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.     

Comparative morphology,anatomy and palynological studies of perennial species of Onobrychis (Fabaceae) in northeast Iran

This study examined morphological, anatomical and palynological characters for six species, two subspecies and two varieties of perennial Onobrychis growing in northeast Iran. In the morphological study, shape, size and color of vegetative organs and legumes were assessed. For the anatomical study, cross sections of mature fresh or dried stems, leaflets and petioles were prepared and stained. In the palynological study, pollen were extracted, acetolysed and observed by SEM. The morphological study indicated that some characteristics varied among species, such as shape of stipules and calyx teeth, color and shape of corolla segments and legumes and an identification key is provided based on these characters. Also, in internal structure some differences were noticed in the shape of petiole cross sections, the thickness of cortex and pith shape in petioles and the arrangement of vessels in stems. Some anatomical characters proved useful to identify species, but not sections. Palynological results showed circular and obtuse triangular pollen in polar view and elliptical pollen in equatorial view with reticulate ornamentation. In addition, a cluster analysis was done based on morphological, anatomical and palynological features. The resulting dendrogram confirmed the taxonomic treatment suggested by Sirjaev and Rechinger.     

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。     

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.     

Pollen morphology of Carthamus L. species in Anatolian flora

In flora of Turkey, there are six species of Carthamus L: Carthamus dentatus (Forssk.) Vahl., C. glaucus M. Bieb. subsp. glaucus, C. lanatus L., C. tenuis (Boiss. & Blanche) Bornm., C. persicus Desf. ex Willd. and C. tinctorius L. Within these species, C. tinctorius L. is an alien species for the Turkish flora. The pollen grains belonging to six taxa collected from different locations of Anatolia and examined both under light microscope (LM) and under scanning electron microscope (SEM). As a result of these examinations and measurements, pollen grains of taxa are radially symmetrical, isopolar, oblate-spheroid, spheroid, tricolporate rarely tetracolporate, echinate. There are high similarities among taxa but some differences are recorded in size and spin length. According to cluster analyses of Carthamus species’ pollens, C. glaucus and C. tenuis have the most similar pollens. C. dentatus grouped with these two. C. persicus placed as the closest relative of C. tinctorius in the dendrogram.     

Pollen morphology of some taxa of Vicia L. subgenus Vicia (Fabaceae) from Turkey

In this study, the pollen morphology of 11 taxa belonging to Atossa (Alef.) Asch. & Graebner, Hypechusa (Alef.) Asch. & Graebner, Peregrinae Kupicha, Wiggersia (Alef.) Maxted, Vicia L. and Narbonensis (Radzhi) Maxted sections of the genus Vicia L. subgenus Vicia (Fabeae, Fabaceae) naturally growing in Turkey has been studied using Light Microscopy (LM) and Scanning Electron Microscopy (SEM) to evaluate the taxonomic relevance of pollen characters. Twelve morphometric characters are analysed with one-way analysis of variance and Tukey’s honestly significant difference test for multiple comparisons. Of the taxa studied V. narbonensis var. narbonensis (sect. Narbonensis) has the largest pollen grains (P = 51.98 μm × E = 30.52 μm) and V. lathyroides (sect. Wiggersia) has the smallest pollen grains (P = 27.71 μm × E = 20.14 μm). The pollen grains are subprolate to perprolate (P/E = 1.16–2.11), but the prolate shape occurs in the majority of the taxa. The regular pollen grains of all taxa are trizonocolporate, isopolar, and released in monads. Ornamentation of the mesocolpium is psilate-perforate in V. lathyroides (sect. Wiggersia), reticulate-rugulate in V. narbonensis var. narbonensis (sect. Narbonensis), (micro)reticulate in V. sericocarpa var. sericocarpa (sect. Hypechusa), V. sativa subsp. sativa (sect. Vicia) and V. grandiflora var. grandiflora (sect. Vicia), and reticulate-perforate in the remaining taxa. The apocolpium and colpus area are psilate or perforate in all taxa except V. sericocarpa var. sericocarpa (sect. Hypechusa) and V. sativa subsp. sativa (sect. Vicia), which exhibit the obscurely reticulate-perforate pattern. Several palynological features have taxonomic importance in sectional level: polar axis, equatorial diameter, pollen shape (P/E ratio), colpus length, colpus width, size of pori, porus length/width ratio, lumina diameter, muri thickness and ornamentation. The results also indicate that pollen characters can be useful in distinguishing the examined taxa.     

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.     

Pollen morphology of Lathyrus (Leguminosae) taxa belonging to Lathyrus, Orobastrum and Cicercula sections from Turkey

The pollen morphology of 20 wild taxa belonging to Lathyrus (Syn: Eulathyrus), Orobastrum (Taub.) Boiss. and Cicercula (Medic.) Gren. & Godr. sections of Lathyrus L. grown in Turkey (L. rotundifolius Wild. subsp. miniatus (Bieb. ex Steven) P.H. Davis, L. grandiflorus Sibth. & Sm., L. saxatilis (Vent.) Vis., L. vinealis Boiss. & No?, L. inconspicuus L. var. inconspicuus, L. inconspicuus var. stenophyllus (Boiss.) Rech. f., L. tauricola P.H. Davis, L. woronowii Bornm., L. hierosolymitanus Boiss., L. cassius Boiss., L. gorgoni Parl. var. gorgoni, L. pseudo-cicera Pamp., L. sativus L., L. blepharicarpus Boiss., L. stenophyllus Boiss. & Heldr., L. belinensis Maxted & Goyder, L. phaselitanus Hub.-Mor. & P.H.Davis, L. chrysanthus Boiss., L. chloranthus Boiss., and L. trachycarpus (Boiss.) Boiss were examined by light microscopy (LM) and scanning electron microscopy (SEM) in this study. The pollen grains were 3-zonocolporate, spheroidal, subprolate, and prolate (P/E?=?0.99–1.48) types, and were medium in size (equatorial view: rectangular or elliptical-obtuse-convex; polar view: circular, triangular and quinquangular-obtuse-convex). The smallest pollen grains belonged to L. tauricola (P?=?30.94/E?=?31.20) and the largest to L. grandiflorus (P?=?50.60/E?=?36.40). The ornamentation was reticulate and reticulate-granulate in the mesocolpium, and usually psilate in the apocolpium. Some photographs included in this work were taken using both LM and SEM.     

安徽母草属植物花粉形态特征的研究

利用光镜和扫描电子显微镜首次对安徽母草属(Lindernia)六种植物的花粉形态进行了观察研究.结果表明该属花粉粒多为近球形, 具3 沟, 表面光滑.探讨了花粉形态特征的分类学意义,支持属级水平的分类.     

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.     

The fine morphology of pollen grains from the pollen chamber of a supposed ginkgoalean seed from the Middle Jurassic of Uzbekistan (Angren locality)

Pollen grains of Cycadopites-type were found in the pollen chamber of a supposed ginkgoalean seed Allicospermum sp. from the Middle Jurassic deposits of Uzbekistan (Angren locality). The pollen grains were studied with help of LM, CLSM, SEM, and TEM. All pollen grains show the identical morphology and exine ultrastructure allowing us to suppose the same botanical affinity. The pollen morphological data do not contradict the ginkgoalean interpretation of the seed; therefore, the pollen grains and the seed most probably did belong to the same parent plant. The pollen grains are monosulcate, the non-apertural surface is nearly psilate, with low short elements, which are occasionally scattered over the surface or more densely distributed. The aperture and adjacent areas appear to bear more distinct sculpturing. The ectexine is composed of a prominent solid tectum, a thin infratectum, and a thin foot layer. The infratectum is formed of one row of alveolae, which are more voluminous laterally, where the ultrastructure is more easily understandable. The endexine is multilamellate, although it is evident only in some regions of stained sections. Towards the aperture the ectexine becomes gradually thinner; over the aperture no sublayers can be discerned within the ectexine. The ectexine of the apertural region repeatedly varies in thickness, reflecting a sculpturing surface of this region. The obtained data contribute to the knowledge about the exine ultrastructure of ginkgoaleans; nonetheless, a TEM study of ginkgoalean pollen grains extracted from pollen organs is still highly desirable. We also considered pluses and minuses of CLSM: it failed to substitute SEM, since the surface pattern under study was too fine, but demonstrated the general morphology of the pollen grains under study better than conventional LM. The possibility of viewing virtual sections of any area of the pollen grain was profitable for later interpretation of TEM sections. CLSM would give better results in interpreting relatively large palynological objects with distinct sculptural elements, a complicated architecture, variously arranged appendages, or possessing cameras.     

Pollen morphology and its taxonomic significance of the monotropoideae (Ericaceae)

A palynological survey, including LM, SEM and TEM is presented for eight genera, nine species and 76 samples of the Monotropoideae which is composed of 10 genera and 13 species. On the basis of the aperture number and shape, the following six pollen types are recognized: 1) 3-colp(oroid) ate—Allotropa, 2) 2-colpor(oid)ate—Monotropsis andHemitomes, 3) 4-colpor(oid)ate—Pterospora, Sarcodes andPleuricospora, 4) 2- and 3-colpor(oid)ate—Monotropa hypopitys, 5) 3-colporate and-porate —Monotropa uniflora, 6) 3- to 4- porate—Monotropastrum humile. Relationships among taxa within the Monotropoideae are illustrated on the palynological characters including the aperture type, exine sculpture and structure. Both pollen grains with two and four apertures have evolved independently from pollen grains with three apertures, according to the infraspecific variation of aperture numbers and the usual occurrence of three-aperturate pollen grains in the Ericaceae. 3-colp(oroid)ate pollen ofAllotropa is the most primitive and occupies an isolated position in the subfamily, on the other hand 3- to 4-porate pollen ofMonotropastrum humile is most advanced in the subfamily, probably even in the Ericaceae. A reduced sexine inMonotropastrum humile appears to be a specialized pollen character. Infraspecific geographical difference in palynological characters is revealed in two species ofMonotropa for the first time.M. hypopitys is basically characterized by pollen grains with two apertures in the New World vs. three apertures in the Old World.M. uniflora is basically characterized by 3-porate pollen in the New World vs. 3-colporate in the Old World. In both species pollen grains with more primitive characters usually occur in the Old World.     

The comparative pollen morphology of genera Matricaria L. and Tripleurospemum Sch. Bip. (Asteraceae) in Turkey

Pollen morphology of four Matricaria species and 28 Tripleurospermum species was investigated with light microscopies (LM) and scanning electron microscopies (SEM). Pollen slides were prepared using Wodehouse technique. Measurements were based on 20 or more pollen grains per specimen. For SEM studies, dried pollen grains were transferred on aluminum stubs and coated with gold for 4 min in a sputter-coater. The pollen grains of Matricaria and Tripleurospermum are radially symmetric and isopolar. The pollen grains of the Matricaria are oblate-spheroidal with the polar axes 16.6–31.2 μm and the equatorial axes 18.7–23.9 μm. Tripleurospermum is oblate-spheroidal, suboblate and prolate-spheroidal with the polar axes 15.6–32.2 μm and the equatorial axes 17.7-38.5 μm. The pollen grains of Tripleurospermum are operculate and tricolporate. Matricaria is operculate and usually tricolporate or rarely syncolporate, tricolpate and tetracolporate. The pollen grain of both taxa shows echinate ornamentation. The spines are commonly conical with a broadened base and a tapered apical portion. The spine length varies between 1.8–4 μm in Tripleurospermum and 2.3–3.3 μm in Matricaria. The width of spines varies between 2.8–4.6 μm in Tripleurospermum and 2.4–3.6 μm in Matricaria. Inter-spinal area shows granulate–perforate, reticulate–perforate, rugulate–perforate ornamentations and the tectum surrounding the spine base is micro perforate. Overall exine thickness ranges from 2.8 to 4.8 μm in Tripleurospermum, 3.6 to 5.2 μm in Matricaria. Intine is thicker under pores in Tripleurospermum (0.3–0.62 μm) than in Matricaria (0.6–0.8 μm). Inter-spinal ornamentations, pollen shape and the numbers of perforations at the spin base have been observed as important morphological characters.     

Structural basis of harmomegathy: evidence from Boraginaceae pollen

Upon release from the anther, pollen grains can be exposed to dry environment and dehydrate. To survive in dry conditions, the pollen wall possesses the ability to fold itself due to water loss-harmomegathic mechanism. Apertures seem to function as the primary elements of harmomegathy as they are more elastic than the remainder of the pollen wall. Contribution of other sporoderm structures, surface features, and pseudocolpi in harmomegathy are usually not considered in palynological studies. The nature of pseudocolpi has not been properly understood until now, partly because of common use of acetolysis method as a standard procedure. Different structures involved in the harmomegathy mechanism were studied in Cryptantha celosioides, Cryptantha coryi, Heliotropium europaeum, Myosotis palustris, Rindera bungei, and Rindera tetraspis. Scanning electron microscopy was used to study harmomegathy in hydrated and dehydrated pollen grains. In addition, transmission electron microscopy was used to elucidate the ultrastructural basis of pseudocolpi and other harmomegathic structures with special attention to intine structure. Our data reveal that additional flexibility of the pollen wall in Boraginaceae is provided by pseudocolpi, rugulate surface, tectate–columellate ultrastructure, and a transverse groove. Curious triangular polar poroid areas are described in M. palustris.     

A combined light and scanning electron microscopy study

The uppermost Eocene Florissant Formation, Rocky Mountains, Colorado, has yielded numerous insect, vertebrate and plant fossils. Three previous comprehensive palynological studies investigated sections of lacustrine deposits of the Florissant Formation and documented the response of plant communities to volcanic eruptive phases but overall found little change in plant composition throughout the investigated sections. These studies reported up to 150 pollen and spore phenotypes. In the present paper, we used a taxonomic approach to the investigation of dispersed pollen and spores of the Florissant Formation. Sediment samples from the shale units containing macrofossils were investigated using light microscopy (LM) and scanning electron microscopy (SEM). The general picture of the palynoflora is in agreement with previous studies. However, the combined LM and SEM investigation provides important complementary information to previous LM studies. While a fairly large amount of previous pollen determinations could be confirmed, the purported taxonomic affinities of several pollen phenotypes need to be revised. For example, pollen referred to as Podocarpus or Podocarpidites sp. belongs to the Pinaceae Cathaya, Malus/Pyrus actually belongs to Dryadoideae, pollen of the form genus Boehlensipollis referred to as Proteaceae/Sapindaceae/Elaeagnaceae or Cardiospermum belongs to Sapindaceae but not to Cardiospermum, and pollen of Persicarioipollis sp. B with previously assumed affinities to Polygonaceae actually belongs to Thymelaeaceae. Pandaniidites and one type of Malvacipollis cannot be linked with Pandanaceae and Malvaceae. A few taxa are new records for Florissant (Ebenaceae: Diospyros; Mernispermaceae; Trochodendraceae: Tetracentron). In general, SEM investigations complement the LM palynological studies and improve the identification of dispersed pollen and spores and enable integration of data from dispersed fossil pollen into a wide range of comparative morphological, taxonomic, evolutionary, biogeographic and phylogenetic studies.     

Pollen morphology of some selected plant species from Horton Plains,Sri Lanka

The pollen morphology of four species: Gaultheria leschenaultii (Ericaceae), Psychotria zeylanica (Rubiaceae), Rubus leucocarpus (Rosaceae) and Symplocos bractealis (Symplocaceae) from Horton Plains, Sri Lanka was investigated using LM and SEM. Psychotria zeylanica is endemic to Sri Lanka and the other three taxa are common in Horton Plains. Gaultheria leschenaultii has tetrads with 3-colporate, coarsely granular to granulo-rugulate pollen grains; in the polar region and at the colpus margin the exine is smooth with micro-rugulate structure. Psychotria zeylanica has 3- (or 4-) colporate, reticulate-spinulose to granular pollen grains. Rubus leucocarpus has 3-colporate, striate-perforate pollen grains and Symplocos bractealis has 3- (or 4-) colporate pollen grains with a verrucate-perforate exine. Pollen grains of these four species are very indicative of human impact and palaeoclimate.     

Pollen morphology of Strobilanthes Blume (Acanthaceae) in China and its taxonomic implications

The pollen morphology of 37 Chinese species of Strobilanthes Blume (Acanthaceae), a large and taxonomically problematic genus, was investigated by LM and SEM. Two distinct pollen shape classes, ellipsoid and spheroidal, recognised by earlier workers were found. Eight pollen types could be distinguished on the basis of variation in ornamentation, and the number, distribution and type of apertures. Five of these fall within twenty two pollen types reported from Strobilanthes in southern India and Sri Lanka. Three new pollen types are also described, further extending the palynological diversity of the genus. From the results presented, the characters of palynological diversity and the related taxonomic implications are discussed.     

A comparative study on two closely relative Tulipa L. taxa from NE Anatolia

This study presents observations on the anatomical, palynological and ecological features of Tulipa gumusanica Terzio?lu and its morphologically similar relative, T. armena Boiss. var. armena, in order to clarify their similarities and differences. We found that these taxa have some important differences with regard to anatomical, palynological and ecological features, as well as morphological traits. General anatomical traits of both examined taxa are similar, both having isolateral leaves with distinct hypodermis and a stem with distinct monolayer collenchyma close to the epidermis. However, some anatomical characters such as mesophyll width, average number of stomata on lower epidermis, and epidermal cells on upper epidermis are found to be important in delimiting these taxa. In addition, considerable differences have been observed in pollen shape and size. The species differ ecologically in that T. gumusanica prefers slightly acidic soil with low organic content in the woodland, whereas T. armena var. armena prefers slightly alkali soil with high organic content in steppe vegetation.     

Combined cryo- and thin sectioning (TEM) in the elucidation of mimosoid tetrad/polyad ultrastructure

Cryosections (freeze-sections) of tetrads/polyads and their subsequent examination in the SEM complement traditional methods (LM, SEM, and TEM) by providing both a dynamic three-dimensional overview of polyad morphology and clarifying aperture morphology, internal wall characters and tetrad/polyad cohesion mechanisms. Cryosections of Dinizia tetrads reveal that cohesion is maintained through adhesion/fusion of the apices of clavate elements in localized regions of the subproximal and proximal walls — a feature not discernible using traditional SEM preparations. Cryosections of the globose 20-grained polyads of Parkia species reveal small central grains and triradiate-shaped proximal apertures — features unobservable or misinterpreted using traditional LM, SEM and TEM methods. In Anadenathera polyads, cryosections have clarified the nature and distribution of small gaps present in the lateral and proximal walls of individual grains.