COMPARATIVE POLLEN MORPHOLOGY AND TAXONOMIC AFFINITIES IN CYCADALES

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

Pollen characteristics of New Zealand Gnaphalieae (Compositae) and their taxonomic significance II. Tem

Pollen ultrastructure has been investigated in 35 taxa of Gnaphalieae (Compositae), predominantly from New Zealand. Pollen grains of all taxa examined are tectate‐columellate and caveate. The characters regarded as particularly distinctive include variability of columellar form and internal organisation of the columellae. The importance of pollen characters in the classification of the New Zealand Gnaphalieae is discussed. The species of Haastia, characterised by senecioid pollen and internal tecta, do not belong to the Gnaphalieae. In the species with helianthoid pollen and sub‐columellae five different columellae types can be distinguished.     

POLLEN MORPHOLOGY AND SYSTEMATIC RELATIONSHIPS AMONG NEW WORLD SPECIES IN TRIBE ANTIRRHINEAE (SCROPHULARIACEAE)

The morphology of mature pollen grains among 12 New World genera in tribe Antirrhineae has been examined with light and scanning electron microscopy. Pollen from 29 American species have mean polar diam that range from 17 to 26 μm, have a tectate structure with perforate, microreticulate, or reticulate surface pattern, and are subspheroidal to prolate, and trizonocolporate with fusiform or narrowly oblong colpi that are free or occasionally fused at the poles. Among the genera, pollen size is positively associated with style length but not haploid chromosome number. Three major morphological categories are recognized based on grain shape, exine structural pattern, and colpus shape. Two of these morphotypes each characterize individual genera (Mohavea, Linaria). The other morphological category includes ten New World genera and has been divided into three subcategories based on variation in lumina diam and intermittent occurrence of fused colpi. Only Antirrhinum and Pseudorontium have species or populations with pollen in more than one subcategory. Concordance of morphological, anatomical, chromosomal, and palynological data suggests that Linaria and Mohavea are best placed in separate subtribes from other native American genera. Compared to Old World taxa in tribe Antirrhineae, the most common pollen type (subspheroidal, microreticulate, apocolpate, fusiform colpi) may be the most primitive. Pollen grains with fused colpi (Antirrhinum p.p., Pseudorontium) and perforate (Mohavea) or reticulate (Holmgrenanthe) structural patterns probably represent derived types.     

Pollen morphology of Chamaebuxus (DC.) Schb., Chodatia Paiva and Rhinotropis (Blake) Paiva (Polygala L., Polygalaceae)

Polygala L. is a large and highly diverse genus with complex taxonomy, but pollen morphological information for this taxon is scarce. In the present study, pollen characters have been used to assess the taxonomic delimitation and phylogenetic relationships of three newly established subgenera of Polygala: Chamaebuxus, Chodatia and Rhinotropis (sensu Paiva). The pollen morphology of 22 species has been examined using light microscopy and scanning electron microscopy of acetolysed material. The pollen of 15 of the species is examined for the first time. The pollen grains are isopolar, radially symmetrical, tectate and, typically, polyzonocolporate with numerous colpi running parallel to the polar axis, and an endocingulum around the equator. Two pollen types can be distinguished: Type I, which includes species belonging to Rhinotropis, and Type II, which includes species from Chamaebuxus and Chodatia. The two pollen types are described and the pollen of the three studied subgenera is illustrated. Despite the low infrageneric morphological diversity observed within the genus Polygala, quantitative characters of pollen grains support the current classification of the subgenera Chamaebuxus, Chodatia and Rhinotropis, and reveal a closer relationship between the first two taxa. Pollen characters are shown to be a useful and informative tool for assessing taxonomic position and phylogenetic relationships within Polygalaceae, especially at higher taxonomic levels.     

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.     

Pollen morphology of the Pavetteae (Rubiaceae,Ixoroideae) and its taxonomic significance

Pollen grains of the tribe Pavetteae (Rubiaceae, subfamily Ixoroideae) are examined using LM and SEM. Grains are 3‐ or 4‐colporate and (semi‐) tectate (in one Versteegia species atectate). Sexine patterns vary between perforate, microreticulate, reticulate, rugulate and striato‐reticulate. Supratectal elements are sometimes present. The variation in pollen morphology in the Pavetteae allows to recognize seven pollen types, the distribution of which is useful to evaluate generic delimitations and relationships within the tribe. Pollen characters corroborate the close relationships between the genera Coleactina, Dictyandra and Leptactina and between Homollea, Homolliella and Paracephaelis. All the genera of the tribe proved to be stenopalynous (the species examined possess the same pollen type), except Pavetta, Rutidea, Versteegia and Tarenna which are eurypalynous. In the huge genus Pavetta the existing infrageneric classification is supported pollen morphologically. Pollen morphology further indicates that the genus Tarenna is badly delimited and strongly in need of a revision. The small genus Versteegia is in need of further taxonomic and palynological study to understand the pollen morphological variation encountered here. At a higher rank, pollen morphology also does not contradict the recent division of the Pavetteae in the Ixoreae (a stenopalynous tribe with presumably primitive pollen) and the Pavetteae sensu stricto (eurypalynous).     

POLLEN MORPHOLOGY OF AMPHIANTHUS,ARTANEMA, CURANGA,GLOSSOSTIGMA, AND PEPLIDIUM (SCROPHULARIACEAE-GRATIOLEAE)

Pollen of the scrophulariaceous genera Amphianthus, Artanema, Curanga, Glossostigma, and Peplidium was examined by light and scanning electron microscopy. All have single, isopolar, radially symmetrical pollen grains with equally spaced meridional colpi and columellate exines. Intergeneric differences are established mainly on the differential correlation of colpus number, colpus morphology, and sculpturing of the mesocolpium. The colpi vary in length and range in number from 3 to 6. The colpus membrane can be variously granular, verrucate, rugulate, or spinulose. The exine varies from semitectate and microreticulate or reticulate to tectate and sparsely perforate. It may be psilate or ornamented with supratectal processes ranging from small granules to relatively long spinules or bacules. All the genera examined, with the possible exception of Artanema vs. Amphianthus, are distinguished from one another by pollen morphology, and the grains of both Curanga and Glossostigma have morphotypes not previously reported in tribe Gratioleae. Final taxonomic assessment of the pollen data in the genera examined must await further comparative studies in the Gratioleae and other tribes, but the morphological diversity observed is clearly of potential significance in evaluating specialization levels and clarifying current conflicting interpretations of relationships in these taxa.     

Trends of Specialization in Pollen of Flacourtiaceae with Comparative Observations of Cochlospermaceae and Bixaceae

Pollen from 17 genera of Flacourtiaceae was examined by SEM and TEM; twelve of them represented the primitive tribes Berberidopsideae and Oncobeae. Small, minutely structured grains are widespread in this family, and have often been lumped together as one morphological type. A study of the relative thickness of the tectal layers, and of the ar-rangement and size of columellae and perforations indicates that not all minutely structured grains are homologous in development. Most genera of Berberidopsideae and Oncobeae can be termed microperforate tectate, as the muri are x2-5 as wide as the lumina and both features are less than one micrometer in diameter. Berberidopsis pollen is unique in the Flacourtiaceae in being tricolpate and in having a rough, thick tectum infrequently traversed by microperforations 0.03 p. in diameter. Tendencies toward perreticulate structure occur in Mocquerysia. Development of a high infratectal space is illustrated by Casearia nitida and an intectate exine in the related Samyda dodecandra. Pollen of Cochlospermum and Bixa are distinct from Flacourtiaceae and show independent trends of specialization.     

Appomattoxia ancistrophora gen. et sp. nov., a new Early Cretaceous plant with similarities to Circaeaster and extant Magnoliidae

A new genus and species of fossil angiosperm (Appomattoxia ancistrophora) is established based on well-preserved fruiting units and associated pollen from the Early Cretaceous (Early or Middle Albian) Puddledock locality in the Potomac Group sequence of Virginia, eastern North America. Fruiting units are small, unilocular, and with a single, pendulous, orthotropous seed. The fruit surface is characterized by densely spaced unicellular spines with hooklike tips, which probably functioned in biotic dispersal. Pollen grains adhering to the stigmatic area of many specimens are monocolpate and tectate with granular to columellate infratectal structure, and are similar to dispersed grains assigned to Tucanopollis and Transitoripollis. Comparison of fossil Appomattoxia ancistrophora with extant plants reveals an unusual combination of characters that includes similarities with some magnoliid taxa, particularly Piperales (Piperaceae, Saururaceae) and Laurales (Chloranthaceae), as well as the monotypic ranunculid family Circaeasteraceae. Appomattoxia ancistrophora differs from extant Piperales in having a pendulous rather than erect ovule, and differs from extant Circaeaster in details of the fruit wall, as well as the presence of monosulcate rather than tricolpate pollen.     

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

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

Pollen morphology and its relation with meiotic irregularities in ten species of Campuloclinium (Eupatorieae,Asteraceae)

Pollen grains of ten species of Campuloclinium (Eupatorieae, Asteraceae) are described and illustrated using light and scanning electron microscopy. The species included in this study are C. burchelli, C. campuloclinioides, C. chlorolepis, C. hirsutum, C. irwinii, C. macrocephalum, C. megacephalum, C. parvulum, C. purpurascens and C. riedelli. Pollen grains of Campuloclinium are typically radially symmetric, echinate, tectate, oblate-spheroidal to prolate-spheroidal (P/E ratio: 0.94–1.10). Pollen is mostly triaperturate, but tetraaperturate or biaperturate pollen also occurs in some species. The pollen diameter was highly variable ranging from 10.54 µm to 36.18 μm. The significance of the variation in type and size of pollen is discussed in relation to diverse meiotic abnormalities and possible reproductive apomictic processes within the polyploid species.     

螺序草属(茜草科)的花粉形态

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

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 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 of the Aceraceae

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

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.     

Studies in Montia L. and Claytonia L. and Allied Genera

The pollen morphology of several genera in Portulacaceae is described. Particular attention has been paid to the genera of the subfamily Montioideae, as a stage of continued monographical studies. Among genera especially dealt with are Claytonia, Montia, Crunocallis, Naiocrene, Neopaxia, Mona, Maxia, Limnalsine, and Montiastrum. In the taxonomical treatment of these genera the pollen morphology has proved to afford many important additional characters.

The pollen grains of Claytonia are distinguished from those of the remainder in being 3-colpate. The grains of the Claytonia-type have many similarities with those of Lewisia, a genus of the subfamily Portulacoideae. The other genera of Montioideae have pantocolpate pollen grains. Among these genera several different pollen types are distinguished, chiefly with regard to the sexine structures and the aperture membranes. The Montiastrum-type is especially interesting, with tholate grains, a particular pollen type not met with in any other genus in the family. The pollen morphology of some genera in the Portulacoideae is also treated. In some species in Calandrina and Talinum pantotreme pollen grains are observed with apertures transitional between pori and colpi. The apertures of the pantotreme grains are arranged in characteristic patterns.

Particular attention has been given to the variation of the pollen morphological characters. This variation has been examined with regard to the differences between different populations of the same species as well as between different species. The greatest variation has been observed in the shape and size of the grains. The structure and sculpture and thickness of the sexine and the aperture membranes are less variable. Some polyploid taxa are connected with the occurrence of pollen grains with divergent and varying aperture numbers.

In a survey of the genera the taxonomical results of the investigation are presented with particular regard to the pollen morphology. The new genus, Maxia Ö. Nilss., is described. One new species, Montia clara Ö. Nilss., is described and some new combinations are made.

Pollen morphological diagnoses are given for 46 different taxa. The aperture conditions for 96 different species are presented.     

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.     

Rosularia globulariifolia var. pallidiflora comb. et stat. nov. (Crassulaceae)

Rosularia globulariifolia (Fenzl) A. Berger and R. pallidiflora (Holmboe) Meikle were originally described as species in Umbilicus DC. in 1842 and 1914, respectively. In the present study, morphological and palynological features were examined in detail to determine the status of these taxa. The general characteristics, seed structure and micromorphology, pollen measurements and pollen characteristics were established. Morphologically, the two taxa cannot easily be distinguished. Pollen grains of both species are tricolporate, subspheroidal and eutectate, ornamentation is psilate and exine structures are similar. Based on morphological and palynological similarities, and the fact that both taxa are found in the same habitats, we propose that R. pallidiflora should be reduced to the rank of variety as Rosularia globulariifolia (Fenzl) A. Berger var. pallidiflora (Meikle) Minareci & K. Yildiz comb. et stat. nov.