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题名: 壳斗科栎属青冈亚属的形态解剖、分类、分布及其系统演化
作者: 邓敏
学位类别: 博士
答辩日期: 2007-01-11
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: 周浙昆
关键词: 广义栎属 ; 青冈亚属 ; 壳斗科 ; 系统学 ; 分类学
学位专业: 植物学
中文摘要: 本文对青冈亚属(Quercus subg. Cyclobalanopsis)的胚胎学、果实形态解剖学、花粉学、叶形态解剖学、分子系统学进行了研究;在此基础上,对国内外青冈亚属植物标本及模式进行查阅及研究,对青冈亚属进行了分类修订,主要研究结果如下: 1. 胚胎学、果实形态解剖学 首次报道青冈亚属胎胚学特征:花药壁基本型,由五至六层形成(宿存的表皮,具纤维加厚的药室内层、2-3层中层和一层腺质型绒毡层)、小孢子发生同时型,四分体排列四面体形;成熟花粉粒二细胞或三细胞。倒生胚珠二层珠被,厚珠心,内外珠被共同形成珠孔,单孢源至多孢源,大孢子四分体排列直线形或“T“型,其中只有一个发育形成功能型大孢子,雌配子体发育蓼型,极核位于卵器下部,在受精前溶合,游离核型胚乳。 子房内被毛,胚珠在子房中所占体积,胎座和子房横隔的伸长三个特征表明青冈亚属与sect. Cerris(特别是subsect. Lepidobalanus)有最近的亲缘关系,与栎亚属其它类群关系较远。在壳斗科中,胎座的演化从伸长向明显短缩演化。花柱体积的进一步增大是对壳斗目中从授粉至受精过程滞后的适应。明显伸长的子房横隔为青冈亚属与sect. Cerris的共衍征。支持青冈亚属作为广义栎属下分类群的系统地位。 对壳斗科胚胎、果实及花序的形态分支分析表明,广义栎属为一自然单系,青冈亚属是广义栎属的基出类群,可能与sect. Cerris subsect. Lepidobalanus有较近的关系。三棱栎与栎属的关系并不接近。 2. 花粉学研究 青冈亚属花粉外壁纹饰可划分细颗粒,小刺状及指状(I)、颗粒状(II)、聚颗粒状(III)三种。花粉大小和形态与栎亚属中sect. Cerris较为接近,与sect. Quercus. Sect. Lobatae, sect. Protobalanus具有明显疣状、瘤状或聚疣状、瘤状的花粉外壁区别明显,暗示青冈亚属植物与栎亚属中sect. Cerris有较近的亲缘关系。I型花粉为栎属花粉中的原始式样,通过I型花粉指状或棒状突起的愈合形成II,III型花粉,在其它落叶栎属中则通过在I型表面的不匀均加厚形成疣状、瘤状或聚疣状、瘤状外饰。 3. 叶形态分支分析 青冈亚属基于叶片形态划分为复合毛基及单毛基两类。在复合毛基类中,下表皮均具有明显的光滑的蜡鳞,明显的二级间脉,二级脉间距不规则与栎属中sect. Cerris中的种类较为接近,单毛基类中气孔均具十字交叉形的蜡鳞,无或极少二级间脉,二级间脉规则有一定差别。简单毛基星状毛和复杂毛基星状毛不能划归为一类,其间不具演化关系。复杂毛基仅出现于sect. Helferiana, sect. Lepidotricha和sect. Gilva及sect. Cerris的常绿类群中,并且在sect. Gilva和sect. Lepidotricha中还新发现与栎属一致的厚壁单列腺毛和头状毛,暗示这一类群与sect. Cerris有较近的亲缘关系。丁字形毛和典型单列毛则仅分布于sect. Glauca的大部分种类中。乳突和具明显的深波状的表皮细胞则常出现于sect. Semiserrata中。基于对51叶表皮微形态、脉序及果实特征分支分析表明,青冈亚属内存在复合毛基类和单毛基类两个分支。在单毛基类中可以划分为乳突类和丁字形毛-腺毛类。 4. 分子系统学 对51种青冈亚属植物基于ITS的系统学研究支持青冈亚属作为广义栎属分类群的系统位置。在广义栎属可能存在两大分支,即sect. Quercus + sect. Protobalanus与sect. Cerris和subg. Cyclobalanopsis。在后一支中,在青冈亚属中的复合毛基类与栎亚属中的sect. Cerris聚为一支,其它青冈亚属植物位于另一支,虽然只得到较低支持,但是在形态上也得到一些性状的支持,在此无法肯定这一分支的合理性。但表明青冈亚属(特别复合毛基类)与栎亚属中sect. Cerris间具有较其它栎亚属植物更为接近的亲缘关系。叶绿体序列过于均一,无法运用于青冈亚属系统学研究。 5. 分类修订 重新对青冈亚属的组进行划分和调整。新分类系统包括6个组,分别为毛叶青冈组(Sect. Helfereiana),赤皮青冈组(sect. Gilva),青冈组(sect. Glauca),饭甑青冈组(sect. Pachyloma),大叶青冈组(sect. Cyclobalanopsis),吊罗青冈组(sect. Chrysotrichae)。综合形态及系统学研究成果,推论毛叶青冈组和赤皮青冈组可能是青冈亚属的基出类群,与sect. Cerris中的常绿种类(eg. sect. Lepidobalanus)有最近的关系。大叶青冈组中具明显粗壮的雌花序和其它单毛基类群有较明显区别。饭甑青冈组代表单毛基类群中的大果类,叶下表皮具乳突或是上表皮垂周壁明显深波形而与其它类群相区别。青冈组则具特有的丁字形毛和典型单列毛,是青冈亚属中最为进化的类群。吊罗山青冈组形态介于饭甑青冈组及青冈组间,可能为中间的过渡类群。 分类学修订归并20种,提出8个新组合,对云山青冈、青冈、白枝青冈、滇青冈进行了模式选定,初步澄清了青冈组中,青冈复合群及曼青冈复合群的分类混乱。最终确认青冈亚属植物89种(其中7种存疑),9变种,其中国产青冈亚属58种(其中7种存疑),9变种。
英文摘要: The phylogenetic relationships of Quercus subg. Cyclobalanopsis were comprehensively investigated using integrated studies on comparative embryology, fruit anatomy, palynology, leaf anatomy and molecular phylogeny, along with field and herbarium investigations. The taxonomy of Quercus subg. Cyclobalanopsis was revised. The results are summarized as follows: 1. Comparative embryology, fruit anatomy and cladistic analysis Embryologic features of Quercus subg. Cyclobalanopsis are reported for the first time. The anther wall is five to six layered with a basic type of development (including a persistent epidermis, a fibrous endothecium, two to three ephemeral middle layers and a secretory anther tapetum). Simultaneous cytokinesis during microsporogenesis follows meiosis of the microspore mother cell, thereby forming a tetrahedral. The mature pollen grain is two or three-celled. Ovules are bitegmic-crassinucellate and anatropous. The inner and outer integuments together form a micropyle. The sporogeneous cell (sometimes several cells) of the nucellus functions directly as a megaspore mother cell (cells), later developing into one to three tetrads. The tetrad is arranged in a line or a “T” shape, within which only one random megaspore develops into a functional megaspore, which later develops into a polygonum-type embryo sac with ephemeral antipodal cells. Two polar cells occur beneath the egg apparatus and fuse before fertilization. The development of endosperm is of the Nuclear Type and the embryo sac possesses hypostase. Other embryological and fruit characters (ovary cavities, trichomes in locule, size of ovules relative to the ovary cavity, and if aborted ovules are attached to the elongate septum) indicate that subg. Cyclobalanopsis has a closer relation to sect. Cerris, than other taxa in Quercus s.s. Elongation of the placenta, the potential synapomorphy of the two taxa, represents the primitive status in Fagacaeae. Increasing the volume of the style (especially the parenchyma in the style) in pistillate flowers demonstrates a better adaptive strategy to the long term delayed fertilization in the Fagales. These results support the basal position of the subg. Cyclobalanopsis and sect. Cerris in Quercus s.l.. Cladistic analysis based on 25 embryological and fruit characters supported the monophyletic status of Quercus s.l. Subg. Cyclobalanopsis - the basal group and having a closer relationship to sect. Cerris than other sections in subg. Quercus. This was supported by tree topology, but without a bootstrap value. Additional characters related to the comparative anatomy of the cotyledon region, and the pericarp, should be investigated to help understand the phylogeny of Quercus s.l. Trigonobalanus, a basal group within the Fagaceae is systematically closer to Castanea than to Quercus. 2. Palynology The apertures in pollen grains of Quercus subg. Cyclobalanopsis have little taxonomic significance and can be classified into 3 types: uniform rod or spinulate tufts (type I), fine granulate (type II), and verrucate and rugulae (type III). The pollen sculptures of subg. Cyclobalanopsis are similar to those of the section Cerris, but are different from those species having the tuberculate or complex verrucate sculptures of sections Quercus, Lobatae and Protobalanus. All pollen characters support the intragenus status of the Cyclobalanopsis. Two morphological evolutionary trends are discussed. Type I pollen is the primitive type within Quercus s.l.. The other pollen extine is derived either by fusion of the rods, spinulate, or by thickening on special sites on the extine of type I pollen. 3. Cladistic analysis of morphologic characters Two groups can be discriminated within the subg. Cyclobalanopsis: a complex trichome basal cells group (CTB) and a simple trichome basal cell group (STB). On the lower cuticle, CTB presents smooth wax ornamentation, obviously irregular interspaces between secondary veins and frequently presenting intersecondary veins, which is similar to sect. Cerris. The STB group has a cruciate wax scale, regularly spaced secondary veins, and rarely presents intersecondary veins. Simple stellate trichomes and complex trichomes are distinct structures with different lineages and are not related.. The complex trichome basal cells were only found in sections Helferiana, Gilva, Lepidotricha and Cerris. In sect. Gilva and sect. Lepidotricha, the thick-walled uniseriate and bulbous glandular trichomes were also observed for the first time within subg. Cyclobalanopsis. Two-armed no-glandular trichomes and pappilae structures are the synapomorphies to the species in sect Glauca and sect. Semiserrata respectively. Cladistic analysis on 51 cuticle, stomata, venation and acorn characters also demonstrated the two lineages: CTB and STB. Within STB, a pappilae group and a lateral-appressed trichome-uniseries group can be discriminated from tree topology. Additional reweighting of the characters stages and adding more characters on anthers and stigmas are necessary to achieve sufficient bootstrap value. 4. Molecular phylogeny A phylogenetic analysis of 51 species of subg. Cyclobalanopsis, based on nuclear ITS sequences, was used to investigate phylogenetic relationships within the subg. Cyclobalanopsis. The monophyletic status of Quercus s.l. was confirmed and data supported the subgenus status of Cyclobalanopsis. Two clades exist in Quercus s.l.: sect. Quercus+sect. Protobalanus vs. Sect. Cerris+subg. Cyclobalanopsis. Within the latter clade, a complex trichome basal cells lineage and a simple trichome basal cell lineage were detected. The Sect. Cerris is nested within the complex trichome basal cells group indicating the basal position of the complex trichome basal cell group within subg. Cyclobalanopsis. Plastid psbA-trnH did not help to solve the phylogenetic relationships within subg. Cyclobalanopsis, due to its uniform sequences. More data sequences from single or low copies of nuclear DNA are needed to improve the phylogenetic solution. 5. Taxonomic revision The systematic taxonomy of Quercus subg. Cyclobalanopsis was revised based on these results and field observations. Quercus subg. Cyclobalanopsis was divided into six sections: Helferiana,. Gilva, Glauca, Pachyloma,. Oidocarpa, and Chrysotrichae. Section Helferiana and section Gilva are the basal groups within the subgenus with obvious complexed trichome basal cells and primitive venation characters. These two sections have a close relation to sect. Cerris subsect. Lepidobalanus. The Section. Cyclobalanopsis possesses an obvious elongate inflorescence, and has distinct differences from other simple trichome basal cell groups. Section Pachyloma is characterized by large global or oblong acorns, glabrous leaves with pappilaes. Section Glauca, having two-armed no- glandular trichomes and typical uniseriate glandular trichomes, is the most advanced section in subg. Cyclobalanopsis. Section Chrysotrichae shares some characters of sect. Pachyloma and sect. Glauca, indicating that it links the two sections. Eighty-nine species and 9 varieties of Quercus subg. Cyclobalanopsis are recognized worldwide. In China, fifty-eight species and 9 varieties are recognized. Among them, seven species were enumerated as dubious taxa because of the unavailability of type specimens or ambiguous original publication citations. Twenty species were reduced to synonymies. Eight new combinations are proposed. The confused taxonomy in the Q. glauca and Q. oxyodon complexes was clarified.
语种: 中文
内容类型: 学位论文
URI标识: http://ir.kib.ac.cn/handle/151853/196
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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壳斗科栎属青冈亚属的形态解剖、分类、分布及其系统演化.邓敏[d].中国科学院昆明植物研究所,2007.20-25
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