BOTANICAL AFFINITIES OF SOME TRIPROJECTACITES FOSSIL POLLEN

During Late Cretaceous to Oligocene times, fossil pollen of the Triprojectacites group (also known as Aquilapolles or triprojectates), comprised a temporally and environmentally distinctive element of palynofloras in eastern Asia and western North America. Several species of this group serve as biostratigraphic index fossils for this interval. Using electron microscope and numerical analyses, primarily of North American triprojectate pollen, it is possible to recognize the presence of three distinct subgroups. One group, corresponding to the fossil genus Mancicorpus, has no morphologically close modern representative. Characters resembling those in Santalaceae (Santalales, Rosidae) occur in the second group, which is represented by a previously underscribed fossil triprojectate genus. The third group contains retipilate, isopolar pollen and strioreticulate, isopolar pollen. Forms exhibiting the latter morphology are commonly assigned to the genus Integricorpus, while the retipilate morphology characterizes another underscribed genus. This third group may have some phylogenetic connection to Apiaceae (Apiales, Asteridae). Pollen of some other extant families exhibits triprojectate features, although no close fossil representative can be presently identified. The triprojectate morphology is thus interpreted in modern and fossil forms as resulting from convergence rather than close phylogenetic relationships at the group level.     

Pollen morphology of the tribe Rhinantheae (Orobanchaceae) and its systematic significances

Pollen morphology of 36 species representing 14 genera within the tribe Rhinantheae in the family Orobanchaceae was studied and illustrated with light microscopy (LM) and scanning electron microscopy (SEM). Five major pollen types were recognized on the basis of exine ornamentation. Within these major types, minor types (subtypes) were distinguished based on exine surface pattern, size, shape, amb form, colpi and colpus membrane. These types and subtypes are as follows: type I. retipilate: subtype Ia. regular retipilate: (1) pollen size < 27 μm, (2) pollen size > 27 μm, subtype Ib. irregular retipilate; type II. verrucate: subtype IIa. macro-verrucate, subtype IIb. verrucate, subtype IIc. sparse verrucate; type III. retirugulate; type IV. granulate; type V. micro-reticulate. A key to pollen morphology of genera studied within the Rhinantheae was made based on pollen morphology from our study and earlier work. Combining with other sources of information on the Rhinantheae, the systematic relationships of this tribe are discussed. Rhinantheae pollen displays considerable variation between genera and species, with taxonomically significant characters at genus and species level. Palynological characteristics provide evidence for interpreting the conflicting views concerning the “Pterygiella Complex”. The evolutionary trend in exine sculpture of Rhinantheae could be proposed, namely that retipilate sculpturing which is the most widespread type is more primitive than the other types (such as foveolate, granulate, regulate, reticulate, retirugulate and verrucate). The pollen data in present study and the view of Hong (1986), as well as the molecular data from Bennett and Mathews (2006) indicated that Asia and related regions were likely to the origin centre of the tribe Rhinantheae.     

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

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

Pollen ultrastructure in <Emphasis Type="Italic">Aristolochia manshuriensis</Emphasis> and <Emphasis Type="Italic">A. contorta</Emphasis> (Aristolochiaceae)

Pollen ultrastructure has been studied in two relict and rare species of the genus Aristolochia, A. contorta Bunge and A. manshuriensis Kom. (Aristolochiaceae). Both species have inaperturate, spheroidal, sometimes distally monocolpate or distally bicolpate pollen grains. The equatorial and polar axes of pollen grain in A. manshuriensis are 48.5 and 44.0 μm, respectively. The percentage of defective pollen grains in A. manshuriensis is 3.4%. The fossulate, perforated exine is up to 2.3 μm in thickness; the sexine and the nexine are almost equal in thickness. In A. contorta, the equatorial axis of pollen grain is 36.6 μm: the defectiveness percentage, 24.5%. The exine is verrucate, up to 0.3 μm in thickness, while the sexine is two to three times thicker than the nexine. The pollen germination experiments have shown that pollen of A. manshuriensis, in contrast to A. contorta, can germinate in 10–20% sucrose at 22°С. These data and the high percentage of pollen defectiveness in A. contorta indicate that the androecium function in this species is reduced. The reduction of the androecium function is evidenced by a small amount of pollen grains in anthers or empty anthers and a high percentage of defective pollen grains.     

New insights into the pollen morphology of the genus Mutisia (Asteraceae, Mutisieae)

The pollen morphology from 67 collections representing 52 species of Mutisia was analyzed by means of light, scanning electron and transmission electron microscopy. Pollen of Mutisia is comparable to that of the more widespread type found in Mutisiinae. The pollen grains of the genus are characterized by the spheroidal-subprolate to prolate shape, large size, tricolporate aperture, with mesoaperture, exine Mutisia type, with sculpturing microechinate to rugulate or microechinate-rugulate. This basic pollen type is very variable in size, endoaperture shape, and exine thickness of the pollen grains. A new type of arrangement of columellae was observed in pollen of Mutisia section Mutisia (e.g., M. stuebelli, M. microphylla, and M. sodiroi). It is suggested that it might have harmomegathic significance. The pollen morphology and polymorphism are discussed with regard to functional mechanisms and possible evolutionary significance.     

Pollen morphology and ultrastructure of the Bennettitales: In situ pollen of Cycadeoidea

The micromorphology and ultrastructure of in situ pollen from Cycadeoidea dacotensis are described from permineralized specimens collected from the Lower Cretaceous of North America. Pollen grains are ovoid and relatively small, averaging 25 μm in length and 12 μm in width. Grains are monosulcate with the exine typically invaginated in apertural regions. Exine ornamentation ranges from punctate to psilate. The exine averages 0.73 μm in thickness and is composed of a light-staining sexine and a dark-staining nexine. The sexine consists of a thin, homogeneous tectum, typically with a well-defined inner boundary, and a thicker granular infratectum. The infratectal granules are relatively uniform in size, however, variation occurs in the arrangement of granules. In some grains, the sexine appears homogeneous because there is little lacunal space between the individual granules. The granular infratectum is in direct contact with the underlying nexine. The nexine is uniform in thickness in both apertural and nonapertural regions, and it lacks lamellae throughout. Pollen morphology and ultrastructure are compared with those of the bennettitalean genus Leguminanthus and the dispersed genus Monosulcites. In addition, the fine structure of Cycadeoidea pollen is compared to that of the gymnosperm groups with which the Bennettitales are regarded to be most closely related, including Gnetales, Pentoxylales, and Eucommiidites-type pollen-producing plants.     

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.     

Studies on Pollen Morphology of the Genus Corylopsis

The present paper describes the pollen morphology of 11 species of thc genus Corylopsis (Hamamelidaceae). The pollen grains were all examined with light microscope and scanning electron microscope. Pollen grains of the genus are prolate, spheroidal or subprolate. Three size types are recognized: (l) 18-23 μm, found in Corylopsis omeiensis Yang, C.gotoana Makino, C. himalayana Griff and C.platypetala Rehd. et Wils. (2) 24-32μm, found in C. sinensis Hemsl., C.willmottiae Rehd. et Wils., C.veitchiana Bean, C. brevistyla H.T.Chang, C.multiflora Hance, C.pauciflora Sieb. et Zucc. and C.spicata Sieb. Zucc. (3) 34-39μm, found in C. wilsonii Hemsl. and C. cordata Merrill. Pollen grains are 3-colpate and 4-colpate (only infrequently found in Corylopsis sinensis Hemsl.); the exine is 2-layered, 1.25-2.50μm thick, distinctly reticulate, sexine almost equal to nexine in thickness. From the pollen morphology the genus is considered as a natu-ral group.     

中国酸模属植物花粉形态研究

利用光学显微镜、扫描电镜对１５种,透射电镜对２种国产酸模属植物的花粉进行观察与研究。本属花粉为球形或近球形,大小为１５～４０×１８～４０μｍ。具３孔沟、４斜孔沟、散孔沟、散沟等;沟或外沟狭长,几达两极,内陷或不内陷,内孔圆形、稍横长或纵长;表面具皱波状纹饰;外壁厚１．０～３．５μｍ,外层明显厚于内层。在透射电镜下,花粉外壁分３层：覆盖层最厚,具穴状或穿孔,颗粒状或刺状纹饰,内面均匀;柱状层明显,厚,小柱为钝圆锥体或圆柱状,基层和内层不分,最薄,但在萌发孔处明显增厚。根据花粉形态,本属花粉可以分成７个组,它们是：小酸模组、滨海酸模组、酸模组、土大黄组、荒地酸模组、巴天酸模组和钝叶酸模组。对萌发孔分布规律的研究,基本支持Ｗｏｄｅｈｏｕｓｅ（１９３１）的观点,同时据花粉萌发孔演化的规律,认为本属花粉可能是蓼科花粉由少数萌发孔向多数萌发孔演化的过渡类型。     

Pollen morphology and ultrastructure of Marathrum schiedeanum (Podostemaceae)

The Podostemaceae, or river-weeds, comprise 46 genera and 270 species of dicots and are the largest family of strictly aquatic angiosperms. Despite the large size, specialized habitats, and enigmatic morphology of the family, relatively little is known about the palynology of Podostemaceae. In the current paper, pollen morphology and ultrastructure of Marathrum schiedeanum are described. Pollen grains are relatively small, spheroidal, and tricolpate to spiraperturate. The exine has a microechinate ornament, a tectate-granular sexine and a relatively thick nexine in non-apertural regions, and a semitectate sexine and thinner nexine in apertural regions. Although aperture variation occurs in the family, this is the first report of the spiral aperture type in Podostemaceae. The spiraperturate condition appears to be derived in river-weeds, as does the granular pollen wall, which represents a reduction of the typical columellae found in eudicots.     

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.     

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

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

The nature of intraspecific and interspecific genome size variation in taxonomically complex eyebrights

Background and aimsGenome size varies considerably across the diversity of plant life. Although genome size is, by definition, affected by genetic presence/absence variants, which are ubiquitous in population sequencing studies, genome size is often treated as an intrinsic property of a species. Here, we studied intra- and interspecific genome size variation in taxonomically complex British eyebrights (Euphrasia, Orobanchaceae). Our aim is to document genome size diversity and investigate underlying evolutionary processes shaping variation between individuals, populations and species.MethodsWe generated genome size data for 192 individuals of diploid and tetraploid Euphrasia and analysed genome size variation in relation to ploidy, taxonomy, population affiliation and geography. We further compared the genomic repeat content of 30 samples.Key resultsWe found considerable intraspecific genome size variation, and observed isolation-by-distance for genome size in outcrossing diploids. Tetraploid Euphrasia showed contrasting patterns, with genome size increasing with latitude in outcrossing Euphrasia arctica, but with little genome size variation in the highly selfing Euphrasia micrantha. Interspecific differences in genome size and the genomic proportions of repeat sequences were small.ConclusionsWe show the utility of treating genome size as the outcome of polygenic variation. Like other types of genetic variation, such as single nucleotide polymorphisms, genome size variation may be affected by ongoing hybridization and the extent of population subdivision. In addition to selection on associated traits, genome size is predicted to be affected indirectly by selection due to pleiotropy of the underlying presence/absence variants.     

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.     

A Study of Pollen Morphology of Sparganium from China

Pollen morphology of 8 species in genus Sparganium from China has been studied. All of them were examined under the light microscope and SEM. It may draw the following conclusions: The characteristics of pollen grains in genus Sparganium are very similar. They are spheroidal, subspheroidal or sometimes slightly angular in shape, 21.8–38.3μm in diameter, monoporate. Porus is slightly sunken, its margin distinct or indistinct, and some granules on the porus membrane. The thickness of exine is 1.7–2.6μ m. The sexine is about as thick as nexine, sometimes thinner than nexine. It is difficult to distinguish one another under the light microscope, but may be seperated under SEM on the basis of the shape or size of lumina and width or height of muff. In some species small excrescenses can be observed under SEM. Some taxonomists (Rendle, 1953) considered that Sparganium is closely related to Pandanus, but others (Hutchison 1934, Takhtajan 1969) to Typha. According to the data of pollen morphology, Sparganium is more close to Typhaceae than to Pandanaceae, thus we agree to put it into family Typhaceae.     

COMPARATIVE POLLEN MORPHOLOGY AND TAXONOMIC AFFINITIES IN CYCADALES

Scanning and transmission electron microscopy of pollen grains of 29 species, representing the ten extant genera of Cycadales, has provided valuable insight into their relationships. Pollen grains of these taxa are boat-shaped, monosulcate, and bilaterally symmetrical. They range from narrowly to widely elliptical or subcircular when viewed distally, and have an exine surface of psilate, foveolate, or fossulate. Pollen wall ultrastructure of Cycadales is typically tectate with alveolate∗∗∗spongy exine. The nexine is laminated in all genera. Nexine 1 (footlayer) is present in most species as a thin and often discontinuous layer. There is consistent variation in thickness of the sporoderm layers among the genera but relative uniformity within them. Pollen characteristics are well correlated with macro- and micromorphological features, chromosome numbers, geographical distribution, and postulated pollination mode. A close affinity between Encephalartos, Lepidozamia, and Macrozamia is recognized. Pollen characteristics of the genus Bowenia show some similarity with those of the latter group. Except for two species of Macrozamia which are narrowly elliptic, all of the genera have widely elliptic pollen and share a psilate exine surface and the thinnest sexine with nearly identical arrangement of alveoli. Pollen grains of the species in the genus Dioon exhibit a unique morphology but are more similar to Stangeria than they are to those of taxa in Zamiaceae. The circular outline of the grains and the foveolate exine surface are characters shared by these two genera, but several morphological features distinguish Dioon from Stangeria. Ceratozamia and Zamia share a widely elliptic shape, foveolate exine surface and nearly identical sexine, as well as morphological features and chromosome numbers. They differ from Microcycas in sexine thickness, gross morphology and chromosome numbers. The pollen grains of Cycas circinalis and C. revoluta differ in size and structure of the sexine from all other genera and from each other, substantiating their distinct subgeneric delimitations.     

Pollen morphology in delimiting subgenera and species of the genus <Emphasis Type="Italic">Cybianthus</Emphasis> s.l. (Myrsinoideae–Primulaceae)

Pollen grains of species of the subgenera of Cybianthus s.l. were analyzed in order to determine if pollen morphology is consistent with the currently accepted infrageneric classification, and if it can provide characters useful for distinguishing species or groups of species. Acetolyzed pollen grains were analyzed, measured, described and photomicrographed, while non-acetolyzed pollen grains were analyzed using scanning electron microscopy. The pollen grains of all species investigated were isopolar monads, small- to medium-sized, subprolate or prolate, 3-colporate, and rugulate or rugulate-perforate. The different analytical methods (statistical analyses, exploratory analysis of the data and cluster analysis) used demonstrated that pollen morphology is informative in delimiting this genus. However, the pollen characters are generally quite uniform among the subgenres of Cybianthus s.l. Thus, the use of the pollen attributes in the separation of the subordinate species to different subgenera is not effective. Pollen morphology did not give support to current subgeneric systematics, but it did support the genus Cybianthus s.l.     

Pollen morphology of Microstachys (Euphorbiaceae) with emphasis on neotropical species

Abstract

In Euphorbiaceae tribe Hippomaneae is the pantropical genus Microstachys, comprised by 24 species, mostly from Brazil. Palynological studies with several representatives of the genus are scarce. The purpose of this study was to describe the pollen morphology of Microstachys and find possible differentiating characteristics between its species. Ten species of the genus were analysed, represented by 21 herbarium collections. Their pollen grains were acetolysed, measured and photographed under light microscopy (LM) and scanning electron microscopy (SEM). The analysed species possess pollen in monads, of small to medium size, isopolars, amb distincticly trilobed, oblate-spheroidal, prolate-spheroidal or subprolate, tricolporate with narrow and very long colpi, almost fused at the poles sometimes forming an invagination with tapering ends and margo of the colporus wide to very wide and psilate; endoapertures lalongate to very lalongate with tapering ends, presence of a narrow costae distinct in the endoaperture; exine microreticulate homobrochate, walls with simple columella; at the mesocolpus the sexine separates from the nexine forming cavea; and sexine always thicker than the nexine. Microstachys is a stenopalynous genus, despite its species varying in pollen size, shape and width of the cavea, margo of the colporus, and length of the endoapertures, demonstrating taxonomic value, and indicating a contribution towards future systematic analyses.     

POLLEN MORPHOLOGY OF THE CHLORAEINAE (ORCHIDACEAE: DIURIDEAE) AND RELATED SUBTRIBES

Pollen organization and morphology of the South American Chloraeinae (Orchidaceae) was examined by scanning electron microscopy and compared with that of the remainder of the otherwise Australasian Diurideae. All five genera of the Chloraeinae, Bipinnula, Chloraea, Codonorchis, Gavilea, and Geoblasta, and at least one genus from each of the other subtribes were sampled. The Australasian Diurideae are diverse in pollen organization and morphology. The two genera of the Acianthinae, Corybas and Acianthus, have very different pollen and their classification is questioned. Monad pollen organization of Pterostylis (Pterostylidinae) is reinterpreted as primitive and not secondarily derived. Pollen of the Chloraeinae is uniform in exine morphology and organization. Most species sampled have reticulate pollen which tends to be foveolate distally. The basic pollen unit of all Chloraeinae is the tetrad, except Codonorchis which possesses monads. Pollen morphology and organization of the Chloraeinae is most similar to the Caladeniinae, which supports the contention that the Chloraeinae including Codonorchis should be retained in the Diurideae.