凤仙花属总状花序组的花粉形态及其分类学意义

总状花序组隶属于凤仙花亚属凤仙花属,以具多花的总状花序、侧生萼片2枚、子房5室、果实为线状圆柱形、种子多数、迈卵形、种皮表面具指状突起等特征区别于该亚属的其他类群。前人对凤仙花花粉形态的研究表明凤仙花属的花粉形态对于属下分类及种间界定具有重要的分类学价值。而对于总状花序组来说,有关花粉形态方面的研究少有报道,为了进一步探讨该组花粉S的形态特征及其分类学意义,该研究应用扫描电镜技术对总状花序组的21种植物的花粉进行了观察。结果表明:总状花序组植物的花粉粒均具4条萌发沟,两侧对称,极面观为矩圆形、长矩圆形或近方形;长赤道面观多为椭圆形至矩圆形,短赤道面观为椭圆形;长、短赤道轴比值多在1.4~2.4范围内;表面具网状纹饰,网眼在花粉粒表面均匀分布或赤道面的网眼较极面观的小,网脊边缘光滑或具波状突起,网眼内具不同密度的颗粒状突起。该组的花粉形态与宏观形态性状相关性较小,但花粉极面观形状、长短赤道轴的比值、网眼的分布、网脊边缘形状、网眼内颗粒物密度等特征具有种水平上的稳定性和特异性,对该组植物种间界定具有重要的意义。     

Pollen Morphology of Impatiens L. (Balsaminaceae) and Its Taxonomic Implications

Two forms of pollen morphology of Impatiens L. (Balsminaceae) have been described before: 3-colpate and 4-colpate pollen grains. Pollen grains of 13 species of Impatiens on Mt. Omei were examined under SEM for the first time and both forms were observed here, which imply the diversity of species in this region. Of the two forms, 3-colpate one is quite similar to that of the genus Hydrocera Blume (Balsaminaceae) in external morphology. This relation also exists between the 3-colpate pollen group and Hydrocera in habit, habitat, and distribution, etc. It is supposed that the group with 3-colpate pollen type is a natural one, and may have a close relationship with Hydrocera. The pollen morphology is of important significance in classification of the taxa and evaluation of systematic position in the fami-ly balsaminaceae.     

New Materials for Chinese Brassica (Cruciferae)

The present study deals with external morphology, chromosome number, pollen morphology and seed anatomy of three species of Brassica in China. The results are summarized as follows: 1. Brassica xinjiangensis is described as a new species based on the chromosome number n=9, and 3-4-colpate and reticulate pollen grains. In the transverse section of seed, epidermal cells are triangular, semiterete or irregular, subepidermal cells consist of one series, palisade cells terete and equal in length. Leaves are petiolate or not; stems, sepals, pistils and siliquae are usually covered with retrorse-pilose hairs; rostrum stout, 12 mm long. By the above characters this species differs from others in this genus. 2. Relationship between B. nigra and B. xinjiangensis. In the former all leaves are petiolate, and the lower part of stems and leaves usually bristly; sepals, pistils and siliquae are glabrous; pedicels are erect and appressed to the axis of infructescence; rostrum is gracile, 2.5 mm long. The chromosome number is n=8. Pollen grains are 3-colpate; the exine is cerebroid-reticulate under SEM. In the transverse section of seeds, epidermal cells are undulate, subepidermal cells in 1-2 series, palisade cells terete, short and unequal in length. These characteristics are the same as reported by Koch (1833), Darlington and Wylie (1955), Goldblatt (1981), Vaughan et al. (1971), Lan (1986). 3. Taxonomical position of B. celerifolia. It was treated as B. juncea var. celerifolia by Tsen et Lee (1942). According to the chromosome number n=10, 3-4-colporate pollen grains, it deserves specific rank. This species differs from B. juncea, which has the chromo-some number n=18 and 3-4-colpate pollen grains.     

中国木犀属花粉形态研究及其系统学意义

观察了中国木犀属4组21种植物的花粉形态。花粉为圆球形或近长球形,外壁表面均具网状纹饰。大多数种类都具有三孔沟的萌发孔,极少四孔沟,或六萌发孔。内孔明显或不明显,有9种植物同时存在具三孔沟和具三沟二种类型的花粉及中间过渡类型(显脉木犀同时存在具三拟孔沟和具四拟孔沟)。很多特点表明,木犀属花粉的萌发孔在木犀科中处于由简单萌发孔向复合萌发孔过渡的阶段。根据网眼从沟间区中央到沟边是否变细,可将本属花粉分为两大类型,与分组有一定的对应关系。综合一些花粉特征,可发现本属花粉形态差异较明显,与一般形态差异有一定的相关性。花粉形态在木犀属具有一定的系统学意义。     

国产淫羊藿属花粉形态研究

本文用光学显微镜和扫描电镜对我国淫羊藿属Epimedium 14种花粉进行比较深入的研究,试图通过花粉形态的研究为该属的植物分类和原植物鉴定提供参考依据。     

Studies on the Pollen Morphology and Seed Coat of the Genus Cistanche (Orobanchaceae) in China

In the present paper pollen grains of 5 species and seed coat of 4 species of the genus Cistanche (Orobanchaceae) from China were examined by scanning electron microscope As a result, two types of the pollen exine sculpture are distinguished: (1) tuberculate, C. salsa and C. lanzhouensis; (2) rugulate or fine reticulation formed by the fusion of rugae, C. sinensis, C. deserticola and C. tubulosa. Chinese Cistanche was classified into two sections based on the gross morphology by the present author (Zhang, 1984). C. lanzhouensis and C. sinensis were included in one section. This classification is in conflict with the pollen type, which indicates that the characters of pollen grains and external morphology in this genus have evolved at different rates. The differences in pollen morphology of Chinese Cistanche can serve as characters for delimitating species. According to our abservation, C. lanzhouenis Z. Y. Zhang differs from the other members of Cistanche not only in external morphology but also in pollen morphology. Pollen grains are subprolate in this species, while prolate in the other four species, and exine sculpture is tuberculate. The present study provides the establishment of this new species (Zhang, 1984) with palynological evidence. The seed coat sculpture in Chinese Cistanche are constantly alveolate, but there are some slight differences, by which the sculpture can be divided into two types: (1) testa cells have or have not thickenings on the inner anticlinal walls; cavernulous sculpture is present on outer periclinal wall of some cells: C. sinensis; (2) testa cells have striate thickenings on the inner anticlinal walls: C. salsa, C. deserticola and C. tubulosa. The characters of seed coatof some significance for the delimitation of species.     

中国丁香属花粉形态的研究

本文利用扫描电子显微镜和光学显微镜,研究了丁香代表植物11种和2变种的花粉形态,并对花粉的形态特征用电子计算机进行了主分量分析,同时将计算结果作了二维与三维排序,结果表明:(1)丁香属植物花粉形态在光学显微镜下种间区别甚微,而在电镜下的区别则很明显,可做为分种的依据,尤其对形态上难以区分的种类更为适用;(2)所做花粉除欧丁香外,其余花粉均为三孔沟型,对前人的工作作了一定的修正;(3)通过主分量分析,指出了花粉孔的长宽比、沟长及网眼大小是本属花粉分类的主要成分;(4)试图通过主分量的二维和三维排序,为属下分组、分种提供新的观点和依据。     

十四种凤仙花属植物种皮微形态特征及其分类学意义

通过扫描电镜对凤仙花属(Impatiens)14种植物的种子表皮微形态特征进行了观察。根据种皮表面纹饰及衍生物的特点,将该属植物的种皮微形态特征划分为4种类型,即光滑型、颗粒型、网状型和隆起型,后两者又可细分为若干亚型,其中负网状亚型和网状-丘状隆起亚型为首次报导。研究表明,14种凤仙花属植物种子表皮微形态特征差异显著,它们作为属内种的分类鉴别特征具有重要价值;种子表皮的微形态学特征与其植物体的部分表型特征具有相关性,在一定程度上反映了属内类群的分化。     

Micromorphology of fruits and seeds of Iranian Geranium (Geraniaceae), and its systematic significance

The fruit and seed micromorphology of 22 species of Geranium, representing the eight sections of the genus represented in Iran (G. sectt. Dissecta, Geranium, and Tuberosa of subgen. Geranium; sectt. Batrachioidea, Divaricata, Lucida, Ruberta and Trilopha of subgen. Robertium), have been examined by scanning electron microscopy (SEM). Macro‐ and micromorphological characters, including fruit and seed shape, size, color, hair type and density, mericarp ornamentations, hilum position, seed coat pattern, epidermal cell shape, and anticlinal and periclinal cell walls, are presented. Two microsculpturing patterns are recognized on the mericarp surface: reticulate and pusticulate. The micromorphology of the seed coat showed four distinctive cell patterns. The seed epidermis is constructed either of polygonal, elongated polygonal, or square to rectangular cells. The polygonal type is the most common among the studied species, but the variation in testa cell characters, their size and shape, may provide further information and useful diagnostic characters at specific and infraspecific rank. The shape and color of the seeds are, however, of little systematic value. Fruit characters were found to be important for separating taxa at infrageneric rank and our results show that the species can be separated into subgenera and sections based on fruit morphology.     

Investigation on Pollen Morphology of Aconitum L.

Pollen morphology of 62 species in Aconitum L. was investigated with light and scanning electron microscopies. The pollen grains in this genus are 3-colpate and exhibit spinulate surface pattern in all the species. According to the shape of pollen grains in equatorial view Aconiturn L. may be divided into three types. Type Ⅰ pollen grains are wide elliptic; Type Ⅱ are narrow-elliptic and type Ⅲ are rectangular or close to square in shape. The pollen characteristics among species are very similar. The morphological information of the pollen grains shows that species of Aconiturn L. are a natural taxon.     

Studies on the Pollen Morphology of the Genus Brassica (Cruciferae) in China

The present paper reports the pollen morphology of 36 species and varieties of Brassica L. in China mostly for the first time. They were examined both under LM and SEM. Pollen grains of Brassica L. in China are long-spheroidal, subspheroidal, spheriodal, oblate, rarely reniformis or subsquare. According to the aperture characters pollen grains may be divided into four types: 2,3,4-colpate, 3-colpate, 3,4-colpate, 3,4-colporate. The exine sculpture may also be divided into three types: 1, reticulate, 2, cerebroid-reticulate, 3, foveolatereticulate. Detailed characters of pollen grains for each taxon are presented in Table 1. The 2,3,4-colpate type of pollen grains may be considered as primitive one and the 3,4-colpate or 3,4-colporate type of pollen grains is the most advanced in Brassica. The tricolpate aperture and reticulate exine have been reported by the previous authors, whereas, 3,4-colporate and cerebroid-reticulate or foveolate-reticulate pollen grains are newly reported.     

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.     

A Taxonomic Study on the Genus Polygala L. in China

The present paper deals with the history of taxonomy, groos morphology, pollen morphology, distribution, system and taxonomic treatment of the genus Polygala L. from China. Three subgenera, 4 sections, 41 species and 8 varieties are recognized in this treatment, of which 2 sections are described as new. The system of Chinese species of Polygala L. is proposed as follow: Subgen. 1. Chamaebuxus (DC.)Duchartre (Typus: Polygala chamaebuxus L.) Sect. 1. Arillus S. K. Chen, sect. nov. (Typus: Polygala arillata Buch.-Ham. ex D. Don). This section consists of about 20 species, distributed in the tropics and subtropics of Asia and Africa, with 7 species found in South China. Subgen. 2. Pseudosemeiocardium (Adema) J. Chetek et B. Krisa( Typus: Polygala furcata Royle). Sect. 1. Villososperma C. Y. Wu et S. K. Chen (Typus: Polygala wattersii Hance ). There are 3 species in this section, distributed in S. China and N. Vietnam. Sect. 2. Saxicola S. K. Chen, sect.nov. (Typus: Polygala saxicola Dunn). This section consists of 8 species, among which P. tricornis Gagnep. and P. saxicola Dunn are distributed in both Vietnam and China, and the other 6 species are endemic to China. Sect. 3. Pseudosemeiocardium ( Typus: Polygala furcata Royle). This section consists of about 7 species, distributed in SW China, extending from Indo-China Peninsula, southward to Malay, the Philippines, Indonesia and New Guinea, westward to the southern slope of the Himalayas and N. India, northward to Japan. Subgen. 3. Polygala (Typus: Polygala vulgaris L. ). 400 species or more belong to this cosmopolitan subgenus. Most species are found in America and Africa, with 18 species discovered in China. Taxonomic evidence: The characters of flowers, fruits and seeds, the indumentum of seeds and presence or absence of caruncles are stable and also correlated with one another. There is a certain combination of the above-mentioned characters in a given group, which provide reliable evidence for the infrageneric division. A good example is the Polygala arillata group, grown under the tropical and subtropical forests, which has large, yellow flowers, with sepals fallen after blossom. The flower has carinas with a cristate appendix, which is of 2-9 long narrow pieces. The seeds are spherical, with a hooded caruncle. Its fruits are of annuloid stripes. (Fig. 1). The Polygala tenuifolia Willd. group, cosmopolitan in distribution, has a middle-sized, purple or yellow flower with persistent sepals. Cristate appendix of carina is usually fimbricate. The seed in the group is oblong or subovoid, with a trilobate caruncle. Carpels of the fruits of this group are without annuloid stripes(Fig. 3). The third group, Polygala saxicola group, is between the two as indicated above. The group has a small, yellow or occasionally purplecolored flower, with sepals fallen after blossom or sometimes with a persistent outer sepal. The cristate appendix of carina is lamellar or cucullate. The seed of the group is sometimes with pubescences and hooded caruncles sometimes with hirtoselike villoses and no caruncle (Fig. 2) pollen morphology and systematic treatments: The pollen morphology of the domestic Polygala seems to support division of Polygala L. into three groups as follows: A. Pollen grains subspheroidal, 10-19-colpate, colpi usually rather wide, relatively psilate in the polar area; B. Pollen grains reniform, 22-26-colpate, sculptural in the polar area; C. Pollen grains long-spheroidal or subspheroidal, 9- 23-colpate, with long and deep colpi, which usually extend to the relatively psilate polar area. The pollen morphology is naturally correlated with the morphology of flowers, fruits and seeds. Based on these, three subgenera and four sections no subdivision in Subgen. Polygala)are recognized.     

Pollination ecology and pollination system of Impatiens reptans (Balsaminaceae) endemic to China

BACKGROUND AND AIMS: China is one of the centres of geographical distribution of Impatiens L. Studies of the pollination ecology of this genus in China have, until now, been unreported. Impatiens reptans, a species endemic to China, was studied. The aims were to examine the pollination ecology and pollination system of this species, to compare its pollination ecology with other Impatiens species growing in Sumatra and Japan, and to discuss possible reasons for its limited distribution. METHODS: The pollination ecology of I. reptans was studied by carrying out continuous observations within three naturally growing populations. Its pollination system was studied using different pollination methods, marking and counting pollen grains, assessing pollen viability and observing pollinator behaviour. KEY RESULTS: The flowering phase of the protandrous I. reptans lasted for 89 d. The life span of an individual flower was 3.6 d. Primary pollinators were honey-bees and bumble-bees. Secondary pollinators were diurnal hawk moths and butterflies. Bombus briviceps and Bombus sp. were nectar gatherers. The mean nectar sugar concentration was 29.5 %, and the mean value of sucrose/glucose + fructose was 0.82. The proportion of seed set ranged from 0.857 to 0.873. Distances that seeds were ejected ranged from 0.58 to 1.17 m. Percentage seed germination under controlled conditions was 23.1. Pollen viability was highest on the day of anthesis and thereafter decreased. Ratios of pollen : ovules ranged from 958.8 to 970.6. CONCLUSIONS: Impatiens reptans reproduces by means of cross-pollination. Its dependence on a specialized habitat, a narrow environmental niche, a low percentage of seed germination, and habitat loss could be reasons for its limited distribution and endemism.     

Former diversity of Ephedra (Gnetales): evidence from Early Cretaceous seeds from Portugal and North America

BACKGROUND AND AIMS: The extant species of the seed plant group Gnetales (Ephedra, Gnetum and Welwitschia) have been considered a remnant of a much greater, now extinct, diversity due to the pronounced differences in form and ecology among the genera. Until recently, this hypothesis has not been supported by evidence from the fossil record. This paper adds to the expanding information on Gnetales from the Early Cretaceous and describes coalified seeds from Barremian-Albian localities in Portugal and USA. METHODS: The fossils were extracted from sediment samples by sieving in water. Adhering mineral matrix was removed by chemical treatment. Seeds were investigated using light and scanning electron microscopy. Morphology and anatomy of the seeds were documented and compared with those of extant species. KEY RESULTS: The fossils share characters with extant Ephedra, for example papillae on the inner surface of the seed envelope and in situ polyplicate pollen grains that shed the exine during germination. They differ from extant Ephedra seeds in morphological and anatomical details as well as in their smaller size. Two new species of Ephedra are described together with one species assigned to a new genus of Gnetales. Other Ephedra-like seeds, for which pollen and critical morphological details are currently unknown, are also present in the samples. CONCLUSIONS: These Cretaceous seeds document that key reproductive characters and pollen germination processes have remained unchanged within Ephedra for about 120 million years or more. There is sufficient variety in details of morphology to suggest that a diversity of Ephedra and Ephedra-like species were present in the Early Cretaceous flora. Their presence in Portugal and eastern North America indicates that they were widespread on the Laurasian continent. The fossil seeds are similar to seeds of Erdtmanithecales and this supports the previously suggested relationship between Erdtmanithecales and Gnetales.     

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.     

Systematic implication of seed micromorphology in Arenaria (Caryophyllaceae) and allied genera

Variations in seed micromorphology of 64 taxa of Arenaria and allied genera were examined using scanning electron microscopy (SEM) in order to evaluate their diagnostic value for systematic studies in the genus and providing additional evidence on delimitation of natural groups. Significant diversity was found in seed coat morphology both among and within genera. Size, color, and shape of seeds appear to be of low importance at species level but provide useful evidence at generic rank. Cerastium and Stellaria are well distinguished from Arenaria by serrate seed margins and solid columellae on testa cells. The sculpturing pattern renders most characters potentially applicable in taxonomy and phylogeny assessment in Arenaria. The studied taxa show four basic types of sculpturing, viz. colliculate, rugose, papillate, and smooth. The colliculate type which is characteristic for the core group of Arenaria (A. subg. Arenaria) can be further divided into five subtypes. The smooth seed surface with indistinct cell boundaries represents a synapomorphy for A. subg. Leiosperma, whereas the winged seeds along with rugose surface characterize subg. Odontostemma. Otherwise seed micromorphology rarely provides adequate evidence for separation of natural groups, especially at sectional rank. We hypothesize that the seeds are highly polymorphic in certain groups of taxa showing recent adaptive radiation, such as the annual species of Arenaria, and do not provide strong support for natural groups within the genus.     

Impatiens quadriloba sp. nov. (Balsaminaceae) from Sichuan,China

Impatiens quadriloba K. M. Liu et Y. L. Xiang sp. nov. collected from the Siguniang Mountains Nature Reserve in northwestern Sichuan, China, is described and illustrated. The seed‐coat and pollen morphology of the new species are described and diagnostic morphological characters that distinguish the new species from the related I. falcifer Hook. f. are discussed.     

The Restoration of the Genus Cyclocodon ( Campanulaceae) and Its Evidence from Pollen and Seed-coat

The genus Cyclocodon Griff. was reduced by C. B. Clarke in 1881 into a section of the genus Campanumoea. Our LM and SEM observations on pollen morphology show that the pollen of all the three species in the former is 3-colporate with the exine sparsely high-spinulose, whereas that of the two species in the latter is 5～8-colpate with the exine relatively densely short-spinulose. SEM observations on seed-coat indicate that the primary ornamentation of the seed-coat of the two species in Campanumoea (s. str. ) is characterized by regular and polygonal areoles which are much larger than the radial walls in diameter and by the bead-like secondary ornamentation on the radial walls, while that of the three species in Cyclocodon is characterized by irregular-shaped areoles which are nearly equal to the radial walls in diameter and by the rope-like secondary ornamentation on the radial walls. Thus, the pollen morphology is closely correlated with the seed-coat morphology. Taking the characteristics of pollen, seed-coat and gross morphology into consideration, the genus Cyclocodon is restored, separate from Campanumoea (s. str. ). One new combination, Cyclocodon celebicus (B1.)Hong, is made in the present paper. The genus Cyclocodon is con-sidered closely related to Platycodon rather than to Campanumoa.