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题名: 东亚地区帚菊木族的系统学与生物地理学
作者: 林娜娜
学位类别: 博士
答辩日期: 2007-06-08
授予单位: 中国科学院昆明植物研究所
授予地点: 昆明植物研究所
导师: 李德铢
关键词: 帚菊木族 ; 孢粉学 ; 比较胚胎学 ; 叶表皮解剖 ; 分子系统学 ; 生物地理 ; 东亚
学位专业: 植物学
中文摘要: 帚菊木族(Tribe Mutisieae Cass.)是菊科较基部的一个类群,中南美洲是其主要分布中心,东亚地区亦为该族的分布中心之一,分布有栌菊木属群(Nouelia group)、帚菊属群(Pertya group)、扶郎花属复合群(Gerbera-complex)三个关键类群。本文利用孢粉学、比较胚胎学、叶表皮解剖学和分子系统学等多学科的研究手段,对这几个关键类群进行了系统研究,主要探讨了这些类群的属间及部分属的种间关系。在综合已有各分支学科资料的基础上,利用化石资料对东亚帚菊木族各类群的起源、分异时间进行校正,并对它们的地理分布格局的形成进行了初步的探讨。主要研究结果如下: 1孢粉学 对东亚地区帚菊木族8属27个种1个变种及产于美洲的Gochnatia 1个种进行了光镜和电镜研究,其中22种是首次研究报道。研究该族的8个属中,花粉粒通常对称,等极,三孔沟。赤道面观球形或近长球形,极面观三裂圆形。在扫描电镜下观察,花粉外壁纹饰可分为三种类型,即:粗颗粒状、细颗粒状和刺状。研究结果表明,Macroclinidium 的花粉特征与其他7个属有显著的区别。东亚帚菊族花粉形态特征在种间变化较小,但其对于研究属间关系提供了十分有用的依据,对于进一步了解该族的进化历史具有重要意义。 2比较胚胎学 主要对亚洲产白菊木属(Leucomeris D. Don)和栌菊木属(Nouelia Franch.)中的代表种(Leucomeris decora D. Don和Nouelia insignis Frach.)的大小孢子从产生,发育直至成熟的整个过程进行了详细观察,在结合已知的兔儿风属(Ainsliaea DC.)和Gerbera胚胎学资料基础上,对其胚胎学特征进行比较和分析,探讨它们之间的亲缘关系。研究结果表明,白菊木属和栌菊木属的胚胎学特征较为相似,均只具有一列孢原,花药壁三层,每层只有单层组成,单核腺质绒毡层,倒生胚珠,单珠被薄珠心,八核蓼形胚囊。但它不同之处在于,其各个分化时期的发生和持续时间稍有差异。结果支持白菊木和栌菊木互为姐妹群的关系,同时,它们的胚胎学特征具有一定的原始性,表明了白菊木和栌菊木可能为古老的类群的后裔。 3 叶表皮解剖 选取了12属 31个种在光镜和电镜下进行观察。研究发现气孔器普遍存在于叶的下表皮,少数种上表皮也有分布,气孔器主要分为星状(staurocytic)和无规则(anomocytic)两种类型。叶表皮细胞种间区别不明显,属间较为一致,叶表皮细胞为多边形和不规则形,垂周壁平直、弓形、浅波状或波浪状。气孔器表面观为椭圆形和近圆形,气孔外拱盖内缘近平滑至浅波状。其中Macroclinidium具有双层气孔外拱盖。大部分种叶表面光滑无蜡质或角质,部分种叶表面具薄片状或层状蜡质。 4分子系统学 基于对东亚地区帚菊木族及相关类群的ITS、ndhF、trnL-F、rpl16四个DNA分子序列的单独和联合分析,结果发现东亚的白菊木与分布于产于美洲的Gochnatia关系较远,应恢复其原有属名为Leucomeris D. Don,白菊木和栌菊木属为姐妹群关系,在系统树上较为孤立,与南美的Stifftia J. C. Mikan属位于帚菊木族的基部位置。兔儿风属是一个单系类群,但其属下分组有待商榷。基于前面分子系统学和形态学的证据,蚂蚱腿子属(Myripnois Bunge)应并入帚菊属(Pertya Sch. Bip.)中,为其华帚菊系(Ser. Sinenses Ling)的成员之一,这样Pertya group减少为三个属。对Gerbera complex分析结果支持Jefrey (1967)和吴征镒等(2002)的观点,即Piloselloides应该从Gerbera中分离出来,恢复为单独的属,而Gerbera和Leibnitzia的属间关系未得到很好的解决,目前倾向于将这两个属作合并处理,但这有待于进一步扩大取样范围进行深入研究来进行验证。 5生物地理学 使用分子钟对东亚分布的帚菊木族的起源、分异时间进行较正,并在此基础上探讨了这些类群现在地理分布格局形成的过程。研究认为帚菊木族起源于始新世晚期,为新热带起源。Nouelia group祖先类群从起源地沿北美,通过当时尚存的白令陆桥或北大西洋陆桥向亚洲扩散,第四纪冰期在我国西南部避难所得以保存,并在冰期后期分化为两个属,与南美的几个较原始的属为平行演化关系。Gerbera complex的祖先在向亚洲及非洲迁徙过程中分为两支,一支后期分化为南美现代的Chaptalia、Trichocline等属,另一支则迁移至非洲,并在迁移途中发生分化,部分种属在第四纪冰期在我国西南地区得以保存,并在冰期后向北扩散。东亚现存的Pertya group与非洲的一些属有共同的祖先类群,在第四纪冰期后逐渐向北扩散,并因多变的气候发生辐射演化。
英文摘要: The tribe Mutisieae Cass. is considered as one of the most important tribes for understanding the systematic relationships in the Compositae as it is a basal lineage in the family. In southern America the greatest concentration of genera and species is found along the southern Andes, however there are still many key genera distributed in eastern Asia including the Nouelia group, Pertya group and Gerbera complex. Phylogeny of Mutisieae in eastern Asia was reconstructed by means of integrative methods involving palynology, comparative embryology, anatomy, and molecular systematics which were based on four DNA sequences. The emerging time and the diverging time of those genera were evaluated using fossil records and the molecular clock. Phylogenetic relationships among genera and some species were addressed, mainly based on the results of molecular systematics. Biogeography of those genera was discussed. Major results of the dissertation were summarized as follows: 1、Palynology Pollen morphology of 27 species and one variety representing seven Asian genera and the southern American Gochnatia of the tribe Mutisieae Less. in the sense of Bremer (Compositae) was investigated by scanning electron microscopy and light microscopy. The pollen grains are usually radically symmetrical, isopolar, and tricolporate. The shape varies from spheroidal to subprolate in equatorial view, and is three-lobed circular in the polar view. Based on exine sculpturing observed under SEM, the pollen grains can be divided into three types, i.e. (i) type I, with macrogranulate sculpture; (ii) type II, with a finely granulate sculpture, and (iii) type III, with spiny sculpture. The palynological data showed that the Macroclinidium Maxim. was remarkably distinguished from the other seven genera. The variation of pollen characteristics within the Mutisieae in Asia is little at the species level, but it proves some useful information for studying the relationships among genera. Further study with broader sampling at the genric level is needed for further understanding of the evolutionary history of the Mutisieae. 2、Comparative embryology We described and compared the characters of embryological development from both Asian species ie. Nouelia insignis and Leucomeris decora (=Gochnatia decora), in order to infer ontogenetic transitions that occurred during the early phases. The analyses indicate that N. insignis and L. decora resemble each other, in having only one row of microspore mother cells in each microsporangium, three layers of anther wall and each layer comprising sole layer of cells. The feature of one row of microspore mother cells is somewhat unique compared with microsporangia in other angiosperms and is likely an apomorphy of the common ancestor of N. insignis and L. decora. They also share the following features. Glandular tapetum is with one nucleus. The ovule is anatropous, unitegmic and tenuinucellar. Eight-nucleate embryo sac is of “Polygonum” type. However, they differ from each other mainly in their androecium developments, among which, the tapetum of N. insignis crushed earlier than that of L. decora. Both successive type and simultaneous type of cytokinesis present in the meiotic divisions of microspore mother cells in N. insignis, but only successive type in L. decora. The results show that many of these characters are similar to each other and support the sister relationship of N. insignis and L. decora. Some embryological characters revealed that both L. decora and N. insignis are from an archaic lingeage. 3、Leaf epidermis Leaf epidermis of 31 species belonging to 12 genera of Mutisieae from both eastern Asia and southern America was investigated. It was shown that the stomatal apparatus was mainly staurocytic and anomocytic and presented in abaxial epiderimis of all species and in the adaxial epidermis of some species. The leaf epidermal cells were with little difference among species but can serve as a criterion for distinguishing genera. The leaf epidermal cells were usually irregular or polygonal, with the anticlinal walls straight, arched, sinuolate, sinuous to sinuate. The major shapes of stomatal apparatus were elliptical or nearly rounded, the inner margin of the outer stomatal rim ranged from nearly-smooth to sinuolate or erose, among which the outer stomatal rim of Macroclinidium were two pairs and quite different from other species. The surface is mostly smooth and no waxy or corneous types, however there are some species with surface covered with ridges of wax or waxy types. 4、Molecular phylogeny Molecular phylogeny of eastern Asian Mutisieae was elucidated using ITS、ndhF、trnL-F and rpl16 sequences. It was shown there were no close affinites between the eastern Asian Leucomeris and the southern American Gochnatia, and therefore Leucomeris should be resumed. Leucomeris decora and Nouelia insignis are sister to each other and occupy an isolated position in the basal lineage together with some south American genera, and this basal lineages were the so-called “Stifftia Group”. Ainsliaea was supported as a monophyletic group and formed a natural genus; however the sectional divisions within the genus were artificial as none of the sections was supported in the all MP trees. Based on the morphological and molecular evidence, we suggested that Myripnois should be merged into Pertya and therefore only three genera were included in Pertya group. The results support the viewpoint of Wu (2002) and Jefrey (1967) that Piloselloides should be separated from Gerbera and resumed. However, the relationship of Gerbera and Leibnitzia was not resolved. Based on the morphological and molecular evidence, the viewpoint of Hansen (1990) was somewhat supported that the two genera should be merged. However more samples were needed to support this viewpoint. 5、Biogeography Using a molecular clock, the relative divergent time of major groups in the eastern Asian Mutisieae was estimated. Based on calibration results and the statistical analyses of the geographical distribution, the formation of patterns of the geographical distribution of those groups were discussed. The result implied that the tribe Mutisieae is of neotropic origin. The ancester of Nouelia group was dispersed into the southwest of China along North American and Beringia or North Atlantic landbridge, then surviveing in there during the Quaternary Glaciation and diverged into two genera afte the Quaternary Glaciation. The Gerbera complex was diverged in two major routes, the first one dispersing into the Central America and South America and evolving Chaptalia、Trichocline etc., the other lineage migrating through Beringia into East Asia and Africa with some genera differentiated in the Southwest of China after the Quaternary Glaciation then migrating toward the north later. The Pertya group was distributed in East Asia, which may have the same ancestral group with some African genera and formed an eruptive divergency due to the climate change after Quaternary Glaciation.
语种: 中文
内容类型: 学位论文
URI标识: http://ir.kib.ac.cn/handle/151853/296
Appears in Collections:昆明植物所硕博研究生毕业学位论文_学位论文

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东亚地区帚菊木族的系统学与生物地理学.林娜娜[d].中国科学院昆明植物研究所,2007.20-25
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