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     

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.Wu)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 Hayata)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 Grey.)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×vzrians)),京云铁角蕨(A.×jingyunense(=A.pekinense×yunnanense))和吉多铁角蕨(A.× kidoi(=A.pekinense×wudangen.se)).另外3个天然三倍体新杂种也被发现并给出了推断的亲本,他们是华武铁角蕨(A.×huawuense(=A.sarelii×wudangense)),泸云铁角蕨(A.×luyunense(=A.lushanense×yunnanense))和细变铁角蕨(A.×tenuivarians(=A.tenuicaule×varians)).本文阐明了华中铁角蕨复合体成员间在网状进化中的错综复杂关系,同时表明等位酶比较结合细胞学观察是揭示多倍体和杂种起源的有效工具.     

江苏铁角蕨的系统位置和多倍体起源

运用分子系统发育分析的方法研究了江苏铁角蕨（Asplenium kiangsuense）的系统位置及其与庐山铁角蕨（Agulingense）的关系,并探讨了该类群可能的多倍体起源方式。结果显示：江苏铁角蕨与庐山铁角蕨可能为同源四倍体,是组成倒挂铁角蕨复合体（Anormale complex）的成员之一;二者在形态特征与基因序列方面均表现一致,接受英文版中国植物志的处理,即把庐山铁角蕨处理为江苏铁角蕨的异名。     

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.     

铁角蕨科的多倍化与物种多样性形成

铁角蕨科植物约800种,广布世界各地,主产热带和亚热带地区。本文统计了铁角蕨科188种植物的细胞学资料,发现167种植物具有多倍化现象,占总种数的88.8%,表明该类群植物中普遍存在多倍化事件。具有多倍化现象的物种中,90种只有1种细胞型,占总种数的47.9%;77种具有种内多倍性,即种下存在多个细胞型组合,占总种数的41.0%。多倍体细胞倍性极其丰富,有三倍体、四倍体、六倍体、八倍体、十倍体、十二倍体以及十六倍体。本文还对铁角蕨科多倍化现象与其物种多样性形成的关系进行了讨论,同时对铁角蕨科多倍化研究中存在的问题进行了探讨。     

云南铁角蕨与泸山铁角蕨的关系:来自叶绿体rbcL、trnL-F和rps4-trnS序列的证据

云南铁角蕨Asplenium yunnanense与泸山铁角蕨Asplenium lushanense的分类是一个悬而未决的问题。本文对云南铁角蕨与泸山铁角蕨及近缘类群的叶绿体rbcL基因和trnL-F、rps4-trnS基因间隔区序列进行PCR扩增和序列分析;并与已经发表的铁角蕨属植物的相应序列进行比较,发现云南铁角蕨与泸山铁角蕨的rbcL基因和trnL-F、rps4-trnS基因间隔区序列之间均未表现出差异,因此叶绿体DNA序列的证据不能将云南铁角蕨与泸山铁角蕨分开,从而从分予系统学方而证明云南铁角蕨与泸山铁角蕨是一对亲缘关系非常近的物种。     

广西蕨类植物孢子形态的研究Ⅱ.铁角蕨属

利用光学显微镜和扫描电子显微镜对广西产9种铁角蕨属植物即拟狭翅铁角蕨、黑柄铁角蕨、剑叶铁角蕨、长生铁角蕨、岭南铁角蕨、石生铁角蕨、阴湿铁角蕨、半边铁角蕨和细裂铁角蕨植物的孢子形态进行了观察,对每个种的形态特征进行了详细描述,其中5种铁角蕨属植物孢子的表面纹饰为首次报道。结果如下:9种铁角蕨属植物孢子都为单裂缝,两侧对称,极面观为椭圆形或近圆形,赤道面观为肾形、椭圆形或近圆形。主要纹饰类型有翅脊状纹饰、翅状纹饰和脊状纹饰类型。讨论了各种间的差异。为铁角蕨属的分类和系统演化研究提供孢粉学的资料。     

中国铁角蕨科铁角蕨属一新记录杂交种——东方铁角蕨

该研究报道了采自山东省泰安市泰山的铁角蕨科铁角蕨属一中国新记录植物——东方铁角蕨(Asplenium×akaishiense Otsuka),并对其形态特征进行了描述。该物种形态介于过山蕨(A. ruprechtii Sa. Kurata)与细茎铁角蕨(A. tenuicaule Hayata)之间,可能是二者的自然杂交种。     

基于孢子形态和分子证据探讨鳞盖蕨属(碗蕨科)系统分类

孢粉学是解决植物分类中疑难类群物种微形态分化的重要方法, 随着分子系统学的发展, 结合这两门学科的优势可以更加有效地解决疑难类群的分类学问题。鳞盖蕨属(Microlepia)是一个分类困难的疑难类群, 采用孢粉学与分子系统学一一对应的方法, 以及居群取样方式, 选取280份样本, 联合4个叶绿体片段(rbcL、trnL-F、psbA-trnH和rps4), 采用最大似然法和贝叶斯法构建该属的系统发生关系, 在此基础上对凭证标本中100份材料的孢子进行观察和分析。综合分子系统学和孢粉学的研究结果, 得出结论: (1) 在形态学研究中广泛被接受的15个物种得到了单系支持, 并厘清了分类困难的复合群; (2) 发现边缘鳞盖蕨(M. marginata)可能存在隐性种; (3) 建议恢复过去归并处理为异名的瑶山鳞盖蕨(M. yaoshanica)、罗浮鳞盖蕨(M. lofoushanensis)、四川鳞盖蕨(M. szechuanica)以及滇西鳞盖蕨(M. subspeluncae); (4) 提出鳞盖蕨属可能存在杂交现象; (5) 提出鳞盖蕨属完整的属下分类建议。     

蕨类植物孢子囊形态 I. 鳞始蕨科

孢子囊是蕨类植物的繁殖器官, 其形态在蕨类植物的分类和系统发育研究上具有重要意义。用次氯酸钠溶液处理新鲜成熟的孢子囊, 在光学显微镜下获得清晰的孢子囊图像, 系统研究了中国鳞始蕨科4属13种孢子囊的形态特征。结果表明, 鳞始蕨科孢子囊呈椭球形, 孢子囊柄由3列细胞构成, 环带类型为垂直环带。通过分析孢子囊形态数据探讨了中国鳞始蕨科属内及属间差异。结果表明, 乌蕨属(Odontosoria)、香鳞始蕨属(Osmolindsaea)、达边蕨属(Tapeinidium)和鳞始蕨属(Lindsaea)的孢子囊环带细胞数依次减少, 囊蒴体积、唇细胞数和囊壁细胞数的变化由大(多)到小(少)依次为乌蕨属、达边蕨属、香鳞始蕨属和鳞始蕨属。孢子囊属内差异最大的是阔片乌蕨(Odontosoria biflora)与乌蕨(O. chinensis)以及香鳞始蕨(Osmolindsaea odorata)与日本鳞始蕨(O. japonica); 而达边蕨属和鳞始蕨属的属内差异则很小。研究结果为揭示鳞始蕨科系统发育关系提供了形态基础, 特别是提出阔片乌蕨和乌蕨以及香鳞始蕨和日本鳞始蕨在孢子囊形态上的差异值得进一步研究。     

Phylogenetic and biosystematic relationships in four highly disjunct polyploid complexes in the subgenera Ceterach and Phyllitis in Asplenium (Aspleniaceae)

Phylogenetic studies using DNA sequences of two chloroplast regions, rbcL and trnL-F, demonstrate that the proposed genus Ceterach is a small clade within the large genus Asplenium, and sister to the Phyllitis clade. The Ceterach clade is characterised by irregular anastomosing veins and often densely scaled leaf blades. Its taxonomic status as a group nested within Asplenium is confirmed, and it is accepted here as a subgenus with seven species. The Ceterach clade comprises four lineages that correspond to disjunct polyploid complexes: the A. aureum clade forming a polyploid complex (4×, 6×, 8×) in Macaronesia, the A. ceterach clade forming a polyploid complex (2×, 4×, 6×) in the Mediterranean Basin, the A. paucivenosum clade (4×, 6×) in central Asia, and the A. dalhousiae clade (2×) with a disjunct distribution in the Himalaya, Yemen and Eritrea, and southwestern North America. Asplenium paucivenosum is sister to all other members of the Ceterach clade, whereas A. dalhousiae is sister to the A. aureum clade that includes tetraploid A. aureum, hexaploid A. lolegnamense, and octoploid A. parvifolium. Asplenium ceterach and its variations – including the hexaploid A. ceterach subsp. mediterraneum subsp. nov. first described below – form a monophyletic unit, sister to a clade consisting of A. aureum and A. dalhousiae. Asplenium cordatum from Africa and A. haugthonii from the isolated atlantic island of St. Helena are not members of the Ceterach clade, which suggests that leaf blades with dense indumenta have evolved at least twice within asplenioid ferns. The allotetraploid species A. hybridum has the chloroplast DNA from A. ceterach, and therefore the latter species is the maternal ancestor of the former. The other parent of this hybrid species is A. sagittatum that is nested within the sister clade of Ceterach, the Phyllitis clade comprising A. sagittatum and A. scolopendrium. The findings suggest that the current distribution of Ceterach is either the result of long-distance dispersal or represents fragmented relicts of a previously more widely distributed species.     

Integrating coalescent-based species delimitation with ecological niche modeling delimited two species within the Stewartia sinensis complex (Theaceae)

Accurate species delimitation is the key to precise estimation of species diversity and is fundamental to most branches of biology. Unclear species boundaries within species complexes could lead to the underestimation of species diversity. However, species delimitation of species complexes remains challenging due to the continuum of phenotypic variations. To robustly examine species boundaries within a species complex, integrative approaches in phylogeny, ecology, and morphology were applied to the Stewartia sinensis complex (Theaceae) endemic to China. Multispecies coalescent-based species delimitation using 572 nuclear ortholog sequences (anchored enrichment) supported reciprocal phylogenetic monophyly of the northern lineage (NL) and southern lineage (SL), which were not sister clades. Niche equivalency and similarity tests demonstrated significant climatic niche differentiation between NL and SL with observed Warren et al.'s I = 0.0073 and Schoener's D = 0.0021. Species distribution modeling also separated their potential distribution. Morphometric analyses suggested significant interlineage differentiation of multiple traits including the ratio of length and width, leaf width, and pedicel length, although overall similarity did not differ. Based on the integrative species concept, two distinct species were proposed with legitimate names of Stewartia gemmata for SL and S. sinensis for NL. Our empirical study of the S. sinensis complex highlights the importance of applying multiple species criteria, in particular the underappreciated niche differentiation, to species delimitation in species complexes pervasive in plants.     

Phylogenetic relationships and the maternal donor of Roegneria (Triticeae: Poaceae) based on three nuclear DNA sequences (ITS,Acc1, and Pgk1) and one chloroplast region (trnL-F)

Roegneria is a polyploid perennial genus in the tribe Triticeae. Some species of Roegneria are morphologically similar to genus Elymus and have been classified in Elymus. To investigate the delimitation and phylogenetic relationships of Roegneria, nuclear (ITS, Acc1, and Pgk1) and chloroplast (trnL–trnF) DNA regions were sequenced for 38 allopolyploid species and 32 diploid species of Triticeae. Phylogenetic analyses of nuclear DNA revealed that all Roegneria species were included in the St and Y genome clades, and that the Y genome was closely related to the V and Xp genomes. The chloroplast DNA dataset showed that Roegneria species were grouped with Pseudoroegneria species. The Pseudoroegneria species from the Middle East (P. libanotica and P. tauri) and Central Asia (P. strigosa) were more closely related to Roegneria species. The results suggested that: (i) the species containing the St and Y genomes should be segregated from Elymus and treated as a distinct genus, Roegneria, based on the genomic constitution; (ii) P. libanotica, P. tauri, and/or P. strigosa potentially served as the maternal donor of the St genome in Roegneria; (iii) The Y genome of Roegneria originated from a diploid Y genome species, and the V and Xp genomes may have contributed to Y genome formation; (iv) among Roegneria species of previously uncertain genomic constitution, R. seriotina was tetraploid and possessed the StY genomes, E. calcicolus was hexaploid with the StYH genomic constitution and should be classified in Campeiostachys, R. glaucifolia possessed the StStY genomes, and R. tschimganica had the genomic constitution St₁St₂Y.     

Experimental hybridization of Angiostrongylus mackerrasae, Bhaibulaya, 1968 and Angiostrongylus cantonensis (Chen, 1935)

Angiostrongylus mackerrasae and A. cantonensis are closely related and their status as distinct species was tested by experimental hybridization. Hybridization between A. mackerrasae and A. cantonensis was observed and the morphology of the F₁ hybrids was found to be intermediate between that of the parental species which suggests strongly that A. mackerrasae and A. cantonensis are both valid species. However, male F₁ hybrids were sterile, whereas females were fertile; no evidence of parthenogenesis or other form of asexual reproduction were observed in either species. It is suggested that A. mackerrasae and A. cantonensis may cross breed in nature.     

中国南方变绿红菇近似种类“青头菌”的物种多样性

本研究以我国西南、华中和华东部分地区变绿红菇近似种类“青头菌”为研究对象,使用形态学和rDNA ITS区段核苷酸序列分析相结合的方法进行鉴定。结果表明,“青头菌”多数子实体属于变绿红菇Russula virescens复合群,其余子实体为绿桂红菇R. viridicinnamomea、红边绿菇R. viridirubrolimbata和本文描述的新种拟壳状红菇R. pseudocrustosa。在形态特征和ITS核苷酸序列方面,我国变绿红菇近似种R. aff. virescens与欧洲的变绿红菇R. virescens和北美的小变绿红菇R. parvovirescens均存在一定差异。变绿红菇近似种类在亚洲的分布范围较广,物种准确划分尚需进一步深入研究。     

中国薄叶卷柏复合群的物种划分

卷柏属(Selaginella)是石松类植物中最大的属, 也是分类难度较大的类群之一。该属的物种划分主要基于形态特征, 但许多近缘种在形态上很难区分。近年来, 已有大量分子证据被用于各植物类群的分类学研究, 但目前未发现一套适合卷柏属物种鉴定的分子标记。薄叶卷柏复合群(S. delicatula group)是卷柏属下鉴定较为困难的类群, 包括了薄叶卷柏(S. delicatula)、黑顶卷柏(S. picta)和瓦氏卷柏(S. wallichii) 3个物种, 主要分布于亚洲的热带和亚热带地区。为了探讨薄叶卷柏复合群内物种的亲缘关系和评估不同分子标记在卷柏属分类学研究中的应用价值, 本研究对该复合群物种进行广泛取样, 共收集到73个个体, 并选取3个叶绿体基因(rbcL, psbA和atpI)和2个核基因(26S nrDNA和pgiC)片段进行系统树的构建及叶绿体单倍型分析。研究发现, 基于叶绿体和核基因构建的系统发生关系存在冲突: 叶绿体基因树上薄叶卷柏个体分为两个分支(A和B), 薄叶卷柏B分支与薄叶卷柏A-S. picta分支呈姐妹关系, 并且rbcL单倍型分析结果也表明薄叶卷柏A和B两个分支存在明显分化; 而核基因结果则支持该复合群3个物种各自的单系性, 其中, S. delicatula分支与S. picta分支为姐妹群, S. wallichii与S. delicatula-S. picta分支为姐妹关系。在对复合群分布区大量标本的观察以及野外群体调查的基础上, 评估了植株茎和枝的分枝方式、孢子叶、营养叶(侧叶、中叶和腋叶)和孢子表面纹饰等形态性状的分类学价值。结果表明, 薄叶卷柏A和B分支的样本仅在植株分枝方式和大孢子表面纹饰上存在差异, 但无法依靠小孢子表面纹饰、孢子叶穗和营养叶形态等特征进行区分。基于现有证据, 薄叶卷柏复合群至少可划分为薄叶卷柏、黑顶卷柏和瓦氏卷柏3种, 但彻底澄清该复合群的物种划分还需要获取模式标本产地的材料和细胞学证据。最后, 建议在未来卷柏属的分类学研究集中于该属分类复杂的复合群, 结合使用形态学、细胞学、分子生物学(同时使用核基因和叶绿体分子标记)及地理分布等整合证据来进行物种划分。     

黑木耳复合群中种类学名说明

黑木耳在我国已经有1 000多年栽培历史,是我国最重要的栽培真菌之一,多年来我国黑木耳的学名一直使用模式产地为欧洲的Auricularia auricula-judae。最近的研究表明A. auricula-judae实际为一复合种,该复合种在全球范围内有5个种。其中A. auricula-judae仅分布于欧洲;美洲有2个种,生长在针叶树上的美洲木耳A. americana和生长在阔叶树上的尚未命名的木耳。中国该类群有3个种：自然分布和栽培最广泛的黑木耳的学名应为A. heimuer,此外,短毛木耳A. villosula在我国东北也广泛分布,并有少量栽培,生长针叶树上的木耳为美洲木耳A. americana,主要分布于中国东北和华北。     

Out of the Middle East: New phylogenetic insights in the genus Tamarix (Tamaricaceae)

Tamarix is one of the taxonomically most complex genera among the angiosperms, and there is little consensus regarding its infrageneric classification. Here we present the most complete phylogenetic reconstruction of the genus to date. This includes a DNA phylogenetic tree based on nuclear ribosomal ITS, and a plastid DNA phylogeny based on three intergenic spacers (trnS‐trnG, ndhF‐rpl32, and trnQ‐rps16). In total, both nuclear and plastid phylogenetic analyses include more than 70 samples of 39 species from 27 countries, which represent close to 60% of the diversity of the genus. Two complementary trees, based only on one plastid marker, are also included. The first, based on trnS‐trnG, is used to increase the number of species related to T. amplexicaulis. The second, based on ndhF‐rpl32, is used to investigate the separation between T. tetrandra and T. parviflora. The incongruence between the available infrageneric classifications and the molecular results is confirmed. A reticulate evolution is inferred from the trees, showing characters such as vaginate leaves appearing at different stages along the evolutionary history of the genus. The presence of T. canariensis outside the Canary Islands is cast into doubt, and all such records from NW Africa and Europe are here considered to belong to T. gallica. The results also suggest independence of T. karelinii from T. hispida, and T. parviflora from T. tetrandra. Relationships between a number of species are still not resolved, and additional studies will be needed to further refine the complex taxonomy of Tamarix.