Pollen morphology of Dioscorea (Dioscoreaceae) and its relation to systematics

This paper adds new data on the pollen and orbicule morphology of 61 Dioscorea L. (Dioscoreaceae) species to the survey of Schols et al . (2001). The results indicate that pollen characters may be significant in infrageneric systematics in Dioscorea . Pollen and orbicule characters are described based on observations with light microscopy, and scanning and transmission electron microscopy, and are critically evaluated and discussed in the context of existing hypotheses of systematic relationships within the genus. Pollen is mostly disulculate (sometimes monosulcate) with a perforate, microreticulate, striate, gemmate, rugulate, or cerebroid perforate sexine. The basal section Stenophora is one of the few sections with monosulcate pollen. Brachyandra , Cardiocapsa , and Seriflorae , three Malagasy sections, are characterized by striate pollen. Pollen morphology strongly supports section Enantiophyllum as a monophyletic group. The correlation between pollen size and tuber type, as suggested previously by P. Su (1987), is confirmed by our data. As found in our earlier survey, orbicules in Dioscorea are mostly spherical with a smooth or spinulose surface.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 375–390.     

Systematic relevance of pollen and orbicule characters in the tribe Hillieae (Rubiaceae)

Pollen and orbicule morphology of 26 species in the tribe Hillieae is described based on observations by light, 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 tribe. Pollen is 3-zonocolporate with a perforate, microreticulate, reticulate or eutectate sexine. In the two species of Blepharidium , however, the pollen has one, four or five apertures. These pollen morphological data were incorporated into an existing macromorphological matrix of the group and cladistically analysed. The resulting phylogenies indicate that it may be appropriate to reduce Cosmibuena to a subgenus of the genus Hillia , while Blepharidium should be removed from Hillieae because of the deviating pollen type that is unique to the Rubiaceae. All species investigated produce orbicules, which are mostly spherical and possess an electron-lucent core that is sometimes characteristically flattened.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 303−321.     

半日花花粉形态种内变异及其花粉地理研究

用扫描电镜在个体水平上研究了产自两个地区的半日花花粉,并首次发现花粉种内变异与其地理分布之间的显著相关,半日花花粉形态在种内个体之间存在差异,以餐壁纹饰明显分为穿孔型和条纹型花粉;按形状和大小又可分为近球形较小和长球形较大花粉。外壁纹饰与形状、大小所得类型基本吻合。本种花粉,因所处地理环境的不同而外壁纹饰也有明显差异,条纹型花粉分布于新疆天山地区,而穿孔型花粉出现在内蒙古伊盟桌子山,并认为花粉外壁     

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 morphological support for the Catesbaeeae-Chiococceae-Exostema-complex (Rubiaceae)

The pollen morphology of the Catesbaeeae-Chiococceae-Exostema complex as recently treated by Delprete (1966) was examined with LM and SEM. The group is remarkably stenopalynous; typical representatives have medium sized, 3-colpate pollen with a perforate tectum covered with microspines. The inner nexine ornamentation is pronounced and offers more variation than the sexine pattern. A typology of the inside structures is presented based on LM observations and SEM observations of sectioned grains. Orbicules are common and numerous in the Catesbaeeae and Exostema-group; for most genera of the Chiococceae confirmation is needed of orbicule presence. All orbicules observed are relatively large (1-4 μm) and spiny. Pollen and orbicule morphology proved to be a powerful tool to delimit the Catesbaeeae-Chiococceae-Exostema complex. The overall delimitation of the complex is corroborated with our pollen data. The genera Mastixiodendron and Placocarpa, however, can be excluded from the complex based on their pollen morphology. Mastixiodendron has 3-colporate, perforate pollen without microspines and the endocolpi are fused into an endocingulum. Pollen of Placocarpa is reticulate and 3-colporate with perpendicular endocolpi.     

Palynological variation in Balsaminoid Ericales. II. Balsaminaceae, Tetrameristaceae, Pellicieraceae and general conclusions

BACKGROUND AND AIMS: The objective of this study is to examine the palynological diversity of Balsaminaceae (two genera/+/-1000 species), Tetrameristaceae (two genera/two species) and Pellicieraceae (one genus/one species). The diversity found will be used to infer the systematic value of pollen features within the balsaminoid clade. METHODS: Pollen morphology and ultrastructure of 29 species, representing all families of the balsaminoid clade except Marcgraviaceae, are investigated by means of light microscopy, scanning electron microscopy and transmission electron microscopy. KEY RESULTS: Balsaminaceae pollen is small to medium sized with three to four apertures, which can be either colpate or porate, and a sexine sculpturing varying from coarsely reticulate to almost microreticulate. Tetrameristaceae pollen is small sized, 3-colporate, with a heterobrochate reticulate sculpturing and granules present in the lumina. Pellicieraceae pollen is large sized, 3-colporate with long ectocolpi and a perforate sexine sculpturing with large verrucae. Furthermore, Pelliciera is characterized by the occurrence of aggregated orbicules, while orbicules are completely absent in both Balsaminaceae and Tetrameristaceae. Balsaminaceae pollen differs from the other balsaminoid families due to the occurrence of colpate or porate grains with an oblate to peroblate shape, a very thin foot layer and a lamellated endexine. CONCLUSIONS: From a pollen morphological point of view, Balsaminaceae are completely different from the other balsaminoid families. Therefore, no pollen morphological synapomorphies could be defined for the balsaminoid clade. However, various pollen features were observed that could indicate a possible relationship between Tetrameristaceae, Pellicieraceae and Marcgraviaceae. Despite the palynological similarities in the latter three families, it remains unclear to what extent they are related to each other.     

Palynological diversity and major evolutionary trends in Cyperaceae

Pollen and orbicule morphology of 84 species, representing 52 genera from all tribes and subfamilies are investigated, in order to assess the systematic value of palynological data and to determine palynological evolutionary trends in Cyperaceae. A total of 90% of the species are examined for the first time with scanning electron microscopy. Pollen grains of Cyperaceae are oblate spheroidal to perprolate in shape, inaperturate to polyporate with opercula or pontopercula on pori or colpi. We distinguished seven different sexine ornamentation patterns. Orbicules occur in all species investigated. Pollen morphological variation within Cyperaceae is considerable and includes dispersal unit; number, location and degree of differentiation of apertural zones; and sexine ornamentation patterns. In subfamily Mapanioideae both tribes can be characterized by palynological synapomorphies. However, in subfamily Cyperoideae, the observed pattern of variation does not fit the most recent molecular phylogeny due to high levels of homoplasy and polymorphism in major pollen characters.     

Systematic implications of pollen morphology in Elsholtzia (Elsholtzieae–Lamiaceae)

Pollen morphology of 18 species (32 specimens) representing all three currently recognized sections of the genus Elsholtzia (Elsholtzieae–Lamiaceae) was investigated in detail using light, scanning electron and transmission electron microscopy. Elsholtzia pollen grains are small to medium in size (P=20–50 μm, E=16–45 μm), mostly prolate‐spheroidal to prolate, and rarely oblate‐spheroidal to subprolate in shape (P/E=0.95–1.46), hexacolpate (the amb more or less circular or ellipsoid) with granular aperture membranes. Three distinct types of sexine ornamentation are observed in Elsholtzia: perforate, rugulose‐bireticulate, and bireticulate. The bireticulate pattern is the most common, showing a wide range of morphological variation among species. The exine thickness varies from 1.2 to 2.5 μm. Among the present palynological data, the variability observed in sexine ornamentation of Elsholtzia could be of systematic significance. Current infrageneric classification of the genus Elsholtzia is partially congruent with pollen morphological data.     

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.     

亚洲茜草科粗叶木属及其相关属植物花粉形态研究

利用光学显微镜和扫描电镜研究了茜草科粗叶木属Lasianthus 16种2亚种、1变种及相关的5属5种的花粉形态。粗叶木属的花粉属于广孢型, 单粒。一般中等大小, 绝大多数为圆球形, 少数为近长球形或长球形。花粉形态特征, 特别是在萌发孔和外壁纹饰上表现出多样化。根据孔沟的数目或是否具有内孔, 可以将萌发孔分为(3-)4-(-5)孔沟和3孔。在所观察的这些种中, 萌发孔以3-4孔沟为主要类型, 比例为62.4%。外壁纹饰可分为细网状、粗网状和穴状。有部分种的花粉极面有穴状纹饰, 其余均为网状纹饰。网眼一般椭圆形、近圆形、三角形或者不规则形。少数外壁纹饰网脊上有颗粒状雕纹或模糊的颗粒, 网脊轮廓线呈波浪形, 一般凸出且平滑。大部分种的花粉具有沟膜, 沟膜上具有瘤状突起或小颗粒状, 沟边缘一般较平滑, 或粗糙, 有的种具有沟桥。     

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。     

Comparative pollen morphology of Glechoma and Marmoritis(Nepetinae,Lamiaceae)

The pollen morphology of 13 taxa(34 specimens)of the genera Glechoma L.,and Marmoritis Benth.was investigated in detail using light,scanning electron,and transmission electron microscopy.Pollen grains of all studied taxa are small to large in size(P = 32.5-60.4μm,E = 20.2-50.5μm),prolate-spheroidal to prolate in shape and mostly hexacolpate(the amb more or less circular or rarely ellipsoid)with granular membranes.The sexine ornamentation of Glechoma is bireticulate; the muri of the primary reticulum are irregularly circled,and lumen size is short.In contrast,the sexine surfaces of the Marmoritis pollen tend to more elongate or wider at the muri of the primary reticulum than those of the Glechoma.The pollen wall stratification of selected taxa(three from Glechoma and one from Marmoritis)is characterized by unbranched columellae,and continuous or distinctly discontinuous endexine based on transmission electron microscopy observation.The results of Glechoma and Marmoritis reveal rather similar pollen morphological features,however,fine details of sexine ornamentation are characteristic to differentiate the pollen taxa.Although these differences may be useful in establishing the taxonomic boundary between two genera,they are too weak to segregate diagnostic characters.     

Studies on Pollen Morphology of Pulsatilla Mill.

Pollen grains of 18 species of Pulsatilla in Ranunculaceae distributed in Asia and Europe were examined by LM and SEM, and exine ultrastructure of tricolpate pollen grains of P. chinensis and of pantoporate pollen grains of P. campanella was examined by TEM. Pulsatilla pollen is divided into four major types based on the aperture character, i.e. tricolpate, di- and tricolpate, pantocolpate and pantoporate. The revolutionary trend of pollen types is as follows: tricolpate→pantocolpate→pantoporate. Surface spinulate and perforate. According to density and size of sptnulae and distribution of perforation, the pollen grains of the genus can be divided into two groups. Thin sections of P. chinensis and P. campanella show endexine thickened at colpi and ora. Ektexine consists of a foot layer, a collumellae layer and a continuous, perforate tectum. The columallae layer is thicker than foot layer and tectum. Pollen morphology of Pulsatilla is similar to that of Anemone, but different in the distribution of spinules and perforation. Pollen information supports Wang’s view about systematic arrangment of species of Pulsatilla in China.     

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 genus Rubus from China

Pollen morphology of 103 species belonging to twenty-three subsections of seven sections of Rubus L. was examined with SEM. The pollen grains are usually 3-colporate, spheroidal, subspheroidal, prolate and perprolate, though occasionally rhombic and hexagonal, 16.0~38.0 μm × 12.0~30.0 μum in size. The exine ornamentation can be divided into rugulate, striate, cerebroid and perforate-reticulate types, which include 11 subtypes: rugulate-subpsilate, rugulate-striate, rugulate-perforate, striate, striate-perforate, striate-reticulate, cerebroid, cerebroid-perforate, perforate, perforate-reticulate and reticulate. The pollen morphology is of some significance for a better classification of the genus Rubus, but seems to be of little help for a better understanding of thephylogeny of the genus.     

中国悬钩子属花粉形态观察

通过扫描电镜观察了中国悬钩子属7组23亚组共103个种的花粉形态。本属花粉为圆球形、近 球形、长球形和超长球形,少数为菱形和六边形,具三孔沟,大小多为16.0～38.Oμm×12.0～30.0μm,外壁纹饰可分为皱波型、条纹型、脑纹型和穿孔网型等类型,进而可分为皱波-近光滑状、皱波-条纹状、皱波-穿孔状、条纹状、条纹-穿孔状、条纹-网状、脑纹状、脑纹-穿孔状、穿孔状、穿孔-网状及网状等11个亚型。分析结果认为,悬钩子属的花粉形态特征对该属种的区分有一定的参考价值,但要合理地阐释其系统发育意义,仍需积累更多的信息。     

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).     

Comparative pollen morphology and ultrastructure of Mentheae subtribe Nepetinae (Lamiaceae)

This study provides new pollen data of 52 representative species belonging to all 12 genera in the currently classification of the subtribe Nepetinae, and considers the possible presence of orbicules for the first time. Pollen morphology and ultrastructure were investigated with light, scanning electron and transmission electron microscopy. Nepetinae pollen is small to large (P = 16–65 µm, E = 17–53 µm), oblate to prolate (P/E = 0.7–1.6) in shape and mostly hexacolpate (sometimes octocolpate). The exine stratification in all taxa studied is similar and characterized by unbranched columellae and a continuous, granular endexine. Sexine ornamentation in the Nepetinae is bireticulate, microreticulate or perforate. In perforate and microreticulate pattern a tendency towards a bireticulum could be recognized due to trace of secondary tectal connections. The bireticulate pattern is most common with variations of primary muri and secondary reticulum. In Hymenocrater and Schizonepeta the observed variation of sexine ornamentation is particularly valuable at the generic level. Pollen data support Lophanthus and Nepeta as very closely allied and Lallemantia is clearly distinct from Dracocephalum. The formerly suggested infrageneric relationships within Dracocephalum and Nepeta are only partly corroborated by palynological characters. Orbicules are absent in the Nepetinae.     

Comparative pollen morphology of Glechoma and Marmoritis (Nepetinae,Lamiaceae)

Abstract The pollen morphology of 13 taxa (34 specimens) of the genera Glechoma L., and Marmoritis Benth. was investigated in detail using light, scanning electron, and transmission electron microscopy. Pollen grains of all studied taxa are small to large in size (P= 32.5–60.4 μm, E= 20.2–50.5 μm), prolate‐spheroidal to prolate in shape and mostly hexacolpate (the amb more or less circular or rarely ellipsoid) with granular membranes. The sexine ornamentation of Glechoma is bireticulate; the muri of the primary reticulum are irregularly circled, and lumen size is short. In contrast, the sexine surfaces of the Marmoritis pollen tend to more elongate or wider at the muri of the primary reticulum than those of the Glechoma. The pollen wall stratification of selected taxa (three from Glechoma and one from Marmoritis) is characterized by unbranched columellae, and continuous or distinctly discontinuous endexine based on transmission electron microscopy observation. The results of Glechoma and Marmoritis reveal rather similar pollen morphological features, however, fine details of sexine ornamentation are characteristic to differentiate the pollen taxa. Although these differences may be useful in establishing the taxonomic boundary between two genera, they are too weak to segregate diagnostic characters.