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

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

POLLEN MORPHOLOGY IN THE GENUS MIMULUS (SCROPHULARIACEAE) AND ITS TAXONOMIC SIGNIFICANCE

The pollen morphology of 117 species and varieties of Mimulus was examined by light and scanning electron microscopy. Five major and 8 more tentative, minor types were found based on the differential correlation of aperture type, exine morphology, pollen grain diameter and other characters: type 1—synaperturate, usually ±spiraperturate, exine perforate to microreticulate with supratectal processes; type II—trocolporate, exine microreticulate (IIa and IIb, supratectal processes absent; IIa, mean polar axis 16–19 μm; IIb, mean polar axis 25–35 μrn; IIc, supratectal processes present); type III—tricolpate, colpus membrane ±psilate. exine with supratectal processes (IIIa, exine microreticulate and 1.4–2.0 μm thick, polar axis ≥ 30 μm; 111b, exine densely perforate and 2.2–2.8 μm thick, polar axis ≤ 23 μm); type IV—tricolpate, colpus covered with spinulose granules (operculate), exine microreticulate with supratectal processes; type V—5–7 stephanocolpate (Va and Vb, colpus margins ±straight and nongranular; Va, exine microreticulate with supratectal spinules; Vb, exine perforate with supratectal spinules or spinulose verrucae; Vc, colpus margins ragged and granular, exine microreticulate with supratectal processes). The pollen data correlate well with geographical and macromorphological data and, where the latter are ambiguous, often provide important clues toward the resolution of conflicting interpretations of infrageneric classification and generic delimitation.     

POLLEN MORPHOLOGY IN THE GENERA MONTTEA AND MELOSPERMA (SCROPHULARIACEAE)

Pollen of the scrophulariaceous genera Melosperma and Monttea was studied with light and scanning electron microscopy. Both have single, radiosymmetric, isopolar, trizoniaperturate pollen grains with usually ruptured colpus membranes, tectate-perforate to semitectate exines with a columellate interstitium and pitted or perforate nexines. Intergeneric differences in pollen morphology are established based on the differential correlation of sculpturing characters. Comparisons of the pollen of Melosperma and Monttea with that of the tribe Minuleae sensu stricto reveals no difference in the case of Melosperma and the existence of intermediate morphologies and “reticulate” similarities in the case of Monttea; the taxonomic significance of this morphological intergradation is considered. The transfer of Oxycladus aphyllus to Monttea is consistent with the pollen evidence, but proposed affinities between Monttea and tribe Antonieae of the Loganiaceae are not supported.     

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.     

Pollen Morphology of the Subtribe Hyoscyaminae (Solanaceae)

The pollen morphology of 12 species representing 6 genera was examined under the light; scanning and transmission electron microscope. According to form of aperture; sculpture and structure of exine; the palynological relationships among the 6 genera are discussed. 1. Scopolia Jacq. (plate 1:1-4) Pollen grains subspheroidal or irregular; 43.7-49.6 μ in diameter; nonaperturate; Exine about 1.2-1.5μ thick; verrucate; with verrucae unequal in size; spinulose or granular. Stratification indistinct; with no columella layer visible under TEM. 2. Anisodus Link et Otto (plate 1: 5-12) Pollen grains subspheroidal; 43.4-48.7 μ in diameter; nonaperturate. Exine 1.5-1.8 μ thick; verrucate; with verrucae unequal in size; spinulose or smooth. Stratification indistinct under TEM. 3. Atropanthe Pascher (plate 2: 1-4) Pollen grains subspheroidai or spheroidal; triangular-circular in polar view; oblong in equatorial view; 47.2-49.3 μ ×39.8-47 μ; 3-colpate; colpi wide or narrow; colpus membranes finely granular. Exine with irregular short-stick sculpture; 2.3μ thick; distinctly 2-layered; columella layer clearly visible; sexine thicker than nexine under TEM. 4. Przewalskia Maxim. (plate 2: 5-8) Pollen grains subspheroidal; triangular-circular in polar view; oblong in equatorial view; 39.2-40.3 μ × 36.4-42.8 μ ; tricolporate; ora lalongate; colpus membranes granular. Exine reticulate; 2-layered; 2u thick; columella layer distinct; sexine and nexine equal in thickness under TEM. 5. Physochlaina G. Don (plate 2: 9-12; plate 3: 1-12) Pollen grains polymorphic; usually subspheroidal; 36.5-39.4μ in diameter; aperture varying from nonaperturate (P. macrophylla); 3-4 rugose (P. physaloides) to 3-colporate (P. praealta). Exine 1.5-2μ thick; reticulate; smooth or with spinulose or verrucous processes under SEM; distinctly 2-layered; or without stratification; columella layer differentiated or not. 6. Hyoscyamus Linn. (plate 4: 1-9) Pollen grains subspheroidal; circular-triangular in polar view; oval in equatorial view; 39.9-41.8 μ × 38.7-4.2 μ; tricolporate. Ora orbicular; colpus membranes finely granular or with striate processes. Exine reticulate or striate-rugulate; 2-2.3μ thick; distinctly 2-layered; columella layer developed. The dendrogram (Fig. 1) indicates probable phylogenetic relationships among the sixgenera in the subtribe Hyoscyaminae.     

A Note on Remarkable Exines in Acalyphoideae (Euphorbiaceae)

As part of an extensive study of pollen of Euphorbiaceae that combines transmission electron microscopy with scanning electron microscopy, distinctive exines are reported and documented for certain Acalyphoideae. Cheilosa and Neoscortechinia, which comprise the tribe Cheiloseae, are the only Acalyphoideae with an echinate tectum, but their apertures and exine structure do not support a relationship to Oldfieldioideae. In Ditaxis, one of the three mesocolpia is much smaller than the other two and the pollen can be easily distinguished from all other Euphorbiaceae. In SEM, the tectum of Pycnocoma appears almost complete, but in TEM the exine consists of irregular, mostly discrete tectal elements that narrow to points (=columellae) at the interface with the threadlike footlayer. The operculate grains of Alchornea and Boquillonia have exines with a poorly differentiated double layer of columellae in the mesocolpium, but nearer the endoaperture the lower tier becomes greatly elongated and appears to rest directly on the endexine. Plukenetia polyadenia has a complete tectum and a thick exine with a predominant infratectum of large, occasionally branched columellae that nearer the tectum are combined with densely spaced granules. Plukenetia penninervia has a reticulate tectum of crenate muri and short, sparse columellae. The pollen diversity in Acalyphoideae suggests that the subfamily, many tribes and even genera may not be monophyletic.     

COMPARATIVE POLLEN MORPHOLOGY AND TAXONOMIC AFFINITIES IN CYCADALES

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

POLLEN MORPHOLOGY AND DETAILED STRUCTURE OF FAMILY COMPOSITAE,TRIBE CICHORIEAE. II. SUBTRIBE MICROSERIDINAE

A survey of pollen morphology of 40 species representing eight genera of the primarily North American subtribe Microseridinae reveals seven of the eight genera to have caveate, echinolophate, tricolporate grains, Picrosia being the only taxon with echinate pollen. Sectioned grains reveal the exine to consist of an ektexine and endexine. The ektexine, composed of spines, columellae, and foot layer appears to be of two basic types, one with six or seven levels of horizontally anastomosing columellae which are reduced to a single columellar layer under the paraporal lacunae and the second, a bistratified ektexine not reduced to a single layer below the paraporal lacunae. Sectioned exines of Pyrrhopappus are unusual, having very large columellae fused to the foot layer below ridges and highly reduced columellae under lacunae. Endexine organization is similar in most of the genera. Exceptions to this are Pyrrhopappus and some species of Agoseris, which have an “endexine 2” layer. Subtribe Microseridinae is essentially stenopalynous. The pollen data support most of the relationships suggested by Stebbins in his classification. The genera Agoseris, Microseris, Nothocalais, and Phalacroseris seem to form a natural group while Krigia and Pyrrhopappus form another cohesive series. The position of Picrosia, as an advanced offshoot of Pyrrhopappus, is not supported by the pollen data.     

Selagineae and Manuleeae, two tribes or one? Phylogenetic studies in the Scrophulariaceae

 Based on results from a cladistic analysis of molecular characters using parsimony and Bayesian inference, the tribe Selagineae (Scrophulariaceae, sometimes Selaginaceae or Globulariaceae), is found to have arisen within Manuleeae (Scrophulariaceae). The inclusion of Selagineae into Manuleeae is therefore proposed. The result is corroborated by morphological characters, and indicates that the typical Selagineae condition with one ovule per ovary locule has arisen several times within the Manuleeae. Two former Manuleeae taxa are excluded from Manuleeae; Antherothamnus and Camptoloma. The former shows affinity with Scrophularia and Verbascum, and the latter groups with Buddleja and related genera. The controversy between parsimony and likelihood based methods are briefly discussed as is the possibility of historical bias in prior studies. Received February 9, 2001 Accepted June 1, 2001     

A PALYNOLOGICAL INVESTIGATION OF THE GENUS TOURNEFORTIA (BORAGINACEAE)

The pollen morphology of 65 species of Tournefortia L. was examined by light, scanning, and transmission electron microscopy. Four distinct morphological types were found: Type I, grains subprolate with 3-colporate apertures alternating with three pseudocolpi and the ektexine psilate; Type II, grains subprolate, the poles expanded, 3-colporate with the ektexine psilate polarly and verrucate equatorially; Type III, grains spheroidal with three porate apertures and the ektexine clavate; Type IV, grains suboblate with 3–4 colporate apertures and the ektexine psilate to finely rugose. Type I was found in 46 species representing the most diverse group of tournefortias. Type II was found in 12 species from the Old and New Worlds representing two related groups. Type III was found in five closely related species from the American tropics. Type IV was found in two species from South America, but differences in both the pollen and sporophyte indicate that this grouping may be artificial. Pollen morphology does not support the separation of those species treated as Messerschmidia and confirms the close relationship of Tournefortia and Heliotropium. The striking morphological divergence of Tournefortia pollen raises the question of its adaptive significance.     

Pollen morphology of some <Emphasis Type="Italic">Paronychia</Emphasis> species (Caryophyllaceae) from Turkey

The Pollen morphology of 13 taxa 11 of which are endemics belonging to Paronychia Miller (Caryophyllaceae) viz., P. agryloba, P. angorensis, P. arabica subsp. euphratica, P. carica, P. cataonica, P. chinonea, P. condensata, P. davisii, P. dudleyi, P. galatica, P. kurdica subsp. kurdica, P. kurdica subsp. montis-munzur and P. mughlaei from Turkey has been investigated by light (LM) and scanning electron microscopy (SEM). LM observations show that pollen grains are usually radially symetrical, isopolar, pantoporate, polygonal (6-gonal) or polygonal-spheroidal. Tectum is psilate or punctate. Tectal surface sparsely-densely spinulose. The numbers of pores are between 6 and 12. On the basis of pollen sizes, P. davisii was the biggest pollen type (23.45 μm) and P. kurdica subsp. kurdica (16.2 μm) was the smallest pollen types. According to exine sculpturing, pollen size and spinule numbers per 1 μm², three pollen types were distinguished.     

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 morphology of some selected plant species from Horton Plains,Sri Lanka

The pollen morphology of four species: Gaultheria leschenaultii (Ericaceae), Psychotria zeylanica (Rubiaceae), Rubus leucocarpus (Rosaceae) and Symplocos bractealis (Symplocaceae) from Horton Plains, Sri Lanka was investigated using LM and SEM. Psychotria zeylanica is endemic to Sri Lanka and the other three taxa are common in Horton Plains. Gaultheria leschenaultii has tetrads with 3-colporate, coarsely granular to granulo-rugulate pollen grains; in the polar region and at the colpus margin the exine is smooth with micro-rugulate structure. Psychotria zeylanica has 3- (or 4-) colporate, reticulate-spinulose to granular pollen grains. Rubus leucocarpus has 3-colporate, striate-perforate pollen grains and Symplocos bractealis has 3- (or 4-) colporate pollen grains with a verrucate-perforate exine. Pollen grains of these four species are very indicative of human impact and palaeoclimate.     

POLLEN MORPHOLOGY OF THE DILLENIACEAE AND ACTINIDIACEAE

Pollen of 53 species of Dilleniaceae and Actinidiaceae was examined by light microscopy, scanning electron microscopy, and a selected group in transmission electron microscopy. Dilleniaceae pollen ranges from tricolpate, tricolporate, tetracolpate, and incipiently inaperturate. Tricolpate types occur only among the Old World subfamily Dillenioideae and the compound aperturate (3-colporate) condition is restricted to the subfamily Tetraceroideae. Within the Dilleniaceae the tricolpate pollen type with elongated apertures is considered primitive, having given rise to the 3-colporate and 4-colpate conditions. The striking pollen dimorphism in the Neotropical species of Tetracera, all of which are androdioecious, is documented; however, in contrast to previous reports, pollen from bisexual flowers appears to be incipiently inaperturate and not pantoporate. The inaperturate condition is interpretated as an early stage in the evolution of outcrossing. Pollen morphology does not support a close relationship between Dilleniaceae and Actinidiaceae. Pollen morphological differences that can be noted between these families are: tectum complete and predominantly psilate or psilate-granular in Actinidiaceae, tectum incomplete, punctate to reticulate in Dilleniaceae; an equatorial bridge of ektexine over the endoaperture usually present in Actinidiaceae, absent in Dilleniaceae; columellae reduced in Actinidiaceae, columellae usually well-developed in Dilleniaceae. Pollen morphology does not argue against a close relationship between Actinidiaceae and Theaceae.     

Pollen Morphology in the Order Centrospermae

Pollen of 190 species from 16 families has been examined by light and scanning electron microscopy. Three basic pollen types were found: 3-colpate, pantoporate, and pantocolpate, all with a spinulose and tubuliferous/punctate ektexine. The palynological data emphasize the close relationship of the betalain families, and support their association with the anthocyanin families. Caryophyllaceae and Molluginaceae. Pollen morphology reinforces the placement of the Cactaceae and Didiereaceae in the order, and would support the inclusion of Dysphania in the Chenopodiaceae. The pollen grains of the Achatocarpaceae, Bataceae, Gyrostemonaceae and Theligonaceae, none of which had the spinulose and tubuliferous/punctate ektexine, had no counterparts in the remaining taxa examined. Thus, the evidence from palynology does not support their inclusion in the Centrospermae and indicates that their placement should be re-examined.     

Pollen morphology of some taxa of Vicia L. subgenus Vicilla (Schur) Rouy (Fabaceae) from Turkey

The pollen morphology of the ten taxa belonging to Cassubicae Radzhi, Cracca Dumort., Ervilia (Link) Koch, Lentopsis Kupicha, Trigonellopsis Rech. f. and Variegatae Radzhi sections of the genus Vicia L. subgenus Vicilla (Schur) Rouy (Fabeae, Fabaceae) in Turkey has been examined by use of light microscopy and scanning electron microscopy. Twelve morphometric characters are analysed with one-way analysis of variance and Tukey’s honestly significant difference test for multiple comparisons. Polar axis and equatorial diameter of pollen grains range from 24.64 to 44.19 and from 15.90 to 30.42 μm, respectively. The pollen grains are prolate–spheroidal to perprolate (P/E = 1.09–2.07), but the prolate shape occurs in the majority of the taxa. The regular pollen grains of all taxa are trizonocolporate, isopolar, and released in monads. Ornamentation of the mesocolpium is psilate–perforate in V. tenuifolia subsp. dalmatica and V. tenuifolia subsp. tenuifolia (sect. Cracca), reticulate–rugulate in V. villosa subsp. villosa (sect. Cracca), and reticulate–perforate in the remaining taxa. The apocolpium and colpus area are reticulate–perforate in V. hirsuta (sect. Cracca), V. ervilia (sect. Ervilia) and V. lunata subsp. grandiflora (sect. Trigonellopsis), and psilate, psilate–slightly perforate or perforate in the other taxa. The results obtained from analyses of pollen grains from the studied taxa of the subgenus Vicilla reveal significant differences in general morphology and some morphometric attributes, and ornamentation types. However, it is concluded that pollen morphology has proved to be uninformative for differentiating the subgenera Vicilla and Vicia.     

Intraspecific variation in the pollen exine sculpture of Jaborosa runcinata Lam. (Solanaceae)

The variability of ectexine sculpture in the pollen of Jaborosa runcinata (Solanaceae) was studied with light and scanning electron microscopy. Three main ectexine variants are recognized: incomplete reticulate, a combination of incomplete reticulate and gemmate‐granular, and gemmate‐granular. A range of transient conditions were also noted between the two extremes. Apertures vary from a basically triporate‐aspidate arrangement to pollen grains which appear to have a rudimentary colpus. The sculptural variability of the ectexine, and a tendency towards a colpus, in J. runcinata are interesting as intraspecific variation does not appear to be common within Solanaceae.     

Pollen morphology in the ericoid clade of the order Ericales,with special emphasis on Cyrillaceae

Pollen morphology and pollen wall structure in 23 species representing all families of the ericoid clade of order Ericales, i.e. Actinidiaceae, Sarraceniaceae, Roridulaceae, Clethraceae, Cyrillaceae, and Ericaceae, was investigated by means of light microscopy and SEM. The pollen wall of all taxa was found to consist of two solid strata, a tectum and foot layer, separated by a more or less narrow infratectum with granular elements. The three genera of Cyrillaceae were found to differ from each other. In Cyrilla and Cliftonia the pollen is psilate, often more than 20 mum in diameter, spheroidal with an indistinct colpus rim; in contrast, pollen of Purdiaea are rugulate and slightly different between species, smaller and often less than 20 mum in diameter, oblate with a projecting rim reaching over the colpus. In several characteristic features, pollen of Purdiaea is more similar to pollen of Clethra (Clethraceae), than to pollen of Cyrilla and Cliftonia .     

中国槲寄生属花粉形态的研究

首次系统地报道中国槲寄生属10种植物花粉形态。通过光学显微镜和扫描电镜观察,国产槲寄生属种类可按花粉的外壁纹饰分为两大类群,与形态上划分相一致。类群Ⅰ(8种)具不显著或近光滑的外壁纹饰,这一类型又可分为2个亚型:纹饰在花粉各个部位基本上是一致的和纹饰在赤道和极、沟两侧不一致的;类群Ⅱ(2种)外壁具显著的杆状纹饰。但花粉性状对形态上相近的种类鉴定意义不大。还讨论了槲寄生属花粉的可能的演化趋势。