Pollen Morphology of Parnassia L. （Parnassiaceae） and Its Systematic Implications

The pollen morphology of 28 species of Parnassia L. was investigated with light microscopy and scanning electron microscopy (SEM). The shape of pollen grains in this genus varies from subspheroidal to prolate in equatorial view and is three-lobed circular in the polar view. Pollen grains are usually radially symmetrical, isopolar, tricolporate or syntricolporate, with reticulate sculpture. The pollen characteristics among species are fairly similar to each other. Morphological information regarding the pollen grains shows that Parnassia is a natural genus. Based on exine ornamentation observed under SEM, three types of pollen grains were recognized: (i) type I, with foveolate-reticulate sculpture; (ii) type II, with a finely reticulate sculpture; and (iii) type III, with a coarsely reticulate sculpture. Most sections of this genus have one type of sculpture of pollen morphology, but Sect. Nectarotrilobos has three types of sculpture and Sect.Saxifragastrum has two types of sculpture. All three types of sculpture can be found in Southwest China,with species with the longest (Parnassia delavayi Franch.) and shortest (Parnassiafaberi Oliv.) colpi,implying that Southwest China is the center of diversification of the genus.     

中国梅花草属植物的叶表皮特征及其系统学意义

利用光学显微镜和扫描电镜对梅花草属Parnassia 30种植物的叶表皮进行了观察。结果表明:气孔器普遍存在于叶的下表皮,少数种的上表皮也有分布,均为无规则型。叶表皮细胞形状为多边形或不规则形;垂周壁式样可区分为近平直、浅波状和波状。在扫描电镜下,叶表皮气孔器外拱盖内缘为近平滑、浅波状或波状;一些种的保卫细胞两端有加厚;角质膜条纹状,有的条纹隆起,有的条纹上附有颗粒或小孔穴。气孔器类型及下表皮细胞形状的一致性表明梅花草属是一个自然分类群;sect. Saxifragastrum叶表皮特征具有多样性显示该组可能是一个复合群;突隔梅花草P. delavayi属于subsect. Xiphosandra,其气孔下陷,与其细胞学特征相似,支持独立为一组;此外,气孔器的分布、保卫细胞两端加厚、气孔器外拱盖内缘形态以及角质膜等特征对该属部分种的区分有一定的参考价值。     

A Systematic and Evolutionary Study of Zizania L. (Gramineae) Pollen Morphology

This paper deals with pollen morphology of Zizania L. and its relatives. A total of 7 genera, 13 species, 3 varieties and 1 form were examined under light microscope and scanning electron microscope. The results are as follows: 1. The genus Zizania belongs to tribe Oryzeae as shown by pollen characters, i, e. subspheroidal to ovoid in shape, monoporate, exine two-layered, with minute granules under LM. 2. The evolutionary trend of these taxa seems to be from minute granules free (Zizania latifolia, Z. texana, Zizaniopsis milicea and Oryza sativa) to minute granules aggregated in a group of 2-4 (many) (Zizania aquatica, Z. palustris, Leersia hexandra etc.). The genus Zizania may be derived from the ancient stock which has also given rise to the genus Oryza, and therefore parallel evolution may have taken place in Oryzeae, i. e. from perennial species to annual species in Zizania in one line, and from the genus Oryza to Leersia, Chikusichloa etc. in the other. 3. The characters of pollen morphology under LM and SEM support the division of the genus, Zizania into 4 species, 2 subspecies in the world, i. e. Z. latifolia (Griseb.) Turcz. ex Stapf, Z. texana Hitchc., Z. aquatica subsp. aquatica, Z. aquatica subsp. brevis (Fassett) S. L.Chen, Z. palustris subsp. palustris, and Z. palustris subsp. interior (Fassett) S. L. Chen.     

梅花草属四个种的新异名

将丽江梅花草(Parnassia lijiangensis Ku)处理为凹瓣梅花草(P．mysorensis Heyne ex Wight et Am．)的异名;彝良梅花草(P．yiliangensis Ku)处理为大叶梅花草(P．monochorifolia Franch．)的异名;思茅梅花草(P．simaoensis Y．Y．Qian)处理为鸡眼草(P．wightiana Wall．ex Wight ＆ Am．)的异名;龙胜梅花草(P．longshengensis Ku)处理为宽叶梅花草(P．dilatata Hand．-Mazz．)的异名。     

Pollen Morphology of the Platycodonoid Group (Campanulaceae s. str.) and Its Systematic Implications

In this study, we examined the pollen morphology of the platycodonoid group in Campanulaceae s. str. using a scanning electronic microscope. We used pollen grains of 25 accessions representing 24 species of the Codonopsis complex (including Campanumoea, Cyclocodon, Leptocodon, and all three subgenera of Codonopsis), which is extremely controversial among authors for taxonomic treatment. Pollen morphology of all the other genera in the group observed by previous authors is taken into account in our discussion. A total of nine pollen types with two subtypes in the group were recognized and named for the first time. Molecular and morphological data imply that each pollen type corresponds to a natural group at generic level, and thus the mergence of Leptocodon with Codonopsis and the restoration of Cyclocodon as a separate genus are justifiable, and Codonopsis subg. Pseudocodonopsis, subg. Obconicicapsula, and two species of Codonopsis subg. Codonopsis (C. purpurea and C. chimiliensis) may be better classified as three independent genera separate from the core Codonopsis.     

中国梅花草属( 梅花草科) 一些种类的订正

在标本研究和野外观察的基础上, 对中国分布的梅花草属( Parnassia) 的一些种类进行了修订, 将披针瓣梅花草( P. lanceolata) 、窄瓣梅花草( P . angustipetala) 与德格梅花草( P. degeensis) 归并入云南梅花草( P. yunnanensis) ; 将贵州梅花草( P. petimenginii ) 归并入峨眉梅花草( P. faberi ) ; 将矮小梅花草( P. humillis ) 归并入三脉梅花草( P. trinervis) ; 将青海梅花草( P . qinghaiensis) 归并入高山梅花草( P. cacuminum); 将康定梅花草( P . kangdingensis) 归并入类三脉梅花草( P. pusilla) 。     

中国梅花草属(梅花草科)一些种类的订正

在标本研究和野外观察的基础上,对中国分布的梅花草属(Parnassia)的一些种类进行了修订,将披针瓣梅花草(P.lanceolata)、窄瓣梅花草(P.angustipetala)与德格梅花草(P.degeensis)归并入云南梅花草(P.yunnanensis);将贵州梅花草(P.petimenginii)归并入峨眉梅花草(P.faberi);将矮小梅花草(P.humillis)归并入三脉梅花草(P.trinervis);将青海梅花草(P.qinghaiensis)归并入高山梅花草(P.ca-cuminum);将康定梅花草(P.kangdingensis)归并入类三脉梅花草(P.pusilla)。     

Pollen Morphology of Impatiens L. (Balsaminaceae) and Its Taxonomic Implications

Two forms of pollen morphology of Impatiens L. (Balsminaceae) have been described before: 3-colpate and 4-colpate pollen grains. Pollen grains of 13 species of Impatiens on Mt. Omei were examined under SEM for the first time and both forms were observed here, which imply the diversity of species in this region. Of the two forms, 3-colpate one is quite similar to that of the genus Hydrocera Blume (Balsaminaceae) in external morphology. This relation also exists between the 3-colpate pollen group and Hydrocera in habit, habitat, and distribution, etc. It is supposed that the group with 3-colpate pollen type is a natural one, and may have a close relationship with Hydrocera. The pollen morphology is of important significance in classification of the taxa and evaluation of systematic position in the fami-ly balsaminaceae.     

Applying plant DNA barcodes to identify species of Parnassia (Parnassiaceae)

DNA barcoding is a technique to identify species by using standardized DNA sequences. In this study, a total of 105 samples, representing 30 Parnassia species, were collected to test the effectiveness of four proposed DNA barcodes (rbcL, matK, trnH-psbA and ITS) for species identification. Our results demonstrated that all four candidate DNA markers have a maximum level of primer universality and sequencing success. As a single DNA marker, the ITS region provided the highest species resolution with 86.7%, followed by trnH-psbA with 73.3%. The combination of the core barcode regions, matK+rbcL, gave the lowest species identification success (63.3%) among any combination of multiple markers and was found unsuitable as DNA barcode for Parnassia. The combination of ITS+trnH-psbA achieved the highest species discrimination with 90.0% resolution (27 of 30 sampled species), equal to the four-marker combination and higher than any two or three marker combination including rbcL or matK. Therefore, matK and rbcL should not be used as DNA barcodes for the species identification of Parnassia. Based on the overall performance, the combination of ITS+trnH-psbA is proposed as the most suitable DNA barcode for identifying Parnassia species. DNA barcoding is a useful technique and provides a reliable and effective mean for the discrimination of Parnassia species, and in combination with morphology-based taxonomy, will be a robust approach for tackling taxonomically complex groups. In the light of our findings, we found among the three species not identified a possible cryptic speciation event in Parnassia.     

中国15种变豆菜属植物的花粉形态及系统学分析

利用光学显微镜和扫描电子显微镜对变豆菜属15种植物的花粉形态进行观察。结果表明,变豆菜属15种植物的花粉大小为（28.51~54.29） μm~（12.23~28.98） μm,极轴与赤道轴比值（Polar axis/Equatorial axis）为1.48~2.46,具三孔沟。花粉粒赤道面观有近矩形、赤道收缩形、长椭圆形等类型,其中近矩形和赤道收缩形占多数;极面观为圆形或三角圆形。在扫描电镜下,其外壁表面纹饰为网状。我们研究结果支持在伞形科变豆菜亚科中,变豆菜属植物的花粉具有中等进化、比较进化的形态特征,花粉形态可为个别物种的分类处理提供孢粉学证据。     

茴芹属16种植物的花粉形态及系统学分析

利用光学显微镜和扫描电镜对茴芹属16种植物的花粉形态进行了观察,其中12种为首次报道。结果表明,茴芹属16种植物花粉大小为（19.75～33.03）μm×（11.52～17.41）μm,极轴与赤道轴的比值（P/E）为1.40～2.28,体积大小指数（P×E）^1/2为16.71～23.97,属于中型花粉,具三孔沟,萌发孔位置为边萌发孔。花粉粒赤道面观有近矩形、赤道收缩形、椭圆形和近菱形等类型,其中近矩形和赤道收缩形占多数;极面观为三角形或三角圆形。扫描电镜下,其外壁表面纹饰赤道面观为脑纹状,极面观近光滑或为穴纹状。在伞形科芹亚科中,茴芹属植物的花粉具有中等进化、比较进化的形态特征。研究认为,花粉形态可为个别物种的分类处理提供孢粉学证据。     

山小橘属 (芸香科) 花粉形态及其系统学意义

利用扫描电镜（SEM）对芸香科山小橘属（Glycosmis）12个分类群16个样品的花粉形态特征（包括花粉大小、萌发孔及外壁纹饰等）进行了观察研究,其中光叶山小橘（G.craibii var. glabra）、长果山小橘（G.erythrocarpa）、锈毛山小橘（G.esquirolii）、长叶山小橘（G.longifolia）、亮叶山小橘（G.lucida）、海南山小橘（G.montana）、少花山小橘（G.oligantha）、小花山小橘（G.parviflora）和华山小橘（G.pseudoracemos）等9个分类群的花粉形态为首次报道。结果表明,国产山小橘属植物的花粉均为单粒花粉,近球形或长球形,花粉大小范围为(15.90~23.92)×(14.56~22.00)μm,极面观为圆形、三裂圆形或近三角形,极轴与赤道轴比值（P/E）范围为1.03~1.31,均为3孔沟;根据花粉外壁纹饰可以将国产种类划分为条网状、复条网状、条纹-穿孔纹饰以及孔穴状纹饰等四种类型,其中小花山小橘不同来源花粉的外壁纹饰差异较大,至少存在5种类型：(1) 极面穴状,赤面条网状或穿孔,(2) 极面穴状,赤面均匀分布圆形穿孔,(3) 具复条网状,(4) 条网状兼具不规则小穿孔,(5) 条网状兼近圆形穿孔纹饰。研究发现山小橘属植物的花粉形态特征具有较高的多样性,为该属植物的分类及系统学研究提供了可用的孢粉学资料。     

国产省藤属植物的花粉形态学

对国产棕榈科省藤属(Calamius L.)15种植物的花粉进行了光学和扫描电镜观察,其中12种为首次报道。省藤属的花粉均为两沟型花粉,外壁覆盖层多为网状纹饰(大喙省藤C．macrorrhynchus)或具穿孔(华南省藤C.rhabdocladus)。首次发现省藤属花粉的外壁纹饰存在穿孔和外壁疣状突起的类型(阔叶鸡藤C.pulchellus)以及皱波状突起的类型(长鞭省藤C.flagellum)。花粉的大小、形状、外壁纹饰、外壁是否有突起,外壁厚度和网状纹饰网眼的大小,对于省藤属的种级分类有较大的意义。     

阿拉善荒漠驼蹄瓣属7种植物的花粉形态研究

用扫描电镜对阿拉善荒漠驼蹄瓣属(ZygophyllumL.)7种植物的花粉形态特征进行了观察分析.结果表明,在扫描电镜下,每种植物花粉粒的表面纹饰、萌发孔类型等性状符合驼蹄瓣属的主要特征;而花粉粒的大小、P/E值等细微特征存在明显差异.     

凤仙花属（Impatiens L.）10种植物花粉形态的扫描电镜观察

利用扫描电镜观察了10种凤仙花属（Impatiens L.）植物的花粉形态。结果表明：本属花粉为单粒花粉,呈长圆形至长矩圆形,大小为20.3～46.7 μm,具角萌发孔,网状纹饰,网眼明显;根据花粉网状纹饰中网眼内是否具颗粒状突起可将其分为2类：（1）网眼内无或近无颗粒状突起,黄金凤（I. siculifer）和婺源凤仙花（I. wuyuanensis）的花粉纹饰属于这一类型;（2）网眼内有明显颗粒状突起,其余8个种的花粉纹饰均属于该类型。研究表明,花粉特征,特别是花粉粒网状纹饰中网眼内有无颗粒状突起及颗粒状突起的形态等特征,在凤仙花属内常具种水平上的可见变异,因而可作为种类划分的依据,它们在分类学上的价值应予以关注。     

中国西南地区当归属植物花粉形态及其系统进化分析

用光学显微镜和扫描电镜研究了分布于中国西南地区的当归属(Angelica L.)16种植物的花粉形态,其中有7种植物的花粉形态为首次报道。研究结果表明当归属植物花粉粒较大,分化明显,包含了伞形科花粉的原始、中等进化和进化的类型,其形态以菱形、椭圆形、近长方形、赤道收缩形为主,萌发孔为角孔或边孔,从花粉的这种多类型证明该地区的当归属植物既起源古老同时又在不断分化着。从孢粉学角度初步分析了当归属的属下系统,为该属植物种的划分及系统演化关系的探讨提供依据。     

山小橘属(芸香科)花粉形态及其系统学意义

利用扫描电镜(SEM)对芸香科山小橘属(Glycosmis)12个分类群16个样品的花粉形态特征(包括花粉大小、萌发孔及外壁纹饰等)进行了观察研究,其中光叶山小橘(G.craibii var.glabra)、长果山小橘(G.erythrocarpa)、锈毛山小橘(G.esquirolii)、长叶山小橘(G.longifolia)、亮叶山小橘(G.lucida)、海南山小橘(G.montana)、少花山小橘(G.oligantha)、小花山小橘(G.parviflora)和华山小橘(G.pseudoracemos)等9个分类群的花粉形态为首次报道.结果表明,国产山小橘属植物的花粉均为单粒花粉,近球形或长球形,花粉大小范围为(15.90～23.92)×(14.56 ～ 22.00) μm,极面观为圆形、三裂圆形或近三角形,极轴与赤道轴比值(P/E)范围为1.03 ～1.31,均为3孔沟;根据花粉外壁纹饰可以将国产种类划分为条网状、复条网状、条纹-穿孔纹饰以及孔穴状纹饰等四种类型,其中小花山小橘不同来源花粉的外壁纹饰差异较大,至少存在5种类型:(1)极面穴状,赤面条网状或穿孔,(2)极面穴状,赤面均匀分布圆形穿孔,(3)具复条网状,(4)条网状兼具不规则小穿孔,(5)条网状兼近圆形穿孔纹饰.研究发现山小橘属植物的花粉形态特征具有较高的多样性,为该属植物的分类及系统学研究提供了可用的孢粉学资料.     

Pollen Morphology and Its Taxonomic Significance of Caragana Fabr. (Fabaceae) from China

Pollen grains of thirty one species and two varieties of the genus Caragana Fabr. were observed under LM and SEM.The results revealed that the pollen exine ornamentation of the species examined could be divided into two types: perforate and reticulate.The later type consists of two subtypes according to lumina size and muri width. In the first subtype the lumina is round with d iameter equal to muri width,and the exine is perforate in two polar areas,while in the second subtype lumina is irregular or polygonal,with diameter sm aller than muri width.The pollen volume was divided into four types.The genus Caragana is a natural group which was indicated by the general resemblance of pollen morphology. In this genus the pollen morphological classification was not in concordance with the arrangement of the sections and series except Ser.Pygmeae and some species. The pollen variation at infraspecific level was obvious,especially the species distributed in Qinghai Xizang Plateau, such as C.jubata, C.bicolor and C. erinacea . The evolutionary trend of pollen morphology is from perforate to reticulate, which is congruous with the evolution from the pinnate leaf group to the almate leaf group in the genus.     

杜鹃花族花粉形态及其系统学意义

利用光学显微镜和扫描电子显微镜对杜鹃属（Rhododendron L.）和仿杜鹃属（Menzie-sia Smith.)26种植物的花粉形态进行了观察和比较研究,其中大部分种为首次报道。杜鹃属和仿杜鹃属的花粉均为四合花粉,呈正四面体排列。四合体为圆角三角形,连接紧密。杜鹃属四合体平均直径在30.29-62.45μm之间,仿杜鹃属四合体直径为26.38-32.19μm,单粒花粉为球形或近球形,具三沟孔。在四合体上,相邻花粉粒上的沟相接,有时可见粘丝存在。杜鹃属和二鹃属花粉大小和花粉粒外壁纹饰均不同,具有明显的属间差异。在扫描电镜下,杜鹃属亚属内组（或亚组）之间花粉粒具有不同的外壁纹饰,可为分类学研究提供孢粉学方面的证据,具有重要的分类学意义。花粉分析在杜鹃属内不能作为分亚属的主要特征,但可用于某些种的分类学研究。     

中国木犀属花粉形态研究及其系统学意义

观察了中国木犀属4组21种植物的花粉形态。花粉为圆球形或近长球形,外壁表面均具网状纹饰。大多数种类都具有三孔沟的萌发孔,极少四孔沟,或六萌发孔。内孔明显或不明显,有9种植物同时存在具三孔沟和具三沟二种类型的花粉及中间过渡类型(显脉木犀同时存在具三拟孔沟和具四拟孔沟)。很多特点表明,木犀属花粉的萌发孔在木犀科中处于由简单萌发孔向复合萌发孔过渡的阶段。根据网眼从沟间区中央到沟边是否变细,可将本属花粉分为两大类型,与分组有一定的对应关系。综合一些花粉特征,可发现本属花粉形态差异较明显,与一般形态差异有一定的相关性。花粉形态在木犀属具有一定的系统学意义。