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.     

Some evidence for infrageneric classification in Ziziphora L. (Lamiaceae:Mentheae)

The genus Ziziphora L. (Lamiaceae) consists of annual herbs, except Ziziphora clinopodioides, and is represented by six taxa in the flora of Turkey: Z. clinopodioides, Z. capitata, Z. persica, Z. tenuior, Z. taurica subsp. taurica and Z. taurica subsp. cleonioides. They are strong aromatic herbs and important medicinal plants. This study has been conducted to determine the palynological and karyological features of these taxa. The pollen morphology and exine structure of Ziziphora species were investigated by light microscope and scanning electron microscope. Pollen grains were determined as hexacolpate, medium in size and a circular or elliptic amb. The shape of pollen grains varies from oblate to prolate spheroidal. Ornamentation types of pollen grains are bireticulate, microreticulate or perforate. Somatic chromosome number has been determined and counted as 2n = 16, 18. Results indicate that the pollen characters and chromosome numbers of the genus are valuable for taxonomic applications and may be useful for infrageneric classification.     

Pollen Morphology of the Aceraceae

Pollen of 40 Acer taxa and one species of Dipteronia were examined by light, scanning and transmission electron microscopy. Pollen grains in both genera are tricolpate or tricolporate of prolate to nearly spheroidal shape. Based on sexine pattern, four major pollen types were recognized and include striate grains, rugulose grains, microreticulate grains, and granular grains. Although the family appears to be rather stenopalynous, several useful taxonomic variations occur.     

Distyly in Polygonum jucundum Meisn. (Polygonaceae)

Pollen grains of tribe Sanguisorbeae (Rosaceae, Rosoideae) were examined using scanning electron microscopy to identify useful characters, test taxonomic and phylogenetic hypotheses among genera, and elucidate pollen character evolution based on a molecular phylogeny. Aperture number, aperture structure, pollen shape, and exine sculpturing were variable within Sanguisorbeae and were used to delineate six pollen types. Four types (I–IV) were observed only in subtribe Sanguisorbinae whereas two types (V–VI) were found only in subtribe Agrimoniinae. Pollen grains of tribe Sanguisorbeae were generally subprolate to spheroidal in shape, had operculate or pontoperculate apertures, and had three apertures, except for Margyricarpus (tetraperturate). Exine sculpturing within Sanguisorbinae represented variations of striate, verrucate, rugulate, and perforate patterns often with microechinate sculpturing. Striate exine patterns and prolate shapes characterized the pollen of the Agrimoniinae, except for the microechinate-verrucate pattern and subprolate to spheroidal shapes observed in Hagenia. Pollen characters are most useful at the generic level and, when mapped on to a molecular phylogenetic tree of the tribe, are concordant with a monophyletic Agrimoniinae and a clade comprising Margyricarpus + Acaena + Polylepis + Cliffortia + Sanguisorba in the Sanguisorbinae. Outgroup comparison indicated that operculate colpi, three apertures, and polymorphism for striate or microverrucate exines represented primitive states for tribe Sanguisorbeae.     

Palynological analysis of five selected <Emphasis Type="Italic">Onosma</Emphasis> taxa

Pollen size and exine structure of the following five Onosma L. species from Slovakia and Italy were studied by light and scanning electron microscopy: Onosma echioides, O. tornensis, O. visianii, O. arenaria, and O. pseudoarenaria. Among the taxa studied, O. echioides has the smallest pollen grains in average (10.5–17.5 × 8.0–14.5 μm), and O. pseudoarenaria the largest grains in average (15.3–20.5 × 12.3–16.3 μm). Pollen grains of all 5 species are small sized, 3-syncolporate, subprolate, heteropolar, with ovate equatorial outlines and circular to rounded triangular polar outlines; the tectum is microechinate. A positive correlation is found between pollen size and chromosome number. The value of pollen characters for taxonomic purposes and the position of the taxa studied within the genus Onosma are discussed.     

Pollen morphology in the family Asphodelaceae (Asphodeleae,Kniphofieae)

Pollen of 72 species from 7 genera of the Asphodelaceae was studied by means of LM and SEM in order to investigate their taxonomic relationships. The pollen of all the genera investigated is heteropolar, monosulcate, non-operculate and differs in size, form and exine ornamentation. The species of Eremurus are homogenous palynologically and studies of this genus do not support the segregation of the genera Henningia, Selonia, and Ammolirion. The pollen of Bulbinopsis mainly differs from the pollen of Bulbine species in having a perforate-areolate exine surface (B. bulbosa) but the genera are close in their other palynological features. The genera Asphodelus and Asphodeline have the largest, almost spheroidal grains with the thickest exine but differ in details of exine ornamentation. The species of Kniphofia are very close palynologically.     

COMPARATIVE POLLEN MORPHOLOGY AND TAXONOMIC AFFINITIES IN CYCADALES

Scanning and transmission electron microscopy of pollen grains of 29 species, representing the ten extant genera of Cycadales, has provided valuable insight into their relationships. Pollen grains of these taxa are boat-shaped, monosulcate, and bilaterally symmetrical. They range from narrowly to widely elliptical or subcircular when viewed distally, and have an exine surface of psilate, foveolate, or fossulate. Pollen wall ultrastructure of Cycadales is typically tectate with alveolate∗∗∗spongy exine. The nexine is laminated in all genera. Nexine 1 (footlayer) is present in most species as a thin and often discontinuous layer. There is consistent variation in thickness of the sporoderm layers among the genera but relative uniformity within them. Pollen characteristics are well correlated with macro- and micromorphological features, chromosome numbers, geographical distribution, and postulated pollination mode. A close affinity between Encephalartos, Lepidozamia, and Macrozamia is recognized. Pollen characteristics of the genus Bowenia show some similarity with those of the latter group. Except for two species of Macrozamia which are narrowly elliptic, all of the genera have widely elliptic pollen and share a psilate exine surface and the thinnest sexine with nearly identical arrangement of alveoli. Pollen grains of the species in the genus Dioon exhibit a unique morphology but are more similar to Stangeria than they are to those of taxa in Zamiaceae. The circular outline of the grains and the foveolate exine surface are characters shared by these two genera, but several morphological features distinguish Dioon from Stangeria. Ceratozamia and Zamia share a widely elliptic shape, foveolate exine surface and nearly identical sexine, as well as morphological features and chromosome numbers. They differ from Microcycas in sexine thickness, gross morphology and chromosome numbers. The pollen grains of Cycas circinalis and C. revoluta differ in size and structure of the sexine from all other genera and from each other, substantiating their distinct subgeneric delimitations.     

Pollen morphology of the genera Polygonum s. str. and Polygonella (Polygoneae: Polygonaceae)

The pollen of 30 taxa (27 species, one subspecies and two varieties) in two genera, viz Polygonum s. str. and Polygonella was investigated with LM and SEM, and some selected taxa with TEM. In all genera investigated the pollen is prolate to spheroidal, and the aperture is mostly tricolporate, rarely panto-hexacolporate (especially Polygonum section Polygonum). The exine sculpturing pattern is the most variable feature. Three types of exine can be recognized. Type 1 (Avicularia-Type, sensu Hedberg) - All species of section Polygonum and section Tephis share the smooth tectate exine with spinules, sometimes the surface is more or less rough (Polygonum afromontanum in section Tephis). Type 2 (Pseudomollia-Type, sensu Hong) - Pollen of Polygonum molliaeforme (section Pseudomollia) has the exine, which is verrucose on both poles and nearby the mesocolpium, and mostly psilate around the ectoaperture. Type 3 (Duravia-Type, sensu Hedberg) - Pollen grains of Polygonum section Duravia and Polygonella have the exine which is semitectate-reticulate at the mesocolpium and the poles, and rugulate/reticulate or sometimes foveolate with microspinules around the ectoapertures. The pollen grains in four taxa (viz Polygonum section Pseudomollia, P. section Duravia and genus Polygonella) have a well-marked dimorphism of the ektexine, which is considered to be a synapomorphic condition. The differences of pollen grain between the genus Polygonella and Polygonum section Duravia are almost non existent and clearly interrelated. It is therefore postulated that the similarity in pollen of both taxa is not the result of convergency, but is interpreted as a homology. It is noteworthy that the pollen of Polygonum molliaeforme (section Pseudomollia) appears as intermediate between the Avicularia-type and the Duravia-type, and is well supported the value of separated section for its own. Additionally, in TEM, some exine ultrastructures (e.g. columellae, foot layer, endexine) appear to be valuable characters for comparison between/among taxa. The systematic potentialities of the pollen data of the studied taxa at various systematic levels are also discussed.     

蜘蛛抱蛋亚族的花粉形态

本文研究并报道了百合科蜘蛛抱蛋亚族的花粉形态,主要结论如下：1．开口箭属含两种花粉类型即：(1)单槽、椭圆形;(2)无萌发孔、球形。具单槽、椭圆形花粉的种类含单型性核型,花形较小,柱头小而花柱短,雄蕊高于柱头;无萌发孔、球形花粉的种类则为单型性核型,花形较大,柱头常高度膨大,雄蕊常着生在花被筒基部。2．万年青属花粉为椭圆形,具单槽;蜘蛛抱蛋属花粉则为球形,无萌发孔;开口箭属花粉一部分与万年青属相似,另一部分则与蜘蛛抱蛋属雷同。3．蜘蛛抱蛋亚族中最原始的花粉类型为单槽椭圆形,具孔状或网状外壁纹饰。球形,无萌发孔,外壁具皱波状、瘤状或芽孢状纹饰的花粉为派生类型。4．蜘蛛抱蛋亚族种下花粉形状、萌发孔类型和外壁纹饰基本稳定,可作为分种特征。     

Pollen Morphology in the Subtribe Aspidistrinae (Liliaceae s. l.)

Pollen morphology in the subtribe Aspidistrinae is reported and the main viewpoints are summarized as follows: (1) Two major types of pollen grains, i.e., monosulcate,ellipsoidal pollen and inaperturate, spheroidal pollen, are identified in the genus Tupistra. The species with monosulcate, ellipsoidal pollen usually have monomodal karyotypes, brightcolored flower, smaller stigma, and stamens inserted at upper or middle part of perianth tube, while the inaperturate and spheroidal pollen is always found in the species with bimodal karyotypes, dingy-colored flower, swollen stigma and stamens inserted at the base of perianth tube. (2) Pollen grains in the genus Rhodea are monosulcate and ellipsoidal, while those in the genus Aspidistra are inaperturate and spheroidal, correspondingly similar to the second type of pollen grains in the genus Tupistra. (3) The most primitive pollen in the subtribe Aspidistinae is regarded as monosulcate and ellipsoidal, having perforate or reticulate exine sculpture. The inaperturate, spheroidal pollen with verrucate, gemmate or rugulate exine sculpture is considered derived; (4) Unlike those reported in other groups of the family Liliaceae, the infraspecific pollen shape, aperture type and exine sculpture in Aspidistrinae are basically stable and may serve as a taxonomic character.     

螺序草属(茜草科)的花粉形态

首次对螺序草属(Spiradiclis B1.)的11种和1变型共16份植物的花粉样品在光镜和电镜下进行了观察。结果表明,本属植物花粉为单粒,小型和中型花粉,近球形至椭球形,辐射对称,常具3孔沟,偶为4孔沟,外壁表面纹饰为孔状。花粉外壁由不连续的具穿孔的覆盖层、柱状结构和基层组成,内外壁明显。此外,本属植物的花粉具有花粉2型和“芽状”结构的现象。螺序草属植物的花粉形态在属下分类上有一定的局限性,但是在个体的系统发育上却可以提供极有价值的信息,如具有花粉单型的种类比花粉二型的种类进化。     

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 of Xyridaceae Systematic (Poales) and its Potential

Xyridaceae are a predominantly tropical family of five genera that exhibit two pollen morphologies often considered to be of taxonomic importance. Xyris comprises about 95% of the species and is characterized by medium to large, elliptic, sulcate pollen grains. The other pollen class is spheroidal grains without an evident aperture. Many of the species with spheroidal grains have remarkably large and ornamented pollen found to be species specific in earlier research. A scanning electron microscopy investigation of 23 taxa representing all genera with spheroidal pollen revealed new data to further distinguish the genera based on pollen characters. Reliable specific pollen characters need to be evaluated in a statistical study.     

东北杜鹃花属植物花粉形态的研究

利用光学显微镜和扫描电子显微镜对东北杜鹃花属(Rhododendron Linn.)6种, 1变种, 1变型的花粉形态进行了观察和比较研究。该属花粉粒均为四合花粉。四合花粉呈正四面体排列。花粉粒近球形。每粒花粉均具3沟,在四合花粉粒上,相邻花粉粒上的沟相接,有些种可见沟内的萌发孔。光镜下可见花粉外壁为内外两层,厚度约相等。不同的种在花粉粒大小上有区别。在扫描电镜下观察可见不同的种花粉表面具有不同的纹饰。即该属在花粉形态上具有种的特性。     

Taxonomic implications of pollen exine morphology in infrageneric classification of Scutellaria (Lamiaceae)

Scutellaria L. in Lamiaceae subfamily Scutellarioideae is a subcosmopolitan genus with about 360 species. According to the latest subgeneric classification, two subgenera are recognized: S. subgen. Scutellaria and S. subgen. Apeltanthus. We studied pollen grains from 29 species of Scutellaria belonging to both subgenera and from both the Old and the New World to evaluate their taxonomic importance for the infrageneric classification of the genus. Pollen grains were acetolysed and studied by scanning electron microscope (SEM) and light microscope (LM). All examined pollen grains are isopolar, tricolpate, suboblate, oblate spheroidal, prolate spheroidal to subprolate. The exine is mainly bireticulate perforate, but with variations in lumina shape and size as well as perforation size and uniformity. Based on the exine sculpturing, two subtypes are recognized within the bireticulate pollen type. In subtype I the primary reticulum of the exine consists of angular muri; the lumen has an irregular polygonal shape and the perforations of the secondary reticulum are more or less uniform and of the same size. This pollen subtype was observed in species of subgenus Scutellaria, sectt. Scutellaria p.p., Anaspis, Perilomia and Salazaria. In subtype II the primary reticulum of the exine consists of a more or less curved murus; the lumen is rounded or obtusely angular and the secondary reticulum consists of one or more large central perforations surrounded by some smaller ones. This pollen subtype was observed in species of subgenus Scutellaria sectt. Scutellaria p.p., Salviifoliae and in all members of subgenus Apeltanthus. The results of this study reveal that the exine ornamentation is a diagnostic character useful for the infrageneric classification of Scutellaria.     

POLLEN MORPHOLOGY OF THE TRIBE SWARTZIEAE (SUBFAMILY PAPILIONOIDEAE: LEGUMINOSAE). 1. INTRODUCTION AND ALL GENERA EXCLUDING ALDINA AND SWARTZIA

The pollen morphology of 9 of the 11 genera of the tribe Swartzieae is described together with that of Holocalyx and Cyathostegia, two genera recently removed from the tribe based on macromorphology. The pollen is small, spheroidal to subprolate, primarily tricolporate with a perforate tectum and generally typical of the Leguminosae. Nonetheless, many of the genera have distinctive pollen morphology. Baphiopsis is 6-colporate. The genera Harleyodendron, Lecointea and one species of Exostyles have supratectal spinules. Bocoa viridiflora has striate/rugulate ornamentation very distinct from the other species of the genus Bocoa. The exine stratification is varied but Candolleodendron has a very thick endexine and narrow foot layer. The pollen of African taxa does not differ significantly from that of South American taxa. Pollen morphology does not clarify the taxonomic relationships of the tribe and provides little evidence to assist in positioning Holocalyx and Cyathostegia.     

水鳖科9属15种植物花粉形态的研究

应用光学显微镜、扫描电镜和透射电镜对水鳖科Hydrocharitaceae 9属15种植物的花粉形态进行 了观察。水鳖科植物花粉为圆球形至近椭球形,无萌发孔或偶为单沟萌发孔,外壁纹饰通常为小刺状纹 饰,刺密集或稀疏,花粉表面具瘤状、疣状、颗粒状、皱波状突起或光滑。外壁由覆盖层、柱状层和基层组 成。覆盖层厚或较薄,柱状层小柱发育不明显,基层薄。水鳖科植物在花粉大小、纹饰类型、刺的长短、 密度、形态、萌发孔的有无以及花粉壁的结构等方面表现出了较为明显的差异,这些特征对探讨类群间 关系具有较重要意义。由于黑藻属Hydrilla和Stratiotes属花粉较为特殊,支持将它们各自作为一个独立 的族处理。水鳖科植物花粉外壁纹饰和结构特点表明该科与水雍科Aponogetonaceae、泽泻科Alismataceae 和花蔺科Butomaceae等近缘,而该科植物花粉大多无萌发孔等则反应了该科与茨藻目Najadales植物有密切联系。     

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.     

国产爵床科山牵牛属6种、叉柱花属和老鼠簕属各1种植物的花粉形态

报道了国产爵床科Acanthaceae山牵牛属Thunbergia(山牵牛亚科Thunbergioideae)6种、叉柱花属Staurogyne(瘤子草亚科Nelsonioideae)和老鼠簕属Acanthus(老鼠簕亚科Acanthoideae)各1种植物在扫描电镜下的花粉形态。山牵牛属植物的花粉粒为圆球形,均具螺旋状萌发孔,外壁纹饰以光滑或颗粒状为主,偶具棒状突起。具螺旋状萌发孔被认为是该科独特而较原始的花粉特征。叉柱花属的花粉粒为圆球形,具3孔沟,外壁平滑。老鼠簕属的花粉粒为长球形,具3沟,外壁具细网状纹饰或具小穿孔。花粉形态特征支持传统上将上述3属置于3个不同亚科的处理。     

Pollen morphology of the East Asiatic genus Chelonopsis (Lamioideae: Lamiaceae) and allied genera,with reference to taxonomic implications and potential pollination ecology

Pollen grains of 15 taxa of the genus Chelonopsis (14 spp. and 1 variety) from China and Japan and 6 species of the closely related genera Bostrychanthera (1 species) and Gomphostemma (5 species) were examined by light and scanning electron microscopy. Of these, the pollen morphology of 18 taxa was studied for the first time. Pollen grains were found to be tricolpate with polar lengths of 20.8–30.0 μm and equatorial widths of 17.5–27.3 μm. The basic shape of the pollen in most taxa is subprolate or prolate-spheroidal, but spheroidal, subprolate-spheroidal, oblate-spheroidal, and prolate-subprolate grains are also found in some species. The surface is generally reticulate or suprareticulate in Chelonopsis and granulate in Bostrychanthera. In comparison with those of Chelonopsis and Bostrychanthera, the pollen grain surfaces of Gomphostemma are more diverse. In Chelonopsis, pollen is taxonomically useful at the sectional level, and some grains provide enough characters for species delimitation. The potential pollination ecology of cellular hairs on the anthers of Chelonopsis and Bostrychanthera is also briefly discussed.