Pollen morphology and taxonomy of Atraphaxis (Polygoneae,Polygonaceae)

Pollen grains of 32 species of Atraphaxis L. including six species previously attributed to Polygonum L. s.str. were investigated with LM (acetolysed pollen) and SEM (dry pollen). Pollen grains of all studied species are spheroidal to oblong-spheroidal, mostly tricolporate, rarely 4-6-loxocolporate, ellipsoidal in equatorial view, rounded-trilobed or trilobed in polar view, with striate or striato-perforate, reticulato-foveolate, or reticulato-perforate sporoderm ornamentation. Exine sculpture is the most variable feature, differing in width, distinctness, and spacing of the striae, which are divided by grooves with small pits or perforations in rows. The most distinct variants of sporoderm ornamentation are connected by a full range of transitions. Striato-perforate ornamentation is common in species with the perianth typical for Atraphaxis, whereas reticulato-foveolate or reticulato-perforate sporoderm ornamentation was found in species with the perianth of Polygonum-type.     

Exine ultrastructure of in situ peltasperm pollen from the Rhaetian of Germany and its implications

Morphology and exine ultrastructure of pollen grains of Triassic peltasperms have been studied for the first time. Pollen grains of Antevsia zeilleri from the Rhaetian of Germany are of the Cycadopites-type and monosulcate; the sculpturing is the same in the apertural and non-apertural areas. The proximal exine includes a row of lacunae covered by a solid, thick tectum and underlined by a foot layer. Pillars are hanging from the tectum between the lacunae. The exine is thinning to a homogeneous layer in the apertural region. The latter is bordered by thicker alveolate areas of the exine, in places resembling a saccus-like ultrastructure. The endexine includes white-line-centred lamellae. The exine ultrastructure is compared with that of pollen of Permian peltasperms. Although pollen types ascribed to Permian peltasperms are completely different in their general morphology, a transformation can be hypothesized by ultrastructural data from Permian Vesicaspora into Triassic Cycadopites extracted from pollen sacs of Antevsia. Comparison with Cycadopites of non-peltaspermalean (Ginkgoalean, Cycadophyte) and unknown affinities has been accomplished. The exine ultrastructure is distinctive enough to differentiate among peltaspermalean, cycadalean and bennettitalean Cycadopites; some ultrastructural features are shared with pollen of modern Ginkgo biloba. More ultrastructural data are needed as well as numerous sections of pollen grains are necessary to reveal original unchanged ultrastructure.     

Pollen morphology of endemic species of the Gonospermum Less., Lugoa DC. and Tanacetum L. complex (Asteraceae: Anthemideae) in the Canary Islands (Spain), and its taxonomical implications

The pollen of nine endemic taxa of the Gonospermum Less., Lugoa DC. and Tanacetum L. complex and of T. parthenium L. (introduced species from the Canary Islands) was studied using light and scanning electron microscopy with the aim of extending the knowledge of their morphology and to investigate whether pollen morphology may help clarifying the controversial taxonomy of these taxa. Twenty six populations in the Canary Islands (Spain) were studied and the following parameters were measured: polar axis (P), equatorial diameter (E), polar exine thickness (Ex1), length (L) and thickness (A) of the spines in the meridional optical section, exine thickness at the centre of the mesocolpium in the interapertural area (Ex2) and distance between spines (De) in the equatorial optical section. The studies show that the pollen of the three genera is very similar regarding shape, apertures, and ornamentation, but may be distinguished by size and exine thickness.     

THE ULTRASTRUCTURE OF SAHNIA POLLEN (PENTOXYLALES)

The micromorphology and fine structure of in situ pentoxylalean pollen are described from the holotype of Sahnia laxiphora Drinnan and Chambers 1985 collected from the Lower Cretaceous (Valanginian-Aptian) of Victoria, southeastern Australia. Pollen grains are ovoid, monosulcate, and relatively small, averaging 26 μm in length. Exine ornamentation is psilate. The sporoderm is two-parted with the sexine staining lightly throughout and approximately six times the thickness of the more darkly staining nexine. The exine over the sulcus is typically strongly invaginated, and may or may not include an extremely thin sexine layer. The outer part of the sexine is homogeneous, while the inner part is composed of relatively large granules separated by irregular lacunae of various sizes; lacunae are most pronounced at the sexinenexine interface. Faint lamellae characterize the nexine in both apertural and nonapertural regions. Granular orbicules are often associated with the exine surfaces and also occur appressed to pollen sac walls along with lamellated tapetal membranes. Sporoderm ultrastructure is compared to that of nonsaccate pollen of other groups, and particularly to pollen of Bennettitales, Gnetales, angiosperms, and similar plants, to which the Pentoxylales have been thought to be closely related. Although Sahnia laxiphora pollen is not identical to that of any of these taxa, the strongest similarity is with pollen of Bennettitales.     

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.     

FINE STRUCTURE OF MISTLETOE POLLEN. III. LARGE-FLOWERED NEOTROPICAL LORANTHACEAE AND THEIR AUSTRALIAN RELATIVES

Pollen of all large-flowered neotropical loranthaceous genera and related Australian taxa (Nuytsia, Atkinsonia) were examined in the light, scanning, and transmission electron microscopes. Trilobate, deeply concave, peroblate or oblate pollen shapes are basic within the complex; triangular, slight convex, suboblate shapes are more derived features. The non-fixiform pollen of Atkinsonia is unique within the family. Most of the large-flowered genera possess exclusively syncolpate aperture types. However, among Psittacanthus species tricolpate, diplosynrugate and diplorugate apertures are also present, representing derived types for the complex. The inaperturate pollen of Atkinsonia appears to have developed independently within the family. Pollen sculpturing is typically non-uniform, i.e., there are pronounced sculpturing differences in polar and equatorial regions. Uniformly sculptured pollen is restricted to Atkinsonia, Ligaria and some species of Psittacanthus. Ultrastructurally, most exine modifications have occurred in the equatoral ektexine. Here the basic organization ranges from essentially columellaless to columellate, the latter the more derived condition. Endexine is typically thick and stratified in polar areas, thin and lamellate in peripheral and apertural regions. Similar pollen morphologies of Nuytsia and Gaiadendron support the idea of a transoceanic evolutionary connection between the Old and New Worlds. Pollen characters show Gaiadendron to be the most primitive and Psittacanthus the most advanced among the large-flowered neotropical genera.     

中国紫草科厚壳树亚科的花粉形态及其系统学意义

为探讨厚壳树亚科在紫草科中的系统位置,用扫描电子显微镜研究了国产厚壳树亚科Ehretioideae的厚壳树属Ehretia、轮冠木属Rotula和基及树属 Carmona 3属12种植物的花粉形态,其扫描电镜资料属首次报道。从本文并结合文献报道的双柱紫草属 Coldenia的花粉资料,可得知该亚科花粉为长球形 、近球形,稀球形,赤道部位具有不同程度的膨大;萌发孔类型为三孔沟和三拟假沟二者交替排列,稀具三孔;外壁表面具微弱的小穿孔,稀光滑或皱波状纹饰。厚壳树亚科Ehretioideae与破布木亚科     

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.     

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.     

Studies on Pollen Morphology and Taxonomy of Lespedeza and Its Allied Genera from NE China

The pollen morphology of 11 species and 1 variety in the genus Lespedeza and its allied genera (Campylotropis, Kummerowia) from NE China was examined under light and scanning electron microscopes. 1. Lespedeza Michx. (plate 1:1-6; 2:1-6; 3:1-6; 4:1-2) Pollen grains prolate, rarely subprolate or spheroidal, elliptic or rarely suborbicular in equatorial view, 3-lobed-rounded in polar view, tricolporate, colpus margins smooth or jagged. Polar axis 20.7-33.1μm long, equatorial axis 15.4-20.9 μm long. Exine reticulate or foveolate, lumina verrucose or smooth under SEM. 2. Campylotropis Bge. (plate 4:3-4) One species in NE China, C. macrocarpa (Bge.) Rehd. Pollen grains prolate, elliptic in equatorial view, 3-lobed-rounded in polar view, 3-colporate, colpus linear, 25.1μm long, 1.79μm broad, colpus margins jagged, with a series of verrucae equal in size along one side visible under SEM. Polar axis 19.7μm long, equatorial axis 14.6μm long. Exine reticulate, lumina nearly rounded, verrucose at periphery under SEM. 3. Kummerowia Schindl. (plate 4:5-6) Pollen grains spheroidal, oblate or prolate, elliptic in equatorial view, obtuse-triangular in polar view, tricolporate, colups linear, 25.1μm long, 2.01μm broad, colpus margins sinuate. Polar axis 24.7-27.9μm long, equatorial axis 19.7-26.6μm long. Exine reticulate or subreticulate, lumina nearly rounded, with verrocae visible under SEM. According to the pollen morphology of Lespedeza and its allied genera, the division of Lespedeza (s. lat.) into Lespedeza (S. str.), Campylotropis and Kummarowia by Schindler (1912) is reasonable. The subdivision of Lespedeza (s. str.) into Sect. Macrolespedeza and Sect. Lespedeza by many botanists, and the treatment of Lespedeze juncea (L. f.) Pers. var. inschanica Maxim. as an independent species (i.e. Lespedeza inschanics (Maxim). Schindl.) are also supported by the pollen morphology shown in the present work.     

Upatoia barnardii gen. et sp. nov., an araucarian pollen cone with in situ pollen from the Late Cretaceous (Santonian) of Georgia,USA

Upatoia barnardii gen. et sp. nov., a conifer pollen cone from the Late Cretaceous (Santonian) Eutaw Formation of Upatoi Creek, Georgia, USA, is known from lignified and fusainised mesofossils that preserve its three-dimensional structure. The cone consists of numerous helically arranged microsporophylls, each composed of a thin stalk and distal lamina. Three elongate pollen sacs are attached to the base of the lamina. Pollen grains isolated from the pollen sacs are relatively large (52 – 75 μm), spheroidal to ellipsoidal in outline, lack sacci, and have a thickened equatorial exine that is often strongly folded. Pollen of Upatoia barnardii indicates a close relationship to extant Araucariaceae. Microsporophylls of U. barnardii confirm suggestions from previous studies of fossil material that some Mesozoic Araucariaceae had only three pollen sacs per microsporophyll, in contrast to extant species that often have more than ten pollen sacs per microsporophyll.     

Pollen morphology of the aquatic Brazilian endemic genus Castelnavia Tul. & Wedd. (Podostemaceae)

The utility of pollen characteristics for taxonomic and phylogenetic analysis of most genera of Podostemaceae has barely been examined. Herein we report on the diversity of pollen structure in Castelnavia (Podostemaceae), a genus of six species endemic to central and southern Brazil. Pollen grain was shed as three-colpate, planaperturate monads and ranged from 12.5 to 22.5?μm for the polar axis and from 12.5 to 20.0?μm for the equatorial axis. All species examined had polar and radially symmetrical pollen grains. Grains from Castelnavia fluitans, Castelnavia multipartita, and Castelnavia noveloi had an oblate spheroidal shape (PD/ED), those from Castelnavia monandra and Castelnavia pendulosa were subprolate, and those from Castelnavia princeps were prolate spheroidal. The polar view was subcircular for C. monandra and subtriangular for the other species. A fastigium was observed in C. fluitans and C. pendulosa whereas a costa was present in C. multipartita and C. princeps, with the colpus margin not thickened in the former species but prominently thickened in the latter. Castelnavia monandra and C. noveloi did not have a fastigium or costa. For all species spicules occurred on the exine (microechinate sexine) and were most conspicuous on the colpus membrane. There was little correspondence between variation in pollen structure and species relationships supported by phylogenetic analysis.     

Pollen morphology and ultrastructure of Marathrum schiedeanum (Podostemaceae)

The Podostemaceae, or river-weeds, comprise 46 genera and 270 species of dicots and are the largest family of strictly aquatic angiosperms. Despite the large size, specialized habitats, and enigmatic morphology of the family, relatively little is known about the palynology of Podostemaceae. In the current paper, pollen morphology and ultrastructure of Marathrum schiedeanum are described. Pollen grains are relatively small, spheroidal, and tricolpate to spiraperturate. The exine has a microechinate ornament, a tectate-granular sexine and a relatively thick nexine in non-apertural regions, and a semitectate sexine and thinner nexine in apertural regions. Although aperture variation occurs in the family, this is the first report of the spiral aperture type in Podostemaceae. The spiraperturate condition appears to be derived in river-weeds, as does the granular pollen wall, which represents a reduction of the typical columellae found in eudicots.     

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.     

中国钩藤属9种植物的花粉形态

利用光学显微镜和扫描电镜对中国钩藤属(Uncaria Schreb)9种植物的花粉形态进行了观察.结果表明,钩藤属植物的花粉为单粒,小型,辐射对称,赤道面观圆球形至近长球形,极面观3-裂圆形,具3孔沟,萌发沟裂缝状,长达两极.萌发沟中央有"内壁加厚突出"结构.外壁具条纹至细网纹饰,表面光滑无附属物.钩藤属花粉大小、形状、萌发沟和纹饰类型都较接近,属于单一型花粉.钩藤属植物的花粉形态特征是茜草科(Rubiaceae)中较为原始的类型.     

Effects of X-rays and Colchicine on Pollen of Trichosanthes anguina L. (Cucurbitaceae)

Pollen grains of M, and C, generation plants of Trichosanthes anguina L. were studied after treatment of seeds with X-rays and colchicine respectively. Pollen sterility increased with increase in X-ray dose and colchicine concentration. The polar axis of X-irradiated populations was shorter than that of controls. But colchicine treatment resulted in larger pollen grains than the controls, although the exine did not increase as much. Ex-perimentally-produced tetraploids had larger pollen grains than diploids and triploids. The triploids were characterised by changes in pore morphology. Of 293 treated plants examined, 11 aberrant plants were isolated in the M, and C, generations. These plants had large and small fertile pollen grains, and also showed treatment effects in external morphology and chromosomal aberration during meiosis. The changes in polar diameter were not associated with variation in chromosomal material.     

Pollen ultrastructure in <Emphasis Type="Italic">Aristolochia manshuriensis</Emphasis> and <Emphasis Type="Italic">A. contorta</Emphasis> (Aristolochiaceae)

Pollen ultrastructure has been studied in two relict and rare species of the genus Aristolochia, A. contorta Bunge and A. manshuriensis Kom. (Aristolochiaceae). Both species have inaperturate, spheroidal, sometimes distally monocolpate or distally bicolpate pollen grains. The equatorial and polar axes of pollen grain in A. manshuriensis are 48.5 and 44.0 μm, respectively. The percentage of defective pollen grains in A. manshuriensis is 3.4%. The fossulate, perforated exine is up to 2.3 μm in thickness; the sexine and the nexine are almost equal in thickness. In A. contorta, the equatorial axis of pollen grain is 36.6 μm: the defectiveness percentage, 24.5%. The exine is verrucate, up to 0.3 μm in thickness, while the sexine is two to three times thicker than the nexine. The pollen germination experiments have shown that pollen of A. manshuriensis, in contrast to A. contorta, can germinate in 10–20% sucrose at 22°С. These data and the high percentage of pollen defectiveness in A. contorta indicate that the androecium function in this species is reduced. The reduction of the androecium function is evidenced by a small amount of pollen grains in anthers or empty anthers and a high percentage of defective pollen grains.     

Palynological diversity and major evolutionary trends in Cyperaceae

Pollen and orbicule morphology of 84 species, representing 52 genera from all tribes and subfamilies are investigated, in order to assess the systematic value of palynological data and to determine palynological evolutionary trends in Cyperaceae. A total of 90% of the species are examined for the first time with scanning electron microscopy. Pollen grains of Cyperaceae are oblate spheroidal to perprolate in shape, inaperturate to polyporate with opercula or pontopercula on pori or colpi. We distinguished seven different sexine ornamentation patterns. Orbicules occur in all species investigated. Pollen morphological variation within Cyperaceae is considerable and includes dispersal unit; number, location and degree of differentiation of apertural zones; and sexine ornamentation patterns. In subfamily Mapanioideae both tribes can be characterized by palynological synapomorphies. However, in subfamily Cyperoideae, the observed pattern of variation does not fit the most recent molecular phylogeny due to high levels of homoplasy and polymorphism in major pollen characters.     

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.