,温州六道木与糯米条的关系密切 ,单花六道木与大花六道木差异很小 ,黄花六道木与日本产的近缘种组成一单系分支。本文认定 ,根据形态特征鉴定浙江永嘉的标本为黄花六道木 (AbeliaserrataSieb .etZucc .)是正确的 ,属中国分布新记录。文章最后补充描述了中国产的黄花六道木。黄花六道木在中国的发现进一步证明中国植物区系与日本植物区系的密切关系。     

从叶绿体DNA trnL-F序列论双参属的归属问题

双参属Triplostegia Wall．ex DC．由分布于东南亚地区的2个种组成,为多年生草本植物。它的归属一直存在争议,有时置于川续断科Dipsacaceae或败酱科Valerianaceae,有时单立一科,即双参科Triplostegiaceae。本研究对广义川续断目Dipsacales s．l．的21种植物(分别来自于败酱科、川续断科、双参属、刺参属Morina、广义忍冬科Caprifoliaceae s. l．、五福花科 Adoxaceae)和外类群人参Panax schin-seng Nees．的叶绿体 DNA trnL-F区进行了测序,并建立系统发育树状图。结果显示,败酱科、川续断科、双参属、刺参属和广义忍冬科的4个属(双盾木属Dipelta、虫胃实属Kolkwitzia、六道木属Abelia和北极花属Linnaea)形成 了一个单系群并得到了很强的支持(100％ bootstrap);双参属与川续断科有更近的关系,建议作为一个亚科置于川续断科;广义忍冬科为一多系类群;而刺参属与其他广义川续断目类群之间的关系尚不能确定。     

川续断目的花粉演化

以APGⅢ定义的川续断目(Dipsacales)忍冬科(Caprifoliaceae)和五福花科(Adoxaceae)为研究对象,基于已有的川续断目分子系统树和花粉形态数据,分析了忍冬科25属和五福花科5属的花粉形态多样性,采用简约法(Fitch Parsimony)推测花粉祖征和演化式样,寻找共衍征和分类性状.选择花粉萌发孔数目、萌发孔类型、花粉形状、大小和外壁纹饰五个关键性状推断了其演化式样.研究表明:三孔沟、近球形、较小花粉、网状纹饰是川续断目花粉祖征.长球型、中等和较大花粉,以及刺状纹饰是忍冬科的共衍征,支持忍冬科和科下分支为单系群.忍冬科花粉刺状纹饰和五福花科花粉网状纹饰明显将两个科区分开来.通过追溯性状演化分析,支持将七子花属(Heptacodium)置于忍冬族(Caprifolieae),以及Zabelia置于刺续断科的观点.     

北方优良观花灌木──锦带花

锦带花(Weigela florida)为忍冬科锦带花属落叶灌木。枝茎开张,幼枝有2列短柔毛。单叶对生,椭圆形,缘有锯齿,叶面脉上有毛,背面具柔毛。聚伞花序着生于短枝顶及叶腋,萼片披针形,花冠漏斗状钟形,花色玫瑰红,夏季开花,花期长,果熟10月。锦带花的花期正值春花凋零、夏花不多之际,花色艳丽如锦。关于锦带花,《宋.王禹珃海仙花诗并序》中说,海仙花者世谓之锦带,维扬人传云,初得于海州山谷间,其枝长而花密,若锦带然。”诗的最后又说“……锦带为名俚且俗,为君呼作海仙花。”除以上名称,锦带花还有文官花、五色海棠等异名。它是华北、东北地区重…     

罗霄山脉种子植物区系的特有现象与残遗现象

罗霄山脉种子植物区系非常丰富, 是连接华东、华南、华中植物区系的重要通道, 区系组成具有明显的古老性和过渡性特征。本文针对该区域种子植物的特有现象和残遗现象, 分析了其组成特点及形成原因。结果表明: (1)罗霄山脉共有中国特有科3科, 特有属55属, 特有种1,624种, 罗霄山脉区域特有种43种7变种; 特有属中以古特有属为主, 并以温带成分占优势(达55.91%), 体现出明显的古老性和孑遗性。(2)罗霄山脉共有孑遗属165属, 以热带亚洲分布、东亚-北美间断分布、东亚特有及中国特有成分为主, 包括木本属132属。(3)受中新世气候波动以及第四纪冰期的影响, 许多北方热带植物群的常绿成分、北极-第三纪的落叶成分, 在罗霄山脉得以保存下来成为古特有和孑遗成分。特有现象、残遗现象的分析结果表明罗霄山脉是一个重要的生物避难所, 其对中国东部植物区系的保存和重新扩散具有重要意义。     

浙江天台山七子花群落优势种群结构及物种多样性研究

七子花(Heptacodium miconioides)为我国特产的落叶小乔木,属忍冬科的单种属植物,现资源极少.在浙江省天台山分布着一小片以七子花为优势的群落,作者曾对该群落特征和种群结构作了初步的研究[4,5].本文通过对该群落优势种群的年龄结构和物种多样性进行分析,探讨该群落优势种群的年龄动态和物种多样性特征,为七子花群落的保护与持续利用提供科学依据.     

七子花的解剖学研究

研究了七子花根、茎、叶和芽的解剖学结构,并与忍冬科荚Mi属的珊瑚树进行比较。结果表明,两种植物具有一些共同特点：茎中髓部发达;异面叶,栅栏组织细胞都有两层,气孔大多集中于下表皮,叶表面有腺毛。两者也存在一些明显差异：七子花根毛明显少于珊瑚树,甚至未见;幼茎形状、茎中维管柱形状也明显不同;七子花栅栏组织细胞排列较珊瑚树整剂。     

基于ITS序列探讨忍冬属的系统发育关系

以七子花(Heptacodium miconioides)为外类群,运用MEGA软件对20种忍冬属植物进行系统发育分析,采用邻接法(NJ)和最大简约法(MP)构建系统发育树,从分子系统学角度探讨忍冬属下的亲缘关系.结果表明:(1)在NJ和MP系统树中,没有形成系统树的基部分支,忍冬亚属(Subg.Chamaecerasus)和轮花亚属(Subg.Lonicera)没有形成姐妹群关系.(2)在各系统树中,囊管组内的各种没有聚为一支,故认为对囊管组的划分应进一步探讨.(3)忍冬属ITS区(ITS1+ITS2)的信息位点达到11.0%,信息位点比较丰富,证明ITS序列可以为解决忍冬属植物的系统发育问题提供较强的证据.     

Occurrence of plastids in the sperm cells of Caprifoliaceae: biparental plastid inheritance in angiosperms is unilaterally derived from maternal inheritance

It is widely held that organelles inherit from the maternal lineage. However, the plastid genome in quite a few angiosperms appears to be biparentally transmitted. It is unclear how and why biparental inheritance of the genome became activated. Here, we detected widespread occurrence of plastids in the sperm cells (a cellular prerequisite for biparental inheritance) of traditional Caprifoliaceae. Of the 12 genera sampled, the sperm cells of Abelia, Dipelta, Heptacodium, Kolkwitzia, Leycesteria, Linnaea, Lonicera, Symphoricarpos, Triosteum and Weigela possessed inheritable plastids. The other genera, Sambucus and Viburnum, lacked plastids in sperm cells. Interestingly, such exclusion of plastids in the sperm cells of some Caprifoliaceae appeared to be associated with the divergence of Dipsacales phylogeny. Closer examination of Weigela florida revealed that both plastids and plastid DNA were highly duplicated in the generative cells. This implies that the appearance of plastids in sperm cells involved cellular mechanisms. Because such mechanisms must enhance the strength of plastid transmission through the paternal lineage and appear ubiquitous in species exhibiting biparental or potential biparental plastid inheritance, we presume that biparental plastid genetics may be a derived trait in angiosperms. This is consistent with our extended phylogenetic analysis using species with recently discovered modes of potential plastid inheritance. The results show that basal and early angiosperms have maternal plastid transmission, whereas all potential biparental transmission occurs at terminal branches of the tree. Thus, unlike previous studies, we suggest that biparental plastid inheritance in angiosperms was unilaterally converted from the maternal transmission mode during late angiosperm evolution.     

Phylogeny of the Dipsacales s.l. based on chloroplast trnL-F and ndhF sequences

Sequences of the chloroplast trnL-F region and 3(') end ndhF gene were used to elucidate phylogenetic relationships and the delimitation of families within Dipsacales s.l. Parsimony analyses of individual and combined data were conducted using maximum parsimony method. The most parsimonious tree based on combined trnL-F and 3(') end ndhF data set recognizes seven major clades of Dipsacales s.l. with the following relationships: Apiales (Adoxaceae ((Diervillaceae, Caprifoliaceae s.str.) (Linnaeaceae (Morinaceae (Dipsacaceae, Valerianaceae))))). Both Sambucus and Viburnum have close relationships with Adoxaceae, supporting their inclusion in this family. Caprifoliaceae s.l. (excluding Sambucus and Viburnum) is polyphyletic, and comprises three clades or families, i.e., Linnaeaceae (Abelia, Dipelta, Kolkwitzia, and Linnaea), Diervillaceae (Weigela and Diervilla) and Caprifoliaceae s.str. (Heptacodium, Leycesteria, Lonicera, Symphoricarpos, and Triosteum). This study focuses on the systematic position of Heptacodium, Triplostegia, and Morinaceae; and suggests that Heptacodium is closely related to the other Caprifoliaceae s.str.; Triplostegia is a sister to Dipsacaceae; Morinaceae, which has an affinity with Dipsacaceae, is possibly a sister group with Dipsacaceae-Valerianaceae clade. Our results are highly congruent with those of and.     

Eastern Asian endemic seed plant genera and their paleogeographic history throughout the Northern Hemisphere

We review the fossil history of seed plant genera that are now endemic to eastern Asia. Although the majority of eastern Asian endemic genera have no known fossil record at all, 54 genera, or about 9%, are reliably known from the fossil record. Most of these are woody (with two exceptions), and most are today either broadly East Asian, or more specifically confined to Sino-Japanese subcategory rather than being endemic to the Sino-Himalayan area. Of the "eastern Asian endemic" genera so far known from the fossil record, the majority formerly occurred in Europe and/or North America, indicating that eastern Asia served as a late Tertiary or Quaternary refugium for taxa. Hence, many of these genera may have originated in other parts of the Northern Hemisphere and expanded their ranges across continents and former sea barriers when tectonic and climatic conditions al-lowed, leading to their arrival in eastern Asia. Although clear evidence for paleoendemism is provided by the gymnosperms Amentotaxus, Cathaya, Cephalotaxus, Cunninghamia, Cryptomeria, Glyptostrobus, Ginkgo, Ketel-eeria, Metasequoia, Nothotsuga, Pseudolarix, Sciadopitys, and Taiwania, and the angiosperms Cercidiphyllum, Choerospondias, Corylopsis, Craigia, Cyclocarya, Davidia, Dipelta, Decaisnea, Diplopanax, Dipteronia, Em-menopterys, Eucommia, Euscaphis, Hemiptelea, Hovenia, Koelreuteria, Paulownia, Phellodendron, Platycarya, Pteroceltis, Rehderodendron, Sargentodoxa, Schizophragma, Sinomenium, Tapiscia, Tetracentron, Toricellia,Trapella, and Trochodendron, we cannot rule out the possibility that neoendemism plays an important role espe-cially for herbaceous taxa in the present-day flora of Asia, particularly in the Sino-Himalayan region. In addition to reviewing paleobotanical occurrences from the literature, we document newly recognized fossil occurrences that expand the geographic and stratigraphic ranges previously known for Dipelta, Pteroceltis, and Toricellia.     

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     

First fossil record of an East Asian endemic genus Sladenia (Sladeniaceae) from its modern range: Implications for floristic evolution and conservation biology

Fossil records of endemic plants play an important role in recognizing the floristic history of East Asia and thereby facilitate the conservation of plant diversity in the region. However, the fossil record of many extant East Asian endemic genera remains poorly documented thus far. Here, we report an infructescence fossil of an East Asian endemic genus, Sladenia (Sladeniaceae), from the early Miocene of southeastern Yunnan, China. The fossil is characterized by: (i) dichasial cymes; and (ii) flask‐shaped ovary with dense subparallel ribs on the surface extending from the base to the distal end of the united style. It represents the first fossil record of Sladenia in Asia, showing that the genus was established in the region at least by the early Miocene. Given that a much older fossil record of Sladeniaceae has been reported from Africa and the sister group of Sladenia is distributed only in Africa, Sladenia is not likely of East Asian origin. The present endemic status of Sladenia was possibly achieved by regional extirpation in Africa and taking refuge in East Asia. This case thus supports the “Museum” rather than “Cradle” hypothesis for the genesis of high plant species in the flora of East Asia. A comparison of the present fossil with extant Sladenia infructescence shows morphological stasis from the early Miocene to present. Such evolutionary tardiness might have resulted in the reduced fitness of the genus, which further caused its current endangered situation.     

安息香科的系统位置及地理分布

安息香科为柿目的一成员,包括１１个属,为一自然的分类群．它与山茶科很接近,并可能是从它的祖先类群演化而来的．本文分析其形态特征的演化趋势,认为子房上位,花冠裂片覆瓦状排列,雄蕊为花冠裂片两倍,花序圆锥花序为原始性状,而子房下位．花冠裂片镊合状排列,雄蕊与花冠裂片的同数,花序少花或单花为进化性状．安息香属为本科最大的属,形态变异多样化,既具有最原始性状,为本科原始类群代表,同时又有较多进化性状．其他各属可能是以它作为基干演化而来．从分析各属的分布区类型,本科有７属分布于热带地区,但仅有３属真正分布于热带,其余４属分布于亚热带或其边缘地区．因此,安息香科基本上是一个热带科,但不典型,它可能是从古热带山区的亚热带地区演化而来的．根据全科的属和种的统计,有１１属,１５０余种,间断分布于欧亚和美洲两大陆块上,亚洲有１０属,５７种,主要分布于东亚,在这一地区,以我国秦岭和长江以南至南岭以北及华西南种类最丰富,包括有最原始类群和系统演化各阶段类群;热带南美洲有２属８３种,这一带种类虽丰富,但仅２属及缺乏原始类群．因此,我们称东亚为安息香科的分布和分化中心,而热带南美洲为第二分布中心．根据化石记载结合本科现代分布格局,我们     

一些东亚特有种子植物的化石历史及其植物地理学意义

特有植物在植物区系地理的研究中有重要意义,不同等级的特有植物往往成为不同等级植物区系分区的重要依据。查明东亚特有植物的地史渊源对于揭示东亚植物区的特征与性质,理解中国植物区系及东亚植物发生和演变都有重要的意义。本文在现有资料的基础上,分析总结了东亚植物区系中有化石记录的银杏科、杜仲科、连香树科、大血藤科和昆栏树科等5个特有科,水杉等21个特有属的化石历史。从这些特有植物化石历史的分析可以发现,东亚植物的特有植物从来源上可以分为北极-第三纪,北热带,和就地起源等3种类型,特有植物的来源表明东亚植物区系是一个来源复杂的植物区系。尽管各种特有类群的地质历史各不相同,但是都经历了从广布到分布区逐步缩小,最后形成特有的过程。大部分特有类群形成特有的时间是在上新世末到第四纪初。根据特有类群划分区系等级的原则,东亚现代植物区系最终形成的时间应该是上新世末到第四纪初。     

无量山种子植物区系的特有现象

地处滇中南的无量山地区迄今计有种子植物２０７科,１０２６属,２５４０种,其中东亚特有科１０个,中国特有属２７个,无量山特有种子６７个,这一地区不仅植物种类丰富,而且特有现象较为显著,在对科级持有现象进行较细致的研究后,本文认国无量山具有较多的分类系统上隔离的东亚特有科反映了该地作为东亚古老植物区系的一部分,其地质历史与整个东亚是一致的,且与东亚植物区系的发端密切相关,属的特有现象表明,地处中国特有     

云南省小黑山自然保护区种子植物区系研究

小黑山自然保护区有野生种子植物170科787属2195种。分析表明该区种子植物区系的地理成分复杂,联系广泛。热带性质的属有506属,占总属数的68.6%,温带性质的属有224属,占总属数的30.4%。热带性质的种890种,占总种数的40.9%,温带性质的种有1267种,占总种数的58.3%。表明该地区植物区系具有明显的亚热带性质,同时深受热带植物区系的影响。植物区系来源主要由东亚成分、热带亚洲成分(印度—马来西亚成分)、中国特有成分三部分融合而成。特有现象明显,有东亚特有科5科,中国特有属6属,中国特有种778种,占种总数的35.8%,而云南特有种有355种,占中国特有种的45.6%。保护区还拥有众多的珍稀濒危植物。