Pollen morphology and its taxonomic significance of the monotropoideae (Ericaceae)

A palynological survey, including LM, SEM and TEM is presented for eight genera, nine species and 76 samples of the Monotropoideae which is composed of 10 genera and 13 species. On the basis of the aperture number and shape, the following six pollen types are recognized: 1) 3-colp(oroid) ate—Allotropa, 2) 2-colpor(oid)ate—Monotropsis andHemitomes, 3) 4-colpor(oid)ate—Pterospora, Sarcodes andPleuricospora, 4) 2- and 3-colpor(oid)ate—Monotropa hypopitys, 5) 3-colporate and-porate —Monotropa uniflora, 6) 3- to 4- porate—Monotropastrum humile. Relationships among taxa within the Monotropoideae are illustrated on the palynological characters including the aperture type, exine sculpture and structure. Both pollen grains with two and four apertures have evolved independently from pollen grains with three apertures, according to the infraspecific variation of aperture numbers and the usual occurrence of three-aperturate pollen grains in the Ericaceae. 3-colp(oroid)ate pollen ofAllotropa is the most primitive and occupies an isolated position in the subfamily, on the other hand 3- to 4-porate pollen ofMonotropastrum humile is most advanced in the subfamily, probably even in the Ericaceae. A reduced sexine inMonotropastrum humile appears to be a specialized pollen character. Infraspecific geographical difference in palynological characters is revealed in two species ofMonotropa for the first time.M. hypopitys is basically characterized by pollen grains with two apertures in the New World vs. three apertures in the Old World.M. uniflora is basically characterized by 3-porate pollen in the New World vs. 3-colporate in the Old World. In both species pollen grains with more primitive characters usually occur in the Old World.     

Pollen morphology and its evolutionary significance in Hemerocallis (Liliaceae)

Pollen morphology of 10 species of Hemerocallis was investigated with LM and SEM. In addition to pollen with reticulate sexine, a new verrucate sexine pattern is found in three Chinese endemic species, H. plicata, H. forrestii and H. nana . Between the two typical sexine patterns, there are transitional variants. Also, a new pollen shape, boat-shaped-elliptic, is observed exclusively in H. nana , besides-oblong shape in other species. Phylogenetically, the common reticulate sexine pattern would be more primitive than the verrucate one. The typical reticulate sexine pattern would be basic, from which transitional variants would have derived and finally led to the typical verrucate pattern. Similarly, the common boat-shaped-oblong pollen would be more primitive than the -elliptic one, from which the latter would have derived during phylogenetic development.     

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 Phyllanthus species occurring in the continental United States

Twelve native United States species have been examined. The pollen grains could be divided into five pollen types based on sexine ornamentation, aperture structure, sexine/nexine ratio and sexine 1/sexine 2 ratio. Within these types it was often possible to distinguish the grains of the different species. Two groups of types have been recognized based on the similarities in ornamentation. One group, consisting of the Phyllanthus amarus type, the P. tenellus type and the P. urinaria type, is characterized by the Tilia structure, whereas the second group of types, comprising the P. caroliniensis type and the P. liebmannianus type are microreticulate. The postulation that different subgenera show different pollen types is true in general, but there is one exception: P. niruri, taxonomically belonging to the subgenus Phyllanthus, has pollen grains of the P. caroliniensis type, a type also comprising species from the subgenus Isoclades.     

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.     

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。     

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.     

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.     

STUDIES OF SEED FERN POLLEN: DEVELOPMENT OF THE EXINE IN MONOLETES (MEDULLOSALES)

Monoletes pollen extracted from the seed fern synangium Dolerotheca sclerotica Baxter illustrate four stages in the development of the sporoderm. In the first stage the grains are up to 100 μm long and possess an apparent homogeneous exine in which there is little differentiation between the nexine and sexine. Numerous nexine lamellae and the initiation of sexine expansion mark stage 2 in exine ontogeny. Further expansion of the sexine continues in the third stage until the ratio between the nexine and sexine is approximately 1:5. The final stage in maturation of the sporoderm shows an expanded alveolate sexine with some of the sporopollenin units broken and disorganized. It is at this stage of development that nexine lamellae are most prominent. The formation of sporoderm layers in the fossil grains is compared with pollen grain development in living cycads (Cycadophyta) and a model proposed to account for the apparent early formation of nexine lamellae in Monoletes. The evolution of exine components in early pollen types is discussed.     

A Study on Pollen Morphology and Ultrastructure of Subfamily Castaneoideae (Fagaceae) in China

The present paper describes the pollen morphology of 45 species of the subfamily Castaneoifeae (including genera Castanea, Castanopsis and Lithocarpus) from China. The pollen grains were all examined with light microscope, scanning electron microscope and transmission electron microscope. Pollen grains of the subfamily are prolate, subprolate or perprolate, (14.7-23.1)× (8.4 -18.9) μm in size, 3-colporate, the exine in 2-layered, 0.9-1.9μm thick, indistinctly ornate, striate-rugulose or crass-striate, sexine and nexine almost equal in thickness, the sexine consists of tectum, bacules and endonexine under TEM. On the basis of very similar pollen shape, pollen size, type of aperture and exine structure and also other characteristics of plant morphology of the genera Castanea, Castanopsis and Lithocarpus, the present authors tend to support the opinion that they all fall into the samesubfamily, Castaneoideae.     

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.     

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.     

An Ultrastructural Study of Pollen Wall Ontogeny in Silene Pendula

Pollen of 41 species representing all seven genera of the Neottieae were examined by scanning electron microscopy. Except for Lecanorchis, the genera of the Neottieae constitute a natural group based on pollen morphology. Pollen occurs as single grains in the primitive species and in tetrads in the other species. Most grains are monoaperturate, porate or tenuate, and the tetrads often have irregularly shaped grains. Exine structure varies from tectate-perforate to semitectate. Lecanorchis is anomalous among the Neottieae in that it has 0–5, sunken, relatively small pores. Pollen morphology of this genus indicates that it is probably more closely related to the Gastrodieae than the Neottieae. There are at least four basic phyletic units in the Orchidaceae: the Neottioideae, Apostasioideae, Cypripedioideae, and Epidendroideae. These groups are distinguished by the presence of monads in at least their more primitive members and by their unique pollen types.     

Pollen morphology and ultrastructure of Marathrum schiedeanum (Podostemaceae)

The Podostemaceae, or river-weeds, comprise 46 genera and 270 species of dicots and are the largest family of strictly aquatic angiosperms. Despite the large size, specialized habitats, and enigmatic morphology of the family, relatively little is known about the palynology of Podostemaceae. In the current paper, pollen morphology and ultrastructure of Marathrum schiedeanum are described. Pollen grains are relatively small, spheroidal, and tricolpate to spiraperturate. The exine has a microechinate ornament, a tectate-granular sexine and a relatively thick nexine in non-apertural regions, and a semitectate sexine and thinner nexine in apertural regions. Although aperture variation occurs in the family, this is the first report of the spiral aperture type in Podostemaceae. The spiraperturate condition appears to be derived in river-weeds, as does the granular pollen wall, which represents a reduction of the typical columellae found in eudicots.     

Pollen morphology of Microstachys (Euphorbiaceae) with emphasis on neotropical species

Abstract

In Euphorbiaceae tribe Hippomaneae is the pantropical genus Microstachys, comprised by 24 species, mostly from Brazil. Palynological studies with several representatives of the genus are scarce. The purpose of this study was to describe the pollen morphology of Microstachys and find possible differentiating characteristics between its species. Ten species of the genus were analysed, represented by 21 herbarium collections. Their pollen grains were acetolysed, measured and photographed under light microscopy (LM) and scanning electron microscopy (SEM). The analysed species possess pollen in monads, of small to medium size, isopolars, amb distincticly trilobed, oblate-spheroidal, prolate-spheroidal or subprolate, tricolporate with narrow and very long colpi, almost fused at the poles sometimes forming an invagination with tapering ends and margo of the colporus wide to very wide and psilate; endoapertures lalongate to very lalongate with tapering ends, presence of a narrow costae distinct in the endoaperture; exine microreticulate homobrochate, walls with simple columella; at the mesocolpus the sexine separates from the nexine forming cavea; and sexine always thicker than the nexine. Microstachys is a stenopalynous genus, despite its species varying in pollen size, shape and width of the cavea, margo of the colporus, and length of the endoapertures, demonstrating taxonomic value, and indicating a contribution towards future systematic analyses.     

POLLEN STUDIES IN THE GENUS CORDIA (BORAGINACEAE)

The pollen of 40 species of Cordia L. was studied by light microscopy and scanning and transmission electron microscopy. Three major types were found, 3-porate with a reticulate sexine, 3-colpate to 3-colporoidate with a striatoreticulate sexine, and 3-colpate to 3-colporate with a spinulose sexine. These pollen types are correlated with significant differences in the inflorescence, flower, and fruit structures found in the sporophyte. The following revisions are suggested and discussed: that Cordia sect. Varronia P. Br. be elevated to generic status; that Cordia sebestena L. and a few close allies be treated as generically distinct; and that Cordia myxa L. be conserved as the lectotype for the genus, thus continuing a wide application of the epithet Cordia.     

Aperture Orientation in Laurelia Pollen (Atherospermataceae Syn. Subfamily Atherospermoideae of Monimiaceae)

The development of the encircling aperture of pollen grains or the New Zealand species Laurelia novae-zelandiae is described by means of light and scanning electron microscopy. Some observations are also reported from pollen development and structure in the South American species Laurelia sempervirens, mature pollen of which can be distinguished from that of L. novae-zelandiae. The aperture begins to develop while pollen of L. novaezelandiae is still in tetrads, and passes through the distal and proximal poles of each grain. The widest parts of the mature aperture are at and near what are morphologically the distal and proximal poles of each grain. No such meridionosulcate type of pollen was recorded in a recent survey of pollen aperture types of primitive angiosperms and contradicts a recent report that pollen in the Atherospermataceae had equatorially aligned apertures.     

POLLEN MORPHOLOGY AND EVOLUTION IN HEDYOTIS SUBGENUS EDRISIA (RUBIACEAE)

Primarily on the basis of aperture structure, 31 species of Hedyotis subg. Edrisia (Houstonia) are separable into 5 palynological groups. Group 1 is characterized by a simple os circumscribed by varying nexinous thickenings, groups 2–4 combine this os with a crassimarginate one to form a compound os, while group 5 is known only with the crassimarginate os. The first type is considered primitive, the compound os specialized and more advanced, and the third type reduced and highly advanced. Other characteristics of the pollen, including size and shape, aperture number, thickness of sexine and nexine, and reticulum, are discussed in relation to the apertures, and their probable primitive and advanced expressions are outlined. The data agree with the phyletic trends found in the sporophyte on the basis of results from morphology, chromosome number and size, and distribution, and they support and add to an earlier phylogenetic scheme proposed for the subgenus. The evidence from palynology also supports the treatment of Hedyotis and Houstonia as congeneric.     

THE ULTRASTRUCTURE OF SAHNIA POLLEN (PENTOXYLALES)

The micromorphology and fine structure of in situ pentoxylalean pollen are described from the holotype of Sahnia laxiphora Drinnan and Chambers 1985 collected from the Lower Cretaceous (Valanginian-Aptian) of Victoria, southeastern Australia. Pollen grains are ovoid, monosulcate, and relatively small, averaging 26 μm in length. Exine ornamentation is psilate. The sporoderm is two-parted with the sexine staining lightly throughout and approximately six times the thickness of the more darkly staining nexine. The exine over the sulcus is typically strongly invaginated, and may or may not include an extremely thin sexine layer. The outer part of the sexine is homogeneous, while the inner part is composed of relatively large granules separated by irregular lacunae of various sizes; lacunae are most pronounced at the sexinenexine interface. Faint lamellae characterize the nexine in both apertural and nonapertural regions. Granular orbicules are often associated with the exine surfaces and also occur appressed to pollen sac walls along with lamellated tapetal membranes. Sporoderm ultrastructure is compared to that of nonsaccate pollen of other groups, and particularly to pollen of Bennettitales, Gnetales, angiosperms, and similar plants, to which the Pentoxylales have been thought to be closely related. Although Sahnia laxiphora pollen is not identical to that of any of these taxa, the strongest similarity is with pollen of Bennettitales.