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.     

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.     

中国槲寄生属花粉形态的研究

首次系统地报道中国槲寄生属10种植物花粉形态。通过光学显微镜和扫描电镜观察,国产槲寄生属种类可按花粉的外壁纹饰分为两大类群,与形态上划分相一致。类群Ⅰ(8种)具不显著或近光滑的外壁纹饰,这一类型又可分为2个亚型:纹饰在花粉各个部位基本上是一致的和纹饰在赤道和极、沟两侧不一致的;类群Ⅱ(2种)外壁具显著的杆状纹饰。但花粉性状对形态上相近的种类鉴定意义不大。还讨论了槲寄生属花粉的可能的演化趋势。     

中国桑寄生科的孢粉学研究

系统报道了中国桑寄生科Loranthaceae33种5变种植物的花粉形态,并与澳大利亚 2属6种植物的花粉形态做了比较。通过光学显微镜和扫描电镜观察,国产桑寄生科花粉外壁 纹饰可明显分为两个类型:一种类型为刺状或条状纹饰,另一种为颗粒状纹饰,这与该科的鞘 花族和桑寄生族两个族相吻合。在鞘花族类型中,3合沟、钝刺状或条状纹饰的花粉是基本类 型,合半沟或孔沟形,刺状纹饰的花粉是较进化的类型;在桑寄生族类型中,等极、3合沟、 颗粒状纹饰的花粉是基本类型,异极、副合半沟-合半沟、3沟形和沟形-短沟形或沟孔形、粗 糙或模糊颗粒状纹饰的花粉是较进化类型。根据萌发孔和纹饰可将桑寄生族类型花粉分为3个 类群:类群I包括五蕊寄生属Dendrophtho、梨果寄生属Scurrula、钝果寄生属Taxillus和大苞 寄生属Tolypanthus;类群II仅包括离瓣寄生属Helixanthera;类群III也仅1属,桑寄生属Lor anthus。在这3个类群中,类群I属于基本的类型,属间花粉差别较小,其中梨果寄生属和钝 果寄生属花粉差别最小,显示出较近的亲缘关系;类群II和类群III皆是较进化类型。     

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.     

Pollen morphology of the tribes Naucleeae and Hymenodictyeae (Rubiaceae – Cinchonoideae) and its phylogenetic significance

The tribe Naucleeae has recently been recircumscribed on the basis of both morphological and molecular [ rbcL , trnT-F , internal transcribed spacer (ITS)] evidence, and has been found to be the sister group of the tribe Hymenodictyeae Razafim. & B. Bremer. In order to find pollen morphological support for this new classification, the pollen and orbicules of 65 species, representing 23 Naucleeae and the two Hymenodictyeae genera, were investigated by scanning electron and light microscopy. Naucleeae pollen is very small (< 20 µm) to small (20–30 µm) and its shape in equatorial view is suboblate to spheroidal or, more rarely, subprolate. Three compound apertures are present, each comprising a long and narrow ectocolpus, a circular to slightly lolongate mesoporus, and an often H-shaped endoaperture. The sexine ornamentation is perforate, rugulate, or (micro)reticulate, and supratectal elements are always absent. Apart from the variation in sexine ornamentation, the tribe is rather stenopalynous. The pollen of Hymenodictyeae is very similar to that of Naucleeae. The H-shaped endoapertures often observed probably form a synapomorphy for the clade comprising Naucleeae and Hymenodictyeae. Our pollen morphological observations are not in conflict with the widened delimitation of Naucleeae. Unambiguous pollen support for the recent subtribal or generic concepts of Naucleeae could not be found because of a lack of variation of pollen characters within the tribe. Orbicules are invariably present in the ten Naucleeae taxa investigated. They are spheroidal and smooth or irregularly folded.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 329–341.     

Pollen morphology and systematics of Atripliceae (Chenopodiaceae)

Pollen morphology of 58 species from 17 putative genera of the tribe Atripliceae (Chenopodiaceae) was investigated using light (LM) and scanning electron microscopy (SEM). Morphological variation was analyzed based on a dense sampling of the subtribes Atriplicinae and Eurotiinae, including many of the species in the two largest genera: Atriplex and Obione. The pantoporate pollen grains of Atripliceae are characterized by their spheroidal or subspheroidal shape, flat or moderately vaulted mesoporia with 21–120 pores, tectum with 1–8 spinules and 5–28(?38) puncta per?µm², and 1–13 ectexinous bodies bearing 1–7 spinules each. Taxonomic relevance of the most important pollen morphological characters is discussed (pollen diameter, pore number, pore diameter, interporal distance, spinule and puncta density and ratio, number of ectexinous bodies, and their spinules). Pollen morphological data support the exclusion of Suckleya from the tribe and the recognition of subtribe Eurotiinae, but suggest that it needs to be reviewed. Pollen does not support generic recognition of Atriplex, Neopreissia and Obione and infrageneric subdivisions as currently recognized, and suggests the need to review them. Smaller or monotypic genera, such as Axyris, Ceratocarpus, Endolepis, Krascheninnikovia, Microgynoecium, Proatriplex and Spinacia have distinctive pollen morphological characters that support their generic status. Grayia needs to be reevaluated; although its two species are distinct from all the other species in the study, there are notable differences between each of them, and this suggests they may not form a natural group. Multivariate techniques were employed to investigate if there are discrete patterns of variation within Atripliceae. Principal Component Analyses (PCA) weakly differentiates four groups based on variation in pore number, puncta density per?µm², and ratio between spinule and puncta density per?µm²; species of Ceratocarpus, Haloxanthium, Krascheninnikovia, Manochlamys, Microgynoecium, Spinacia, and some species of Atriplex and Obione are isolated. Preliminary results indicate that pollen data are potentially useful in the classification of the tribe, and further studies will be of taxonomic value.     

Pollen morphology of the Thymelaeaceae in relation to its taxonomy

 Pollen morphology of the four subfamilies of Thymelaeaceae sensu Domke (1934) was examined using LM, SEM and TEM. The variation of the exine architecture allows to distinguish four pollen types and three subtypes. Distribution of pollen types proves to be widely correlated with the occurrence of characters from wood anatomy and flower morphology. If pollen types are connected with subfamilies sensu Domke (1934), Gonystyloideae, Thymelaeoideae and Synandrodaphnoideae are stenopalynous, whereas Aquilarioideae are eurypalynous. Based on pollen morphology as well as on other characters it is evident that Aquilarioideae are not monophyletic. Its genera Aquilaria and Gyrinops are more closely related to Thymelaeoideae and Synandrodaphnoideae than to other genera of Aquilarioideae sensu Domke (1934). The remaining genera of Aquilarioideae Deltaria, Solmsia and Lethedon are most similar to Gonystyloideae and therefore are included in this subfamily, as it was suggested by Airy Shaw (1979). Furthermore, palynological and other characters favour the transfer of Octolepis from Aquilarioideae to Gonystyloideae. The re-circumscribed Aquilarioideae together with Thymelaeoideae and Synandrodaphnoideae are shown to form a monophyletic group. Received August 8, 2001 Accepted December 7, 2001     

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.     

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.     

Pollen wall stratification and pollination

Qualities of the stratified pollen walls were evaluated for their possible role in pollination (pollination modes, and pollen tube formation). The importance of studying pollen grains in their respective natural state is noted. Examples of pollen morphological features specific to pollination vectors are rare and difficult to demonstrate. However, some complex, but significant correlations are reported.     

Pollen grain morphology ofCoris (Primulaceae)

Pollen grain morphology, sculpturing, and wall ultrastructure are investigated in two species ofCoris (Primulaceae),C. monspeliensis L. andC. hispanica Lange. The study includes both acetolysed and unacetolysed pollen. No evidence of any major palynological difference is recorded between these two species, apart from a somewhat larger pollen inC. monspeliensis. However,Coris can be distinguished from the remaining members of thePrimulaceae by the conjunction of relatively large pollen grains, prominent margo, and particular tectal pattern causing a reticulate surface with minute luminal perforations decreasing towards the colpi. From both these distinctive features, and others typically primulaceous, some evolutionary considerations are inferred. Finally, the higher proportion of irregular grains inC. hispanica is interpreted in light of environmental stress.     

Pollen morphology in relation to pollinators in Papilionoideae (Leguminosae)

The surface sculpturing of the pollen of some species of the taxonomically widely separated genera Harpalyce (tribe Brongniartieae), Camoensia (tribe Sophoreae), Millettia (tribe Tephrosieae), and of the monotypic Dahlstedtia (tribe Tephrosieae) which have large red or white flowers adapted for pollination by birds or bats, is coarsely rugulate or verrucate. Related taxa with small insect pollinated flowers have pollen with simple reticulate or perforate surface sculpturing. The exine stratification of Alexa and Castanospermum (tribe Sophoreae), genera with large red bird-flowers, is complex with a layer of tectal columellae and differs from that of other genera in the tribe Sophoreae which have a normal pollen wall structure. These modifications of pollen structure and sculpture appear to be the result of convergent evolution and a secondary adaptation to pollination. The taxonomic and functional significance of the observations are briefly discussed.     

Pollen morphology and ultrastructure of selected species of Magnoliaceae

The pollen morphology and ultrastructure of 20 species, representing eight genera of the Magnoliaceae are described based on observations with light, scanning and transmission electron microscopy. The family represents a homogeneous group from a pollen morphological point of view. The pollen grains are boat-shaped with a single elongate aperture on the distal face. The tectum is usually microperforate, rarely slightly or coarsely rugulose. Columellae are often irregular, but well-developed columellae do occur in some taxa. The endexine is distinct in 14 species, but difficult to discern in the genera Parakmeria, Kmeria and Tsoongiodendron. Within the aperture zone the exine elements are reduced to a thin foot layer. The intine has three layers with many vesicular-fibrillar components and tubular extensions in intine 1. The symmetry of the pollen grains, shape, type of aperture and ultrastructure of the intine show a remarkable uniformity in the family. Nevertheless there is variety in pollen size, ornamentation and the ultrastructure of the exine. The pollen of Magnoliaceae is an example of an early trend of specialization, and supports the view that Magnoliaceae are not one of the earliest lines in the phylogeny of flowering plants.     

Polyploidy and sterility in relation to sex in Dioscorea alata L. (Dioscoreaceae)

Chromosome numbers and fertility studies of 73 male and 30 female flowering germplasm accessions of Dioscorea alata L. were carried out. All males were tetraploids showing the same chromosome number (n=20 or 2n=40) and were pollen fertile (10.9–96.2%), most of them being highly fertile. Among the female the majority were higher ploids (hexa-and octoploids; 2n=60 and 80) and sexually completely sterile. There were only two tetraploid female accessions which were sexually fertile. Pollination studies revealed that seed sterility in D. alata was due to female sterility associated with the occurrence of higher levels of ploidy. The female sex-limited occurrence of higher polyploidy and sterility observed in D. alata is a curious situation among dioecious higher plants.     

Pollen morphology within the Monodora clade,a diverse group of five African Annonaceae genera

Pollen morphology has played a major role in elucidating infrafamiliar‐level systematics and evolution within Annonaceae, especially within the African genera. The Monodora clade is composed of five genera, Asteranthe, Hexalobus, Isolona, Monodora and Uvariastrum, which are restricted to Africa and contain together c. 50 species. A molecular phylogeny of the family showed that the monophyly of the Monodora clade is strongly supported and that it is part of a larger clade of 11 African genera. In order to support classification a detailed survey was made of the pollen morphological variation within the Monodora clade, using scanning and transmission electron microsopy. For the two most species‐rich genera, Isolona and Monodora, a molecular species‐level phylogeny was used to assess the taxonomic usefulness of the pollen characters. The survey showed a wide range of pollen morphological diversity. The most conspicuous variation concerned the occurrence of monads without a thicker outer foliation in the basal exine layer in Isolona in contrast to tetrads with a thicker outer foliation in Asteranthe, Hexalobus, Monodora and Uvariastrum. At the infrageneric level, Hexalobus, Isolona and Monodora showed the largest diversity, with various pollen types based on tectum morphology. Hexalobus is exceptional with three types within only five species. The pollen types defined in this study are hardly useful in characterizing major groups identified within both Isolona and Monodora, but they do illustrate relationships within smaller groups.     

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

The pollen morphology of Adenandra Willd. (Rutaceae: Diosminae) was investigated to determine its taxonomic significance. Pollen of 27 of the 30 infrageneric taxa (representing 16 of the 18 species) was investigated by LM, SEM and TEM. Adenandra differs from all other Diosminae in having 4-colporate rather than 3-colporate pollen grains. This supports the alleged monophyletic status of the genus. Exine morphology and structure, however, is extremely diverse considering the size of the genus, with eight distinct pollen types and four subtypes being discerned. In some members the grains have uniform macroreticulate, striate or striato-reticulate sculpturing, whereas in others the poles have striate, striato-reticulate or reticulate-perforate sculpturing with various types of reticulate or rugulate sculpturing at the mesocolpia. Taxonomic groupings revealed by the pollen characters suggest elationships between taxa that were not previously apparent. It is suggested that evolutionary diversification among species of Adenandra is often more strikingly reflected by pollen morphology than by macromorphology, hence the eurypalynous state of the genus.     

Pollen morphology of Trigonostemon and its relatives (Euphorbiaceae)

The pollen of Trigonostemon and the related genera Dimorphocalyx, Ostodes, Tritaxis and Jatropha (outgroup) has been studied with light microscopy, and scanning and transmission electron microscopy. The two major pollen types within Trigonostemon correlate well with macromorphological characters. Species belonging to the Trigonostemon reidioides type have pollen with ‘croton pattern’ ornamentation, a pistil with deeply divided stigmas (to at least half the length of the stigma arm) and stamens with a protruding appendage on the connective, while species of the Trigonostemon verrucosus type have verrucate (to almost gemmate) pollen, stigmas that are shortly cleft and stamens without an appendage on the connective. Dimorphocalyx, Ostodes, Tritaxis and Jatropha (outgroup) have similar pollen morphology, while Trigonostemon deviates from these genera in the absence of the ‘vertically’ striate ornamentation on the subunits. Therefore, when compared with an existing phylogeny of the Euphorbiaceae, the pollen characters of Trigonostemon appear to be derived. Moreover, because the ‘croton pattern’ ornamentation itself is widely shared by the ‘inaperturate crotonoids’, the loss of that structure in the Trigonostemon verrucosus type pollen is considered a further apomorphy.     

Pollen morphology in the genus Coffea (Rubiaceae) and its taxonomic significance

The pollen morphology of 38 representative species and varieties within the genus Coffea L. is described. Eight pollen types, placed in two major groups, have been identified on the basis of the number of colpi, colpus characteristics, exine morphology and pollen size. Differences in pollen structures of the species studied do not correspond to present taxonomic groupings, but support a current proposal to place some of the species in the genus Paracoffea Leroy. Among the four sections in the genus, the section Eucoffea contains a high degree of pollen polymorphism.