物参属花粉形态及其分类学意义

首次报道了兰科的手参属Ｇｙｍｎａｄｅｎｉａ国产５种植物花粉形态的研究结果,并与邻近的兜被兰属Ｎｅｏｔｔｉａｎｔｈｅ花粉进行了对比研究。前者花上块形状大多不规则,少数为三棱体形;而后者花粉小块大多形状为三棱锥体形,少数形状不规则。手参属花粉外壁表面具很细微的小穿孔、皱波状,小穴和小穿孔与沟渠状纹饰共存,或沟渠状等。而兜被兰属外壁表面纹饰明显,具有５种类型：（１）小穿孔,（２）表面近光滑,（３）沟渠状     

Pollen Morphology of Neottianthe (Orchidaceae) and Its Taxonomic Significance

The genus Neottianthe (Orchidaceae) consists of about 12 species in the world, of which all are found in China and nine are endemic to China. The pollen morphology of this genus is very specific in shape and composition. All pollen grains in Neottianthe are of a compound pollen grain, called massula, in which of many monad pollen grains adhere togeth er. A massula is usually long and narrow trigonous cone in shape. Pollen grains can be divid ed into five types according to ornamentation, of exine surface: (A) pollen surface almost smooth, or very obscurely fossete, (B)perforate, (C) fossulate or foveolate, (D) reticuloid, and (E)typically reticulate. Neottianthe. calcicola, N. monophylla, N.pseudo-diphylax and N. cucullata have been merged into one species, N. cucullata, by some taxonomists. This treatment is not supported by the pollen morphological data, because pollen morphological characteristics of above-mentioned four species are different from each other. Therefore, pollen morphology shows that they are all independent at the specific level. Finally, it must be stressed that N. camptoceras is uniquite in the genus Neottianthe for its pollen shape,size and surface ornamentation of pollen exine.     

中国兜被兰属植物的花粉形态及其分类意义

兜被兰属Neottianthe全世界共约12种,中国均有分布,其中9种为我国所特有。本属花粉形态 和组成都很特殊,它是由许多单粒花粉粘结在一起形成复合花粉——花粉小块,其形状一般为狭长的三 棱锥体形。外壁表面具5种纹饰类型:(A)近光滑,或具十分微弱的小凹陷;(B)外壁具穿孔状纹饰; (c)外壁具沟渠状纹饰;(D)外壁具拟网状纹饰;(E)具典型的网状纹饰。奉属有些种类,例如:N．cal- cicola,N．monophylla,N．pseudo-diphylax和N．cucullata等4种在分类上还存在许多问题,有些分类 学家把N.calcicola,N.monophylla,N.pseudo-diphylax 3种均归并于N．cucullata。孢粉学资料表 明上述4个种花粉特征完全不同,因此,上述4种仍应各自独立为种。最后,强调指出N．camptoceras的花粉形状和外壁纹饰等特征在本属为特殊类型。     

Pollen morphology of some species of Vernonia s. l. (Vernonieae,Asteraceae) from Argentina and Paraguay

Pollen morphology is widely used in taxonomic treatments of tribe Vernonieae, and differences in exine structure and aperture form have been described for many species within the genera comprising the tribe. However, in recent years a number of new species have been described and, in the present paper, we describe in detail the pollen of 24 species of the tribe which are either endemic or uncommon to certain regions of Argentina or of Paraguay. The pollen of these species are radially symmetrical and more or less spheroidal, 3‐colporate or, in one species, 3‐porate; the exine may be tectate or semitectate and microperforate, with echinate lophae or, in the 3‐porate example, psilate lophae. Using a range of characteristics related to size, shape, wall thickness, apertures and tectum surface morphology, five of the six pollen types previously described for tribe Vernonieae, and a subtype, are recognized. Our results support the usefulness of pollen morphology in helping to determine the taxonomic position of species within tribe Vernonieae.     

Pollen Morphology and Epidermal Characters of Leaves in Convallarieae (s. l.)

The tribe Convallarieae (sensu Krause 1930) consists of 7 genera, i.e. Convallaria, Speirantha, Reineckia, Theropogon, Tupistra, Rohdea and Aspidistra, but now generally recognized as two tribes, Convallarieae (the former 4 genera) and Aspidistreae (the rest). Observed in this work were pollen morphology of 17 species and epidermal characters of leaves of 12 species. All the 7 genera are covered in observations. Pollen grains in Convallarieae (s. str.) are all monosulcate and boat-shaped (Plate 1: A-F). The exine is rather uniformly microperforate (Plate 1: A-F); only Theropogon is exceptional in this respect: it has rugulate exine (Plate 1: O, P). Tang and Zhang (1985) have pointed out the heterogeneity of Theropogon in this tribe. Pollen morphology in the tribe Aspidistreae is widely variable. The genera Tupistra and Rohdea were shown to have monosulcate and boat-shaped pollen grains. Their exine is perforate or reticulate (Plate 1: G-N). Pollen grains in the genus Aspidistra, however, are nonaperturate and spheroidal. The exine in the genus varies from crass-rugulate, variously gemmate to tuberculate-baculate (Plate 2; A-H). The pollen morphology of Aspidistra is therefore distinctly different from that of Tupistra and Rohdea, which supports the Nakai's (1936) establishment of the tribe Rohdeae for Tupistra and Rohdea. Therefore, Krause's Convallarieae is reasonably divided into at least three tribes, Convallarieae (Speirantha, Convallaria, Reineckia and Theropogon), Aspidistreae (Aspidistra) and Rohdeae (Rohdea and Tupistra). The pollen characters of all the 7 genera are shown in Table 1. The evolutionary trends of pollen morphology (aperture and exine) in the three tribes are discussed and our major view-points are shown in Fig. 1. Observations on epidermal characters of leaves show that in the Convallarieae (s. 1.) stomatal apparatuses are all anomocytic; cuticular layer on the upper epidermis is mainly striatethickened or rather uniformly thickened (Plate 2: J--P; Plate 3: A-C, F-N), whereas in the genus Convallaria the cuticular layer is squamosely thickened (Plate 2: I; Plate 3: D, E).The epidermal characters of leaves in the 7 genera are summarized in Table 2.     

9种益母草属植物的花粉粒形态

利用光学显微镜、扫描电镜对国产８种１变种及欧美产１种益母草属植物的花粉粒进行观察与研究。本属植物花粉粒椭 球形或近球形,益母草和白花益母草花粉粒两端突起成喙。具３沟,狭长,几达两极,人内陷或内陷。外壁纹纹饰网状,某些种尚具或为细小的穿孔。结果基本支持各种的系统地位。     

中国紫草科厚壳树亚科的花粉形态及其系统学意义

为探讨厚壳树亚科在紫草科中的系统位置,用扫描电子显微镜研究了国产厚壳树亚科Ehretioideae的厚壳树属Ehretia、轮冠木属Rotula和基及树属 Carmona 3属12种植物的花粉形态,其扫描电镜资料属首次报道。从本文并结合文献报道的双柱紫草属 Coldenia的花粉资料,可得知该亚科花粉为长球形 、近球形,稀球形,赤道部位具有不同程度的膨大;萌发孔类型为三孔沟和三拟假沟二者交替排列,稀具三孔;外壁表面具微弱的小穿孔,稀光滑或皱波状纹饰。厚壳树亚科Ehretioideae与破布木亚科     

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.     

Pollen Morphology of the Genus Malus and Its Taxonomic and Evolutionary Significance

The pollen morphology of 26 species and 5 hybrids of the genus Malus was investigated with aid of SEM. It is found that the pollen morphology of Malus is rather similar in shape, size, position and number of aperture and exine sculpture. The characters of polle morphology of sections and series are as follows: Sect. Malus Ser. Baccatae: Striae regular, parallel to colpi, and conjunct at pole; Ser. Pumilae: the same as in Ser. Baccatae, but more or less curved near pole. Sect. Docyniopes: Striae regular and parallel to colpi, but bent near pole. Sect. Chloromeles: Striae irregular, dense and interlock. Sect. Sorbormalus: Striae irregular, sparse and not interlock. Ser. Sieboldiance: pollen grains prolate, with perforation among striae, colpi narrow; Ser. Kansuenses: Pollen grains spheroidal, colpi wide in the middle but narrow at both ends, striae relatively dense and regular, mostly dichotomous, perforation present; Ser. Yunnanenses; Pollen grain spheroidae, colpi wide in the middle but narrow at both ends, striae obviously irregular, less dichotomous, perforation absent. The major evolutionary trend of exine sculpture of pollen may be from densely thin-striate to sparsely striate with perforation. Characters of the exine sculpture of hybrids can be used to recognize the rela-tionship between the parental species.     

Pollen morphology of the subfamily Aurantioideae (Rutaceae)

The Aurantioideae is one of seven subfamilies of the Rutaceae consisting of two tribes, the Clauseneae, containing five genera, and the Citreae, with 28 genera. Each tribe contains three subtribes. The pollen morphology of the subfamily Aurantioideae is described and illustrated for the first time based on light and scanning electron microscopy. Five pollen types have been recognised in the subfamily, based mainly on aperture number and exine ornamentation. The pollen grains show a high degree of intergeneric variation. Pollen grains of Clauseneae are 3-colporate, microstriate or microstriato-reticulate, whereas pollen grains of Citreae are almost always 4/5 colporate with exines varying from microperforate to coarsely reticulate. Congruence between pollen types and the currently accepted classification is discussed, as well as the systematic implications of pollen morphology for the subfamily.     

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.     

Variation of pollen morphology, and its implications in the phylogeny of Clematis (Ranunculaceae)

Clematis s.l. (including Archiclematis and Naravelia) is a genus of approximately 300 species with cosmopolitan distribution. The diversity of its pollen was surveyed in 162 taxa belonging to all infrageneric groups of Clematis s.l. Pollen morphology was investigated by use of scanning electron microscopy to identify useful characters, test taxonomic and systematic hypotheses, and elucidate pollen character evolution on the basis of the molecular phylogeny. Clematis pollen is small to medium (14.8–32.1?μm?×?14.2–28.7?μm), oblate to prolate (P/E?=?0.9–1.4) in shape. The apertures may be tricolpate and pantoporate sometimes with 4-zonocolpate and pantocolpate pollen grains as transitional forms. The tricolpate pollen grains are predominant and occur in all the sections of the genus, whereas pantoporate pollen grains can be found in sect. Tubulosae, sect. Viorna, sect. Viticella, and Naravelia only. Phylogenetic mapping of aperture types reveals that the pantoporate pollen type may be the apomorphy in the genus and evolved several times. The surface ornamentation in all taxa studied is similar and characterized by microechinae evenly distributed on the microperforate tectum. The size and density of spinules on the tectum vary greatly but successive in the whole genus. According to the character syndromes of the ornamentation, separating sect. Brachiata from sect. Meclatis is supported. Though pollen morphology may contribute to investigation of problematic taxa, the taxonomic value of pollen morphology is limited at the species level.     

Pollen morphology and infra-generic evolutionary relationships in some Chinese species of Pedicularis (Scrophulariaceae)

 The pollen grains of 32 Chinese species of Pedicularis representing 13 groups within the genus were investigated using light microscopy and scanning electron microscopy. The genus is eurypalynous, the apertures may be tricolpate, trisyncolpate, or bisyncolpate and five types of surface ornamentation (microscabrate, microrugulate, retipilate, microreticulate and microfoveolate) were observed. Comparison of the different aperture types using light microscopy allows three pollen types to be distinguished. Examination of exine ornamentation with scanning electron microscopy enabled each pollen type to be divided into two or three sub-types (giving a total of seven sub-types). Pollen morphology within the genus is discussed in relation to infrageneric relationships, evolutionary and pollination biology. There is little correlation with the existing infrageneric taxonomy (some taxonomic groups have more than one pollen type, while the same pollen type may be found in several different species-groups), but greater correlation with the corolla morphology. Received January 10, 2002; accepted March 5, 2002 Published online: March 10, 2003     

Pollen Morphology of Microula Benth. and Allied Genera (Boraginaceae)

Pollen grains of 16 species of Microula Benth. and six species of three related genera were examined under LM and SEM, and four of them also under TEM. Pollen grains of Microula and three related genera are dumb-bell-shaped, 3-colporate apertures alternate with three pseudocolpi. Pollen grains are very small,ranging from 12.18 x 7.13 μm to 6.36 x 3.66μm. In general, colpi with os are wider and shorter, rhomboid, but sometimes they are equal to pseudocolpi in length. Colpus margins are regularly or irregularly tooth-like. The surface of colpi is psilate or processed. Ora are circular or lalongated in outline, protruded or not; surface of os membrane is smooth or scabrid. The exine is usually indistinctly layered under LM. The exine surface is psilate, and more or less perforate. TEM examination shows that the pollen wall is differentiated into exine and intine: the exine includes ectexine and endexine, while the ectexine consists of tectum, columellae and foot-layer. However, there are differences in constriction of equatorial area, apertural characters, ornamentation and exinous ultrastructure between these genera. Pollen morphology indicates that the genus Microula Benth. is primitive, directly related to the genus Actinocarya Benth .; the genus Asperugo L. Is more advanced. The genus Eritrichium Schrad. which has two ora or one os and is anisopolar, represents the most advanced group among them. Noteworthily, the diorate phenomenon is found for the first time not onlyin the genus but in the family Boraginaceae.     

安徽母草属植物花粉形态特征的研究

利用光镜和扫描电子显微镜首次对安徽母草属(Lindernia)六种植物的花粉形态进行了观察研究.结果表明该属花粉粒多为近球形, 具3 沟, 表面光滑.探讨了花粉形态特征的分类学意义,支持属级水平的分类.     

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 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.     

青藏高原东缘28种风毛菊属植物花粉形态研究

通过扫描电镜对青藏高原高寒草甸风毛菊属28种多年生草本植物的花粉进行了形态观察和比较,结果显示,该属植物花粉形态较为一致。花粉近球形或长球形,极面观三裂圆形,三孔沟,外壁表面具刺状突起。刺间纹饰可分为4种类型:穴状、颗粒状、网状—颗粒状和网状。花粉形态没有大的差异,但刺间表面纹饰、萌发沟的宽窄、刺的大小、分布密度和花粉大小在种间存在差异,可作为种的分类依据。花粉特征聚类分析结果表明,此28种风毛菊属植物花粉大致划分为明显的5个类群,所包含的种分别隶属于风毛菊属的四亚属:雪兔子亚属、雪莲亚属、附片亚属和风毛菊亚属。     

Pollen Morphology of Parnassia L. （Parnassiaceae） and Its Systematic Implications

The pollen morphology of 28 species of Parnassia L. was investigated with light microscopy and scanning electron microscopy (SEM). The shape of pollen grains in this genus varies from subspheroidal to prolate in equatorial view and is three-lobed circular in the polar view. Pollen grains are usually radially symmetrical, isopolar, tricolporate or syntricolporate, with reticulate sculpture. The pollen characteristics among species are fairly similar to each other. Morphological information regarding the pollen grains shows that Parnassia is a natural genus. Based on exine ornamentation observed under SEM, three types of pollen grains were recognized: (i) type I, with foveolate-reticulate sculpture; (ii) type II, with a finely reticulate sculpture; and (iii) type III, with a coarsely reticulate sculpture. Most sections of this genus have one type of sculpture of pollen morphology, but Sect. Nectarotrilobos has three types of sculpture and Sect.Saxifragastrum has two types of sculpture. All three types of sculpture can be found in Southwest China,with species with the longest (Parnassia delavayi Franch.) and shortest (Parnassiafaberi Oliv.) colpi,implying that Southwest China is the center of diversification of the genus.     

Morphology of pollen and orbicules in some Dioscorea species and its systematic implications

Pollen and orbicule morphology of 35 Dioscorea L. species is described based on observations with light microscopy, and scanning and transmission electron microscopy. Pollen and orbicule characters are critically evaluated and discussed in the context of existing hypotheses of systematic relationships within the genus. Pollen is mostly bisulcate (sometimes monosulcate) with a perforate, microreticulate or striate sexine. Our results indicate that pollen data may be significant at sectional rank. The close relationship between sections Asterotricha and Enantiophyllum proposed by Burkill and Ayensu is supported by pollen morphology as all species investigated share bisulcate, perforate pollen with small perforations and a high perforation density. Macromorphological differences between the two compound-leaved sections Botryosicyos and Lasiophyton are also supported by pollen morphology; pollens of these two sections have very different perforation patterns. Orbicules in Dioscorea are mostly spherical and possess a smooth or spinulose surface. The latter is often correlated with a striate sexine.