Pollen Morphology of Guttiferae in China

The present paper deals with the pollen morphology of 23 species belonging to 6 genera of Guttiferae occurred in China. All pollen grains have been observed under light microscope, and those of 11 species in 5 genera have been examined with SEM as well. The pollen grains of Guttiferae are subspheroidal, spheroidal or prolate, (17–43) × (15–26)μ in size. Mostly 3-eolporate, less frequently 4-colporate or 5-colporate. The exine is 2-layered, but the occassionally demarcation of these layers is generally indistinct. The exine is 0.8–2.6 μ in thickness. The oramentations of all the pollen grains is generlly finely reticulate under the light microscope. Pollen morphology of the 11 species of 5 genera observed under the SEM is stressed. Their exine oramentations may be classified into four major groups, ie. reticulate, finely reticulate, baculate and perforate. Calophyllum: pollen grains reticulate and rugulo-reticulate. Cratoxylon: pollen grains reticulate, with finely and densely granulate over entire muri. Gareinia: pollen grains baculate, reticulate, and exine with perforate oramentation; in reticulate grains, muri with spinulation. Hypericum: pollen grains reticulate, perforate and finely reticulate. Mesua: pollen grains finely reticulate. From the comparison of pollen morphology, Hypericaceae has much connection with Guttiferae (sensu stricto). Because their pollen grains are all 3-eolporate and the exine is reticulate or oerforate under light mieroseope, but it is different between the pollen grains of Hypericum and those of Guttiferae.     

Pollen Morphology and Its Taxonomic Significance of Caragana Fabr. (Fabaceae) from China

Pollen grains of thirty one species and two varieties of the genus Caragana Fabr. were observed under LM and SEM.The results revealed that the pollen exine ornamentation of the species examined could be divided into two types: perforate and reticulate.The later type consists of two subtypes according to lumina size and muri width. In the first subtype the lumina is round with d iameter equal to muri width,and the exine is perforate in two polar areas,while in the second subtype lumina is irregular or polygonal,with diameter sm aller than muri width.The pollen volume was divided into four types.The genus Caragana is a natural group which was indicated by the general resemblance of pollen morphology. In this genus the pollen morphological classification was not in concordance with the arrangement of the sections and series except Ser.Pygmeae and some species. The pollen variation at infraspecific level was obvious,especially the species distributed in Qinghai Xizang Plateau, such as C.jubata, C.bicolor and C. erinacea . The evolutionary trend of pollen morphology is from perforate to reticulate, which is congruous with the evolution from the pinnate leaf group to the almate leaf group in the genus.     

Pollen morphology of the subtribe Cuspariinae (Rutaceae)

The neotropical subtribe Cuspariinae (Rutaceae) comprises as many as 26 genera and over 125 species. Pollen grains from 111 collections representing 71 species and 24 genera were examined by LM, SEM, and TEM. The pollen morphology of this subtribe is very diverse. Grains are mostly 3–6-aperturate and colporate, rarely porate (Spiranthera) or pantocolporate (Almeidea). Exine sculpturing is most commonly reticulate, sometimes perforate, foveolate-perforate, foveolate, foveolate-reticulate, reticulate, striate-reticulate, echinate, clavate, or baculate. The exine structure is columellate and tectate-perforate, columellate and semitectate, or intectate and is stratified into ektexine and endexine. The exine ofLeptothyrsa is distinctive in that the ektexine of the mesocolpium is longitudinally deeply ridged. The pollen ofHortia, characterized by a psilate exine with rare perforations, a very thick foot-layer, and reduced columellae, is unlike that of any member of the Cuspariinae and offers no support for the transfer of this genus from the Toddalioideae. The pollen data correlate with macromorphological characters and are taxonomically useful.     

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.     

山小橘属 (芸香科) 花粉形态及其系统学意义

利用扫描电镜（SEM）对芸香科山小橘属（Glycosmis）12个分类群16个样品的花粉形态特征（包括花粉大小、萌发孔及外壁纹饰等）进行了观察研究,其中光叶山小橘（G.craibii var. glabra）、长果山小橘（G.erythrocarpa）、锈毛山小橘（G.esquirolii）、长叶山小橘（G.longifolia）、亮叶山小橘（G.lucida）、海南山小橘（G.montana）、少花山小橘（G.oligantha）、小花山小橘（G.parviflora）和华山小橘（G.pseudoracemos）等9个分类群的花粉形态为首次报道。结果表明,国产山小橘属植物的花粉均为单粒花粉,近球形或长球形,花粉大小范围为(15.90~23.92)×(14.56~22.00)μm,极面观为圆形、三裂圆形或近三角形,极轴与赤道轴比值（P/E）范围为1.03~1.31,均为3孔沟;根据花粉外壁纹饰可以将国产种类划分为条网状、复条网状、条纹-穿孔纹饰以及孔穴状纹饰等四种类型,其中小花山小橘不同来源花粉的外壁纹饰差异较大,至少存在5种类型：(1) 极面穴状,赤面条网状或穿孔,(2) 极面穴状,赤面均匀分布圆形穿孔,(3) 具复条网状,(4) 条网状兼具不规则小穿孔,(5) 条网状兼近圆形穿孔纹饰。研究发现山小橘属植物的花粉形态特征具有较高的多样性,为该属植物的分类及系统学研究提供了可用的孢粉学资料。     

Pollen morphology of endemic species of the Horton Plains National Park,Sri Lanka

The pollen morphology of 27 endemic and palaeoecologically-important species belonging to 16 families and 20 genera from the mountain rain forests of Sri Lanka was studied using both light (LM) and scanning electron microscopy (SEM). The pollen grains exhibit a wide range of morphological characters. Their sizes (P and E) are in the range 10-53 2 9-60 mum, and they are distributed among seven shape classes. All the taxa have radially-symmetrical pollen grains except for Impatiens (bilateral). The amb varies from rounded to triangular, with intermediate shapes possible. The pollen grains of all the Lauraceae species studied are inaperturate and microspinose, with a thin, fragile, perforated exine. Sarcococca zeylanica (Buxaceae) has pantoporate pollen grains and a Croton -pattern consisting of faintly striate pegs of various shapes attached or enveloping the smooth rings. Colp(or)ate, syncolpate to parasyncolp(or)ate pollen grains with rugulate to perforate exine patterns occur in Eugenia mabaeoides ssp. mabaeoides and Syzygium ssp. (Myrtaceae). Osbeckia walkeri (Melastomataceae) has heterocolpate pollen with fossulate, foveolate to perforate exine pattern. 3-colporate, microreticulate pollen grains are met with in Euonymus revolutus (Celastraceae), and scabrate-microreticulate ones in Calophyllum walkeri (Clusiaceae). Hedyotis lawsoniae (Rubiaceae) pollen is microreticulate to perforate, while pollen of the Elaeocarpus (Elaeocarpaceae) species is 3-colporate, smooth to scabrate-perforate. Casearia thwaitesii (Flacourtiaceae) has 3-colporoidate pollen grains with a smooth, imperforate to occasionally sparsely perforate exine. Mastixia spp (Cornaceae) have a granular-perforate exine pattern, while Adinandra lasiopetala (Theaceae) has a smooth-perforate exine pattern. Triangular 3 (4)-colporate, and indistinctly faintly microreticulate-perforate pollen was found in Rhamnus arnottianus (Rhamnaceae), rugulate-perforate in Isonandra montana (Sapotaceae) and verrucate-perforate in Symplocos elegans (Symplocaceae). Glochidion coriaceum (Euphorbiaceae) has a 5-colporate grain with a coarsely reticulate exine pattern. Pollen grains of Impatiens walkeri and I. thwaitesii (Balsaminaceae) are bilateral, 4-colpate, reticulate and granular. A pollen key based on light microscope is constructed for all the taxa studied.     

Pollen Characters of Old World Gesneriaceae (Cyrtandroideae)

Pollen of 108 species out of 18 genera (from all tribes of the Old World Gesneriaceae [subf. Cyrtandroideae]) was examined using light, scanning and transmission electron microscopy. The pollen grains are small sized, isopolar and 3-colpate or 3-colporate. In equatorial view they are mostly spheroidal (rarely suboblate or oblate), in polar view circular or subangular. The most variable character is the exine structure and -sculpture. The tectum is perforate, microreticulate, coarsely reticulate or rugulate. Further important characters of the tectum are (a) the presence or absence of conical supratectate sculptural elements, and (b) the width of lumina being either equal or exhibiting different at the apo- and mesocolpium. Ten exine types are distinguished. Some genera and species can be well referred to a special exine type, e.g., Aeschynanthus, Epithema, Stauranthera grandiflora; in other genera several exine types occur, e.g., in the large and heterogeneous genus Didymocarpus. The pollen morphology of the two large subfamilies Cyrtandroideae and Gesnerioideae is compared.     

锦鸡儿属花粉形态及其分类意义

在光学显微镜和扫锚电镜下对锦鸡儿属31种2变种植物花粉形态进行了观察。根据外壁表面纹饰,可分：类型I,表面具小穿孔;类型II, 具网状纹饰。 在类型II中据网眼大小和网脊宽度特征又分为两个亚类型。 根据花粉体积大小,可分4个类型。本属植物花粉形态较为一致,表明是一个自然类群。组、 系的花粉形态特征与一般形态变异不一定相对应,即形态上有明显区别的种类,在花粉形态上未必有很大区别,仅一些种类及矮锦鸡儿系 Ser. Pygmaeae的花粉形态相近。 本属花粉形态在种下等级的变异, 特别是青藏高原分布的种类的变异较为明显,例如 C.bicolor,C. jubata,C. erinacea 与各自的近缘种类。花粉形态为分类处理提供了参考。本属植物花粉形态是从表面具小穿孔向具网状纹饰演化,对应于羽状叶类群向掌状叶类群演化。     

Taxonomy of the Genus Oreocharis (Gesneriaceae)

The genus Oreocharis as circumscribed here consists of 27 species including 5 varieties, of which 5 species and 4 varieties are described as new in the present paper. In the work analysed were the external morphology and geographic distribution and examined under SEM were pollen exine of 22 species and seed coat of 16 species. As a result, three types of the corolla, two types of the anther, three types of the pollen exine and three types of the seed coat are distinguished here in the paper. It is discovered that the corolla in the genus is relatively stable, though diverse, and highly correlated with the characters of pollen grains and seeds. The corolla clearly bilabiate but constricted at the throat, occurring in O. auricula, O. cordatula, O. aurantiaca, etc., for an example, is correlated with smooth, reticulate pollen exine and partial tectum and the reticulate and smooth seed coat. For this reason the subdivision of the genus in the paper is mainly based on the characters of the corolla, but combined with those of the anther, pollen and seed coat. The genus is divided into four sections in the present classification. Dasydesmus Craib, based on a single species. O. bodinieri, is reduced here, and the reasons are given. The genus is distributed mainly in the subtropics, and less frequently in the tropics, of China south of 32.5°N and east of 98.5°E, with only two species beyond the border, O. hirsuta in Thailand (only a single locality in Chiengmai) and O. aurea also found in north Vietnam (see Fig. 1, Table 3). Sect. 1. Stomactin (Clarke) Fritsch. Corolla urceolate-tubular, constricted at the throat, with limb distinctly bilabiate; anthers broad-oblong; seed coat reticulate, smooth, rarely minutely tuberculate; pollen exine fine-reticulate, tectum partial and smooth, luminae slightly unequal in size. Sect. 2. Orthanthera K. Y. Pan Corolla campanulate or campanulate-tubular; anthers broad-oblong; seed coat reticulate, muri smooth, rarely spiny-processed; pollen exine fine-reticulate, with partial and smooth tectum and luminae slightly unequal in size, rarely exine insular and fine-tuberculate, tectum perforate. Setc. 3. Oreocharis Corolla thin-tubular; anthers broad-oblong; seed coat densely spinyprocessed, rarely fine-tuberculate; pollen exine insular, densely spiny-processed, rarely finereticulate and smooth, luminae unequal in size. Sect. 4. Platyanthera K. Y. Pan Corolla campanulate; anthers hippocrepiform; seed coat densely spiny-processed; pollen exine fine-reticulate, tectum perforate, luminae small, nearly equal in size. In the section Stomactin, although the constriction of corolla at its throat is a specialized character, the characters of seed coat, pollen grains and anthers are apparently primitive. Therefore it may be said at least that more primitive characters are preserved in the section. In the section Oreocharis, on the contrary, the characters of corolla, seed coat and pollen exine are all advanced. And in the section Platyanthera, the seed coat, pollen (with perforate tectum) and anthers have developed rather specialized characters.     

Taxonomic implications of pollen exine morphology in infrageneric classification of Scutellaria (Lamiaceae)

Scutellaria L. in Lamiaceae subfamily Scutellarioideae is a subcosmopolitan genus with about 360 species. According to the latest subgeneric classification, two subgenera are recognized: S. subgen. Scutellaria and S. subgen. Apeltanthus. We studied pollen grains from 29 species of Scutellaria belonging to both subgenera and from both the Old and the New World to evaluate their taxonomic importance for the infrageneric classification of the genus. Pollen grains were acetolysed and studied by scanning electron microscope (SEM) and light microscope (LM). All examined pollen grains are isopolar, tricolpate, suboblate, oblate spheroidal, prolate spheroidal to subprolate. The exine is mainly bireticulate perforate, but with variations in lumina shape and size as well as perforation size and uniformity. Based on the exine sculpturing, two subtypes are recognized within the bireticulate pollen type. In subtype I the primary reticulum of the exine consists of angular muri; the lumen has an irregular polygonal shape and the perforations of the secondary reticulum are more or less uniform and of the same size. This pollen subtype was observed in species of subgenus Scutellaria, sectt. Scutellaria p.p., Anaspis, Perilomia and Salazaria. In subtype II the primary reticulum of the exine consists of a more or less curved murus; the lumen is rounded or obtusely angular and the secondary reticulum consists of one or more large central perforations surrounded by some smaller ones. This pollen subtype was observed in species of subgenus Scutellaria sectt. Scutellaria p.p., Salviifoliae and in all members of subgenus Apeltanthus. The results of this study reveal that the exine ornamentation is a diagnostic character useful for the infrageneric classification of Scutellaria.     

绣球属(Hydrangea L.)及近缘属花粉形态的研究

为深入研究绣球属植物花粉形态的分类学价值和系统学意义,厘清绣球属与近缘属之间的系统发育关系,该文利用扫描电子显微镜(SEM,scanning electron microscope)对国产绣球属及其近缘属41种绣球花科(Hydrangeaceae)植物的花粉形态以及表面纹饰进行了观察。结果表明:绣球属及其近缘属的花粉为三孔沟;形状多数为长圆体形或近球体形;赤道面观为椭圆形或圆形;极面观多为圆形,少数为三角形或圆三角形。花粉外壁纹饰可分为网状和孔穴状。网眼内的三级纹饰可分为光滑和具颗粒状突起。根据花粉形状和外壁纹饰类型将上述物种划分为4个组,即花粉的形状为长圆体形,表面纹饰为孔穴纹饰;花粉的形状为长圆体形,表面纹饰为网状纹饰;花粉的形状为近球体形,表面纹饰为孔穴纹饰;花粉的形状为近球体形,表面纹饰为网状纹饰。以上可进一步细分为8个类型。上述表明花粉形态证据可为绣球属及其近缘属的属下分类和种间界定提供重要佐证;但结合前人的系统发育重建分析该属植物花粉形态的系统学意义相对有限,如花粉形态证据对于该属及其近缘种属系统发育树上大支的界定难以提供有力的证据。     

The palynology of the genus Rondeletia L. (Rubiaceae-Cinchonoideae-Rondeletieae)

Pollen of 20 species of Rondeletia, covering three subgenera and seven sections or “species groups” of the genus, was studied using light and scanning microscopy. Rondeletia, characterized by binucleate mature pollen grains like numerous other Rubiaceae, was found to be eurypalynous. The pollen is isopolar and small-sized (average equatorial diameters 11.4–22.8 μm) and 3-colporate (rarely 4-colporate). In equatorial view, the pollen grains are mostly ± spheroidal (rarely suboblate); in polar view (ambitus), they are mainly circular or semiangular (intersemiangular in one species only). Three different exine types are distinguishable: exine with 1) tectum minutely perforate, perforations usually isodiametric, 2) tectum with comparatively large, not isodiametric perforations or 3) coarsely reticulate; supratectal sculpturing elements are lacking; the width of the perforations and lumina is either equal or different at the apo- and mesocolpia; in a few species rod-like luminal processes are present. Palynological features of most of the investigated taxa tend to overlap so that palynology is not very helpful in supporting the subgeneric division of Rondeletia. It was, however, noted that pollen morphology, to some extent, supports the sections of subgen. Arachnothrix. Palynologically, the subgen. Rogiera was found to be heterogeneous in that each of the three investigated species is characterized by a different exine type; one of these, R. strigosa, differs from all other Rondeletia species investigated in having a coarsely reticulate exine. R. odorata (subgen. Rondeletia) differs from all other investigated species in its intersemiangular pollen shape.     

Pollen morphology of Stachys (Lamiaceae) in Iran and its systematic implication

Pollen grains of 30 taxa of the genus Stachys (29 spp. and one subsp.), representing 9 of the currently recognized sections and 1 species of the closely related genus Sideritis (Si. montana) distributed in Iran were examined by light and scanning electron microscopy. Twenty-eight taxa are studied for the first time under aspects of pollen morphology. The basic shape of the pollen grains in most taxa studied is prolate-spheroidal, but subprolate, spheroidal and oblate-spheroidal pollen grains can also be found in few species. The grains are usually tricolpate (the amb triangular), but also tetracolpate (the amb circular to more or less square) in some species (S. iberica, S. atherocalyx and Si. montana). The surface is microreticulate (the frequent type), reticulate, perforate, foveolate-psilate or foveolate. The lumina are separated by smooth or sinuate muri which make them polygonal, more or less rounded and elongate. Major pollen morphological features of the taxa studied are compared and discussed on the basis of taxonomical concepts. In some cases, these characters are useful in delimitation of formerly introduced sections while they mostly provide further characters in separating related species from each other. For example, all members of S. sect. Aucheriana are characterized by elongated lumina. Based on the oblate-spheroidal shape of its pollen as well as tetracolpate aperture type, the results of the present study confirm sect. Pontostachys as including S. angustifolia, S. iberica, S. sparsipilosa as well as S. atherocalyx. Our results also suggest that although some species like S. fruticolosa and S. lavandulifolia are morphologically well characterized, they cannot be separated from other species of Stachys based on pollen morphology.     

Pollen morphology of Diosma and Coleonema (Rutaceae: Diosminae) and its taxonomic implications

The pollen morphology of Diosma and Coleonema of the taxonomically problematic subtribe Diosminae (Rutaceae) was investigated to determine its taxonomic relevance. Pollen of 26 of the 28 species of Diosma and of each of the eight species of Coleonema was investigated by LM, SEM and TEM. In pollen of Diosma the exine sculpturing is uniform, and only one pollen type, reticulate, could be distinguished. In contrast, exine sculpturing shows interspecific variation in Coleonema and three distinct types, striate, reticulate and perforate, could be discerned. Species groupings indicated by the pollen types suggest possible relationships between species within Coleonema and with other genera that were not previously apparent. Pollen data support macromorphological evidence that suggests the generic re-classification of the Diosminae.     

国产省藤属植物的花粉形态学

对国产棕榈科省藤属(Calamius L.)15种植物的花粉进行了光学和扫描电镜观察,其中12种为首次报道。省藤属的花粉均为两沟型花粉,外壁覆盖层多为网状纹饰(大喙省藤C．macrorrhynchus)或具穿孔(华南省藤C.rhabdocladus)。首次发现省藤属花粉的外壁纹饰存在穿孔和外壁疣状突起的类型(阔叶鸡藤C.pulchellus)以及皱波状突起的类型(长鞭省藤C.flagellum)。花粉的大小、形状、外壁纹饰、外壁是否有突起,外壁厚度和网状纹饰网眼的大小,对于省藤属的种级分类有较大的意义。     

Pollen morphology of Chinese Glochidion (Phyllanthaceae) and its taxonomic implications

The pollen morphology of twenty‐five Chinese species belonging to two sections of Glochidion and one species from Phyllanthus subgen. Phyllanthodendron was examined using scanning electron microscopy (SEM). The results showed that pollen grains of Glochidion are isopolar, spheroidal to rarely subprolate in shape, (3)4–5(6)‐colporate in aperture pattern, and tectate‐reticulate to tectate‐rugulate in exine ornamentation. Fusion of colpus ends was observed in some pollen grains of G. pseudo‐obscurum and G. wrightii. The pollen grains of Phyllanthus dunnianus are similar to those of Glochidion species in their spheroidal shape and four‐colporate aperture pattern, but are slightly larger in size, and show tectate‐perforate exine ornamentation. In Glochidion, pollen morphology appears to be useful for distinguishing several macromorphologically similar. Additionally, palynological characters provide evidence for the taxonomy of the supergenus Phyllanthus s.l.     

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.     

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.