华中铁角蕨复合体的生物系统学研究

变异铁角蕨系(Series Variantia Ching et S.H.Wn)主产我国,形态变异大。在标本室中,铁角蕨属(AspleniumL.)的鉴定非常紊乱,同一个种名下往往存在着不同的细胞型或杂种。本文通过对该复合体的生物系统学研究,揭示它们在网状进化中亲缘关系的来龙去脉,并对它们的分类学位置提出讨论和处理建议。细胞学、等位酶、形态学和孢粉学证据表明:由3个基本的二倍体祖先种形成了共13个成员组成的华中铁角蕨复合体(Asplenium sarelii com-plex)。华中铁角蕨(Asplenium sarelii Hook.)应当被标定为二倍体,以往文献中所谓的四倍体“华中铁角蕨”实际上是来自二倍体华中铁角蕨和细茎铁角蕨(A.tenuicaule Hayala)杂交后加倍得来的异源四倍体产物,它被另外处理为新种武当铁角蕨(A.wudangense Z.R.Wang et X.Hou)。北京铁角蕨(A.pekinense Hance)是二倍体华中铁角蕨加倍后得来的同源四倍体。泸山铁角蕨(A.lushanense C.Chr.)是云南铁角蕨群中的惟一二倍体祖先种,不应处理为四倍体云南铁角蕨(A.yunnanense Franch)的异名。变异铁角蕨(A.varians Wall .ex Hook.et Grev.)很可能是细茎铁角蕨的同源四倍体。3个天然四倍体新杂种及其起源被发现,它们是:龙门铁角蕨(A.×longmenense(=A.pekinense×vanians)),京云铁角蕨(A.× jin     

中国西南地区铁角蕨科植物叶表皮微形态及其系统学意义

利用光学显微镜对12种铁角蕨科植物即大盖铁角蕨、齿果铁角蕨、黑柄铁角蕨、半边铁角蕨、胎生铁角蕨、西南铁角蕨、岭南铁角蕨、剑叶铁角蕨、江南铁角蕨、巢蕨、狭基巢蕨和水鳖蕨植物的叶表皮微形态进行观察比较。结果表明:12种铁角蕨科植物的叶表皮细胞多为不规则型,垂周壁为浅波状、波状或深波状;叶上下表皮均无毛。它们的气孔器类型有7种,为极细胞型、腋下细胞型、不等细胞型、无规则四细胞型,无规则细胞型、横列型和辐射状细胞型。不同种间叶表皮微形态特征表现出一定差异,对属、种的划分有一定分类学意义。对巢蕨属与铁角蕨属、水鳖蕨属与铁角蕨属的亲缘关系进行讨论,为铁角蕨科植物的系统分类研究提供新的资料。     

膜叶铁角蕨属的形态系统学研究

膜叶铁角蕨属隶属于铁角蕨科,全世界约有30种,我国分布有18种,是该属植物的分布中心之一。到目前为止,膜叶铁角蕨属的物种数目和物种分类还存在很大争议,一些物种的界限和定义还模糊不清,为了得到一个自然的膜叶铁角蕨属分类系统,还需要对膜叶铁角蕨属的物种分类做进一步研究。该文在前人研究的基础上,对膜叶铁角蕨属10种植物的形态特征,包括孢子形态特征、叶柄和根状茎上的鳞片形态特征、叶片形态、羽片形状以及叶脉特征等进行详细观察分析,探讨了各个形态特征间的关系以及膜叶铁角蕨属植物的物种分类。结果表明:膜叶铁角蕨属植物的叶片及羽片等形态特征存在很大区别,叶柄和根状茎上的鳞片以及孢子形态的种间差异虽然不大,但其在大小、形状、颜色等方面的细微差别仍可作为部分种类的鉴定依据。该研究结果为膜叶铁角蕨属植物的物种分类及进一步研究提供了重要依据。     

二倍体华中铁角蕨Asplenium sarelii Hook.的等位酶遗传变异

通过等位酶分析,并和同属的两个二倍体种泸山铁角蕨A.lushanense和虎尾铁角蕨A.incisum比较,研究了二倍体华中铁角蕨A.sarelii的群体遗传结构。分析了9个酶系统共13个位点。华中铁角蕨的群体内遗传变异非常丰富:多态位点百分率P为61.5～69.2％;每个位点的平均等位基因数A为2.5～2.9;预期杂合度He为0.132～0.220。群体间的分化程度较低,群体间变异占总变异的11.2％;泸山铁角蕨和虎尾铁角蕨群体间的分化程度较高,群体间变异分别占总变异的21.0％和37.0％。3个种的种间遗传一致度为0.33～0.47,种内群体间的遗传一致度为0.93～0.99。固定指数F表明华中铁角蕨接近随机交配,而另外两个种为近交。     

凤尾蕨属配子体若干系统学特征新考

通过对凤尾蕨属(Pteris L.)8种,即半边旗(P.semipinnata L.)、傅氏凤尾蕨(P.fauriei Hieron.)、阔叶凤尾蕨(P.esquirolii Christ)、三叉凤尾蕨(P.tripartita Sw.)、蜈蚣草(P.vittata L.)、溪边凤尾蕨(P.excelsa Gaud.)、有刺凤尾蕨(P.setuloso-costulata Hayata)、西南凤尾蕨(P.wallichiana Agardh)孢子的人工培养,观察并总结了该属配子体发育各阶段形态特征,并与以往的观察结果进行比较.最终选取孢子萌发型与配子体发育类型等稳定特征作为判断依据,讨论该属与书带蕨科(Vittariaceae)、中国蕨科(Sinopteridaceae)、铁线蕨科(Adiantaceae)等科的亲缘关系.研究认为蕨类配子体生长点上方边缘细胞分布情况多变,不是稳定的系统学特征.而配子体特定部位边缘细胞的外侧壁形态则具有较高的系统学参考价值.     

中国蕨类植物孢子形态的研究V. 铁角蕨科

利用扫描电镜对国产铁角蕨科Aspleniaceae 8属59种植物的孢子形态进行了观察。铁角蕨科植物孢子为单裂缝,两侧对称,极面观为椭圆形、宽椭圆形或近圆形;赤道面观为肾形、半圆形、宽椭圆形、椭圆形或近圆形。极轴长17-41 μm,赤道轴长23-60 μm。外壁光滑。周壁较厚,由周壁形成孢子的表面纹饰。主要纹饰类型有6种: (1)窗孔状纹饰。铁角蕨属Asplenium的厚叶铁角蕨A. griffithianum、北京铁角蕨A. pekinense、肾羽铁角蕨A. humistratum属于此种类型。(2)脊状纹饰。铁角蕨属的假大羽铁角蕨A. pseudolaserpitiifolium、毛轴铁角蕨A. crinicaule等7种属于此种类型。(3)翅状纹饰。铁角蕨属有14种属于此种类型,如江南铁角蕨A. loxogrammioides、齿果铁角蕨A. cheilesorum等;细辛蕨Boniniella cardiophylla、过山蕨Camptosorus sibiricus、水鳖蕨Sinephropteris delavayi、对开蕨Phyllitis scolopendrium、疏脉苍山蕨Ceterachopsis paucivenosa和药蕨Ceterach officinarum也属于此种类型。(4)翅脊状纹饰。有铁角蕨属的21种和巢蕨属Neottopteris的6种属于此种类型。(5)角状纹饰。阔足巢蕨N. latibasis属此类型。(6)丝毛状纹饰。只有阔基苍山蕨Ceterachopsis latibasis属此类型。从孢粉学的角度对该科的分类和系统演化进行了探讨。     

中国特有蕨属扇蕨属（水龙骨科）的系统学研究

扇蕨属（Neocheiropteris）为中国蕨类植物4个特有属之一。自Christ建属以来,不同的分类学者对本属的范畴以及系统位置一直存有争议。本研究对扇蕨属和盾蕨属进行了形态解剖观察,并对扇蕨属及其近缘类群共14个种的4个叶绿体DNA片段：rbcL、rpS4 ＆ rpS4-trnS IGS、trnL intron ＆ trnL-trnF IGS以及atpB ＆ atpB-rbcL IGS进行测序,应用最大简约法（Maximum-parsimony）、最大似然法（Maximum-likeli-hood）和贝叶斯推断（Bayesian inference）对序列进行联合矩阵分析,并构建系统树。该研究从形态学和分子系统学方面都提出了支持扇蕨属和盾蕨属分立的证据。形态学研究显示扇蕨属和盾蕨属叶片分裂方式、叶被和侧脉特征迥异,支持扇蕨属和盾蕨属的分立;但两属根状茎鳞片的共同特征则反映了两属问的紧密联系。分子系统学研究表明扇蕨属与毛鳞蕨属和鳞果星蕨属构成姐妹群,其集合与盾蕨属构成姐妹群。综合形态学和分子系统学的证据,对扇蕨属的系统位置以及范畴进行了深入探讨,扇蕨属包括扇蕨和三出扇蕨,结合形态学和分子系统学证据对三出扇蕨的系统位置进行了初步探讨。     

骨碎补铁角蕨（铁角蕨科）——海南岛一新分布种

报导了海南岛铁角蕨科一新分布种,即骨碎补铁角蕨(Asplenium ritoense Hayata)。在国内的植物志采用的学名是一不合法名称,即Asplenium davallioides Hook.。应给予纠正。     

中国铁角蕨的种下分类学研究

综合运用孢粉学、细胞学、生态学和形态学的证据,处理了中国产铁角蕨Asplenium trichomanes L． 的种下分类问题,将其划分为4个亚种和1个变种：原亚种A．trichomanes L．ssp．trichomanes,喜钙亚种 A．trichomanes L. ssp．inexpectans Lovis,四倍亚种A．trichomanes L．ssp. quadrivalens D．E．Meyer emend． Lovis,粗轴亚种A．trichomanes L．ssp.pachyrachis(Christ)Lovis et Reichst．和哈如变种A．trichomanes L. var．harovii Moore emend．Midle,并提供了它们在中国的分布情况。查阅研究中国科学院植物研究所 (PE)的标本时发现,一些定名为A．trichomanes L．var．centrochinense Christ(华中变种)的模式标本碎片在形态上和倍性上均不同于已知分类群,认为应给予其新种的分类地位。     

华中铁角蕨孢子纹饰形成的超微结构观察

利用透射电子显微镜对铁角蕨科(Aspleniaceae)华中铁角蕨(Asplenium sarelii Hook.)孢子及其纹饰的形成过程进行观察。结果表明:①华中铁角蕨孢子囊发育为薄囊蕨型;②孢子外壁表面光滑,远极面的外壁厚约0.8~1.1μm,近极面的外壁厚约1.4~1.8μm;③孢子周壁厚度约4~5μm,染色较外壁深,分为内层和外层;内层紧帖外壁表面,其上具柱状、瘤状或疣状突起;外层向外隆起形成脊状纹饰的轮廓,脊的下方具空腔,脊的顶端具翅;④铁角蕨型与鳞毛蕨型孢子外壁和周壁纹饰的形成过程具有相似性;⑤孢子的成熟度对于孢子形态的研究是至关重要的,只有完全成熟的孢子的表面纹饰才是稳定的。     

A subspecific taxonomic study on Asplenium trichomanes L. from China

With the evidence from palynological, cytological, ecological and morphological data, the subspecific taxonomy of Asplenium trichomanes L. from China is carried out. Four subspecies and one variety, i.e.A. trichomanes L. ssp. trichomanes, A. trichomanes L. ssp. inexpectans Lovis, A. trichomanes L. ssp. quadrivalens D. E. Meyer emend. Lovis, A. trichomanes L. ssp. pachyrachis (Christ) Lovis et Reichst. and A. trichomanes L. var. harovii Moore emend. Midle are recognized from China. The distribution of each subspecies and variety is also presented. Some fragments of a type specimen named as A. trichomanes L. var. centrochinense Christ in PE are found to be different from the known taxa of A. trichomanes L. complex both morphologically andcytologically, and therefore are regarded representing a new species.     

Exploring the reticulate evolution in the Asplenium pekinense complex and the A. varians complex (Aspleniaceae)

The Asplenium pekinense complex mainly comprises one diploid, A. sarelii Hook. (rare), one autotetraploid, A. pekinense Hance (best known and very common), and shares two allotetraploids, A. anogrammoides Christ (common but often misidentified) and A. altajense (Komarov) Grubov (rare and endemic) with the A. varians complex. The latter is further constituted by two diploids, A. tenuicaule Hayata (widespread) and A. semivarians Viane & Reichstein (rare), as well as other three tetraploids, A. kansuense Ching (barely known), A. varians Wallich ex Hooker & Greville (well‐known, relatively common, and morphologically variable), and A. kukkonenii Viane & Reichstein (rare and often misidentified). These two species complexes are notorious for their taxonomic difficulty based on general morphology, which is mainly caused by their history of reticulate evolution. Here, we collected most species within the two complexes, and obtained ploidy information by spore size measurement and flow cytometry investigation. Phylogenetic analyses using DNA markers representing maternally inherited chloroplast and biparentally inherited nuclear genomes helped to reconstruct the reticulate evolution history. The present results support previous hypotheses that A. sarelii is the ancestor of both A. pekinense and A. anogrammoides, as well as that A. tenuicaule is the common progenitor of A. anogrammoides, A. varians, and A. kukkonenii. We also unraveled the autotetraploid origin of A. kansuense from A. tenuicaule for the first time, and found that A. altajense shares essentially identical genomes with A. anogrammoides.     

铁角蕨属4种植物配子体发育的比较研究

以腐叶土为基质培养铁角蕨(Asplenium trichomanes L.)、阿尔泰铁角蕨[A.altajense(Kom.)Grubov]、假大羽铁角蕨(A.pseudolaserpitii folium Ching)和细裂铁角蕨(A.tenui folium D.Don)的孢子,用光学显微镜观察并比较它们的配子体发育过程,以期为铁角蕨属的系统学研究提供基础资料.结果显示,4种铁角蕨属植物配子体均具有:孢子两面型、单裂缝、周壁具褶皱、书带蕨型萌发,成熟原叶体心形,原叶体具有毛状体等共同特征,表明铁角蕨属的演化处于较进化的系统位置,但它们的毛状体形态和细胞数目明显不同,可为属内分系提供依据.研究发现,4种铁角蕨属植物配子体的边缘细胞形状以及假根的形态和数量均有差异,其叶绿体能象单细胞一样进行无丝分裂,强光照射下叶绿体向相邻细胞的侧壁集中等现象.     

Apomixis and reticulate evolution in the Asplenium monanthes fern complex

Background and Aims

Asexual reproduction is a prominent evolutionary process within land plant lineages and especially in ferns. Up to 10 % of the approx. 10 000 fern species are assumed to be obligate asexuals. In the Asplenium monanthes species complex, previous studies identified two triploid, apomictic species. The purpose of this study was to elucidate the phylogenetic relationships in the A. monanthes complex and to investigate the occurrence and evolution of apomixis within this group.

Methods

DNA sequences of three plastid markers and one nuclear single copy gene were used for phylogenetic analyses. Reproductive modes were assessed by examining gametophytic and sporophyte development, while polyploidy was inferred from spore measurements.

Key Results

Asplenium monanthes and A. resiliens are confirmed to be apomictic. Asplenium palmeri, A. hallbergii and specimens that are morphologically similar to A. heterochroum are also found to be apomictic. Apomixis is confined to two main clades of taxa related to A. monanthes and A. resiliens, respectively, and is associated with reticulate evolution. Two apomictic A. monanthes lineages, and two putative diploid sexual progenitor species are identified in the A. monanthes clade.

Conclusions

Multiple origins of apomixis are inferred, in both alloploid and autoploid forms, within the A. resiliens and A. monanthes clades.     

Genetic diversity and phylogeography in two diploid ferns,Asplenium fontanum subsp. fontanum and A. petrarchae subsp. bivalens,in the western Mediterranean

Asplenium fontanum subsp. fontanum and A. petrarchae subsp. bivalens are diploid rock ferns of limestone outcrops of the western Mediterranean region. Asplenium fontanum subsp. fontanum occurs from Valencia through northeastern Spain to the Alpes‐Maritimes and Swiss Jura. Asplenium petrarchae subsp. bivalens occurs only on Majorca, in Valencia and possibly in southern Spain. We analysed allozyme and chloroplast genetic marker diversity in 75 populations of A. fontanum subsp. fontanum and 12 populations of A. petrarchae subsp. bivalens sampled from across their respective ranges. The two species show similar levels of species and population genetic diversity to one another and to other diploid European Asplenium taxa. Both are predominantly outbreeding, as indicated by F_IS = 0.108 and 0.167 respectively. Substantial between‐population differentiation results largely from differentiation between regions. Isolation by distance operates over limited geographic ranges, up to 50 km. In A. fontanum subsp. fontanum, the major geographical differentiation between Valencia and the rest of the taxon range probably represents an ancient range fragmentation. A less pronounced differentiation divides populations in the SW from those in the NE of the range, with evidence for a biogeographic link between the eastern Pyrenees and southeastern France. High diversity in the Pyrenees may either represent ancient population differentiation, or a suture zone. In A. petrarchae subsp. bivalens, populations on Majorca exhibit a subset of the genetic diversity present in Valencia, although the two regions are strongly differentiated by differing allele frequencies. Dispersal from the mainland may have founded Majorcan populations, although a role for in situ island survival cannot be excluded.     

Phylogenetic Relationships between Asplenium bourgaei (Boiss.) Milde and A. jahandiezii (Litard.) Rouy Inferred from Morphological Characters and rbcL Sequences

Abstract: Micromorphological analysis reveals that A. jahandiezii, which occurs as a local endemic taxon in the Grand Canyon du Verdon in southern France, shows substantial differences to A. bourgaei, from the eastern Mediterranean region, in the surface structure of perispores, sporangial indehiscence and the size of stomatal subsidiary cells. Nucleotide sequences and genetic distances (0.9 % nucleotide substitutions) of the rbc L gene support the view that both taxa are related but distinct sibling species. Both species form a monophyletic group together with Asplenium fontanum and A. majoricum.     

Biosystematic studies on Adenophora potaninii Korsh. complex (Campanulaceae) V. A taxonomic treatment

Abstract Adenophora is an extremely variable genus, and its taxonomy is very controversial. Of the genus, Adenophora potaninii complex, including A. potaninii, A. bockiana, A. wawreana, A. lobophylla, A. biformifolia, A. polydentata, and A. wawreana var. lanceifolia, is a typical group with different taxonomical treatments due to high level of morphological diversity. We carried out extensive biosystematic studies based on population sampling, transplantation experiments and offspring tests, cluster analysis, and a crossing experiment. The results reveal four main findings. (i) Leaf forms of the A. potaninii complex were extremely polymorphic; the leaf form of A. potaninii and A. bockiana, and that of A. wawreana and A. biformifolia could be found, respectively, on a single population or among the offsprings of a single plant. (ii) Cluster analysis and a crossing experiment indicated that A. bockiana and A. polydentata could not be separated from A. potaninii, nor A. biformifolia from A. wawreana. (iii) Adenophora potaninii and A. wawreana were gradational in morphology and their compatibility value was slightly reduced compared to that within each entity. (iv) Adenophora lobophylla was distinct from the other members of the complex in shape and size of corolla, relative length of style, and shape of capsule. This species was incompatible reproductively with the other members of the complex, but partly compatible with A. stenanthina, a species in another section. Therefore, we recognized only one species with two subspecies in the complex, A. potaninii subsp. potaninii and subsp. wawreana, moved A. lobophylla out of the complex, and reduced all the other names as new synonyms.