Pollen morphology and the bi-reticulate exine of the Phyllanthus species (euphorbiaceae) from mauritius and réunion

The pollen grains of the Phyllanthus species of Mauritius and Réunion have been divided into seven pollen types. The different species and varieties of teh “phillyreifolius” group have close affinities and can be arranged in a continuous morphological series. All grains in this series are characterized by a two-layered or bi-reticulate ornamentation. They have a peripheral reticulum which consists of arch-like muri (in cross-section) and variform lumina and this covers an infra (micro-) reticulum, the structural units of which are connated with the muri of the suprareticulum.Contrary to earlier observations the Phyllanthus amarus type also shows a bi-reticulate ornamentation; this suggests a relationship between this type and the “phillyreifolius” group. The Phyllanthus maderaspatensis, P. casticum and P. tenellus types, however, have eu-reticulate ornamentation. The P. urinaria type resembles the P. amarus type, but its ornamentation is not clearly bi-reticulate and is better termed tectate-perforate. In the P. casticum type the pollen grains show very characteristic ectoapertures and they are of special interest as many Phyllanthus spp. occurring in Africa and Asia belong to the same type.     

四川雅安地区13种凤仙花属植物花粉形态及其分类学意义

采用扫描电镜观察了中国四川雅安地区13种凤仙花属植物的花粉形态,并对其花粉形态进行了主成分分析和聚类分析,基于孢粉学和SRAP聚类结果的比较,讨论不同孢粉学特征对分类结果的影响。结果显示:(1)雅安地区13种凤仙花属植物花粉粒均为中等大小,花粉粒形状从超扁到扁球形变化,花粉粒极面观形状从矩形和椭圆形到四边形、圆形、三角型变化,且只有湖北凤仙、峨眉凤仙和匙叶凤仙是三沟花粉粒,其余种均为四沟花粉粒;外壁纹饰仅扭萼凤仙和齿萼凤仙为细小网状,其余均为网状纹饰。(2)依据不同指标进行的孢粉学聚类均将13种凤仙花属植物划分为3个类群,而SRAP聚类结果则将13种凤仙花属植物划分为5个类群;不同孢粉学聚类结果的相似率只有30.77%。(3)孢粉学聚类并未与分子聚类显示严格的对应关系,以花粉大小和形状、外壁纹饰为指标进行的孢粉学分类与SRAP分类的相似率最高(69.23%)。研究认为,选择不同的花粉形态特征作指标对分类的准确性影响较大,花粉大小和形状、外壁纹饰这2个指标对凤仙花属植物分类具有意义,而萌发沟的特征则很难作为分类的依据,花粉形态特征在分类上有局限性;进行该属植物孢粉学分类需要选择有价值的分类指标,并结合形态学、分子生物学等综合研究。     

POLLEN MORPHOLOGY OF SCHEFFLERA (ARALIACEAE)

Pollen of 48 species of Schefflera was studied by light and scanning electron microscopy. Eight pollen types are distinguished on the basis of sexine sculpturing and structure, apertural shape, and shape and size of the grains. These pollen types correlate well with other data such as the number of carpels, the condition of corolla, infloresence types, and geographical distribution of each species. To determine the evolutionary status of sexine structure, an association between sexine characters and the number of carpels was sought. Assuming polymery of floral parts to be a primitive condition in Araliaceae, the undifferentiated sexine, which occurs in the multicarpellate species, also appears to be primitive. Other unspecialized features include short grains, and non-sculptured, imperforate tectum. Several species in New Guinea and the Solomon Islands belong to this putative ancestral pollen type, from which other types may have been derived.     

Systematic implications of pollen morphology in Elsholtzia (Elsholtzieae–Lamiaceae)

Pollen morphology of 18 species (32 specimens) representing all three currently recognized sections of the genus Elsholtzia (Elsholtzieae–Lamiaceae) was investigated in detail using light, scanning electron and transmission electron microscopy. Elsholtzia pollen grains are small to medium in size (P=20–50 μm, E=16–45 μm), mostly prolate‐spheroidal to prolate, and rarely oblate‐spheroidal to subprolate in shape (P/E=0.95–1.46), hexacolpate (the amb more or less circular or ellipsoid) with granular aperture membranes. Three distinct types of sexine ornamentation are observed in Elsholtzia: perforate, rugulose‐bireticulate, and bireticulate. The bireticulate pattern is the most common, showing a wide range of morphological variation among species. The exine thickness varies from 1.2 to 2.5 μm. Among the present palynological data, the variability observed in sexine ornamentation of Elsholtzia could be of systematic significance. Current infrageneric classification of the genus Elsholtzia is partially congruent with pollen morphological data.     

凤仙花属（Impatiens L.）10种植物花粉形态的扫描电镜观察

利用扫描电镜观察了10种凤仙花属（Impatiens L.）植物的花粉形态。结果表明：本属花粉为单粒花粉,呈长圆形至长矩圆形,大小为20.3～46.7 μm,具角萌发孔,网状纹饰,网眼明显;根据花粉网状纹饰中网眼内是否具颗粒状突起可将其分为2类：（1）网眼内无或近无颗粒状突起,黄金凤（I. siculifer）和婺源凤仙花（I. wuyuanensis）的花粉纹饰属于这一类型;（2）网眼内有明显颗粒状突起,其余8个种的花粉纹饰均属于该类型。研究表明,花粉特征,特别是花粉粒网状纹饰中网眼内有无颗粒状突起及颗粒状突起的形态等特征,在凤仙花属内常具种水平上的可见变异,因而可作为种类划分的依据,它们在分类学上的价值应予以关注。     

Morphology of pollen and orbicules in the tribe Spiraeeae (Rosaceae) and its systematic implications

This study provides pollen data for 38 representative taxa belonging to all nine genera in the current classification of the tribe Spiraeeae (Rosaceae) including the monotypic Korean endemic genus Pentactina, and considers the distribution of orbicules for the first time. Pollen morphology and wall stratification were investigated using light, scanning electron and transmission electron microscopy. Spiraeeae pollen grains are small to medium in size (P = 6.9–34.0 μm, E = 7.1–28.0 μm), oblate to prolate in shape (P/E = 0.66–1.48) and tri-colporate. Spiraeeae pollen is generally characterised by striate sexine ornamentation, but four ornamentation types are recognised based on the length and direction of the ridge patterns. The observed variation in sexine ornamentation is particularly valuable at the generic level. The exine stratification of the representative Spiraeeae studied is similar and characterised by unbranched columellae and a continuous endexine. Orbicules are present in three genera of the tribe (Luetkea, Sibiraea and Xerospiraea). Orbicule distribution patterns indicate that the absence of orbicules is a synapomorphic condition of the more derived clade, comprising Pentactina + Petrophytum + Kelseya + Spiraea.     

Pollen morphology of alpine butterworts (Pinguicula L., Lentibulariaceae)

The pollen morphology of Pinguicula alpina, P. arvetii, P. grandiflora subsp. grandiflora, P. grandiflora subsp. rosea, P. hirtiflora, P. leptoceras, P. poldinii, P. reichenbachiana, and P. vulgaris, belonging to the Alpine flora, was studied.The pollen grains, coming from different populations, were investigated using light microscopy and scanning electron microscopy. The pollen size, the shape (P/E ratio), the number of colpori and the exine ornamentation are, for Pinguicula, important diagnostic characters.Pinguicula pollen grains are medium sized (∼ 30 μm), trinucleate, isopolar, radially symmetric. The shape of the grains is variable from oblate spheroidal to prolate spheroidal and they are (4)-5-9-(10)-zonocolporate. The prevalent ornamentation is rugulate-microreticulate, P. alpina has a rugulate-reticulate ornamentation and only P. hirtiflora has a perforate ornamentation.A pollen key, based on micromorphological data, is presented.     

Comparative structure of the pollen in species of Passiflora: insights from the pollen wall and cytoplasm contents

Pollen grains of three Brazilian species of Passiflora (P. elegans, P. suberosa and P. haematostigma) belonging to different subgenera were studied with respect to the wall and cytoplasm. New data were obtained on pollen wall histochemistry, cytoplasm contents and organelle inheritance. The structure of the pollen wall layers differed in all the species; P. elegans shares characters with those found in other species from the same subgenus. The exine foot layer is structured and evident only in P. haematostigma and is not structured in P. elegans. The pollen grains have pollenkitt with lipid components. The cytoplasm of the vegetative cell contains dissolved and non-dissolved polysaccharides. The generative cell contains plastids and mitochondria in all the species analyzed, and consequently has the potential for paternal or biparental extranuclear inheritance. Aspects of the evolution of the characters of the species are discussed in the light of a recent phylogeny of the group, with a focus on the three subgenera.     

Pollen diversity in the genus Ptilostemon (Asteraceae,Cardueae) from Italy and its taxonomic and palynoecological implications

The results of a study on pollen morphology of the species of the genus Ptilostemon growing in Italy: Ptilostemon niveus, P. greuteri, P. gnaphaloides, P. casabonae, P. strictus, P. stellatus by light and scanning electron microscopy were presented and discussed including in relation to their taxonomic position. The exine shows two different ornamentation patterns: echinae and scabrae. This last ornamentation pattern together with other features was previously described in the genus Ptilostemon. Of 6 species analysed, P. stellatus is the only one that shows a scabrate ornamentation of exine. Two pollen types were recognized through the exine ornamentation, length of ectocolpus and polar outline: P. stellatus type and P. niveus type. A multivariate analysis (principal component analysis and linear discriminant analysis) was carried out with the aim of examining potential morphological characters which could be used to identify taxa. The data suggest that several characteristics can be used to delimit the species in particular the exine ornamentation. The results of our studies support the actually subgeneric classification, and several features of the pollen grains analysed seem to have a palynoecological role and an important taxonomic significance. A dichotomous key based on palynological data is also given.     

Pollen Morphology of Cyclobalanopsis and Its Relation to Quercus

The present paper deals with pollen morphology of 19 species belonging to Cyclobalanopsis and of 31 species belonging to Quercus from China. All pollen grains were examined under light microscope and scanning electron microscope, and those of 4 species of Cyclobalanopsis and of 8 species of Quercus were examined under transmission electron microscope. Pollen grains of Cyclobalanopsis are spheroidal or subspheroidal, 25.2(18.9-31.5) ×23.1 (16.8-27.3) μm in size. Mostly 3-colporoidate, rarely 3-colpate or 3-colporate. The exine is 2-layered, 1.1-1.9 μm thick, sexine thicker than nexine, finely granulate under LM, granulate, spinulate, verrucate under SEM. Pollen grains of Cyclobalanopsis and of evergreen members of Quercus are very similar to each other in size, aperture and ornamentation of exine, but there are some differences in pollen grains between Cyclobalanopsis and deciduous members of Quercus. Therefore, Pollenmorphology supports the treatment of Cyclobalanopsis as a subgenus of Querc     

Pollen morphology of the genus Iris L. (Iridaceae) from Croatia and surrounding area: taxonomic and phylogenetic implications

In Croatia and the surrounding area, the genus Iris is represented with about 20 Alpine-Dinaric, Mediterranean and Pannonian taxa from the subgenera Iris and Limniris. We researched pollen morphology of all taxa by using scanning electron microscopy. All pollen grains are sulcate with few palynological features, which could have taxonomic importance on different classification levels: shape of dry pollen grains; shape, outline and approximate size of hydrated pollen grains, approximate size of the sulcus, and ornamentation of the sulcus membrane and of the exine. At least four pollen types were recognised and taxonomically delimited to the series level: two characterise the subgenus Iris, section Iris (series Elatae and Pumilae), and two the subgenus Limniris, section Limniris (series Laevigatae, Sibiricae and Spuriae). Taxonomic implications of pollen morphology of their critical groups and taxa have been evaluated. Possible phylogenetic implications of pollen morphology of the genus Iris on the Southern European area were suggested: the subgenus Iris is more advanced than the subgenus Limniris; I. graminea seems to be the most primitive member of the subgenus Limniris on the Southern European territory; and pollen morphology of the population of the subspecies I. sibirica subsp. erirrhiza from the Mountain Bjelolasica could represent a link between the subgenera Limniris and Iris in the territory of Southern Europe. The possible pathway of evolution of the genus Iris on the territory of Southern Europe was suggested: from the subgenus Limniris, through the “linked taxon” I. sibirica subsp. erirrhiza from the Bjelolasica Mountain, to the subgenus Iris, series Pumilae, and finally to the series Elatae.     

Taxonomic significance of pollen morphology for species delimitation in <Emphasis Type="Italic">Psidium</Emphasis> (Myrtaceae)

Previous studies reported Psidium as one of the most difficult genera to delimit within the American Myrtaceae. Even though palynology has improved the taxonomy of Angiosperms, information about the usefulness of pollen morphology for taxonomic purposes in Myrtaceae remains contradictory. Here, we investigate the significance of pollen morphology for Psidium taxonomy with specific focus on its usefulness for determining species groups of taxonomic significance. Pollen traits observed by light and scanning electron microscopy were quantified and examined using cluster and ordination analyses. Average size of pollen grains was visualized by boxplots. Pollen grains of Psidium are isopolar, oblate, peroblate or oblate-spheroidal, 3-syncolporate or 4-syncolporate. The sexine ornamentation is rugulate, granulate or spinulose-granulate and differs between the mesocolpium and apocolpium. Cluster analysis revealed four distinct groups: Psidium cauliflorum (G1) and Psidium oligospermum (G3) as single-species groups; Psidium brownianum, P. oblongatum, P. ovale, P. sartorianum, P. guajava, Psidium sp. 1, Psidium sp. 2 (G2), and Psidium cattleianum, P. longipetiolatum, P. guineense, P. myrtoides (G4). Supported by ordination analysis, three traits better explained these groups: type of exine ornamentation, size of P-EV and pollen shape. The used approach efficiently distinguished related species, as well as explained species groups of taxonomic significance suggesting pollen morphology to be a significant source of information for taxonomic studies in Psidium.     

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.     

New insights into the pollen morphology of the genus Gunnera (Gunneraceae)

Pollen grains from ten species of Gunnera, chosen to represent the six different subgenera in the genus, were examined using light and scanning electron microscopy. The aim of the study was to explore characters that have the potential to define different types of pollen within Gunnera and to study the evolution of these characters in light of the phylogeny of the genus. According to our results, there are three main types of pollen in the examined species of Gunnera. Type 1, unique for the South American species G. herteri (subgenus Ostenigunnera), is characterised by an imperfect reticulum with sinuous undulating-creasted muri. A reticulum with equidimensional polygonal lumina is typical for the plesiomorphic type of pollen (type 2) present in subgenera Gunnera, Misandra and Panke. Lastly, pollen grains of subgenera Pseudogunnera and Milligania are characterised by a reticulum with lumina of variable shape and size (type 3). In G. macrophylla (subgenus Pseudogunnera), the lumina in the apocolpia are of a different shape and size from the lumina in the mesocolpia (type 3a), while in G. dentata, G. monoica and G. cordifolia (subgenus Milligania), the lumina are identical in the apocolpium and the mesocolpium (type 3b).

The identification of pollen types will possibly allow the interpretation of the different specimens of Tricolpites reticulatus, the fossil species believed to be allied to the extant Gunnera.

In addition to the revision on the pollen of Gunnera, a brief comparison between the pollen of this genus and its sister group Myrothamnus is reported.     

Pollen morphology and phylogenetic relationships in neotropical Phyllanthus (Euphorbiaceae)

In order to provide new insights into phylogenetic relationships among the neotropical taxa of Phyllanthus , 28 illustrations are provided of the pollen grains of 22 selected species studied from 11 sections of the subgenera represented in the neotropics. Special attention has been given to subgenus Conami because of its variability in pollen morphology: of eight species illustrated, the apertures are diploporate colpi in three species and pores in five species; exine ornamentation is vermiculate in two species and pilate in the other six species. The six species in the neotropical sections Pityrocladus and Microglochidion (subgenus Emblica ) are characterized by prolate grains with an increased number of colpi (4–8). Of particular interest are species in which the pollen exine is clypeate (with exine shields); clypeate pollen grains are illustrated in two species of subgenus Xylophylla and in one species of section Cyclanthera that has unique exine shields with single central pila. The pollen of the one Brazilian phylloclade-bearing species illustrated (in section Choretropsis ) has 3-colporate grains with reticulate exine, typical for subgenus Phyllanthus , and very different from the clypeate grains of the West Indian phylloclade-bearing species in section Xylophylla . This pollen evidence clearly demonstrates homoplasy in the origin of phylloclades in Phyllanthus . Pollen morphological data suggest that the neotropical taxa of Phyllanthus have arisen following colonization from Africa (subgenus Kirganelia ) and Asia (subgenus Emblica ). © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 325–338.     

Has Pollination Mode Shaped the Evolution of Ficus Pollen?

Background

The extent to which co-evolutionary processes shape morphological traits is one of the most fascinating topics in evolutionary biology. Both passive and active pollination modes coexist in the fig tree (Ficus, Moraceae) and fig wasp (Agaonidae, Hymenoptera) mutualism. This classic obligate relationship that is about 75 million years old provides an ideal system to consider the role of pollination mode shifts on pollen evolution.

Methods and Main Findings

Twenty-five fig species, which cover all six Ficus subgenera, and are native to the Xishuangbanna region of southwest China, were used to investigate pollen morphology with scanning electron microscope (SEM). Pollination mode was identified by the Anther/Ovule ratio in each species. Phylogenetic free regression and a correlated evolution test between binary traits were conducted based on a strong phylogenetic tree. Seventeen of the 25 fig species were actively pollinated and eight species were passively pollinated. Three pollen shape types and three kinds of exine ornamentation were recognized among these species. Pollen grains with ellipsoid shape and rugulate ornamentation were dominant. Ellipsoid pollen occurred in all 17 species of actively pollinated figs, while for the passively pollinated species, two obtuse end shapes were identified: cylinder and sphere shapes were identified in six of the eight species. All passively pollinated figs presented rugulate ornamentation, while for actively pollinated species, the smoother types - psilate and granulate-rugulate ornamentations - accounted for just five and two among the 17 species, respectively. The relationship between pollen shape and pollination mode was shown by both the phylogenetic free regression and the correlated evolution tests.

Conclusions

Three pollen shape and ornamentation types were found in Ficus, which show characteristics related to passive or active pollination mode. Thus, the pollen shape is very likely shaped by pollination mode in this unique obligate mutualism.     

Polymorphism in pollen of Salvia leucantha (Lamiaceae)

LM and SEM study of the pollen from single specimen of Salvia leucantha Cav., gathered from Kumaon mountains in Western Himalaya have shown significant variations in aperture number and type, shape and size of pollen grains, hitherto not reported. Normal pollen is 6-colpate but 4, 5, 7, 8, 9, 10, 11-colpate to spiraperturate also occur to make it a heterogenous assemblage. In shape, pollen varies from oblate, suboblate, oblate-spheroidal, prolatespheroidal to subprolate, whereas, in size it ranges from 15 μm to 40 μm. Furthermore, besides the marked variations in monads, the polymorphism is also expressed in terms of dyads and triads. Exine ornamentation however, does not vary, it being reticulate-retipilate sexine pattern in all the different types under LM and showing double sculpturing under SEM.     

中国芸香科植物的花粉形态

本文用光学显微镜对我国21属52种2变种芸香科(Rutaceae)植物的花粉形态进行了详细地观察研究,其中对6属6种进行了扫描电镜观察。本科花粉具沟孔,是花粉单类型的科之一。根据萌发孔的数目和外壁纹饰特点,本文将芸香科的花粉分为具3孔沟和4-6孔沟两个花粉类型,并根据各属花粉形态特征,作出了花粉形态分属检索表。此外,本文还对花椒(Zanthoxylum L.)和Fagara两属的合并问题及花椒与吴茱萸(Evodia Scop.)两属之间的亲缘关系等问题进行了讨论。