Systematic importance of pollen morphological features of selected species from the genus Rosa (Rosaceae)

The aim of this study is verification of the taxonomic usefulness of the pollen grain features studied, based on pollen morphology of 32 wild species from all 4 subgenera and all 10 sections of the genus Rosa, mainly for delimitation of subgenera, sections, and species. The measurements and observations were carried out with both light microscopy and scanning electron microscopy. Only correctly formed pollen grains (30 per specimen) were measured, and 960 pollen grains were examined in total. They were analyzed for 14 quantitative features of pollen grains and exine sculpturing and the following qualitative ones: outline, shape, and operculum structure. Our study revealed that the diagnostic features of pollen grains studied were: exine sculpture, length of polar axis, and pollen shape (P/E ratio). On the basis of the above characters, five species were isolated and the remaining ones were included in several groups isolated on the basis of exine sculpture types. The following three exine sculpture types occurred in the species studied: granular-verrucate (in R.?stellata), striate-psilate (in R.?multibracteata and R.?multiflora), and striate (the remaining species). R.?banksiae is characterized by small pollen grains, while R.?setigera has strongly elongated pollen with P/E ratio >1.5. Exine sculpture features considered to be diagnostic should be treated as auxiliary because they fail to differentiate individual species, although they can be helpful in distinguishing groups of species of similar exine sculpture. The arrangement of the species examined on a dendrogram only slightly corroborates division of the Rosa genus into subgenera and sections currently adopted in taxonomy (Rehder 1940). An interesting result was reported for the species studied from the Caninae (R.?agrestis, R.?canina, R.?dumalis, R.?jundzillii, and R.?rubiginosa) section which, despite hybrid nature, with the exception of R.?villosa, grouped in the same, most separated group of species.     

Palynological survey of the subtribe Elephantopinae (Asteraceae, Vernonieae)

Despite the taxonomic value of pollen morphology within the tribe Vernonieae, a detailed study of the pollen of the subtribe Elephantopinae is still lacking. The pollen morphology of ten species, representing three of the four genera of the subtribe, Elephantopus, Pseudoelephantopus and Orthopappus, was studied with LM, SEM, and TEM. The pollen of all the species studied was found to be echinolophate, although the differences in aperture features and both sculpture and exine structure allowed recognizing two pollen types. The species of Elephantopus (except for E. elongatus) and Pseudoelephantopus share the regular or more or less regular ridges pattern, the colporate condition but with the ectoapertures little developed, and the acaveate exine structure. From the sculpture and the apertures, the pollen of Caatinganthus harleyi was found to be similar to that of Elephantopus and Pseudoelephantopus. The pollen morphology of the monotypic Orthopappus, which is shared with that of E. elongatus, was further investigated for the first time. We found that it differs from that of the other species in having an irregular pattern of ridges, colporate condition but with a well-developed ectoapertures, and caveate exine. Additional studies of the exine structure and apertures features, coupled with molecular phylogeny, are needed to understand the evolution of pollen characters and re-evaluate the intergeneric relationships within the tribe.     

中国紫金牛属圆齿组花粉形态研究及其分类学意义

利用扫描电子显微镜对紫金牛属圆齿组20种1变种植物的花粉形态进行了研究,并比较分析了腋序组2种及锯齿组1种的花粉。经观察花粉为近球形或近扁球形,具3孔沟,除锯齿组的月月红外,其余种均形成合沟。外壁纹饰可分为四类:穴网状、细网状、皱波状及细颗粒状(带刺突)。结果显示,花粉形态特征可作为属下分组及组下分类处理的依据。     

Pollen morphology,ultrastructure and taphonomy of the Neuradaceae with special reference to Neurada procumbens L. and Grielum humifusum E.Mey. ex Harv. et Sond.

Pollen morphology and sporoderm ultrastructure of modern Neurada procumbens L. and Grielum humifusum E.Mey. ex Harv. et Sond. were studied using light (LM) and electron (SEM and TEM) microscopy. Additionally late Holocene pollen of the Grielum-type was studied using LM. Systematic and ecological aspects have been discussed for the family Neuradaceae. The pollen grains of the studied species are characterized by similarities in size, shape, aperture type and differences in exine sculpture (reticulate semitectate exine in Neurada and finely reticulate to foveolate in Grielum) and sporoderm ultrastructure. The cavea in the exine is situated between the ectexine and endexine which are connected near the aperture region only. A combination of the palynological characters of the Neuradaceae (semitectate exine, rather loose columellae, interrupted foot layer, the cavea in the exine) increases the pollen plasticity, allowing considerable changes of the pollen grain volume but still remains insufficient to survive sharp fluctuations in hydration level.     

Pollen morphology of some endemic Turkish Centaurea L. (Asteraceae, section Phaloletis) and their taxonomic value

In this study, the detailed pollen morphological structures of some endemic Turkish species of Centaurea amaena Boiss., C. antalyense H. Duman & A. Duran, C. aphrodisea Boiss., C. hierapolitana Boiss., C. luschaniana Heimerl, C. lycia Boiss., C. tossiensis Freyn. Et Sint., and C. wagenitzii Hub.-Mor. (Asteraceae, section Phaloletis) were studied under light microscopy (LM) and scanning electron microscopy (SEM) for the first time. LM and SEM investigations showed that the pollen grains of eight taxa are more or less spheroidal-subprolate, the amb triangular and tricolporate. The exine sculpture is tectate, microechinate-scabrate in the pollen of Centaurea taxa. Spinules are less dense in Centaurea amaena, C. antalyense, C. hierapolitana, and C. lycia, but they are more dense in C. aphrodisea, C. luschaniana, C. tossiensis, and C. wagenitzii. Spinule dimensions are different from each other. The exine has one layer of columellae beneath the spines. We determined all taxa that have the Helianthoid type. Exine sctructure and sculpture as well as spine density and dimensions in Asteraceae are the most reliable characteristics for discriminating taxa.     

Pollen Morphology of the Genus Malus and Its Taxonomic and Evolutionary Significance

The pollen morphology of 26 species and 5 hybrids of the genus Malus was investigated with aid of SEM. It is found that the pollen morphology of Malus is rather similar in shape, size, position and number of aperture and exine sculpture. The characters of polle morphology of sections and series are as follows: Sect. Malus Ser. Baccatae: Striae regular, parallel to colpi, and conjunct at pole; Ser. Pumilae: the same as in Ser. Baccatae, but more or less curved near pole. Sect. Docyniopes: Striae regular and parallel to colpi, but bent near pole. Sect. Chloromeles: Striae irregular, dense and interlock. Sect. Sorbormalus: Striae irregular, sparse and not interlock. Ser. Sieboldiance: pollen grains prolate, with perforation among striae, colpi narrow; Ser. Kansuenses: Pollen grains spheroidal, colpi wide in the middle but narrow at both ends, striae relatively dense and regular, mostly dichotomous, perforation present; Ser. Yunnanenses; Pollen grain spheroidae, colpi wide in the middle but narrow at both ends, striae obviously irregular, less dichotomous, perforation absent. The major evolutionary trend of exine sculpture of pollen may be from densely thin-striate to sparsely striate with perforation. Characters of the exine sculpture of hybrids can be used to recognize the rela-tionship between the parental species.     

Pollen morphology in relation to the taxonomy and phylogeny of some native Greek Aegilops species

The pollen morphology of some native Greek Aegilops species is investigated in LM (quantitative pollen characters) and SEM (exine sculpture) using acetolysed material. Furthermore, quantitative data are subjected to a multivariate analysis. The tetraploid Ae. cylindrica and its diploid parent Ae. caudata show a distinct morphological affinity as regards their quantitative pollen profile and the features of the exine sculpture. The tetraploid Ae. triuncialis is morphologically divergent from both its parents Ae. caudata and Ae. umbellulata due to the very large values of its quantitative pollen characters. However, the SEM survey of the exine sculpture indicates a rather high degree of similarity between Ae. triuncialis and Ae. umbellulata. No significant differences have been found between the two varieties of Ae. caudata (caudata and polyathera) concerning the quantitative pollen characters as a total or the morphology of the exine sculpture. The resultant clustering of the taxa on the basis of the quantitative pollen characters as well as the recorded similarities of their exine pattern are related to their sectional classification based on cytogenetical and morphological data.     

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.     

Palynological survey of subtribe Pithecocteniinae (Bignonieae,Bignoniaceae)

The pollen morphology of subtribe Pithecocteniinae was reviewed. Thirty species of the six genera currently recognized, namely Amphilophium, Distictis, Distictella, Glaziovia, Haplolophium and Pithecoctenium, were considered. All the species surveyed fell into one of the two pollen groups: (1) inaperturate, spheroid pollen; and (2) stephanocolpate, prolate pollen. The former group included the studied species of Distictis, Distictella and Pithecoctenium, the latter species of Amphilophium, Glaziovia and Haplolophium. The variation of exine sculpture and thickness did not show any taxonomic relationships. An argument for considering pollen features, together with other morphological characters, to elucidate monophyletic units within Pithecocteniinae is presented. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 155–162.     

樟科植物花粉形态研究

樟科植物为我国南方常绿阔叶林重要组成成分,我国约有20属420余种。过去对于樟科花粉形态的研究仅限于在光学显微镜下进行观察记载,以台湾学者王仁礼的研究最为全面,共记载了台湾产14属40余种的花粉形态。本文作者以国产樟科为主,应用扫描电镜及光学显微镜对该科花粉形态进行了更为全面系统的研究,共观察记载了22属150余种,对各属的花粉形态特征均进行了描述;首次报道了樟科花粉有小穿孔的存在,并依据花粉的形状、外壁纹饰及小穿孔存在与否将樟科花粉划分为7个类型,即:1．厚壳桂型(Cryptocarya,Neocimamomum);2．檬果樟型(Caryodaphnopsis);3．木姜子型(Litsea,Lindera,Neolitsea);4．月桂型(Laurus);5．鳄梨型(Persea,Machilus,Syndiclis,Nothaphoebe, Beilschemiedia,Phoebe,Alseodaphne,Octea);6．檫木型(Sassafras,Cinnamomum,Umbellularia,Actinodaphne,Dehaasia);7．无根藤型(Cassytha)。文中还讨论了樟科花粉的多样性、有关属之间的亲缘关系和分类问题以及同邻近的肉豆蔻科、莲叶桐科的关系。     

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。     

Ultrastruktur und systematische Bedeutung des Pollens beiBocageopsis,Ephedranthus, Malmea undUnonopsis (Annonaceae)

The four genera investigated show solitary sulcate pollen grains. The structure and sculpture of the sporoderm is very similar inBocageopsis andUnonopsis, and supports the close relationship of both genera; their flower morphology also is very similar. In contrast, the sporoderm ofEphedranthus, and especially that ofMalmea, is different in some characters and suggests more remote relationships; this is also confirmed by differences in flower morphology. Within this group of genera a distinct exine progression can be recognized from non-columellate (Malmea) to granular (Unonopsis guatterioides), further to somewhat irregularly (Bocageopsis, someUnonopsis species), and finally to very regularly columellate (Ephedranthus). The sculpture of the tectum varies from a reticulum with large lumina (Malmea) to one with small performations (Ephedranthus). Within theAnnonaceae the genusMalmea is among the most primitive in respect to pollen structure. The sulcus of the four genera is very large and runs over 1/3 of the pollen grain. It is characterized by a reduction of the exine and a bulgy thickening of the intine.

     

A light, scanning electron, and transmission electron microscopic study of pollen wall architecture in the subtribe Gnaphaliinae (Gnaphalieae, Asteraceae)

To better understand relationships within the Asteroideae, the pollen morphology and exine structure of 10 genera and 15 species of Gnaphaliinae were investigated using light, scanning electron, and transmission electron microscopy. All taxa have a Gnaphalioid pattern of exine with an evidently rough foot layer. The tectal complex consists of three main layers that differ in morphology and thickness: a tectum, a median columellar layer, and an internal interlaced sub-columellar layer. The apertural system consists of an ectoaperture, a mesoaperture, and an endoaperture, which intersect, respectively, the tectal complex, the foot layer and the upper part of the endexine, and the inner part of the endexine. On the basis of pollen characteristics, especially those of the spines, all the species examined may be gathered into two groups, one including Helichrysum foetidum, H. italicum, Plecostachys serpyllifolia, and Pseudognaphalium luteo-album and the other including the other 11 species of Gnaphaliinae now investigated. In addition, because Gnaphalieae has been proposed as sister group to several tribes and clades of Asteroideae, a comparison was made between exine pattern in the Gnaphalieae and that in its putatively related tribes. On the basis of previous phylogenetic studies and our present pollen data we suggest that either Astereae, Astereae-Anthemideae clade, or Heliantheae s.l.-Anthemideae clade are the best candidates for the sister taxon/sister group of Gnaphalieae.     

Morphology,reserves and pollen viability of some Solanum Sect. Cyphomandropsis species

The morphology, energy reserves and viability of three species from Solanum Sect. Cyphomandropsis (Solanum adelphum Morton, S. glaucophyllum Desf. and S. stuckertii Bitter), which grow in Argentina were studied. The pollen grains are tricolporate and their characters are homogeneous. The only differences found among the species were the pollen grain size and the sculpture distribution. The surface features of exine were not readily visible under light microscope, but with SEM the sculpture appeared granulate. Analyses of the cytoplasm of pollen grains show the presence of oils. The average viability was high in two of the studied species (S. adelphum and S. glaucophyllum). An interesting difference was observed between the first and second day flowers of S. stuckertii. The second day flowers had high fertility but those from the first day had a very low fertility. In the latter case, an inner placentoid tissue of the anther hinders pollen release until the entire development is completed. The P/O ratio is linked to the breeding system.     

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 inParis and its related genera

Pollen grains ofParis, Medeola, andScoliopus have been examined with light, scanning electron, and transmission electron microscopies.P. quadrifolia L.,P. verticillata Bieb.,P. delavayi Franch. andP. polyphylla Smith have monosulcate pollen characterized by psilate, foveolate or reticulate exine sculpture. In contrast to the former species,P. japonica (Fr. et Sav.) Franch. andP. tetraphylla A. Gray have monosulcate pollen with gemmate (rarely rugulate) exine.Medeola has monosulcate pollen with reticulate exine that is distinct from that ofParis. Scoliopus has monosulcate pollen characterized by a peculiar reticulate exine pattern. The palynological evidence suggests thatParis andTrillium are closely related to each other, andMedeola andScoliopus should be separated fromParis andTrillium.     

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 ofPyrola and its taxonomic significance

Pollen from 152 collections representing 16Pyrola species was examined with LM, SEM and TEM. The genusPyrola is stenopalynous in having tetrads and 3-colporoidate grains. However, a continuous and serial variation in the exine sculpture and the tetrad diameter (D) was revealed within the genus. Křísa's system ofPyrola (1971) was re-examined in the light of these palynological characters which may have a taxonomic significance, as well as the external morphology. P. minor (only member of the subgenusAmelia), which is characterized by having small (D=30–35 μm) pollen tetrads with verrucate sculpture must best be placed near sectionPyrola of the subgenusPyrola. P. faurieana must be segregated from sectionChlorantha and together withP. media placed intermediate between the subgenusAmelia and sectionPyrola of the subgenusPyrola based on their larger (D=ca. 40 μm) pollen tetrads with verrucate sculpture. Two series of sectionChlorantha excludingP. faurieana are differentiated in the exine sculpture. Therefore, they are regarded as the distinct taxonomic groups. SectionScotophylla is characterized by having psilate sculptured pollen tetrads, which indicates that its sectional rank is sufficiently asserted. The evolutionary trend in the exine sculpture from verrucate through rugulate to psilate is suggested withinPyrola.     

POLLEN MORPHOLOGY IN THE GENUS ERYTHRONIUM (LILIACEAE) AND ITS SYSTEMATIC IMPLICATIONS

Pollen morphology was examined in 40 samples from 19 species in the genus Erythronium (Liliaceae) using scanning (SEM) and transmission electron microscopy (TEM). Pollen grains are monosulcate monads, except for a plant of E. oregonum which has dyad grains. North American and European species are characterized by reticulate exine sculpture composed of various degrees of wavy muri and variably-sized lumina. The most distinctive feature of exine sculpture in the genus is exhibited by the Asian species E. japonicum, which has a unique reticulate pattern composed of striate muri and a distinctive exine structure without columellae. The distinctness of E. japonicum pollen suggests that it has specialized in isolation from species in North America and Europe.     

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).