Systematic importance of pollen morphological features of selected species from the genus Rosa (Rosaceae)

The aim of this study is verification of the taxonomic usefulness of the pollen grain features studied, based on pollen morphology of 32 wild species from all 4 subgenera and all 10 sections of the genus Rosa, mainly for delimitation of subgenera, sections, and species. The measurements and observations were carried out with both light microscopy and scanning electron microscopy. Only correctly formed pollen grains (30 per specimen) were measured, and 960 pollen grains were examined in total. They were analyzed for 14 quantitative features of pollen grains and exine sculpturing and the following qualitative ones: outline, shape, and operculum structure. Our study revealed that the diagnostic features of pollen grains studied were: exine sculpture, length of polar axis, and pollen shape (P/E ratio). On the basis of the above characters, five species were isolated and the remaining ones were included in several groups isolated on the basis of exine sculpture types. The following three exine sculpture types occurred in the species studied: granular-verrucate (in R.?stellata), striate-psilate (in R.?multibracteata and R.?multiflora), and striate (the remaining species). R.?banksiae is characterized by small pollen grains, while R.?setigera has strongly elongated pollen with P/E ratio >1.5. Exine sculpture features considered to be diagnostic should be treated as auxiliary because they fail to differentiate individual species, although they can be helpful in distinguishing groups of species of similar exine sculpture. The arrangement of the species examined on a dendrogram only slightly corroborates division of the Rosa genus into subgenera and sections currently adopted in taxonomy (Rehder 1940). An interesting result was reported for the species studied from the Caninae (R.?agrestis, R.?canina, R.?dumalis, R.?jundzillii, and R.?rubiginosa) section which, despite hybrid nature, with the exception of R.?villosa, grouped in the same, most separated group of species.     

Palynological study of Veronica Sect. Veronica and Sect. Veronicastrum (Scrophulariaceae) and its taxonomic significance

Pollen grains from 30 taxa, mainly from the Western Mediterranean area belonging to the perennial sections of the genus Veronica (Scrophulariaceae), were studied by LM and SEM. A close relationship was found between pollen size and ploidy level. Palynological characters cannot be regarded as essential taxonomic tools within the groups studied, although they have systematic value in some specific cases. The monophyly of Veronica subsect. Multiflorae is supported by a common and exclusive sculpture of exine. Delimitation of the genus is discussed and hypothetical evolutionary trends concerning pollen characters are addressed.     

旌节花科及其相关类群花粉形态的研究

对旌节花科及其相关类群五桠果科、猕猴桃科、水东哥科、金莲木科、省沽油科和山茶科等7科19属37种花粉进行了光镜（LM）和扫描电镜（SEM）的观察和比较。其中有些种类是第一次观察和报道。研究表明,上述各科花粉多为近球形至球形,少数近扁球形或近长球形。极面观三裂圆形或近三裂圆形。直径20－30（－35μm）,但厚皮香亚科和省沽油科银鹊树属Tapiscia的花科特别小,仅12.5-20μm。3孔沟,罕有3沟或4孔沟。外萌发孔（沟）相对地较长而宽,内孔多数较发达。花粉外壁纹饰变化大,从模糊而粗糙的细突起到皱波状纹饰,细孔状纹饰,蜂巢状－穴状纹饰,穴－网状纹饰,网状纹饰和颗粒状纹饰等。从形状、大小、萌发孔类型和外壁纹饰的比较看,虽然上述各科花粉均在不同程度上具有一定的相似性,但其中以旌节花科、省沽油科和山茶科中厚皮香科的花粉具有更多的相似性,尤其表现在外壁纹饰特征上。就孢粉学而言,也许旌节花科与省沽油科和山茶科中的厚皮香亚科的关系相对地较为密切。不过,其孢粉学特征所表现出的亲缘关系似乎不如在分子生物学中所表现出的明显。     

部分中国传统月季花粉形态研究

利用扫描电镜观察了15个中国传统月季品种的花粉形态。结果表明:传统月季花粉为单粒花粉,呈长球形或超长球形（P/E为1.92～2.25）,大小为37.59～51.95 μm×17.02～25.33 μm。赤道面观椭圆形或长矩形,极面观三裂圆形,具三孔沟,沿极轴方向等间距环状分布。外壁纹饰条纹型,覆盖层具穿孔,在品种间具有相似性,但变异丰富,划分为4种类型。聚类分析结果表明,花粉大小和外壁纹饰特征在反映传统月季品种的类别上与形态分类基本一致。根据花粉形态演变规律可推断,样品中‘四面镜’可能为最原始的品种类型,‘月月粉’、‘月月红’及‘匍匐红’等品种较为进化。     

Palynological survey of the subtribe Elephantopinae (Asteraceae, Vernonieae)

Despite the taxonomic value of pollen morphology within the tribe Vernonieae, a detailed study of the pollen of the subtribe Elephantopinae is still lacking. The pollen morphology of ten species, representing three of the four genera of the subtribe, Elephantopus, Pseudoelephantopus and Orthopappus, was studied with LM, SEM, and TEM. The pollen of all the species studied was found to be echinolophate, although the differences in aperture features and both sculpture and exine structure allowed recognizing two pollen types. The species of Elephantopus (except for E. elongatus) and Pseudoelephantopus share the regular or more or less regular ridges pattern, the colporate condition but with the ectoapertures little developed, and the acaveate exine structure. From the sculpture and the apertures, the pollen of Caatinganthus harleyi was found to be similar to that of Elephantopus and Pseudoelephantopus. The pollen morphology of the monotypic Orthopappus, which is shared with that of E. elongatus, was further investigated for the first time. We found that it differs from that of the other species in having an irregular pattern of ridges, colporate condition but with a well-developed ectoapertures, and caveate exine. Additional studies of the exine structure and apertures features, coupled with molecular phylogeny, are needed to understand the evolution of pollen characters and re-evaluate the intergeneric relationships within the tribe.     

中国特有属牛筋条属的花粉形态与其系统位置

从发育的角度研究了中国特有单种属DichotomanthesKurz及与其系统学研究有关的外类群Prinsepiautilis的花粉形态 ,扫描电镜观察显示Dichotomanthes花粉粒自脱离四分体胼胝质膜开始至成熟二核花粉粒不同发育时期 ,花粉形态和外壁纹饰未见变化 ,仅花粉体积随成熟度增加而有所增大。而Prinsepiautilis ,其花粉粒刚脱离四分体时形状和成熟花粉明显不同 ,成熟花粉极面观为三裂圆形 ,赤道面观为圆形 ,外壁具清晰的平行条纹 ,但幼嫩花粉粒的形状很特别 ,极面观为深三裂圆形 ,赤道面观亦见花粉在两条沟之间下陷而沟部外突 ,明显为角萌发孔花粉 ,且花粉体积较成熟者小 ,而外壁纹饰同成熟者相比无根本性差异。前述两种植物花粉在不同成熟期体积有明显差异 ,而外壁纹饰在不同成熟期不存在质的变化并相对稳定 ,说明花粉外壁纹饰这一性状在蔷薇科中具有较为重要的分类学意义。DichotomanthesKurz具典型Rosaceae花粉的三孔沟结构 ,外壁具条纹 -穴状纹饰。将其孢粉学特征同Rosaceae 4个亚科有关类群的同类资料相比较 ,并结合其它形态解剖与细胞学等研究结果 ,支持将Dichotoman thes置入Maloideae下而不赞同将其另立亚科或置于Prunoideae之下。此外 ,由于Prinsepiautilis的花粉在其发育初期具角萌发孔花粉 ,与Cunoniacea     

Pollen morphology of some endemic Turkish Centaurea L. (Asteraceae, section Phaloletis) and their taxonomic value

In this study, the detailed pollen morphological structures of some endemic Turkish species of Centaurea amaena Boiss., C. antalyense H. Duman & A. Duran, C. aphrodisea Boiss., C. hierapolitana Boiss., C. luschaniana Heimerl, C. lycia Boiss., C. tossiensis Freyn. Et Sint., and C. wagenitzii Hub.-Mor. (Asteraceae, section Phaloletis) were studied under light microscopy (LM) and scanning electron microscopy (SEM) for the first time. LM and SEM investigations showed that the pollen grains of eight taxa are more or less spheroidal-subprolate, the amb triangular and tricolporate. The exine sculpture is tectate, microechinate-scabrate in the pollen of Centaurea taxa. Spinules are less dense in Centaurea amaena, C. antalyense, C. hierapolitana, and C. lycia, but they are more dense in C. aphrodisea, C. luschaniana, C. tossiensis, and C. wagenitzii. Spinule dimensions are different from each other. The exine has one layer of columellae beneath the spines. We determined all taxa that have the Helianthoid type. Exine sctructure and sculpture as well as spine density and dimensions in Asteraceae are the most reliable characteristics for discriminating taxa.     

Pollen morphology,ultrastructure and taphonomy of the Neuradaceae with special reference to Neurada procumbens L. and Grielum humifusum E.Mey. ex Harv. et Sond.

Pollen morphology and sporoderm ultrastructure of modern Neurada procumbens L. and Grielum humifusum E.Mey. ex Harv. et Sond. were studied using light (LM) and electron (SEM and TEM) microscopy. Additionally late Holocene pollen of the Grielum-type was studied using LM. Systematic and ecological aspects have been discussed for the family Neuradaceae. The pollen grains of the studied species are characterized by similarities in size, shape, aperture type and differences in exine sculpture (reticulate semitectate exine in Neurada and finely reticulate to foveolate in Grielum) and sporoderm ultrastructure. The cavea in the exine is situated between the ectexine and endexine which are connected near the aperture region only. A combination of the palynological characters of the Neuradaceae (semitectate exine, rather loose columellae, interrupted foot layer, the cavea in the exine) increases the pollen plasticity, allowing considerable changes of the pollen grain volume but still remains insufficient to survive sharp fluctuations in hydration level.     

Pollen morphology in relation to the taxonomy and phylogeny of some native Greek Aegilops species

The pollen morphology of some native Greek Aegilops species is investigated in LM (quantitative pollen characters) and SEM (exine sculpture) using acetolysed material. Furthermore, quantitative data are subjected to a multivariate analysis. The tetraploid Ae. cylindrica and its diploid parent Ae. caudata show a distinct morphological affinity as regards their quantitative pollen profile and the features of the exine sculpture. The tetraploid Ae. triuncialis is morphologically divergent from both its parents Ae. caudata and Ae. umbellulata due to the very large values of its quantitative pollen characters. However, the SEM survey of the exine sculpture indicates a rather high degree of similarity between Ae. triuncialis and Ae. umbellulata. No significant differences have been found between the two varieties of Ae. caudata (caudata and polyathera) concerning the quantitative pollen characters as a total or the morphology of the exine sculpture. The resultant clustering of the taxa on the basis of the quantitative pollen characters as well as the recorded similarities of their exine pattern are related to their sectional classification based on cytogenetical and morphological data.     

Studies on the Pollen Morphology and Seed Coat of the Genus Cistanche (Orobanchaceae) in China

In the present paper pollen grains of 5 species and seed coat of 4 species of the genus Cistanche (Orobanchaceae) from China were examined by scanning electron microscope As a result, two types of the pollen exine sculpture are distinguished: (1) tuberculate, C. salsa and C. lanzhouensis; (2) rugulate or fine reticulation formed by the fusion of rugae, C. sinensis, C. deserticola and C. tubulosa. Chinese Cistanche was classified into two sections based on the gross morphology by the present author (Zhang, 1984). C. lanzhouensis and C. sinensis were included in one section. This classification is in conflict with the pollen type, which indicates that the characters of pollen grains and external morphology in this genus have evolved at different rates. The differences in pollen morphology of Chinese Cistanche can serve as characters for delimitating species. According to our abservation, C. lanzhouenis Z. Y. Zhang differs from the other members of Cistanche not only in external morphology but also in pollen morphology. Pollen grains are subprolate in this species, while prolate in the other four species, and exine sculpture is tuberculate. The present study provides the establishment of this new species (Zhang, 1984) with palynological evidence. The seed coat sculpture in Chinese Cistanche are constantly alveolate, but there are some slight differences, by which the sculpture can be divided into two types: (1) testa cells have or have not thickenings on the inner anticlinal walls; cavernulous sculpture is present on outer periclinal wall of some cells: C. sinensis; (2) testa cells have striate thickenings on the inner anticlinal walls: C. salsa, C. deserticola and C. tubulosa. The characters of seed coatof some significance for the delimitation of species.     

Pollen morphology of the family Cistaceae in Egypt and its systematic significance

The pollen morphology of 11 species (including two subspecies and two varieties) belonging to two genera (Helianthemum and Fumana) of the family Cistaceae in Egypt was studied using light and scanning electron microscopy.Pollen grains of the studied taxa were found to be radially symmetrical and tricolporate.Pollen size,shape,apertures,and exine ornamentation characteristics were valuable parameters among the studied taxa.The largest pollen size was recorded in H.salicifolium and the smallest one observed in H.kahiricum subsp,schweinfurthii.Pollen shape in the Egyptian taxa varied from (sub-)prolate to prolate spheroidal,but F.arabica is different in having sub-oblate grains.The pollen data confirm that H.lippii and H.sessiliflorum are very closely related species.Pollen sculpture was useful in distinguishing between H.vesicarium var.vesicarium and H.vesicarium var.ciliatum.Three main pollen types of exine ornamentation were recognized:retipilate; reticulate to verrucate; and striate.Based on palynological data,a key for the studied taxa is suggested.     

Pollen morphology of the genus Gagea (Liliaceae) in Iran

Pollen grains of 26 species of the genus Gagea distributed in Iran were examined by light and scanning electron microscopy. Detailed pollen morphological characteristics are given for these species. Among the studied species, the newly described G. iranica together with G. olgae and G. graminifolia possess the smallest pollen grains, and the widely distributed G. lutea the largest ones. Our studies show that the sculpturing of exine provides valuable characters for separating the species, sometimes even for closely related ones, and delimitation of natural groups within the genus. The exine of the genus Gagea is in most cases perforated upon tectum or rarely tectate-columellate. The muri are solid or structured, compound, simpli-, dupli- or pluricolumellate. It seems that the structure of muri is very important in recognizing natural groups within the genus. Tectal perforations vary from <0.2 to 2.0 μm in diameter among the studied species. Regarding sculpturing of the exine in proximal face, four basic types of pollen grains can be distinguished: reticulate, microreticulate, foveolate and perforate. Within the reticulate type there is sufficient variation in exine structure at distal face to describe three subtypes: reticulate, microreticulate and perforate. A diagnostic key is given for all studied taxa based on palynomorphological characters. For a limited number of populations of selected Iranian species of Gagea, further aspects of pollen biology were studied. It seems that populations with ploidy levels other than diploidy, show a low percentage of pollen fertility. Moreover, the rate of pollen fertility is correlated with the manner of the reproduction in certain species.     

Pollenmorphologie Von Borago Officinalis L.

The pollen morphology of the 16 recognized species of the genus Asphodelus L. (Asphodelaceae) has been investigated by light and scanning electron microscopy. Four pollen types can be distinguished on the basis of variation in size of the polar, equatorial longitudinal and transverse axes, as well as in exine structure and sculpturing, which correlate with the subdivision of the genus into sections. Correlations between pollen size and some biological characters are made. Phylogenetic implications are suggested, and trends of pollen evolution in the genus are indicated.     

槭树科花粉形态及其系统学意义

报道了槭树科(Aceraceae)槭属21组33种和金钱槭属2种植物的花粉形态。本科花粉近球形至长球形,极面观为三裂圆形。从花粉萌发孔类型看,金钱槭属Dipteronia具三孔沟,槭属（Acer)除4组具三孔沟外,其余均为三沟。从花粉外壁纹饰看,金钱槭属2个种和槭属的大多数种为条纹状,罕为细条纹－拟网状和皱波状。通过花粉形态分析,并结合其它方向的证据,我们认为：（1）Sect.Palmata,Sect.Spicata和Sect.Microcarpa可能是槭属中与金钱槭属关系最密切的类群;（2）A.distalum和A.nipponicum代表了近缘的两个单种组;（3）A.pseudoplatanus不同于Sect.Acer的其它4个种而与A.saccharum可能存在更为密切的关系;（4）Sect.Carpinifolia(细条纹－拟网状外壁纹饰）和Sect.Negundo(皱波状外壁纹饰）则可能代表了槭属中最特化的类群;（5）按槭树科花粉三孔沟到三沟的演化规律,Dipteronia较Acer原始。     

山茶属的花粉形态及其分类学意义

本文对山茶属(Camellia L.)18个组27个代表种的花粉进行了光学显微镜、扫描电镜和大部分进行了透射电镜的观察,并对花粉大小、形状、外壁纹饰和超微结构等作了综合的比较。结果表明,其形状和大小差异不很明显。花粉壁的层次、厚度和不同层次的比例,如复盖层与柱状层以及外壁外层与外壁内层的比例等也没有显著的差异。但花粉壁的纹饰特征在扫描电镜下显示出较大的差异,具有较重要的分类学价值。根据其纹饰特征,可以把山茶属花粉大致区分为六个类型:皱疣状纹饰;粗瘤状纹饰;颗粒状至皱-颗粒状纹饰;穴-网状纹饰;皱沟状纹饰及皱波-念珠状纹饰。孢粉学上区分的类型不完全与各个学者从植物形态上划分的组相一致。其中管蕊茶 Camellia lanceolata的纹饰非常特殊(粗瘤状),其外壁表面很象牛心果(Annona reticulata)的外形特征,为其重新分类提供孢粉学依据。     

Stylar polymorphism, reciprocity and incompatibility systems in Nymphoides montana (Menyanthaceae) endemic to southeastern Australia

Heterostyly is a stylar polymorphism that has been shaped by the evolution of floral characters adapted for efficient pollen transfer. Different types of stylar polymorphism are described in which the discrete characterization of the exact polymorphic type (e.g., distyly vs. stigma-height dimorphism) requires detailed floral measurements (e.g., sex-organ reciprocity). In clonal and aquatic Nymphoides montana, although the presence of two floral morphs that contain styles of two lengths has been previously reported, no studies have quantitatively estimated the level of reciprocity and/or described the stylar condition. Morphological variations and incompatibility relationships were explored between the two morphs in three southeastern Australian populations. In this study, one population is characterized as stigma-height dimorphism (i.e., two morphs with discrete variation in stigma height but little variation in anther height), whereas the other two populations are typical distylous (i.e., two morphs for reciprocal stigma and anther height). Nymphoides montana is dimorphic in a wide range of ancillary characters, including corolla size, stigma size, shape and papillae morphology, and pollen size, number and exine sculpture. Following glasshouse pollinations, full incompatibility systems were observed in the distylous populations, whereas the stigma-height dimorphic population showed between-morph variation in the extent of incompatibility. Despite the variation in sex-organ reciprocity and incompatibility, other lines of evidence appear to assure the maintenance of the stylar polymorphism in the N. montana study populations. All populations are nearly isoplethic (i.e., both morphs in equal frequencies), which is indicative of balanced polymorphisms that appear to be maintained by legitimate pollen transfer between the morphs.     

Pollen Morphology of the Genus Malus and Its Taxonomic and Evolutionary Significance

The pollen morphology of 26 species and 5 hybrids of the genus Malus was investigated with aid of SEM. It is found that the pollen morphology of Malus is rather similar in shape, size, position and number of aperture and exine sculpture. The characters of polle morphology of sections and series are as follows: Sect. Malus Ser. Baccatae: Striae regular, parallel to colpi, and conjunct at pole; Ser. Pumilae: the same as in Ser. Baccatae, but more or less curved near pole. Sect. Docyniopes: Striae regular and parallel to colpi, but bent near pole. Sect. Chloromeles: Striae irregular, dense and interlock. Sect. Sorbormalus: Striae irregular, sparse and not interlock. Ser. Sieboldiance: pollen grains prolate, with perforation among striae, colpi narrow; Ser. Kansuenses: Pollen grains spheroidal, colpi wide in the middle but narrow at both ends, striae relatively dense and regular, mostly dichotomous, perforation present; Ser. Yunnanenses; Pollen grain spheroidae, colpi wide in the middle but narrow at both ends, striae obviously irregular, less dichotomous, perforation absent. The major evolutionary trend of exine sculpture of pollen may be from densely thin-striate to sparsely striate with perforation. Characters of the exine sculpture of hybrids can be used to recognize the rela-tionship between the parental species.     

Pollen morphology in relation to pollinators in Papilionoideae (Leguminosae)

The surface sculpturing of the pollen of some species of the taxonomically widely separated genera Harpalyce (tribe Brongniartieae), Camoensia (tribe Sophoreae), Millettia (tribe Tephrosieae), and of the monotypic Dahlstedtia (tribe Tephrosieae) which have large red or white flowers adapted for pollination by birds or bats, is coarsely rugulate or verrucate. Related taxa with small insect pollinated flowers have pollen with simple reticulate or perforate surface sculpturing. The exine stratification of Alexa and Castanospermum (tribe Sophoreae), genera with large red bird-flowers, is complex with a layer of tectal columellae and differs from that of other genera in the tribe Sophoreae which have a normal pollen wall structure. These modifications of pollen structure and sculpture appear to be the result of convergent evolution and a secondary adaptation to pollination. The taxonomic and functional significance of the observations are briefly discussed.     

A palynological study of the genus Nepeta L. (Lamiaceae)

The pollen morphology of forty taxa of the genus Nepeta L. was studied and documented in detail using light microscopy (LM) and scanning electron (SEM) microscopy in this study. Nepeta L. pollen grains are small to large (P = 18.64–63.46 μm, E = 15.62–46.33 μm), suboblate to perprolate (P/E = 0.86–2.09) in shape and hexacolpate (very rarely tetracolpate) with granular membranes. Alternate position of colpi occurs in six Nepeta taxa, N. nuda ssp. glandulifera, N. concolor, N. crinita, N. congesta var. cryptantha, N. stricta var. stricta and N. sibthorpii ssp. tumeniana. In examinations of exine ornamentation with SEM, two types of pollen grains were recognized: (1) type I, with microreticulate sculpture; and (2) type II with bireticulate sculpture, type I and II to be divided into two and five subtypes, respectively. In the two taxa with microreticulate pattern, N. pilinux and N. sulfuriflora, a tendency towards a bireticulum could be recognized due to traces of secondary tectal connections. The bireticulate exine ornamentation is characterized with varying characteristics of the primary muri and secondary reticulum. Pollen morphology within the genus is compared with infrageneric relationships.     

国产黄耆属簇毛黄耆亚属花粉形态及其系统学意义

观察了国产黄耆属簇毛黄耆亚属6组12种植物的花粉形态。这些花粉的外壁表面纹饰可分为网状纹饰,穿孔或小沟渠状,以及穴网状纹饰3种类型。所有种类都具三孔沟的萌发孔,根据内孔明显与否可分为两类。虽然萌发孔类型与分组并没有严格的对应关系,但综合一些花粉特征,仍然发现6个组间花粉形态差异较为明显,与一般的形态差异有一定的对应性。特别是Sect．Robusti比较特殊,根据花粉大小等特征,推测它和Sect.Phyllolobium等有可能是本亚属的原始类群。Sect.Sesbanella的两个近缘种Astragalus heydei和A.hendersonii,花粉形态却很有区别。Sect.Bibacteola种类较多,有24种,花粉形态呈多样性。总之,花粉形态在簇毛亚属中具有一定的系统学意义。