The pollen morphology of Phyllanthus (Euphorbiaceae) section Choretropsis

The pollen grain morphology of the eight South American species of the genus Phyllanthus , subgenus Phyllanthus , section Choretropsis , endemic to Brazil, was investigated using light and scanning electron microscopy. The high homogeneity in pollen shape, as well as aperture number and configuration, allowed the definition of a basic pollen type for this section, viz. prolate spheroidal or subprolate, zonocolporate with 4–5 invaginated colpi, sexine reticulate. Two subtypes have been observed, based on pollen grain apertures and exine sculpture. The first subtype is characterized by straight and narrow colpi provided with costae, lalongate endoapertures and a reticulum of rectilineous muri as in P. choretroides and P. spartioides . The second subtype has wider colpi without costae, circular endoapertures and a reticulum with curvilineous muri as in P. flagelliformis , P. goianensis and P. scoparius , or curvilineous-rectilineous muri as in P. angustissimus , P. gladiatus and P. klotzschianus . It was demonstrated that the pollen shape and the number of colpi varied within and between some of the species. The results suggest that pollen grain morphology, in association with plant architecture and the presence of phylloclades, constitute additional evidence to delimit the species of the section Choretropsis and to further understand their evolutionary origin within the genus Phyllanthus.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 243–250.     

A NEW LYGINOPTERID POLLEN ORGAN WITH ALVEOLATE POLLEN EXINES

A new lyginopterid pollen organ is described based upon specimens occurring in a single coal ball from the Providence, Kentucky locality. Seven to nine beaked sporangia are fused together at their proximal ends forming a common synangial chamber; synangia are joined together in clusters of two or three. In situ prepollen is similar to Cyclogranisporites and Verrucosisporites sporae dispersae. The thick exine has a lamellate nexine and a prominent alveolate sexine.     

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.     

Pollen of African Spermacoce species (Rubiaceae) Morphology and evolutionary aspects

Pollen morphology of 43 African species of the genus Spermacoce has been investigated by scanning electron and light microscopy. The genus is eurypalynous, which is reflected in the remarkable variation of almost all pollen characters. The average equatorial diameter (E) ranges from 15.8 w m to 115.5 w m. Grains are colporate or pororate. The number of apertures varies from 3 up to more than 25. The majority of species has apertures situated only at the equator (being zonoaperturate), but a few species have pantoaperturate grains. The endoaperture is generally an endocingulum, often with a secondary lolongate or lalongate thinning at the ectocolpus; endocolpi and endopores are also observed. The sexine is usually perforate, but eutectate, foveolate, and (micro)reticulate tecta were also found. Supratectal elements are present as granules, microspines or spines. The inner nexine surface is granular, often with irregular grooves (endocracks). Among native African species, nine pollen types are recognized mainly on the basis of pollen size, aperture morphology and tectum peculiarities. In two of the pantoaperturate types, apertures are in a configuration not yet recorded for the angiosperms in general. Some evolutionary trends are proposed that await verification by further systematic study. Pollen morphological characters have a high taxonomic value in the genus Spermacoce . They provide almost unique identification marks for the species, which enables sharpening of species boundaries. Small groups of related species often share the same pollen type. The genus Borreria , previously separated from Spermacoce on the basis of its fruit morphology only, is not supported by pollen data.     

Dimorphic exine sculpturing in two distylous species ofDyerophytum (Plumbaginaceae)

Light and SEM observations on the pollen ofDyerophytum africanum andD. indicum have revealed marked differences in exine features. These distylous species also have dimorphic pollen. In the short-styled individuals of both species, the sexine and nexine are of equal thickness, and the clava-like sexinous processes are short without marked projections. In the long-styled individuals, the sexine is thicker than the nexine, the clavae are higher than broad with an apical spinule. Pollen size and apertures are identical in both morphs. — Palynological evidence is presented for relationships betweenDyerophytum andCeratostigma, Plumbago andAegialitis. Moreover, the genusDyerophytum exhibits pollen morphological similarities with some species ofLinum (Linaceae).     

Pollen morphology and ultrastructure of the Bennettitales: In situ pollen of Cycadeoidea

The micromorphology and ultrastructure of in situ pollen from Cycadeoidea dacotensis are described from permineralized specimens collected from the Lower Cretaceous of North America. Pollen grains are ovoid and relatively small, averaging 25 μm in length and 12 μm in width. Grains are monosulcate with the exine typically invaginated in apertural regions. Exine ornamentation ranges from punctate to psilate. The exine averages 0.73 μm in thickness and is composed of a light-staining sexine and a dark-staining nexine. The sexine consists of a thin, homogeneous tectum, typically with a well-defined inner boundary, and a thicker granular infratectum. The infratectal granules are relatively uniform in size, however, variation occurs in the arrangement of granules. In some grains, the sexine appears homogeneous because there is little lacunal space between the individual granules. The granular infratectum is in direct contact with the underlying nexine. The nexine is uniform in thickness in both apertural and nonapertural regions, and it lacks lamellae throughout. Pollen morphology and ultrastructure are compared with those of the bennettitalean genus Leguminanthus and the dispersed genus Monosulcites. In addition, the fine structure of Cycadeoidea pollen is compared to that of the gymnosperm groups with which the Bennettitales are regarded to be most closely related, including Gnetales, Pentoxylales, and Eucommiidites-type pollen-producing plants.     

Pollen Morphology and Wall Stratification in Rubus L. (Rosaceae) in the Iberian Peninsula

A palynological study using LM and SEM was made of the morphology, ornamentation, apertural system and structure of the pollen wall, of the 18 species of the genus Rubus which are present in the Iberian Peninsula. We demonstrate: a) the stenopalynous character of the genus (isopolar, subprolate, 3-lobate, 3-zonocolporate and striate-perforate, except for R. genevierii whose ornamentation is striate-gemmate-perforate and with a striated margin); b) the lalongate shape of the endoaperture, due to the disappearance or thinning of the nexine; and c) the existence of two structural types, Type I granular and Type II columellar with clearly-defined columellar spaces or compact without intercolumellar spaces.     

Immunocytochemical localization of the allergenic proteins in the pollen of Cryptomeria japonica

Applying an immunocytochemical method, a localization of the protein Cry j I in the Cryptomeria japonica pollen, which is the major allergen responsible for Japanese cedar pollinosis, is investigated with the monoclonal and polyclonal antibodies produced from the protein. The protein that reacts to the polyclonal antibody localizes on the sexine, nexine, between nexine and intine layers, orbicles, cell wall of a generative cell, Golgi body and Golgi vesicles. The allergenic protein contained in the exine and orbicles of Japanese cedar pollen can diffuse or dissolve easily from there into the mucus covering of the eye and nose, causing a response in less than 1 min after exposure. Since the orbicles have a diameter of about 430 nm, they can pass easily through the pores of most protective masks to reach the sensitive tissues of the patient. The proteins react to the monoclonal antibodies (J1BO1 and J1BO7) and localize on the Golgi body, sexine, nexine and orbicles (but not between the nexine and intine layers), and on the generative cell wall. In the young pollen grain, numerous allergenic protein particles contained in the orbicles and sexine layer, but there is only a small amount of the protein between the nexine and intine layers, since the intine layer is not yet complete at this stage. More will be accumulated there during developmental maturation. The allergenic protein is also found on the tapetal materials remaining in the young anther. Since the materials forming the exine layer and orbicles come from tapetal tissue, it is assumed that some of the allergenic protein is produced in the tapetum and localized in the orbicles and pollen wall during maturation, and that the rest of the allergenic protein is produced in the Golgi body in the mature pollen grain.     

Palynological variation in balsaminoid ericales. I. Marcgraviaceae

BACKGROUND AND AIMS: Marcgraviaceae are a rather small family of seven genera and approx. 130 neotropical species. This study aims to present a detailed palynological survey of the family in order to comment on the intrafamily relationships and possible correlations with pollinators. METHODS: In total, 119 specimens representing 67 species and all genera are observed using light microscopy and scanning electron microscopy. Furthermore, eight species from five genera are studied with transmission electron microscopy. KEY RESULTS: Our results show that pollen grains of Marcgraviaceae are small (20-35 microm), have three equatorial apertures, granules on the colpus membrane, oblate spheroidal to prolate spheroidal shapes, mainly psilate to perforate ornamentations, and lalongate colpus-shaped thinnings at the inner layer of the exine, and show the presence of orbicules. Based on our fragmentary knowledge of the pollination biology of the family, there are no clear correlations between pollinators and pollen features. CONCLUSIONS: The genus Marcgravia has a high percentage of reticulate sexine patterns and a relatively thin nexine. Sarcopera can be defined by the presence of an oblate spheroidal to even suboblate shape, while Ruyschia and Souroubea typically show prolate spheroidal to subprolate pollen grains. The presence of a thick foot layer in the pollen wall is characteristic of the genera Norantea, Sarcopera and Schwartzia. Pollen features that are taxonomically useful within the family are the shape, sexine sculpturing, and ultrastructure of the pollen wall.     

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 AND SYSTEMATICS OF SIPHONOGLOSSA SENSU LATO (ACANTHACEAE)

Pollen morphology of 15 species of Siphonoglossa and of two closely related groups was investigated. Two tribal-specific pollen types are found within Siphonoglossa sensu lato suggesting that the genus is artificial, composed of taxa belonging in several genera among two tribes (subtribes sensu Bremekamp) of Acanthaceae. Five taxa currently included in an informal subgeneric category of Siphonoglossa have tricolporate, prolate pollen (termed Type I) that is characteristic of Odontonemeae (= Odontoneminae, Justicieae). Pollen of the remaining taxa, belonging in two formal sections of the genus, are mostly 2-porate, bilateral (Type II) with a sexine sculpturing characteristic of Justicieae (= Justiciinae). Pollen of section Siphonoglossa is rather uniform, 2-porate, bilateral with lolongate pores, and seem to delimit a natural group. Taxa of section Pentaloba have a more heterogeneous pollen morphology, mostly 2-porate, bilateral with lalongate pores. Controversial aspects of the interpretation of pollen morphology in Justiciinae are presented and their relevance to this study are examined. Hypothetical trends in the evolution of pollen of Justiciinae are discussed and the application of pollen morphology to taxonomy of the genus is presented, including a recommendation for narrowing the generic concept of Siphonoglossa to the taxa of the type section.     

POLLEN MORPHOLOGY IN THE SELAGINEAE,MANULEAE (SCROPHULARIACEAE), AND SELECTED GLOBULARIACEAE,AND ITS TAXONOMIC SIGNIFICANCE

The pollen grains of 22 genera and 182 species of Selagineae, Manuleae, and selected Globulariaceae were examined by light and scanning electron microscopy of whole and fractured pollen. Grains of the Selagineae and most Manuleae have trizonicolporate aperatures with long, unconstricted ectocolpi; clearly delimited margos; psilate colpus membranes; lalongate transverse endocolpi extending beyond the boundaries of the ectocolpi; thickened endoaperture margins; semitectate, reticulate to microreticulate exines; and psilate, simplicolumellate muri. In the Globulariaceae, Globularia has spinulose, microperforate exines and undifferentiated ectocolpus margins, and both Globularia and Poskea have granular to spinulose colpus membranes, lolongate endoapertures included within the ectocolpi, and unthickened endoaperture margins. These pollen data support postulated relationships between Selagineae and Manuleae, but pollen grains of Selagineae and Globulariaceae are distinct, and pollen morphology does not correlate with proposals combining these taxa in Globulariaceae or Selaginaceae. Pollen morphology in Glumicalyx is indistinguishable from that in most Manuleae and supports transfer of this genus to the Manuleae from the Digitaleae. Differences in pollen morphology supplement established morphological distinctions between Globularia and Poskea, but the pollen evidence does not support published infrageneric classifications in Globularia or postulated relationships between Selagineae and Gratioleae or Selagineae and Lagotis.     

Pollen Morphology of Ribes L. and Its Taxonomic Significance

The pollen morphology of 19 species and 2 varieties in the genus Ribes Linn. was examined under light microscope (LM) and scanning electron microscope (SEM). Pollen grains of the genus are subspheroidal, 15.0-41.67 μm in diameter, more or less circular to obtusely quadrangular in outline with all sides slightly concave, or with alternating concave sides, or elliptic under SEM, 5-, 6-zonocolporate or 5-, 6-pantoporate. Exine usually thin, 1.0-1.5μm, without visible differentiation between nexine and sexine. Exine is psilate or sometimes coarsely or finely granular or verrucate. The most important characteristic of pollen in this genus is the presence of a “rugoid Area” (ectoaperture) around the endoaperture under SEM. It is different from the other taxa of Saxifragaceae (s. 1.). Based on the pollen morphology and the external morphological characteristics, the treatment of the genus Ribes as an independent family (i.e. Grossulariaceae) by many taxonomists is reasonable. The pollen grains of the genus can be divided into 4 types from the materials examined: (1) Grossularia-type. ectoaperture colpate, with two endoapertures, exine psilate. (2) Ribes-type: ectoaperture porous or colpate, with one or two endoapertures, exine mostly psilate. (3) Grossularioides-type: ectoaperture porous, circular, small, with one endoaperture, exine verrucate. (4) Berisia-type: ectoaperture porous, subcircular to circular, large, with one endoaperture exine mostly psilate. The observation made by the present authors supports the subdivision of Ribes into the subgenera Grossularia, Ribes, Grossularioides and Berisia by many taxonomists based on the other morphological characteristics, and the observation by the present authors is in accordance with this view. Pollen information also shows the close relationships among thefour subgenera and therefore they belong to the same genus — Ribes.     

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.     

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.     

Infrageneric variability of pollen morphology in Palaua (Malveae,Malvaceae) and the taxonomic utility of quantitative pollen characters

Palaua (Malvaceae) comprises 15 species endemic to the coastal deserts of Chile and Peru. Previous molecular phylogenetic analyses have shown that this genus is monophyletic and can be subdivided into three clades. In the present study, pollen morphology of all species of Palaua was examined using light and scanning electron microscopy to determine whether it provides additional data in support of the proposed infrageneric groups. The pollen grains are spheroidal, medium to large in diameter, spinose and pantocolporate – the ectocolpi are very short. The tectum is perforate and characterised by granula. In all species the nexine is of a similar thickness in all areas of the pollen grain, while the sexine thickness varies, being notably thicker in the areas where the broad spine bases are sited (‘spine cushions’). Many of the quantitative characters have conspicuous variability, but the variability shows considerable overlap between species. Nonetheless, P. guentheri, P. inconspicua, P. malvifolia and P. modesta are united as a group by having the smallest pollen grains with the smallest apertures, the shortest spines and the shortest interspinal distance. This grouping reflects only partially the suggested infrageneric clades, although it does tend to unite the species with the smallest flowers. A possible link between reproductive resources and pollen size is also considered, as well as the influence of polyploidy. However, the taxonomic utility of these quantitative characters is weakened by species with more than one cytotype.     

Pollen Morphology of Quercus L. in China

The present paper describes the pollen morphology of 31 species of the genus Quercus from China. The pollen grains were all examined with light microscope and scanning electron microscope, and those of some species under transmission electron microscope. Pollen grains of the genus are spheroidal or subspheroidal, 16.8-50.4μm in diameter, 3colporoidate or 3-colpate. The exine is 2-layered, 0.4-1.8μm thick, sexine thicker than nexine, granulated tuberculate or verrucate.     

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.