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.     

Pollen Characters of Old World Gesneriaceae (Cyrtandroideae)

Pollen of 108 species out of 18 genera (from all tribes of the Old World Gesneriaceae [subf. Cyrtandroideae]) was examined using light, scanning and transmission electron microscopy. The pollen grains are small sized, isopolar and 3-colpate or 3-colporate. In equatorial view they are mostly spheroidal (rarely suboblate or oblate), in polar view circular or subangular. The most variable character is the exine structure and -sculpture. The tectum is perforate, microreticulate, coarsely reticulate or rugulate. Further important characters of the tectum are (a) the presence or absence of conical supratectate sculptural elements, and (b) the width of lumina being either equal or exhibiting different at the apo- and mesocolpium. Ten exine types are distinguished. Some genera and species can be well referred to a special exine type, e.g., Aeschynanthus, Epithema, Stauranthera grandiflora; in other genera several exine types occur, e.g., in the large and heterogeneous genus Didymocarpus. The pollen morphology of the two large subfamilies Cyrtandroideae and Gesnerioideae is compared.     

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。     

绣球属(Hydrangea L.)及近缘属花粉形态的研究

为深入研究绣球属植物花粉形态的分类学价值和系统学意义,厘清绣球属与近缘属之间的系统发育关系,该文利用扫描电子显微镜(SEM,scanning electron microscope)对国产绣球属及其近缘属41种绣球花科(Hydrangeaceae)植物的花粉形态以及表面纹饰进行了观察。结果表明:绣球属及其近缘属的花粉为三孔沟;形状多数为长圆体形或近球体形;赤道面观为椭圆形或圆形;极面观多为圆形,少数为三角形或圆三角形。花粉外壁纹饰可分为网状和孔穴状。网眼内的三级纹饰可分为光滑和具颗粒状突起。根据花粉形状和外壁纹饰类型将上述物种划分为4个组,即花粉的形状为长圆体形,表面纹饰为孔穴纹饰;花粉的形状为长圆体形,表面纹饰为网状纹饰;花粉的形状为近球体形,表面纹饰为孔穴纹饰;花粉的形状为近球体形,表面纹饰为网状纹饰。以上可进一步细分为8个类型。上述表明花粉形态证据可为绣球属及其近缘属的属下分类和种间界定提供重要佐证;但结合前人的系统发育重建分析该属植物花粉形态的系统学意义相对有限,如花粉形态证据对于该属及其近缘种属系统发育树上大支的界定难以提供有力的证据。     

蓼科花粉外壁超微结构的研究

用扫描电镜和透射电镜对蓼科6属15种的花粉壁构造的特点进行了观察和研究。结果表明：(1)外壁纹饰有如下几种：颗粒-穿孔,微刺-穿孔,微刺-穿孔-光滑,颗粒-穿孔-光滑,细网状,粗网状,皱块状,鼓锤状;(2)外壁结构分化成两个明显的层次,即外壁外层以及外壁内层。其外壁外层由覆盖层、柱状层和基层组成。然而,由于每一部分发育状况的不同而导致外壁结构有各种变化,如无覆盖层或无外壁内层。     

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。     

The palynology of the genus Rondeletia L. (Rubiaceae-Cinchonoideae-Rondeletieae)

Pollen of 20 species of Rondeletia, covering three subgenera and seven sections or “species groups” of the genus, was studied using light and scanning microscopy. Rondeletia, characterized by binucleate mature pollen grains like numerous other Rubiaceae, was found to be eurypalynous. The pollen is isopolar and small-sized (average equatorial diameters 11.4–22.8 μm) and 3-colporate (rarely 4-colporate). In equatorial view, the pollen grains are mostly ± spheroidal (rarely suboblate); in polar view (ambitus), they are mainly circular or semiangular (intersemiangular in one species only). Three different exine types are distinguishable: exine with 1) tectum minutely perforate, perforations usually isodiametric, 2) tectum with comparatively large, not isodiametric perforations or 3) coarsely reticulate; supratectal sculpturing elements are lacking; the width of the perforations and lumina is either equal or different at the apo- and mesocolpia; in a few species rod-like luminal processes are present. Palynological features of most of the investigated taxa tend to overlap so that palynology is not very helpful in supporting the subgeneric division of Rondeletia. It was, however, noted that pollen morphology, to some extent, supports the sections of subgen. Arachnothrix. Palynologically, the subgen. Rogiera was found to be heterogeneous in that each of the three investigated species is characterized by a different exine type; one of these, R. strigosa, differs from all other Rondeletia species investigated in having a coarsely reticulate exine. R. odorata (subgen. Rondeletia) differs from all other investigated species in its intersemiangular pollen shape.     

Pollen morphology of the subtribe Cuspariinae (Rutaceae)

The neotropical subtribe Cuspariinae (Rutaceae) comprises as many as 26 genera and over 125 species. Pollen grains from 111 collections representing 71 species and 24 genera were examined by LM, SEM, and TEM. The pollen morphology of this subtribe is very diverse. Grains are mostly 3–6-aperturate and colporate, rarely porate (Spiranthera) or pantocolporate (Almeidea). Exine sculpturing is most commonly reticulate, sometimes perforate, foveolate-perforate, foveolate, foveolate-reticulate, reticulate, striate-reticulate, echinate, clavate, or baculate. The exine structure is columellate and tectate-perforate, columellate and semitectate, or intectate and is stratified into ektexine and endexine. The exine ofLeptothyrsa is distinctive in that the ektexine of the mesocolpium is longitudinally deeply ridged. The pollen ofHortia, characterized by a psilate exine with rare perforations, a very thick foot-layer, and reduced columellae, is unlike that of any member of the Cuspariinae and offers no support for the transfer of this genus from the Toddalioideae. The pollen data correlate with macromorphological characters and are taxonomically useful.     

Pollen morphology of endemic species of the Horton Plains National Park,Sri Lanka

The pollen morphology of 27 endemic and palaeoecologically-important species belonging to 16 families and 20 genera from the mountain rain forests of Sri Lanka was studied using both light (LM) and scanning electron microscopy (SEM). The pollen grains exhibit a wide range of morphological characters. Their sizes (P and E) are in the range 10-53 2 9-60 mum, and they are distributed among seven shape classes. All the taxa have radially-symmetrical pollen grains except for Impatiens (bilateral). The amb varies from rounded to triangular, with intermediate shapes possible. The pollen grains of all the Lauraceae species studied are inaperturate and microspinose, with a thin, fragile, perforated exine. Sarcococca zeylanica (Buxaceae) has pantoporate pollen grains and a Croton -pattern consisting of faintly striate pegs of various shapes attached or enveloping the smooth rings. Colp(or)ate, syncolpate to parasyncolp(or)ate pollen grains with rugulate to perforate exine patterns occur in Eugenia mabaeoides ssp. mabaeoides and Syzygium ssp. (Myrtaceae). Osbeckia walkeri (Melastomataceae) has heterocolpate pollen with fossulate, foveolate to perforate exine pattern. 3-colporate, microreticulate pollen grains are met with in Euonymus revolutus (Celastraceae), and scabrate-microreticulate ones in Calophyllum walkeri (Clusiaceae). Hedyotis lawsoniae (Rubiaceae) pollen is microreticulate to perforate, while pollen of the Elaeocarpus (Elaeocarpaceae) species is 3-colporate, smooth to scabrate-perforate. Casearia thwaitesii (Flacourtiaceae) has 3-colporoidate pollen grains with a smooth, imperforate to occasionally sparsely perforate exine. Mastixia spp (Cornaceae) have a granular-perforate exine pattern, while Adinandra lasiopetala (Theaceae) has a smooth-perforate exine pattern. Triangular 3 (4)-colporate, and indistinctly faintly microreticulate-perforate pollen was found in Rhamnus arnottianus (Rhamnaceae), rugulate-perforate in Isonandra montana (Sapotaceae) and verrucate-perforate in Symplocos elegans (Symplocaceae). Glochidion coriaceum (Euphorbiaceae) has a 5-colporate grain with a coarsely reticulate exine pattern. Pollen grains of Impatiens walkeri and I. thwaitesii (Balsaminaceae) are bilateral, 4-colpate, reticulate and granular. A pollen key based on light microscope is constructed for all the taxa studied.     

中国柽柳科植物花粉形态的研究

本文用光学显微镜和扫描电镜观察了分布于我国的4属30种柽柳科植物花粉形态。并用透射电镜研究了柽柳属的代表种, Tamarix elongata外壁内部的超微结构。本科花粉为三沟, 少数属种为三拟孔沟。根据扫描电镜观察, 柽柳属花粉外壁表面具粗网状纹饰和细网状纹饰, 以及界于这两个类型之间的过渡类型。水柏枝属外壁表面具粗网状纹饰, 细网状纹饰, 小穿孔以及小的蠕虫状突起。红沙属和枇杷柴属均为细网状纹饰。 本科柽柳属花粉与杨柳科中柳属的花粉特征较相似, 本文提出了它们之间的区别。     

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.     

A Study on Pollen Morphology of Potamogeton,Zannichellia and Triglochin from China

The pollen morphology of 19 species in the genera Potamogeton, Zannichellia and Triglochin from China was examined under light and scanning electron microscopes. 1. Pollen characteristics (1) Potamogeton L. (Plate 1, 2:1-8) Pollen grains spheroidal, subspheroidal, rarely oblate or ovoid, (17-35)24.41×27.76(18-45)μm in size; anaperturate; exine 1-2μm thick, 2layered or obscure; crass-reticulate, reticulate or finely-reticulate under LM, distinctly reticulate under SEM; muri uniform or not uniform, smooth or granular, rarely with processes; bacula thicker or thinner at the intersection of muri. (2) Zannichellia L. (Plate 2:9) Pollen grains spheroidal, ,(20-26)23.1×24.15(21-27) μm in size; anaperturate; exine 1 μm thick, stratification obscure; nearly reticulate under LM, shallowly reticulate under SEM; luminae shallow, large and irregular, muri very thin and not uniform; bacula absent. (3) Triglochin L. (Plate 2:10-11) Pollen grains spheroidal or subspheroidal; (20-30)23.65×23.7(20-30) μm in size; anaperturate; exine about 1μm thick, stratification obscure, finely reticulate under LM, distinctly reticulate under SEM; luminae rather large or minute; muri uniform, with 1-2 lines of granules, bacula present or absent. 2. Pollen types of Potamogeton L. (1) Thin muri--thick bacula: Potamogeton pectinatus, P. intramongolicus, P. filiformis, P. pamiricus; (2) Uniform muri and bacula: 1) Large pollen subtype: Potamogeton crespus; 2) Medium-sized pollen subtype: Potamogeton heterophyllus, P. perfoliatus, P. distinctus, P. malaianus; 3) Minute pollen subtype: Potamogeton oxyphyllus, P. obtusifolius, P. lucens, P. cristatus, P. gramineus, P. octandrus, P. pusillus     

Pollen Morphology of Guttiferae in China

The present paper deals with the pollen morphology of 23 species belonging to 6 genera of Guttiferae occurred in China. All pollen grains have been observed under light microscope, and those of 11 species in 5 genera have been examined with SEM as well. The pollen grains of Guttiferae are subspheroidal, spheroidal or prolate, (17–43) × (15–26)μ in size. Mostly 3-eolporate, less frequently 4-colporate or 5-colporate. The exine is 2-layered, but the occassionally demarcation of these layers is generally indistinct. The exine is 0.8–2.6 μ in thickness. The oramentations of all the pollen grains is generlly finely reticulate under the light microscope. Pollen morphology of the 11 species of 5 genera observed under the SEM is stressed. Their exine oramentations may be classified into four major groups, ie. reticulate, finely reticulate, baculate and perforate. Calophyllum: pollen grains reticulate and rugulo-reticulate. Cratoxylon: pollen grains reticulate, with finely and densely granulate over entire muri. Gareinia: pollen grains baculate, reticulate, and exine with perforate oramentation; in reticulate grains, muri with spinulation. Hypericum: pollen grains reticulate, perforate and finely reticulate. Mesua: pollen grains finely reticulate. From the comparison of pollen morphology, Hypericaceae has much connection with Guttiferae (sensu stricto). Because their pollen grains are all 3-eolporate and the exine is reticulate or oerforate under light mieroseope, but it is different between the pollen grains of Hypericum and those of Guttiferae.     

Pollen morphology in Cytisus (Papilionoideae,Leguminosae) from Morocco and the Iberian Peninsula

Although it is generally accepted that the tribe Genisteae has rather homogeneous pollen, we have found considerable variation in pollen size and morphology in Cytisus and related genera (Argyrocytisus and Chamaecytisus), which match taxonomic groups defined on morphological grounds. The results of Principal Component Analysis of pollen characteristics show the following: Cytisus Sect. Alburnoides is well-delimited, with the smallest pollen grains and the simplest pattern in the tectum, fossulate-perforate to perforate; No separation of C. villosus (Sect. Cytisus) from Sect. Alburnoides is observed; Cytisus Sect. Spartopsis, with the largest pollen grains and reticulate to reticulate-fossulate ornamentation, forms an isolated group; C. fontanesii (Sect. Heterocytisus) stands apart from other Cytisus species by its homogeneous ornamentation and the perforate instead of psilate-punctate margo. Thus, the pollen data supports its transfer to the separated genus Chronanthos; Pollen ornamentation and morphology also support the separation of Argyrocytisus battandieri and Chamaecytisus mollis from Cytisus.     

Floral ecology of Oreocharis pumila (Gesneriaceae): a novel case of sigmoid corolla

Attraction of pollinators and successful pollen transfer represent the primary targets of selection during flower evolution, leading to repeated evolutionary shifts between pollinators and consequently to the diversification of floral forms. However, most studies in floral evolution focus on the characteristics of flowers with straight corolla tube. Here, we report on an unusual form of sigmoid corolla combined with protandry and herkogamy in a Chinese species of Gesneriaceae, Oreocharis pumila (formerly Opithandra pumila). Contrary to species with sigmoid corollas studied previously, the base of the corolla tube of this species is inclined at an oblique angle downwards before the tube bends forward, and the stigma and anthers are included in the upper part of the corolla tube. The plants were found to be self‐compatible but incapable of autonomous selfing. Successful pollination was found to depend fully on the presence of insect pollinators (Nomia sp.) and pollen grains are the greatest reward for the visitors. Different from the other sigmoid flowers, the sigmoid corolla of O. pumila was not found to favor insect pollinators with long flexible proboscises. A mechanical fit between floral morphology and pollinator was found, in which only small insect visitors with specialized visiting behavior are legitimate pollinators. The protandry combined with herkogamy in the sigmoid corolla tube strongly ensures pollination efficiencies. Oreocharis pumila is the only species with sigmoid corolla in the genus Oreocharis. We hypothesize that such a corolla has arisen through selection due to inadequate pollination in early spring in the mountainous habitat that O. pumila occupies.     

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.     

锦鸡儿属花粉形态及其分类意义

在光学显微镜和扫锚电镜下对锦鸡儿属31种2变种植物花粉形态进行了观察。根据外壁表面纹饰,可分：类型I,表面具小穿孔;类型II, 具网状纹饰。 在类型II中据网眼大小和网脊宽度特征又分为两个亚类型。 根据花粉体积大小,可分4个类型。本属植物花粉形态较为一致,表明是一个自然类群。组、 系的花粉形态特征与一般形态变异不一定相对应,即形态上有明显区别的种类,在花粉形态上未必有很大区别,仅一些种类及矮锦鸡儿系 Ser. Pygmaeae的花粉形态相近。 本属花粉形态在种下等级的变异, 特别是青藏高原分布的种类的变异较为明显,例如 C.bicolor,C. jubata,C. erinacea 与各自的近缘种类。花粉形态为分类处理提供了参考。本属植物花粉形态是从表面具小穿孔向具网状纹饰演化,对应于羽状叶类群向掌状叶类群演化。     

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 (Parnassia faberi Oliv.) colpi,implying that Southwest China is the center of diversification of the genus.