Molecular phylogenetics of Alternanthera (Gomphrenoideae,Amaranthaceae): resolving a complex taxonomic history caused by different interpretations of morphological characters in a lineage with C4 and C3–C4 intermediate species

Alternanthera (Amaranthaceae) is a diverse genus (80–200 species) largely restricted to the American Tropics. With Pedersenia and Tidestromia, it makes up the ‘Alternantheroid clade’ in Gomphrenoideae. Parsimony and Bayesian analyses of nucleotide sequences of nuclear (ITS) and plastid (rpl16, trnL‐F) and morphological characters identify that the capitate stigma of Alternanthera is a synapomorpy within the Alternantheroids. Within Alternanthera, two major clades were resolved, both of which were marked by otherwise homoplasious characters of the gynoecium: Clade A [99% jackknife (JK); 1.0 posterior probability (PP)] with nine species and Clade B (60% JK; 0.98 PP) with 22 species. Four subclades (B1–B4), strongly supported statistically, were identified in Clade B. Previous subgeneric classifications of Alternanthera appear artificial in light of our new molecular phylogenetic analyses. Most major lineages are congruently resolved by nuclear and plastid data but some incongruence between the nrITS and plastid phylogenetic trees suggests hybridization may have played a role in the rampant speciation in Alternanthera. Whereas C₄ photosynthesis appears to have evolved in a single clade, the position of A. littoralis var. maritima (C₃) in this clade may be explained by hybrid speciation rather than a reversal from C₄ to C₃. All C₃–C₄ intermediates belong to a different clade that also contains C₃ species, but species limits, including the widely studied A. tenella, are unclear. The clade including A. tenella and A. halimifolia contains most of the species endemic to the Galápagos whereas A. nesiotes, also endemic to the islands, is nested among widespread American taxa. This suggests that the Galápagos radiation of Alternanthera may have arisen from at least two independent colonization events followed by a subsequent radiation in the former lineage. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 169 , 493–517.     

Multiple origins of a unique pollen feature: stellate pore ornamentation in Amaranthaceae

Stellate pore ornamentation is an unusual feature of angiosperm pollen, so far it is known in only ten genera of Amaranthaceae. The pollen grains of these plants have apertures with large hook‐shaped ektexinous bodies that are stellately arranged. Previous studies interpreted this character complex as a synapomorphy in consideration of its strong specialization. By reconstructing the evolution of stellate pore ornamentation based on phylogenetic trees of Amaranthaceae obtained by parsimony, likelihood, and Bayesian analysis of chloroplast trnK/matK DNA sequences, clear evidence is provided for several independent origins and reversals to less specialized aperture types. In addition to the gomphrenoid genus Pseudoplantago, stellate pore ornamentation evolved several times among achyranthoid genera, which have an African distribution. The most derived apertures, with 5 – 6 large protruding hooks, appear independently in Centemopsis on the one hand, and in Psilotrichum sericeum on the other. In an effort to break down the complex character syndrome of stellate pore ornamentation, we delimited a set of six pollen morphological characters that could be independently traced on the phylogeny. It appears that stellate pore ornamentation was independently derived from apertures with equally spread ektexinous bodies that became hook‐shaped, reduced in number, and symmetrically arranged. Likewise the symmetrically arranged, rectangular ektexinous bodies in Marcelliopsis represent an independent specialization. In view of this pattern of morphological changes, functional significance in the context of specialized insect pollination syndromes and positive selection for stellate pore ornamentation is hypothesized. Stellate pore ornamentation provides an example of a specialized pollen character complex with adaptive significance, and underlines the need for a dense taxon sampling for analyses of character evolution.     

Scanning electron microscopy of pollen grains of some species of the genusArum (Araceae)

A preliminary report is given on the pollen morphology of eight species of the genusArum examined by scanning electron microscopy. The pollen grains are spheroidal and their sporoderm sculpture possesses spines. The species vary mainly in number, shape and distribution of the spines on the pollen surface and these characters can be used to differentiate them.     

Pollen morphology supports the transfer of Wendlandia (Rubiaceae) out of Rondeletieae

Pollen morphology of 27 species, eight subspecies and one variety of Wendlandia was studied using scanning electron microscopy (SEM). Wendlandia pollen are monads, radiosymmetric, small in size, tricolporate (rarely tetracolporate or bicolporate) and spheroidal (rarely subprolate or suboblate) in equatorial view. The compound aperture consists of ectocolpus, mesoporus and endocolpus. In addition, reticulate sexine and granular nexine were observed. The pollen wall ultrastructure of two Wendlandia spp. was examined by transmission electron microscopy (TEM). The exine consists of the tectum, columellae, foot layer and endexine. The endexine is thickened into a costa around the aperture. The intine forms a protruding oncus at the aperture. The palynological characters show a remarkable uniformity among the Wendlandia spp. Differences with Rondeletia, the main genus of tribe Rondeletieae, exist in the exine pattern, the endoaperture and the pollen wall structure. Our observations indicated that the endoaperture type and the structure of the pollen wall of Wendlandia were similar to those of the Gardenieae–Pavetteae–Coffeeae–Octotropideae clade, which provided palynological evidence for a closer relationship of Wendlandia to subfamily Ixoroideae and the transfer of Wendlandia out of Rondeletieae. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 128–141.     

Pollen morphological studies in subfamily Suaedoideae (Chenopodiaceae)

The pollen morphology of two Bienertia species and 28 Suaeda species from the Old World has been studied with light and scanning electron microscopy. Pollen morphological characters used in this study include pollen diameter, pore number, pore diameter, operculum diameter, chord distance, exine thickness, plus number and characteristics of exine spinules and operculum spinules. Using these characters, the pollen of Bienertia is clearly distinguished from that of Suaeda. In addition, pollen data support recent molecular studies which merge the genus Borszczowia into Suaeda. The pollen grains of Suaeda show notable variation; in some cases this allows morphological distinctions to be made between species. Taxonomically, pore number is observed to be a more useful character than pollen diameter. Principal components analysis (PCA) shows only a weak correlation between pollen characters and habit (life form), habitat, mode of photosynthesis or phylogenetic relationships. Interestingly, however, the annual C₃ and C₄ species of section Schoberia are usually characterised by a higher pore number than the perennial C₄ species.     

Pollen morphology and phylogenetic analysis of Eperua Aublet (Detarieae: Caesalpinioideae: Leguminosae)

The diverse pollen morphology of fourteen species of the genus Eperua is described and illustrated using light, scanning electron and transmission electron microscopy. Six pollen types are described and a key for their identification is provided. A cladistic analysis was carried out using macromorphological and palynological characters to form a hypothesis of relationships between taxa. The pollen morphology is discussed with regard to systematic relationships, function and phylogenetic significance of certain pollen morphological structures within the genus.     

成都地区19个木芙蓉品种的孢粉学研究及其分类学意义

为了解成都地区木芙蓉(Hibiscus mutabilis)品种间的亲缘关系,采用扫描电镜观察了19个木芙蓉品种的花粉显微结构,并探讨其分类学意义.结果表明,19个木芙蓉品种的花粉均为大粒,主要为圆球形,具散孔,外壁纹饰均为刺状纹饰,表面具有颗粒状突起.木芙蓉品种间在花粉粒大小、外壁纹饰上具有一定差异,可以作为品种分类...     

Palynological characters of Glycyrrhiza,Glycyrrhizopsis, and Meristotropis (Leguminosae), with special reference to their taxonomic significance

Abstract The pollen morphology of 11 species of the genus Glycyrrhiza L. with one from each of the genera Glycyrrhizopsis Boiss. & Bal. and Meristotropis Fisch. & C. A. Mey. was investigated by scanning electron microscopy. In pollen morphology, the main differences between Glycyrrhizopsis and Glycyrrhiza are: Glycyrrhizopsis—pollen grains 36.63 × 40.42 μm in size, oblate spheroidal in shape; and Glycyrrhiza—pollen grains 24.47–33.18 × 23.82–31.83 μm in size, prolate spheroidal in shape. Glycyrrhizopsis and Glycyrrhiza should be recognized as two distinct genera based on palynological and morphological characters. Meristotropis and Glycyrrhiza are similar in many important palynological and morphological characters, suggesting that the two should be merged. In Glycyrrhiza, two types of pollen grains, 3‐lobed‐circular or subtriangular in polar view, are found in different species, in accordance with morphological differences in the two groups, shedding light on the classification and evolution of the genus.     

Pollen morphology of some species belonging to Codonoprasum and Allium sections of Allium (Liliaceae-Alliaceae) genus

Pollen morphology of 14 Allium L. species grown in Turkey, that belong to the sections Codonoprasum and Allium, were investigated under LM (light microscopy) and by SEM (scanning electron microscopy). However, the pollens of 5 species were investigated under TEM (transmission electron microscopy). Detailed pollen morphological characteristics are given for Allium in the family on the basis of the results presented here together with data from the literature. The genera Allium homogeneous in both aperture type and exine ornamentation. It is suggested that some palynological characters, such as aperture type and the presence of an operculum, could be of taxonomic value at the section level.     

Pollen morphology and fossil record of the feathery mistletoe family Misodendraceae

Misodendraceae is a small family of mistletoes in the order Santalales. Its distribution is restricted to the southern South American temperate forests. The family comprises the sole genus Misodendrum with eight species of hemiparasitic shrubs, mainly parasitising the southern beech Nothofagus. This contribution presents palynological evidence from seven species, using light microscopy and scanning electron microscopy. Pollen grains are consistently small, periporate and echinate, although differences in the length of echini and number and size of pores were noted. Pollen features can be used to distinguish groups of species and, in some cases, individual species. Cluster analysis of pollen characters differentiates two main groups: one includes M. brachystachyum, M. oblongifolium and M. quadriflorum; and the other includes M. gayanum, M. linearifolium, M. punctulatum and M. angulatum. Palynological results are compared with previous systematic studies of the family. The South American fossil pollen record is summarised and characters of the fossil pollen are analysed using UPGMA to test the relationships between extant and fossil species. Miocene pollen resulted similar to species of subgenus Angelopogon while Eocene pollen is disimilar to extant species of Misodendraceae.     

Pollen morphology of Xanthium L. (s.l.) (Asteraceae,Asteroideae, Heliantheae,Ambrosiinae) in the Iberian Peninsula - a palynotaxonomic approach to a poisonous,allergenic and invasive genus

Abstract

The pollen morphology of the Iberian taxa of the genus Xanthium s.l. and of the non-Iberian species X. ambrosioides was studied to help to identify these poisonous, allergenic and invasive taxa and clarify their taxonomic relationships. In addition, the question of the existence and structure of a mesoaperture in the apertural system of Xanthium was also considered. The pollen grains of sections Acanthoxanthium and Xanthium clearly differ by their characters, which supports the separation of both taxa and allows for their distinction based on pollen morphology. It is suggested that the morphological and palynological differences between Xanthium and Acanthoxanthium, together with the lack of hybridisation between the two sections and the monophyly of Acanthoxanthium, could lead to the elevation of the latter to the generic status. The two species of section Xanthium present in the Iberian Peninsula (X. orientale and X. strumarium) and of section Acanthoxanthium are well separated by their palynological features. Likewise, the Iberian subspecies of Xanthium, X. orientale subsp. orientale and X. orientale subsp. italicum are palynologically distinguishable. However, no significant difference could be found between the pollen of X. strumarium subsp. strumarium and of X. strumarium subsp. brasilicum. The pollen data do not support the hypotheses of a hybrid origin for X. strumarium subsp. brasilicum. A mesoaperture intersecting the foot layer and the upper part of the endexine is present in both sections, which is congruent with the results already found for other Asteroideae.     

Using palynomorphological characteristics for the identification of species of Alsinoideae (Caryophyllaceae): a systematic approach

Palynological characters of 18 species belonging to seven genera of the traditional subfamily Alsinoideae including Arenaria, Cerastium, Eremogone, Lepyrodiclis, Minuartia, Sabulina and Stellaria were studied in detail using light and scanning electron microscopy. Pollen grains of subfamily Alsinoideae are subspheroidal or prolate, pantoporate and 20.04 to 51.4 µm in size, prominent and sunken apertures uniformly distributed on the pollen surface. Two types of pollen grain ornamentation were observed, i.e. microechinate-punctate or microechinate-perforate. Echini are present on the surface of the pollen of all investigated species with medium, dense, or sparse echinodensity. These species exhibit variation in polar view, equatorial diameter, number of apertures, exine thickness, diameter of pore, appendages per pore, pore ornamentation, echini arrangement, echinidensity and shape of pollen. Based on qualitative characters, a dichotomous key has been developed for quick and easy identification. The present investigation contributes to the systematic approach using palynological characteristics and correct identification of species for members of subfamily Alsinoideae (Caryophyllaceae).     

Pollen morphology and systematics of palaeotropical Phyllanthus and related genera of subtribe Phyllanthinae (Euphorbiaceae)

The observations of pollen from 27 species of subtribe Phyllanthinae using scanning electron microscopy reveal considerable morphological diversity in palaeotropical Phyllanthus and the related palaeotropical genera Breynia, Glochidion, and Sauropus. The tribe appears to be monophyletic, but the pollen morphology does not support the monophyly of Phyllanthus or Sauropus. Within Phyllanthus, the pollen characters suggest a close relationship between the subgenera Emblica and Phyllanthodendron. They also reveal a surprising morphological congruence between the pollen grains of section Ceramanthus (subgenus Isocladus) and those of subgenus Eriococcus, although it is not clear whether this similarity is homoplastic. The presence of diploporate colpi is a synapomorphy uniting Breynia and Sauropus, but may be homoplastic in Phyllanthus. The affinities suggested by the morphological features of the pollen in the Phyllanthinae are concordant with recent molecular phylogenies. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 591–608.     

Body musculature of Beauchampiella eudactylota (Gosse, 1886) (Rotifera: Euchlanidae) with additional new data on its trophi and overall morphology

This study presents the results of confocal laser scanning microscopy and fluorescence‐labelled phalloidin used to visualize the system of body musculature in Beauchampiella eudactylota. Moreover, the poorly known trophi of B. eudactylota are described based on scanning electron microscopy. In total, four paired longitudinal muscles (musculi longitudinales I–IV) and three circular muscles (musculi circulares I–III) were identified. Among these are the musculus longitudinalis ventralis, the musculus longitudinalis dorsalis and the musculus circumpedalis as documented in previous studies for other rotifer species. Compared to other species, B. eudactylota is characterized by the low number of lateral longitudinal muscles and the absence of some longitudinal muscles (musculi longitudinales capitum) and circular muscles (corona sphincter, musculus pars coronalis). Moreover, scanning electron microscopic data on the trophi of B. eudactylota reveal a number of striking similarities to the trophi in some species of Epiphanidae. This suggests that either (1) these similarities represent plesiomorphic characters present both in Epiphanidae and B. eudactylota or (2) they are synapomorphic features of B. eudactylota and some species of Epiphanidae, which would question the monophyly of Euchlanidae.     

Pollen morphology of Chinese Glochidion (Phyllanthaceae) and its taxonomic implications

The pollen morphology of twenty‐five Chinese species belonging to two sections of Glochidion and one species from Phyllanthus subgen. Phyllanthodendron was examined using scanning electron microscopy (SEM). The results showed that pollen grains of Glochidion are isopolar, spheroidal to rarely subprolate in shape, (3)4–5(6)‐colporate in aperture pattern, and tectate‐reticulate to tectate‐rugulate in exine ornamentation. Fusion of colpus ends was observed in some pollen grains of G. pseudo‐obscurum and G. wrightii. The pollen grains of Phyllanthus dunnianus are similar to those of Glochidion species in their spheroidal shape and four‐colporate aperture pattern, but are slightly larger in size, and show tectate‐perforate exine ornamentation. In Glochidion, pollen morphology appears to be useful for distinguishing several macromorphologically similar. Additionally, palynological characters provide evidence for the taxonomy of the supergenus Phyllanthus s.l.     

POLLEN MORPHOLOGY AND DETAILED STRUCTURE OF FAMILY COMPOSITAE,TRIBE CICHORIEAE. I. SUBTRIBE STEPHANOMERIINAE

A survey of pollen morphology in 20 species representing the 11 genera of the North American subtribe Stephanomeriinae by light, scanning electron, and transmission electron microscopy revealed 10 of the 11 genera to have echinate, tricolporate pollen grains, Lygodesmia being the only genus with echinolophate pollen. Sectioned exines of most of the species examined are similar, being composed of ektexine and endexine. The ektexine surface is composed of spines which typically have globose perforate bases. A cavus occurs as a separation between the basis (foot layer) and the columellae in all of the genera examined except Chaetadelpha. Pollen of the two species of Glyptopleura were found to be strikingly different in exomorphology. Pollen of the putatively self-fertile G. marginata has much shorter spines than the closely related G. setulosa. Atrichoseris, Anisocoma, Calycoseris, Glyptopleura, Pinaropappus, Prenanthella, and most species of Malacothrix have pollen which lack paraporal ridges. The remaining genera, Chaetadelpha, Lygodesmia, Rafinesquia, and Stephanomeria have well-developed ridges of fused spine bases around the apertures. Pollen characters, particularly those of the aperture region, have been found to be systematically useful in the subtribe, therefore acetolyzed material gives more useful information than untreated pollen.     

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.     

Assessing photosystem I and II distribution in leaves from C4 plants using confocal laser scanning microscopy

Images of chlorophyll fluorescence emitted at wavelengths above and below 700 nm were recorded from leaf sections of C₄ species using confocal laser scanning microscopy (LSM). We investigated species exhibiting both NAD-malic enzyme (NAD-ME) C₄ photosynthesis and NADP-malic enzyme (NADP-ME) C₄ photosynthesis. Comparing LSM fluorescence of leaf sections with flow-cytometrically determined fluorescence from individual chloroplasts revealed that LSM fluorescence was distorted by the optical properties of leaf sections. Leaf section fluorescence, when corrected by transmission data derived from light transmission images, agreed with flow cytometry data. The corrected LSM fluorescence yielded information on the distribution of the individual photosystems in the C₄ leaf sections: PSII concentrations in bundle sheath cells were elevated in NAD-ME species but diminished in most of the NADP-ME species investigated. The NADP-ME species, Arundinella hirta, however, showed normal PSII and increased PSI concentration in bundle sheath chloroplasts. Finally, a gradient of PSI was observed within the bundle sheath cells from Euphorbia maculata.     

A palynological study of the genus Mentha L. (Lamiaceae)

The pollen morphology and exine structure of 10 Mentha L . species were investigated using light microscopy and scanning electron microscopy. The pollen grains of all 11 species were hexazonocolpate with granular membranes and a circular amb, varying in shape from prolate-spheroidal to suboblate. Different colpus shapes were recognized in M . ×  dumetorum . The exine was bireticulate in section Pulegium , and reticulate in section Menthae . A correlation was found between pollen size and chromosome number. The results indicate that the pollen characters of the genus Mentha are valuable for taxonomic applications and may be useful for classification.  © 2008 Uludag University. Journal compilation © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 141–154.