The Pollen Morphology in Relation to the Taxonomy and Phylogeny of Fagaceae

The present paper deals with the pollen morphology of 103 species belonging to sixgenera-Castanea, Castanopsis, Lithocarpus, Quercus, Fagus and Trigonobalanus in threesubfamilies-Castaneoideae, Quercoideae and Fagoideae. All pollen grains were examinedunder light microscope and scanning electron microscope, and those of some species were ex-amined under transmission electron microscope. The results may be summarised as follows:1. Pollen morphology of Fagaceae, seems to support division of the family into threesubfamilies. Fagoideae, Castaneoideae and Quercoideae.2. Four types of pollen grains are recognized in Fagaceae:1) Fagus-type (representative genus: Fagus): pollen granis are oblate-sphaeroidal,(31.5-39.9) x (35.7 46.2) μm in size, 3(-4)-colporate, peritreme or goniotreme, granulate-ornate under LM., granulate or verrucate under SEM.2) Trigonobalanus-type (T. doichangensis): pollen grains are suboblate-sphaeroidal,(23.1-29.4) ×(25.2-29.4) μm in size, 3-colporate, goniotreme, obscurely granulate-ornateunder LM, densely granulate or verrucate under SEM.3) Quercus-type (Quercus): pollen grains are subspheroidal-subprolate, (21-44.3)× (16.8-39.9) μm in size. 3-colporoidate (-3-colpate), peritreme, crassgranulate or finely-gra-nulate under LM, tuberculate verrucate or spinate under SEM.4) Castanea-type (including Castanea, Castanopsis, Lithocarpus): Pollen grains areprolate-supraprolate, (14.7-23.1)×(10.5-16.8)μm in size; 3-colporate, peritreme, obscu-rely ornate or subpsilate, under LM, rugulose, striate-rugulate or crass-striate under SEM.3. Pollen grains of Cyclobalanopsis age very similar to those of Quercus, and there-fore we support the treatment of Cyclobalanopsis as a subgenus of Quercus.4. On the basis of shape, type of aperture and exine structure, pollen of Trigonobalanusis distinguishable from those of the other genera in Fagaceae and it may be a new type of Fagaceae;5. On the basis of pollen morphology, morphological characters and geological stratification a scheme of phylogeny of Fagaceae is here presented.     

Pollen morphology of the genus Nothofagus and its taxonomic significance

The present paper describes the pollen morphology of 26 species and 2 varieties in Nothofagus from Australia, New Zealand, New Caledonia, New Guinea and South America. Pollen grains were all examined with light microscope (LM), scanning electron microscope ( SEM ) and transmission electron microscope(TEM). A comparative analysis of pollen exine ultrastructure was made for some species. The results are summarised as follows: Pollen grains are oblate to peroblate, 5～8-short-colpate, rarely 4- or 9-colpate; colpi generally thickened at margins; pollen surface spinulose. The exine ultrastructure of Nothofagus differs considerably from that of the other genera in the Fagaceae. The pollen grains of the species examined here show great differences in shape, size, colpal number and characteristics of colpi at margins and could be divided into three distinct types, i.e. N. brassii type; N. menziesii type and N. fusca type.     

A Comparative study on pollen exine ultrastructure of Nothofagus and the other genera of Fagaceae

The genus Nothofagus is mainly distributed in South America and New Zealand. The present paper describes its pollen exine ultrastructure and compares the exine ultrastructure with that of the other genera of Fagaceae. The pollen grains were examined using ultrathin sectioning technique under transmission electron microscope. The study shows that the pollen exine ultrastructure of Nothofagus differs from that of the other genera of Fagaceae by its exine structure and thickness, type of aperture, and ornamentation. The pollen exine of Nothofagus is thin and possesses granular bacules, regular foot layer and tectum, spinulate ornamentation, and the endexine is usually visible at poral area, and 5～8 colpate. The pollen exine of the other genera of Fagaceae possesses entire bacules, irregular foot layer and tectum, granulate and tuberculate ornamentation, thicker endexine, and is 3-colporate ( 3-colpate or 3-colporoidate). The pollen exine ultrastructure of Nothofagus may belong to primitive type. The pollen exine ultrastructure data support Kuprianova’s opinion that Nothofagus should be separated from Fagaceae and established as a monogenetic family, i.e. Nothofa-gaceae.     

Pollen Morphology of Cyclobalanopsis and Its Relation to Quercus

The present paper deals with pollen morphology of 19 species belonging to Cyclobalanopsis and of 31 species belonging to Quercus from China. All pollen grains were examined under light microscope and scanning electron microscope, and those of 4 species of Cyclobalanopsis and of 8 species of Quercus were examined under transmission electron microscope. Pollen grains of Cyclobalanopsis are spheroidal or subspheroidal, 25.2(18.9-31.5) ×23.1 (16.8-27.3) μm in size. Mostly 3-colporoidate, rarely 3-colpate or 3-colporate. The exine is 2-layered, 1.1-1.9 μm thick, sexine thicker than nexine, finely granulate under LM, granulate, spinulate, verrucate under SEM. Pollen grains of Cyclobalanopsis and of evergreen members of Quercus are very similar to each other in size, aperture and ornamentation of exine, but there are some differences in pollen grains between Cyclobalanopsis and deciduous members of Quercus. Therefore, Pollenmorphology supports the treatment of Cyclobalanopsis as a subgenus of Querc     

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.     

Study on the Pollen Morphology of the Chinese Family Oleaceae

This paper deals with the pollen grains of 11 genera, 22 species in Chinese Oleaceae. A comparative study on the pollen grains was carried out by means of scanning electron microscope and light microscope. The pollen grains of the family Oleaceae are almost spheroidal, oblate and prolate, in polar view tlie majority of the pollen grains are tri-lobulate-circular, the minority are tetra-lobulate-circular and dilobulate-circular. Pollen size: (15-65) × (12- 60)μ, 3-colporate and 3-colpate the minority of the pollen grains 4-colpate and 2- colpate colpus usually distinct and the en doaperature vague. The exine oonsists of two layers, the sexine thicker than the nexine, with big reticulum or fine reticulum, the majority of the reticulum are very distinct. On the basis of the pollen size and exine ornamentation, the pollen grains are divided into following two main types: (1) The larger pellen grains and the exine with big reticulum are found in the genera: Jasminum, Nyctanthes, Ligustrum, and Syringa. (2) The smaller pollen grains and the exine with finer reticulum are found in the genera: Osmanthus, Olea, Chionanthus, Fraxinus, Forsythia, Fontanesia and Linociera. In view of the features of pollen morphology, this family is much similar to the family Itamamelidaeeae. For example, the first pollen type resembles the genus Coryiopsis of the family Hamamelidaceae, but there are some differences between them, the former with a bigger reticulate exine and without any colpal membrane, the latter with a finer reticulate apocolpium and eolpate membrane. The second pollen type is much similar to that of the genera Hamamelis and Loropetalum of family Hamamelidaceae.     

Studies on Pollen Morphology of Podocarpaceae from China

The pollen morphology and ultrastructure of exine of Podocarpaceae in China were examined with light microscope, scanning electron microscope and transmission electron microscope. The pollen grains of Podocarpus have rather large and prominent sacs on both sides of body, and are 53.9-74.8 μm long in total, with their bodies 29.6-45.2 μm long and 19.1-31.3 μm wide. The sacs are smooth on outer surface with perforation, but reticulate inside. On distal view, they are obviously of radial muri from its base. The body is oblate or spheroidal, laddershaped on distal face. The exine of the capis tuberculate, but more distinctly on the margin than in the centre. The pollen grains of Dacrydium are of small and indistinct sacs around body, which are composed of many small bladders. The body is subcircular in outline. Both body and sacs are irrugulate tuberculate under SEM. Examination of thin sections of Podocarpus macrophyllus var. maki with TEM reveals that the exine includes ectexine and endexine. It is interesting to note that foot layer of ectexine possesses lamellar stru cture, but endexine is homogeneous in structure and lighter in colour. This character is specific in the gymnosperms. Based on informations of fossil pollen grains, Podocarpaceae is rather primitive and of ancient origin.     

中国壳斗科栎亚科花粉形态研究

本文应用光学显微镜与扫描电子显微镜观察了壳斗科所属栎亚科2属26种2变种的花粉形态,并以本科其它属一些代表种作对比,试图从孢粉学方面为栎亚科的系统分类提供依据。 花粉外壁纹饰在扫描电镜下可分三种类型,即水青冈型(含水青冈属),栎型(含栎属和三棱栎属),栗型(含栗属、锥属和柯属),花粉的类型与植物形态分类的三个亚科一致。 栎亚科的花粉外壁纹饰在扫描电镜下为颗粒聚集成的种种形状。栎属可区分为四种类型,即颗粒状、颗粒——蠕虫状、聚颗粒及芽孢状,其中类型一、二、三为常绿种类,类型四为落叶种类。花粉纹饰在常绿与落叶种类之间有较明显的差异,而青冈亚属和栎亚属之间却没有界线。因此,栎属仍以包含两个亚属为宜。三棱栎属花粉纹饰则介于栎属的类型二与三之间。     

我国睡莲科花粉形态的研究

本文对我国睡莲科5属11种2变种的花粉形态进行了系统的研究。每种花粉都通过光学显微镜和扫描电镜进行观察和照相。同时对莲和白睡莲二种花粉作了超薄切片,对它们的外型结构进行了透射电镜的观察和照相。并结合有关分类学和孢粉学资料,讨论了本科的原始性、异质性及演化上的问题。     

Pollen Morphology of the Family Zingiberaceae in China—Pollen Types and Their Significance in the Taxonomy

Pollen morphology of 89 species and 3 varieties belonging to 18 genera (out of 150 spcies in 20 genera) of Zingiberaceae in China was studied under both light microscope and scanning electron microscope. Pollen grains of Zingiberaceae are spherical, subspherical, ovoid and prolate, 36-225 μm in size, nonaperturate or aperturate (spiraperturate, porate). Pollen grains are almost not resistant to acetolysis. The wall is composed of a very thin exine and a thick intine. The exine is psilate, spinate, cerebelloid-areolate, striate, verrucate and foveolate. According to the presence or the absence of aperture and differential ornamentations, two types and six subtypes are recognized: I. The type Nonaperturate: (85 species and 3 varieties in 18 genera). Four subtypes can be recognized within the type based on the characteristics of the exine sculpture. These are: (1) The subtype Psilate, in which, the exine is nearly smooth (including: Hedychium, Curcuma, Kaempferia, Caulokaempferia coenobilis, Boesenbergia rotunda, Stahlianthus, Amomum compactum, Etingera, Hornstedtis, Rhynchanthus). (2) The subtype Spinate, which comprises two groups: (A) The group Short-spinate, pollen grains with smaller spines (Globba), (B) The group Long-spinate, pollen grains with longer spines (Alpinia, Amomum, Plagiostachys, Roscoea, Cautleya, Boesenbergia fallax, Caulokaempferia yunnanensis). (3) The subtype Cereblloid-areolate, pollen grains of which are spherical or subspherical, with cerebelloid sculpture (Zingiber Sect. Zingiber). (4) The subtype Striate, pollen grains of which are prolate or oliveshaped, and striate (Zingiber Setc. Cryptanthium). II. The type Aperturate, in which pollen grains are acetilysis-resistant and possess distinct apertures (mixed colpate-porate or forate), including two subtypes: (1)The subtype Mixed colpate and Porate. Pollen grains are both 3-colpate and 1-3-porate, and usually with one long spiral, two short (straight or slightly curved) colpi and 1-3-poris. The exine is verrucate or not, nearly sinuolate (Costus speciosus, C. tonkinensis, C. lacerus). (2) The subtype porate, whose grains are 6-8-porate and exine is foveolate (Costus megalobractea). The taxonomic significance of the pollen types in the family Zingiberaceae is also discussed.     

旌节花科及其相关类群花粉形态的研究

对旌节花科及其相关类群五桠果科、猕猴桃科、水东哥科、金莲木科、省沽油科和山茶科等7科19属37种花粉进行了光镜（LM）和扫描电镜（SEM）的观察和比较。其中有些种类是第一次观察和报道。研究表明,上述各科花粉多为近球形至球形,少数近扁球形或近长球形。极面观三裂圆形或近三裂圆形。直径20－30（－35μm）,但厚皮香亚科和省沽油科银鹊树属Tapiscia的花科特别小,仅12.5-20μm。3孔沟,罕有3沟或4孔沟。外萌发孔（沟）相对地较长而宽,内孔多数较发达。花粉外壁纹饰变化大,从模糊而粗糙的细突起到皱波状纹饰,细孔状纹饰,蜂巢状－穴状纹饰,穴－网状纹饰,网状纹饰和颗粒状纹饰等。从形状、大小、萌发孔类型和外壁纹饰的比较看,虽然上述各科花粉均在不同程度上具有一定的相似性,但其中以旌节花科、省沽油科和山茶科中厚皮香科的花粉具有更多的相似性,尤其表现在外壁纹饰特征上。就孢粉学而言,也许旌节花科与省沽油科和山茶科中的厚皮香亚科的关系相对地较为密切。不过,其孢粉学特征所表现出的亲缘关系似乎不如在分子生物学中所表现出的明显。     

刺五加、无梗五加花粉的亚显微形态学研究

用扫描电子显微镜观察了刺五加和无梗五加花粉的亚显微形态特征。花粉粒的赤道面观呈近圆形,极面观呈三裂圆形,具三孔沟。无梗五加、刺五加雄株的花粉表面纹饰为网状;刺五加两性株的花粉的表面纹饰亦呈网状,但有些花粉网脊不连续。无梗五加的花粉粒大小为26.5μm×27.4μm;刺五加雄株的花粉粒大小为21.1μm×22.1μm;刺五加两性株的花粉粒大小约为21.6μm×23.9μm。研究结果表明,无梗五加和刺五加雄株花粉的亚显微形态特征支持传统的属级和种级水平的分类;刺五加两性株某些花粉粒亚显微形态的独特表现可能与其发育不正常有关。     

Studies On the Pollen Morphology of the Genus Iris in China

In this paper, pollen grains of 32 species of the genus Iris in China were examined under light microscope and scanning elrctron microscope. Pollen grains in Iris of China are navicular or subspheroidal. According to the characters of aperture and shape, pollen grains may be divided into four types: (1) Monocolpate (distal): pollen grains navicular or subspheroidal, exine reticulate. (2) Monocolpate-colpoidal: pollen grains subspheroidal, exine pilate. (3) 2-syncolpate: polen grains subspheroidal or navicular, exine reticulate. (4) No aperture: pollen grains subspheroidal; exine verrucate. The evolutional trends of aperture and exine ornamentation are traced and systematic po-sitions of some species are discussed based on characteristics of pollen grains and other organs.     

Pollen Morphology of Plumbaginaceae in China

The present paper describes the pollen morphology of 30 species belonging to 7 genera of Plumbaginaceae from China. The pollen grains were all examined under light microscope, and those of some species under scanning and transmission electron microscope. The pollen grains of the family are subspheroidal, prolate or oblate, (37.5-74.5)× (40.4-81.9)μ in size, 3-colpate, rarely 4-6-colpate, only pancolpate in Ceratostigma willmottianum. The exine 2-layered, 2.0-7.4μ thick, sexine thicker than nexine, verrucate, reticulate or coarsely reticulate. On the basis of the morphology, two types of pollen grains are distinguished in the family: (1) The pollen grains are 3-(rarely 4-6 ) or pancolpate, the exine verrucate. They are found in the tribe Plumbagineae (inculuding the genera Ceratostigma, Plumbago and Plumbagella). (2) The pollen grains are all 3-colpate, the exine reticulate or coarsely reticulate. They are found in the tribe Staticeae (including the genera Acantholimon, Ikonnikovia, Goniolimon and Limonium ).     

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 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.     

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.     

部分中国传统月季花粉形态研究

利用扫描电镜观察了15个中国传统月季品种的花粉形态。结果表明:传统月季花粉为单粒花粉,呈长球形或超长球形（P/E为1.92～2.25）,大小为37.59～51.95 μm×17.02～25.33 μm。赤道面观椭圆形或长矩形,极面观三裂圆形,具三孔沟,沿极轴方向等间距环状分布。外壁纹饰条纹型,覆盖层具穿孔,在品种间具有相似性,但变异丰富,划分为4种类型。聚类分析结果表明,花粉大小和外壁纹饰特征在反映传统月季品种的类别上与形态分类基本一致。根据花粉形态演变规律可推断,样品中‘四面镜’可能为最原始的品种类型,‘月月粉’、‘月月红’及‘匍匐红’等品种较为进化。     

Studies on Pollen Morphology of the Genus Corylopsis

The present paper describes the pollen morphology of 11 species of thc genus Corylopsis (Hamamelidaceae). The pollen grains were all examined with light microscope and scanning electron microscope. Pollen grains of the genus are prolate, spheroidal or subprolate. Three size types are recognized: (l) 18-23 μm, found in Corylopsis omeiensis Yang, C.gotoana Makino, C. himalayana Griff and C.platypetala Rehd. et Wils. (2) 24-32μm, found in C. sinensis Hemsl., C.willmottiae Rehd. et Wils., C.veitchiana Bean, C. brevistyla H.T.Chang, C.multiflora Hance, C.pauciflora Sieb. et Zucc. and C.spicata Sieb. Zucc. (3) 34-39μm, found in C. wilsonii Hemsl. and C. cordata Merrill. Pollen grains are 3-colpate and 4-colpate (only infrequently found in Corylopsis sinensis Hemsl.); the exine is 2-layered, 1.25-2.50μm thick, distinctly reticulate, sexine almost equal to nexine in thickness. From the pollen morphology the genus is considered as a natu-ral group.     

The Pollen Morphology and Exine Ultrastructure of Paeonia L in China

The pollen morphology of 9 species of Paeonia L. has been investigated with both light microscope and scanning electron microscope. In addition, the exine structure of pollen grains of Paeonia suffruticosa and P. lactiflora was examined by transmission electron microscope. Tricolporoidate aperture is an important character of the pollen grains of the Paeonia. The surface of the exine is characterized by reticulate, foveolate and irregularly tuberculate-foveolate sculpture under the SEM. Thin sections of the pollen of this genus shows that the layers of exine are complete i.e. a perforate rectum to semitectum, columellae and foot layers. The endexine is continuous, considerably thickened in the aperture areas and relatively thin or indistinct in the mesocolpia. Paeonia has been placed in Ranunculaceae. But since the beginning of this century many authors have suggested to separating Paeonia from Ranunculaceae. Pollen marphology supports such separation. In Ranunculaceae most pollen grains are tricolpate or have other types of aperture, and exine with spinules and perforations between them. In electron microscopy, the ektexine contains a foot layer, columellae, and perforate rectum, the columellar layer with two types of columellae; the endexine is generally thin. However, the columellar layer of Paeonia has only monomorphic columellae. Some authors considered that there is a close relationship between Paeonia and the Dilleniaceae, but these also differ in the characters of the pollen grains. In Paeonia the constriction of the colpus in equator is in some degree similar to that of Theaceae (Camellia sasanqua Thunb.), Guttiferae (Hypericum L.), Actinidiaceae and Rosaceae. But in the other respects they are quite different. In sum, the pollen morphology of Paeonia is unique. So the palynological information supports Takhtajan's view that Paeonia should be elevated to a family (Paeoniaceae) or order (Paeoniales).