Pollen morphology of the genera Polygonum s. str. and Polygonella (Polygoneae: Polygonaceae)

The pollen of 30 taxa (27 species, one subspecies and two varieties) in two genera, viz Polygonum s. str. and Polygonella was investigated with LM and SEM, and some selected taxa with TEM. In all genera investigated the pollen is prolate to spheroidal, and the aperture is mostly tricolporate, rarely panto-hexacolporate (especially Polygonum section Polygonum). The exine sculpturing pattern is the most variable feature. Three types of exine can be recognized. Type 1 (Avicularia-Type, sensu Hedberg) - All species of section Polygonum and section Tephis share the smooth tectate exine with spinules, sometimes the surface is more or less rough (Polygonum afromontanum in section Tephis). Type 2 (Pseudomollia-Type, sensu Hong) - Pollen of Polygonum molliaeforme (section Pseudomollia) has the exine, which is verrucose on both poles and nearby the mesocolpium, and mostly psilate around the ectoaperture. Type 3 (Duravia-Type, sensu Hedberg) - Pollen grains of Polygonum section Duravia and Polygonella have the exine which is semitectate-reticulate at the mesocolpium and the poles, and rugulate/reticulate or sometimes foveolate with microspinules around the ectoapertures. The pollen grains in four taxa (viz Polygonum section Pseudomollia, P. section Duravia and genus Polygonella) have a well-marked dimorphism of the ektexine, which is considered to be a synapomorphic condition. The differences of pollen grain between the genus Polygonella and Polygonum section Duravia are almost non existent and clearly interrelated. It is therefore postulated that the similarity in pollen of both taxa is not the result of convergency, but is interpreted as a homology. It is noteworthy that the pollen of Polygonum molliaeforme (section Pseudomollia) appears as intermediate between the Avicularia-type and the Duravia-type, and is well supported the value of separated section for its own. Additionally, in TEM, some exine ultrastructures (e.g. columellae, foot layer, endexine) appear to be valuable characters for comparison between/among taxa. The systematic potentialities of the pollen data of the studied taxa at various systematic levels are also discussed.     

Pollen morphology of Justicia L. (Acanthaceae) from Thailand and its taxonomic value

The acetolysed pollen of 33 species of the genus Justicia in Thailand is investigated using both light and scanning electron microscopy. The pollen of the genus is characterised as being isopolar, bilaterally or radially symmetrical, with mostly prolate or prolate spheroidal shape. Based on characters with high factor loading in the cluster analysis and principal components analysis, the studied species are divided into two major groups; one with 3-colporate with six pseudocolpi and the other with 2-porate or 2–3-colporate with aperture areas. The pollen morphology of each group is described and illustrated. Pollen characters are useful for taxonomic delimitation and relationships among Thai species.     

Comparative pollen morphology of Turkish species of <Emphasis Type="Italic">Petrorhagia</Emphasis> (Caryophyllaceae) and its systematic implications

Pollen morphology of 10 taxa from Turkey, four of them endemic, belonging to the genus Petrorhagia (Caryophyllaceae), P. alpina subsp. alpina, P. alpina subsp. olympica, P. cretica, P. dubia, P. hispidula, P. lycica, P. pamphylica, P. peroninii, P. prolifera and P. saxifraga, has been investigated using light (LM) and scanning electron (SEM) microscopy. Pollen morphology differences among these taxa have been determined. The pollen type of investigated taxa is polyporate (12–22-porate), sphaeroidal, the exine exhibits a tectate structure and a microechinate ornamentation. Petrorhagia alpina subsp. alpina has the smallest pollen grain diameter (18.57 μm) and P. dubia the largest one (37.80 μm). The number of pores ranges from 12 (in P. dubia and P. saxifraga) to 22 (in P. alpina subsp. alpina and P. alpina subsp. olympica and P. hispidula) with a minimum pore diameter of 2.37 μm in P. alpina subsp. alpina and a maximum pore diameter of 4.23 μm in P. peroninii. The exine thickness ranges from 1.73 μm in P. saxifraga to 3.78 μm in P. pamphylica. In this study, the systematic implications are discussed in the light of palynological results.     

CONTRIBUTION OF POLLEN MORPHOLOGY TO SYSTEMATICS OF COLLOMIA (POLEMONIACEAE)

Pollen morphology of 14 species of Collomia (Polemoniaceae) was examined by light microscopy, and by both scanning and transmission electron microscopy. Four distinct pollen types were observed which are based principally upon 1) shape, number and distribution of apertures, and 2) surface sculpturing: Type 1—zonocolporate with striate ridges; Type 2—zonocolporate with striato-reticulate ridges; Type 3—pantoporate with radiate ridges; Type 4—pantoporate with irregularly reticulate ridges. Evaluation of pollen morphology reveals considerable discrepancy with respect to presently accepted sectional classification. Collomia grandiflora of sect. Collomia has a pollen type similar to that of members of sect. Collomiastrum and is now interpreted as representing an independent evolutionary line derived from the latter section. Collomia diversifolia of sect. Courtoisia has a pollen morphology similar to that of sect. Collomia. whereas C. heterophylla of the same section possesses pollen unique within the genus. This last pollen type shows close similarity to the pollen of members of Polemonium, Gilia, Leptodactylon, and Ipomopsis. Pollen of C. tinctoria and C. tracyi of sect. Collomia are anomalous within Polemoniaceae. No significant difference in exine stratification was discernible among the four pollen types.     

POLLEN MORPHOLOGY AND THE RELATIONSHIPS OF SIMMONDSIA CHINENSIS TO THE ORDER EUPHORBIALES

Pollen from Simmondsia chinensis (Simmondsiaceae) was examined in LM, SEM, and TEM. The pollen is shed as monads, triangular in shape in polar view, with a 3-porate aperture type in which the pores are large and poorly defined. The tectum is irregularly scabrate, sometimes forming minute “islands” topped with spinules. In thin section, the endexine is thickened and lamellate in the aperture regions, and narrow in the mesoporus; the foot layer is well-defined but noticeably thicker in the mesoporus; and thin columellae support an essentially complete tectum. The pollen of four genera, Buxus, Pachysandra, Sarcococca, and Styloceras, from the Buxaceae to which Simmondsia has been assigned by some authors, was also examined and illustrated. The pollen morphology of two families frequently aligned with Simmondsiaceae, Euphorbiaceae and Pandaceae, is briefly discussed. For the most part pollen morphology supports the treatment of Simmondsia as a monotypic family, Simmondsiaceae.     

Pollen morphology of Chamaebuxus (DC.) Schb., Chodatia Paiva and Rhinotropis (Blake) Paiva (Polygala L., Polygalaceae)

Polygala L. is a large and highly diverse genus with complex taxonomy, but pollen morphological information for this taxon is scarce. In the present study, pollen characters have been used to assess the taxonomic delimitation and phylogenetic relationships of three newly established subgenera of Polygala: Chamaebuxus, Chodatia and Rhinotropis (sensu Paiva). The pollen morphology of 22 species has been examined using light microscopy and scanning electron microscopy of acetolysed material. The pollen of 15 of the species is examined for the first time. The pollen grains are isopolar, radially symmetrical, tectate and, typically, polyzonocolporate with numerous colpi running parallel to the polar axis, and an endocingulum around the equator. Two pollen types can be distinguished: Type I, which includes species belonging to Rhinotropis, and Type II, which includes species from Chamaebuxus and Chodatia. The two pollen types are described and the pollen of the three studied subgenera is illustrated. Despite the low infrageneric morphological diversity observed within the genus Polygala, quantitative characters of pollen grains support the current classification of the subgenera Chamaebuxus, Chodatia and Rhinotropis, and reveal a closer relationship between the first two taxa. Pollen characters are shown to be a useful and informative tool for assessing taxonomic position and phylogenetic relationships within Polygalaceae, especially at higher taxonomic levels.     

Pollen morphology of the Pavetteae (Rubiaceae,Ixoroideae) and its taxonomic significance

Pollen grains of the tribe Pavetteae (Rubiaceae, subfamily Ixoroideae) are examined using LM and SEM. Grains are 3‐ or 4‐colporate and (semi‐) tectate (in one Versteegia species atectate). Sexine patterns vary between perforate, microreticulate, reticulate, rugulate and striato‐reticulate. Supratectal elements are sometimes present. The variation in pollen morphology in the Pavetteae allows to recognize seven pollen types, the distribution of which is useful to evaluate generic delimitations and relationships within the tribe. Pollen characters corroborate the close relationships between the genera Coleactina, Dictyandra and Leptactina and between Homollea, Homolliella and Paracephaelis. All the genera of the tribe proved to be stenopalynous (the species examined possess the same pollen type), except Pavetta, Rutidea, Versteegia and Tarenna which are eurypalynous. In the huge genus Pavetta the existing infrageneric classification is supported pollen morphologically. Pollen morphology further indicates that the genus Tarenna is badly delimited and strongly in need of a revision. The small genus Versteegia is in need of further taxonomic and palynological study to understand the pollen morphological variation encountered here. At a higher rank, pollen morphology also does not contradict the recent division of the Pavetteae in the Ixoreae (a stenopalynous tribe with presumably primitive pollen) and the Pavetteae sensu stricto (eurypalynous).     

Systematics ofJusticia sect.Pentaloba (Acanthaceae)

The transfer of the four taxa ofSiphonoglossa sect.Pentaloba toJusticia is proposed. It is shown that the taxa of this section were placed inSiphonoglossa primarily because of a single-character phenetic relationship and that they correctly belong inJusticia. In addition to morphology, data from cytology and flavonoid chemistry are also presented that support this intergeneric transfer. A key to the taxa and a detailed taxonomic treatment of the section are provided.     

Die pollenmorphologie der afrikanischen buxus- und notobuxus-arten (buxaceae) und ihre systematische bedeutung

Pollen grains of 8 African species of Buxus and Notobuxus were examined by light and scanning electron microscopy. Three pollen types are recognized based on the exine sculpture and the evolutionary trends of the pollen characters are discussed. Pollen morphology shows the existence of three major systematic groups. These results support the previous classification of an African section Probuxus with two subsections and speak in favor of the generic rank of the Notobuxus species. Together with anatomical and taxonomical data pollen morphology and its evolutionary trends lend support to the suggestions that the centre of origin of the genus Buxus might have been in Africa, while secondary centres of evolution have developed in the neotropical and Eastasian region.     

囊萼紫草属与滇紫草属花粉形态比较研究

本文借助光学显微镜和扫描电镜研究了囊萼紫草属3种和滇紫草属12种植物的花粉形态。囊萼紫草属的花粉为哑铃形或茧形,中等大小,P／E比为1．6一1．67,三孔沟,内孔横长;具小刺状纹饰。滇紫草属的花粉为近长球形或近卵球形,P／E为l—1．23;三孔沟或三合沟孔,内孔一般纵长,具皱波状纹饰,在皱波上具密集的小瘤或微颗粒。从花粉形态的角度,本文支持把囊萼紫草属从滇紫草属(广义)中分离出来的观点。值得注意的是,在滇紫草属的花粉中首次观察到了一种比较少见且特化的花粉即单极三合沟孔的花粉。     

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.     

Distyly and pollen dimorphism in<Emphasis Type="Italic"> Damnacanthus</Emphasis> (Rubiaceae)

We document distyly in Damnacanthus (Rubiaceae) and investigate pollen morphology in the genus. Within nine taxa in Japan (four species, four varieties and one form), five show distyly. Non-distylous taxa of Damnacanthus always have a long style and short stamens. We examined the pollen morphology of all taxa of Damnacanthus in Japan. The pollen grains of Damnacanthus are 3- to 5-orthocolporate or sometimes loxocolporate. Polymorphic pollen is sometimes observed even within a single flower. Pollen dimorphism is associated with distyly; pollen grains from short-styled (thrum) flowers are significantly larger than those from long-styled (pin) flowers, and the muri are smooth in pin flowers whereas the pollen grains of thrum flowers have minute granules on the top and/or on the sides. In non-distylous taxa, in which the style is longer than stamens, the muri are always provided with minute granules. Interestingly, the pollen grains with granules in the non-distylous taxa contrast with the pollen of the distylous taxa, because granulate pollen grains are found only in thrum flowers. Although Damnacanthus is a stenopalynous genus, the number of colpi is useful for delimiting taxa. Based on palynological characters, we conclude that Damnacanthus should be revised. Electronic Publication     

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 and evolution in the genus Orobanche L. s.l. and its allied genera (Orobancheae/Orobanchaceae) in Turkey

The pollen morphology of 120 samples of the tribe Orobancheae, representing four genera and 40 species of Cistanche, Diphelypaea, Orobanche and Phelipanche native to Turkey, has been studied by light and scanning electron microscopy. Pollen of the tribe Orobancheae is typically isopolar, radially asymmetrical, oblate spheroidal or prolate and belongs to one of the following basic pollen types: inaperturate, tricolpate and syncolpate. Also, pollen heteromorphism is widespread among pest species of Orobanche and Phelipanche. Pollen characters display considerable variation among genera and species, but some characters are significant at the levels of genera and sections. The results of the SEM study show that there are various exine ornamentation types in these genera. The obtained results support the division of the traditionally circumscribed genus Orobanche sensu lato into two genera, Phelipanche and Orobanche. The pollen morphological features are discussed on the basis of molecular phylogeny of the taxa.     

Pollen morphology of Astragalus section Hymenostegis (Fabaceae) and evaluation of its systematic implications

Astragalus is with nearly 3000 described species the largest genus of flowering plants. So far analyses of pollen characters have only been conducted for a few species of the groups within the genus. Here we analyse pollen grains of 22 species representative for Astragalus section Hymenostegis using scanning electron microscopy. We found the basic shape of the pollen grains to be oblate-spheroidal and apertures to be tricolpate as for other eudicots. The sculpturing pattern of the exine is micro-reticulate. Pollen grains show low morphological variation among different species of this section, but differences occur between sections of the genus. We conclude that the vast morphological differentiation that occurred during the rapid radiation of section Hymenostegis was not accompanied by comparable differentiation in pollen morphology.     

Pollen morphology of the genus Euphrasia L. (Orobanchaceae) from Pakistan and Kashmir and its taxonomic significance

Pollen morphology of 25 species of the genus Euphrasia L., belonging to the family Orobanchaceae from Pakistan has been examined by light and scanning electron microscope. It is a stenopalynous genus. Pollen grains are usually radially symmetrical, isopolar, tricolpate, sub-prolate to prolate-spheroidal rarely oblate-spheroidal. Sexine slightly thinner than nexine or as thick as nexine. Tectum coarse to fine retipilate or rugulate–retipilate. On the basis of exine ornamentation three distinct pollen types viz., Euphrasia foliosa, Euphrasia multiflora, and Euphrasia incisa are recognized. Pollen morphology is significantly helpful at specific level within the pollen type.     

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 MORPHOLOGY OF THE EUCHARIDEAE (AMARYLLIDACEAE)

Eucharis, Caliphruria, and Urceolina form a monophyletic group of petiolate-leaved, Neotropical Amaryllidaceae ecologically specialized to the understory of primary tropical rain forest below 2,000 m elevation. Pollen morphology of the three genera is surveyed. Pollen grains of all species of Eucharis, Caliphruria, and Urceolina are boat-shaped elliptic, monosulcate, heteropolar, and bilateral in symmetry. Exine sculpturing is semitectate-columellate and reticulate in all species examined. A transformation series in reticulum coarseness and pollen grain size is described. The large pollen grain with coarse reticulum of most Eucharis species is considered ancestral. The fine reticulation of Caliphruria is considered derived and the exine morphology of Urceolina is intermediate. Both of these genera have medium-sized pollen grains. Exine dimorphism common to all Urceolina, but rare in Eucharis and Caliphruria, may be symplesiomorphous among those taxa exhibiting this morphology. The three genera are largely uniform in pollen grain ultrastructure, with completely ektexinous exines. Pollen grain size in Eucharis is not closely correlated with style length. Several wide-ranging species show considerable intraspecific variation in pollen size. Parallelisms in pollen grain evolution among related tribes of Neotropical Amaryllidaceae are discussed.     

An Ultrastructural Study of Pollen Wall Ontogeny in Silene Pendula

Pollen of 41 species representing all seven genera of the Neottieae were examined by scanning electron microscopy. Except for Lecanorchis, the genera of the Neottieae constitute a natural group based on pollen morphology. Pollen occurs as single grains in the primitive species and in tetrads in the other species. Most grains are monoaperturate, porate or tenuate, and the tetrads often have irregularly shaped grains. Exine structure varies from tectate-perforate to semitectate. Lecanorchis is anomalous among the Neottieae in that it has 0–5, sunken, relatively small pores. Pollen morphology of this genus indicates that it is probably more closely related to the Gastrodieae than the Neottieae. There are at least four basic phyletic units in the Orchidaceae: the Neottioideae, Apostasioideae, Cypripedioideae, and Epidendroideae. These groups are distinguished by the presence of monads in at least their more primitive members and by their unique pollen types.     

Pollen morphology of Stachys (Lamiaceae) in Iran and its systematic implication

Pollen grains of 30 taxa of the genus Stachys (29 spp. and one subsp.), representing 9 of the currently recognized sections and 1 species of the closely related genus Sideritis (Si. montana) distributed in Iran were examined by light and scanning electron microscopy. Twenty-eight taxa are studied for the first time under aspects of pollen morphology. The basic shape of the pollen grains in most taxa studied is prolate-spheroidal, but subprolate, spheroidal and oblate-spheroidal pollen grains can also be found in few species. The grains are usually tricolpate (the amb triangular), but also tetracolpate (the amb circular to more or less square) in some species (S. iberica, S. atherocalyx and Si. montana). The surface is microreticulate (the frequent type), reticulate, perforate, foveolate-psilate or foveolate. The lumina are separated by smooth or sinuate muri which make them polygonal, more or less rounded and elongate. Major pollen morphological features of the taxa studied are compared and discussed on the basis of taxonomical concepts. In some cases, these characters are useful in delimitation of formerly introduced sections while they mostly provide further characters in separating related species from each other. For example, all members of S. sect. Aucheriana are characterized by elongated lumina. Based on the oblate-spheroidal shape of its pollen as well as tetracolpate aperture type, the results of the present study confirm sect. Pontostachys as including S. angustifolia, S. iberica, S. sparsipilosa as well as S. atherocalyx. Our results also suggest that although some species like S. fruticolosa and S. lavandulifolia are morphologically well characterized, they cannot be separated from other species of Stachys based on pollen morphology.