Palynological investigations on some Hypericum taxa (Hypericaceae) growing naturally in Turkey

This study examined the detailed pollen morphological structures of sixteen Hypericum taxa (four endemic, twelve non endemic) including eight sections showing the natural distribution in Turkey: H. sect. Ascyreia (H. calycinum L.), H. sect. Heterophyllum (H. heterophyllum Vent.), H. Sect. Taeniocarpium (Hypericum confertum Choisy subsp. confertum, H. venustum Fenzl, H. linaroides Bosse), H. sect. Drosocarpium (H. montbretii Spach, H. bithynicum Boiss.), H. sect. Crossophyllum (H. adenotrichum Spach, H. orientale L.), H. sect. Olympia (H. olympicum L. subsp. olympicum), H. sect. Origanifolia (H. origanifolium Willd., H. avicularifolium Jaub and Spach subsp. depilatum (Freyn and Bornm.) Robson var. depilatum, H. avicularifolium Jaub. and Spach subsp. byzantinum) and H. sect. Hypericum (H. tetrapterum Fries, H. perforatum L., H. triquetrifolium Tura). These taxa were studied under light microscope and scanning electron microscope for the first time. Of the taxa studied, H. tetrapterum has the smallest pollen grains (on average 15.85–17.20 × 15.45–16.05 μm); and H. olympicum subsp. olympicum the largest grains (on average 22.90–23.10 × 21.40–22.10 μm). The pollen grains of 15 taxa are subprolate and one taxon is prolate-spheroidal. The regular pollen grains of all 16 taxa are 3-zonocolporate. Ornamentation is microreticulate; lumina regularly spaced in eight taxa, tectum perforatum; tectal perforations regularly spaced in seven taxa and tectum perforatum; and tectal perforations grouped together in one taxon. Endoaperture is cruciform porus, with short lateral and meridional extensions in ten taxa, cruciform porus, transversally elongated, with very small lateral extensions in one taxon and lalongate colpus in five taxa. Basic pollen types are ten taxa in type X, five taxa in type IV and one taxon in type II.     

Variation of pollen morphology, and its implications in the phylogeny of Clematis (Ranunculaceae)

Clematis s.l. (including Archiclematis and Naravelia) is a genus of approximately 300 species with cosmopolitan distribution. The diversity of its pollen was surveyed in 162 taxa belonging to all infrageneric groups of Clematis s.l. Pollen morphology was investigated by use of scanning electron microscopy to identify useful characters, test taxonomic and systematic hypotheses, and elucidate pollen character evolution on the basis of the molecular phylogeny. Clematis pollen is small to medium (14.8–32.1?μm?×?14.2–28.7?μm), oblate to prolate (P/E?=?0.9–1.4) in shape. The apertures may be tricolpate and pantoporate sometimes with 4-zonocolpate and pantocolpate pollen grains as transitional forms. The tricolpate pollen grains are predominant and occur in all the sections of the genus, whereas pantoporate pollen grains can be found in sect. Tubulosae, sect. Viorna, sect. Viticella, and Naravelia only. Phylogenetic mapping of aperture types reveals that the pantoporate pollen type may be the apomorphy in the genus and evolved several times. The surface ornamentation in all taxa studied is similar and characterized by microechinae evenly distributed on the microperforate tectum. The size and density of spinules on the tectum vary greatly but successive in the whole genus. According to the character syndromes of the ornamentation, separating sect. Brachiata from sect. Meclatis is supported. Though pollen morphology may contribute to investigation of problematic taxa, the taxonomic value of pollen morphology is limited at the species level.     

Pollen morphology of the genus Salvia L. (Lamiaceae) in Turkey

Pollen grains of 30 taxa of the genus Salvia, belonging to sections Salvia, Horminum, Drymosphace, Plethiosphace and Hemisphace from Turkey were examined by light (LM) and scanning electron microscopy (SEM). Detailed pollen morphological characteristics are provided for these taxa. Among the studied taxa, S. verticillata subsp. verticillata from sect. Hemisphace has the smallest pollen grains, and S. pachystachys from sect. Salvia possesses the largest ones. The basic shape of the pollen grains in most taxa is suboblate, oblate-spheroidal or prolate-spheroidal. However subprolate pollen grains are recorded for S. macrochlamys from sect. Salvia. The grains are hexacolpate in all taxa, but in S. recognita from sect. Salvia also octacolpate pollen was found. Three distinct exine sculpturing types exist, reticulate-perforate (the common type), reticulate-granulate and bireticulate. The reticulate-perforate and bireticulate sculpturing patterns can be divided into subtypes based on the number of perforations and the number of secondary lumina in each primary lumen. Pollen morphological characteristics of the taxa studied are compared and discussed on the basis of taxonomical concepts. In some cases, these characters are useful in distinguishing the sections. For instance, the presence of 1-2 large central secondary lumina per primary lumen is a significant character of sect. Horminum separating it from the other sections. As well, the presence of holes on colpus membrane ornamentation can be used as a diagnostic taxonomic character for sectional division between sect. Hemisphace and others. S. ballsiana from sect. Salvia is clearly distinct from the other taxa examined by its unique pollen morphology. Further, for several macromorphologically similar taxa pollen structures provide additional evidence to delimite them from each other.     

POLLEN MORPHOLOGY IN THE SELAGINEAE,MANULEAE (SCROPHULARIACEAE), AND SELECTED GLOBULARIACEAE,AND ITS TAXONOMIC SIGNIFICANCE

The pollen grains of 22 genera and 182 species of Selagineae, Manuleae, and selected Globulariaceae were examined by light and scanning electron microscopy of whole and fractured pollen. Grains of the Selagineae and most Manuleae have trizonicolporate aperatures with long, unconstricted ectocolpi; clearly delimited margos; psilate colpus membranes; lalongate transverse endocolpi extending beyond the boundaries of the ectocolpi; thickened endoaperture margins; semitectate, reticulate to microreticulate exines; and psilate, simplicolumellate muri. In the Globulariaceae, Globularia has spinulose, microperforate exines and undifferentiated ectocolpus margins, and both Globularia and Poskea have granular to spinulose colpus membranes, lolongate endoapertures included within the ectocolpi, and unthickened endoaperture margins. These pollen data support postulated relationships between Selagineae and Manuleae, but pollen grains of Selagineae and Globulariaceae are distinct, and pollen morphology does not correlate with proposals combining these taxa in Globulariaceae or Selaginaceae. Pollen morphology in Glumicalyx is indistinguishable from that in most Manuleae and supports transfer of this genus to the Manuleae from the Digitaleae. Differences in pollen morphology supplement established morphological distinctions between Globularia and Poskea, but the pollen evidence does not support published infrageneric classifications in Globularia or postulated relationships between Selagineae and Gratioleae or Selagineae and Lagotis.     

Plesiomorphic and apomorphic pollen structure characteristics of anthemideae (Asteroideae: Asteraceae)

Anthemideae (Asteroideae: Asteraceae) pollen grains have basal columellae, a structural type called “anthemoid” in earlier publications. To survey structure variation in Anthemideae pollen, we examined freeze-sectioned grains from 45 species within 23 representative genera using scanning electron microscopy (SEM). From resulting data and a literature review, we concluded that: 1) pollen of Anthemideae taxa is qualitatively identical except for Ursinia (grains essentially lack basal columellae) and the Artemisia group (branches of basal columellae are complex and interwoven); 2) the double tectum (a term introduced in this study) is a synapomorphy of Asteroideae and plesiomorphic in Anthemideae; 3) apomorphies of Anthemideae grains include large basal columellae, a thick foot layer, and absence of internal foramina; and 4) Anthemideae pollen is qualitatively different from similar pollen in Lactucoideae, a distinction we recognized by restricting “anthemoid” to Anthemideae grains. Ursinia grains have occasional basal columellae and features resembling rolled-up columellae; we consider these vestiges of a reversal to the plesiomorphic condition. To assess quantitative structural variation, 2,200 image-analysis measurements were taken from 73 SEM micrographs. Intrageneric variation was analyzed by standard deviation, and intergeneric variation by principal components analysis. Compared to other Anthemideae taxa, the structural elements of Artemisia grains have reduced dimensions and variability. Otherwise, structural radiation of Anthemideae pollen has produced a phenetic continuum.     

Structure of the apertural sporoderm of pollen grains inEuphorbia andChamaesyce (Euphorbiaceae)

The tricolporate pollen grains of 38 Mediterranean and Macaronesian species ofEuphorbia L. andChamaesyce S. F. Gray have a special apertural sporoderm not found in the other taxa of theEuphorbiaceae. At the apertural margo the ectexine is thinner because of shorter columellae and the thin, fragmented or even absent foot-layer. Ectexinous granules, mixed with endexinous material, are present near the ora. The endexine is homogeneous and thickened under the colpi (at the end and at the proximity of the end of colpus). Around the ora, the endexine is granulate and lamellar with irregular cavities. The apertural intine presents a characteristic structure with thickenings running along both sides of the colpi. The arrangement and structure of these intinous thickenings depend on the distance from the ora. This special morphology of the intine is present in all taxa studied here. The genusEuphorbia is considered to be the most evolved taxon of this family. The characteristic apertural sporoderm may be an adaptative modification to different physiological conditions, so it may present an apertural mechanism which is more adapted to harmomegathic changes and thus facilitate the germination and the formation of the pollen tube.     

Pollen morphology of Stachys (Lamiaceae) in Iran and its systematic implication

Pollen grains of 30 taxa of the genus Stachys (29 spp. and one subsp.), representing 9 of the currently recognized sections and 1 species of the closely related genus Sideritis (Si. montana) distributed in Iran were examined by light and scanning electron microscopy. Twenty-eight taxa are studied for the first time under aspects of pollen morphology. The basic shape of the pollen grains in most taxa studied is prolate-spheroidal, but subprolate, spheroidal and oblate-spheroidal pollen grains can also be found in few species. The grains are usually tricolpate (the amb triangular), but also tetracolpate (the amb circular to more or less square) in some species (S. iberica, S. atherocalyx and Si. montana). The surface is microreticulate (the frequent type), reticulate, perforate, foveolate-psilate or foveolate. The lumina are separated by smooth or sinuate muri which make them polygonal, more or less rounded and elongate. Major pollen morphological features of the taxa studied are compared and discussed on the basis of taxonomical concepts. In some cases, these characters are useful in delimitation of formerly introduced sections while they mostly provide further characters in separating related species from each other. For example, all members of S. sect. Aucheriana are characterized by elongated lumina. Based on the oblate-spheroidal shape of its pollen as well as tetracolpate aperture type, the results of the present study confirm sect. Pontostachys as including S. angustifolia, S. iberica, S. sparsipilosa as well as S. atherocalyx. Our results also suggest that although some species like S. fruticolosa and S. lavandulifolia are morphologically well characterized, they cannot be separated from other species of Stachys based on pollen morphology.     

A palynological study and molecular analysis of the Desmodium procumbens group (Leguminosae-Papilionoideae) in Mexico

Desmodium diversity has been mainly classified using fruit characters, and because of variability in these and other characters, the taxonomy has sometimes been problematic and thus, there is a need for clarification and taxonomic reassessments. This is the case for the Desmodium procumbens group, where several infraspecific taxa have been named. The phylogenetic relationships among taxa referred to this group is unknown, and the taxonomic status has been recently revised only for some of the taxa. In this study, pollen morphology and ultrastructure of D. neomexicanum, D. procumbens var. procumbens, D. procumbens var. exiguum, D. procumbens var. longipes and D. procumbens var. transversum, belonging to the Mexican Desmodium procumbens group, were examined using light, phase-contrast, scanning electron and transmission electron microscopy, in order to establish palynological characters that might contribute to a better taxonomic resolution. Pollen grains are tricolporate, radiosymmetric, oblate spheroidal to prolate spheroidal shape, with lalongate endoapertures and rugulate exine with foveolae between rugulae. The exine is tectate with a columellate and/or granulate structure. The absence of a fastigium, features of exine sculpture, a colpus membrane and infrastructure mainly dominated by granules differentiate the pollen grains of the varieties exiguum and transversum from the other taxa studied. Pollen traits and sequence data provide evidence for separating D. neomexicanum from species D. procumbens. In general, the result from the pollen analysis is congruent with results obtained from a preliminary molecular analysis of this group.     

A Study on Pollen Morphology of Tribe Rhamneae (Rhamnaceae)in China

Pollen morphology of 25 Chinese species, representing 6 five genera in the tribe Rhamneae of Rhamnaceae, was examined under LM and SEM. Pollen grains are subspheroidal or suboblate, obtuse-triangular in polar view, 3-colporate, with the polar axis (P)10.4-28.7μm long and equatorial axis (E) 11.3-30.5μm long, P/E = 0.81-1.11. Colpi generally narrow and long, ora lalongate, with two ends connected with the thinned part of exine, forming a H-shape. Four thickenings were observed where colpi and ora cross, but sometimes the thickenings were indistinct or formed a ring. The stratification indistinct. The ornamentation of exine obscure or indistinctly reticulate under LM, but rugulose, short-striate, rugulose-foveolate or reticulate under SEM. The exine ultrastructure of Rhamnus ussuriensis was examined under TEM. It consists of imperforate tectum, granular layer, foot layer and endexine. The tectum is rather thick with uneven surface, while the granular layer is rather thin. A key to the genera based on pollen characteristics is provided and the general pollenmorphology of five genera, i.e. Sageretia, Rhamnus, Hovenia, Colubrina and Alphitonia, is described respectively. Based on pollen size, the four thickenings and the ornamentation, the five genera under study may be distinguished from each other. For example, pollen grains are smaller in Sageretia; the four thickenings are larger and distinct in Hovenia and Alphitonia. However, the ornamentation is short-striate under SEM in Hovenia and Alphitonia, while reticulate in the Colubrina and Rhamnus ( Subgen. Rhamnus) .Two different concepts of Rhamnus exist among taxonomists. Heppeler and Suessenguth divided Rhamnus into two subgenera (Subgen. Frangula and Subgen. Rhamnus), whereas Grubov separated Subgen. Frangula from Rhamnus as an independent genus. According to the pollen morphology, the separationis reasonable.     

Beschorneria yuccoides and Asimina triloba (L.) Dun: Examples for proximal polar germinating pollen in angiosperms

Beschorneria yuccoides (Agavaceae) microspores are arranged mostly in planar tetrads. Later on, the pollen grains of the tetrad usually fall apart, but sometimes remain loosely connected by ektexine elements. The ektexine consists of a tectum, of short columellae, and of a thin, discontinuous foot layer. An endexine is absent. The bilayered intine is without any additional thickening that would usually indicate an aperture region. From this point of view the pollen grain might be considered as omniaperturate. The pollen ornamentation is reticulate with wide lumina and robust, smooth muri.

The pollen grains show an indistinct sulcus characterised by a loose reticulate ornamentation. The sulcus is not exactly at the distal pole, but shifted towards the equator. No pollen tubes are formed regularly at the sulcus. Instead, pollen tubes are normally formed at the proximal pollen face. The proximal area, indicating a large germination field, is morphologically and functionally clearly an aperture (a germination zone); however, it does not represent a sulcus. The proximal face of all pollen grains appears as ornamented, with some exine lumps.

Asimina triloba (Annonaceae) pollen is shed in permanent planar or decussate tetrads. The distal sides are microreticulate to foveolate, and do not show an aperture; the psilate proximal sides are the germination areas of A. triloba.

The presence of apertures placed at the proximal pole was reported for distinct taxa of several angiosperm families. For Drosera, Dionaea (Droseraceae) and most probably for the diaperturate Cuphea species (Lythraceae) the existence of polar germination areas can be excluded. However, in some Annonaceae taxa with permanent tetrads (Annona cherimola, Asimina triloba) a situation similar to Beschorneria might be present, and indeed a proximal polar pollen tube is formed. Beschorneria yuccoides, Annona cherimola and Asimina triloba are unequivocal examples of angiosperm pollen with an exactly proximal aperture (germination area).     

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 Periplocaceae from Madagascar

The pollen morphology of eleven (Baroniella Constantin & Galland, Baseonema Schltr. & Rendle, Camptocarpus Decne., Cryptolepis R. Br., Cryptostegia R. Br., Gonocrypta Baill., Harpanema Decne., Ischnolepis Jum. & H. Perrier, Menabea Baill., Pentopetia Decne., and Tanulepis Balf. f.) of the 13 genera of the Periplocaceae from Madagascar have been examined using light-, scanning-, and transmission electron microscopy. All of the genera are characterized by pollen grains arranged in tetrads. The arrangement of the grains may be rhomboidal, decussate or tetragonal. The 4–6 pores present are restricted to the junction area of adjacent grains. Cryptostegia differs from the other genera in that the arrangement of pollen grains is only decussate. In Menabea the tetrads are united into a pollinium. The exine is smooth and consists of a distal stratum (tectum), subtended by a granular stratum consisting of granules of unequal size. Towards the base larger granules are present. In Camptocarpus, Harpanema and Tanulepis the exine is stratified into a distal stratum (tectum), a thin granular stratum and an almost continuous basal stratum (foot layer). The inline is well developed. The pollen grains of tetrads are connected by wall bridges (cross-wall cohesion). The internal walls in Camptocarpus, Harpanema and Tanulepis differ from the other genera in the absence of a tectum. The pollen morphology of the taxa investigated is very similar and of little value for distinguishing the species and genera investigated. The distinctive difference in exine structure between the above mentioned three genera and other genera investigated emphasizes the importance of exine ultrastructure in the Periplocaceae.     

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 of some taxa of Vicia L. subgenus Vicilla (Schur) Rouy (Fabaceae) from Turkey

The pollen morphology of the ten taxa belonging to Cassubicae Radzhi, Cracca Dumort., Ervilia (Link) Koch, Lentopsis Kupicha, Trigonellopsis Rech. f. and Variegatae Radzhi sections of the genus Vicia L. subgenus Vicilla (Schur) Rouy (Fabeae, Fabaceae) in Turkey has been examined by use of light microscopy and scanning electron microscopy. Twelve morphometric characters are analysed with one-way analysis of variance and Tukey’s honestly significant difference test for multiple comparisons. Polar axis and equatorial diameter of pollen grains range from 24.64 to 44.19 and from 15.90 to 30.42 μm, respectively. The pollen grains are prolate–spheroidal to perprolate (P/E = 1.09–2.07), 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. tenuifolia subsp. dalmatica and V. tenuifolia subsp. tenuifolia (sect. Cracca), reticulate–rugulate in V. villosa subsp. villosa (sect. Cracca), and reticulate–perforate in the remaining taxa. The apocolpium and colpus area are reticulate–perforate in V. hirsuta (sect. Cracca), V. ervilia (sect. Ervilia) and V. lunata subsp. grandiflora (sect. Trigonellopsis), and psilate, psilate–slightly perforate or perforate in the other taxa. The results obtained from analyses of pollen grains from the studied taxa of the subgenus Vicilla reveal significant differences in general morphology and some morphometric attributes, and ornamentation types. However, it is concluded that pollen morphology has proved to be uninformative for differentiating the subgenera Vicilla and Vicia.     

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 and localization of Ni in some Ni-hyperaccumulator taxa of Alyssum L. (Brassicaceae)

Pollen morphology of seven Alyssum L. taxa growing on serpentine soils in different places in the European Mediterranean macrobioclimate territory were studied, described and compared. Cluster analysis was performed to show similarity between species and their populations. The shape of the pollen grains varies among the species and among the grains within the same anther. The pollen grains are 3-colpate, prolate, with long and narrow colpi reaching the poles. The ornamentation of the exine varies from micro-reticulate to reticulate between the species. Pollen sterility/fertility was also calculated. The highest percentage of sterile pollen (73.76%) was calculated for Alyssum murale subsp. murale and the lowest (9.54%) for A. bertolonii subsp. bertolonii. All species are representatives of sect. Odontarrhena (C.A. Meyer) Koch well known as Ni-hyperaccumulators. Nickel and other elements present in pollen and stamen were studied by inductively coupled plasma-mass spectrometry. The stamen parts of all species were micromorphologically analyzed by scanning electron microscopy coupled to an energy-dispersive X-ray probe. Accumulation of Ni was detected in the stamens of all studied species and rarely in the pollen grains. The distribution patterns of Ni were similar among the species examined.     

Pollen Morphology in Tribe Lactuceae (Compositae)

The Lactuceae contain two basic pollen types, echinolophate and echinate. Most taxa have echinolophate, tricolporate pollen. Internally, most ektexines are composed of a perforate spiny tectum, several levels of columellae, a cavus, and a foot layer. In lacunae, the columellae are reduced to a single level and the cavus is often absent. Highly modified echinolophate pollen grains are found in Scolymus, Scorzonera and Tragopogon. Scolymus, Catananche, Scorzonera, and Tolpis have distinctive exine stratification patterns. Exines of Catananche and, to a lesser extent, those of Tragopogon contain internal foramina like those found in the Heliantheae. Echinate pollen is found in all subtribes and is probably ancestral. However, some echinate grains are probably derived.     

POLLEN MORPHOLOGY AND SYSTEMATICS OF SIPHONOGLOSSA SENSU LATO (ACANTHACEAE)

Pollen morphology of 15 species of Siphonoglossa and of two closely related groups was investigated. Two tribal-specific pollen types are found within Siphonoglossa sensu lato suggesting that the genus is artificial, composed of taxa belonging in several genera among two tribes (subtribes sensu Bremekamp) of Acanthaceae. Five taxa currently included in an informal subgeneric category of Siphonoglossa have tricolporate, prolate pollen (termed Type I) that is characteristic of Odontonemeae (= Odontoneminae, Justicieae). Pollen of the remaining taxa, belonging in two formal sections of the genus, are mostly 2-porate, bilateral (Type II) with a sexine sculpturing characteristic of Justicieae (= Justiciinae). Pollen of section Siphonoglossa is rather uniform, 2-porate, bilateral with lolongate pores, and seem to delimit a natural group. Taxa of section Pentaloba have a more heterogeneous pollen morphology, mostly 2-porate, bilateral with lalongate pores. Controversial aspects of the interpretation of pollen morphology in Justiciinae are presented and their relevance to this study are examined. Hypothetical trends in the evolution of pollen of Justiciinae are discussed and the application of pollen morphology to taxonomy of the genus is presented, including a recommendation for narrowing the generic concept of Siphonoglossa to the taxa of the type section.     

Pollen morphology of Quercus sect. Ilex and its relevance for fossil pollen identification in southwest China

Pollen grains of 16 modern species of Quercus were examined by scanning electron microscopy (SEM) and light microscopy (LM). The focus of this investigation was on a small group (c. eight species) of alpine, sclerophyllous oaks that may form a monophyletic group within sect. Ilex. The data were combined with published data for Polar axis (P) and Equatorial diameter (E) for more than 80 species. Based on pollen morphology Quercus pollen grains were divided into three types: (1) The Cyclobalanopsis type is smallest (generally < 23 μm) in size, with P/E ratios of approximately 1.08 and a psilate or fine-grained surface; (2) Quercus sect. Ilex type is 23–29 μm, the P/E ratio is approximately 1.24 (prolate) and it has a rugulate pollen surface; (3) the Quercus sect. Quercus type is generally > 29 μm in size, has a P/E ratio of approximately 1.04, and a clearly verrucate pollen surface. Based on a comparison with modern pollen size and ornamentation, fossil pollen grains of the Quaternary Qinghai Lake core can be assigned to the Chinese southwest mountain oak group of Quercus sect. Ilex (in particular Quercus semecarpifolia). In addition, we show that the pollen percentages of Quercus rapidly increased to the highest level during the glacial period. Using SEM for the identification of fossil Quercus pollen grains is essential to ensure the accuracy of differentiating evergreen Quercus types (sect. Ilex versus sect. Cyclobalanopsis) during the last glacial period in southwest China.     

Pollen morphology and ultrastructure of selected species of Magnoliaceae

The pollen morphology and ultrastructure of 20 species, representing eight genera of the Magnoliaceae are described based on observations with light, scanning and transmission electron microscopy. The family represents a homogeneous group from a pollen morphological point of view. The pollen grains are boat-shaped with a single elongate aperture on the distal face. The tectum is usually microperforate, rarely slightly or coarsely rugulose. Columellae are often irregular, but well-developed columellae do occur in some taxa. The endexine is distinct in 14 species, but difficult to discern in the genera Parakmeria, Kmeria and Tsoongiodendron. Within the aperture zone the exine elements are reduced to a thin foot layer. The intine has three layers with many vesicular-fibrillar components and tubular extensions in intine 1. The symmetry of the pollen grains, shape, type of aperture and ultrastructure of the intine show a remarkable uniformity in the family. Nevertheless there is variety in pollen size, ornamentation and the ultrastructure of the exine. The pollen of Magnoliaceae is an example of an early trend of specialization, and supports the view that Magnoliaceae are not one of the earliest lines in the phylogeny of flowering plants.