Seed morphology and its systematic implications for genus Oxytropis DC. (Fabaceae)

In the present study, seeds of 13 Oxytropis DC. species (including two latest synonyms) classified in subg. Oxytropis (sect. Dolichocarpon, sect. Chrysantha and sect. Eumorpha), subg. Euoxytropis (sect. Orobia) and subg. Phacoxytropis (sect. Protoxytropis, sect. Janthina and sect. Mesogaea) from Turkey were examined using light and scanning electron microscopy to evaluate the taxonomic relevance of macro- and micro-morphological seed characters. As a result of the study, species-specific characters have been determined. Seeds exhibit variation in size, shape, surface sculpturing pattern, hilum position and weight. Seeds ranged from 1.61 to 4.04 mm in length and from 1.02 to 2.56 mm in width. Five different seed shapes were recognised, as prolonged semielliptic, reniform, prolonged reniform, quadratic and cardiform, with a length to width ratio ranging from 1.02 ± 0.08 to 1.75 ± 0.27. Rugulate, rugulate–reticulate and lophate sculpturing patterns were observed in the studied species. According to the results, seed characters, such as the size (length, width and the length/width ratio), the shape, the surface sculpturing and the weight appear to have low taxonomic value in distinguishing subgenera, sections and species.     

Seed coat morphology of some Turkish Campanula (Campanulaceae) species and its systematic implications

The seed coat morphology of 10 Campanula species from Turkey, 3 of which are endemic, was studied using scanning electron microscopy. Characteristics of the seeds and their surface morphology were described and compared. Two different types were described based on seed surface features. The Type 1 seed surface was characterized by a reticulate pattern; only C. Olympica belonged to this type. The Type 2 seed surface had a striate testa and was the most common type. It included C. Lyrata subsp. lyrata, C. rapunculoides L. subsp. rapunculoides, C. glomerata L. subsp. hispida, C. involucrata, C. saxonorum, C. persicifolia, C. latiloba subsp. latiloba, C. lactiflora and C. Rapunculus L. var. lambertiana. However, some differences among these species were determined on the basis of their seed surface features at the microstructural level. These features were evaluated as possibly consistent parameters in the delimination of the Campanula taxa studied.     

Reproductive morphology of Sargentodoxa cuneata (Lardizabalaceae) and its systematic implications

The reproductive morphology of Sargentodoxa cuneata (Oliv) Rehd. et Wils. is investigated through field, herbarium, and laboratory observations. Sargentodoxa may be either dioecious or monoecious. The functionally unisexual flowers are morphologically bisexual, at least developmentally. The anther is tetrasporangiate, and its wall, of which the development follows the basic type, is composed of an epidermis, endothecium, two middle layers, and a tapetum. The tapetum is of the glandular type. Microspore cytokinesis is simultaneous, and the microspore tetrads are tetrahedral. Pollen grains are two-celled when shed. The mature ovule is crassinucellate and bitegmic, and the micropyle is formed only by the inner integument. Megasporocytes undergo meiosis resulting in the formation of four megaspores in a linear tetrad. The functional megaspore develops into an eight-nucleate embryo sac after three rounds of mitosis. The mature embryo sac consists of an egg apparatus (an egg and two synergids), a central cell, and three antipodal cells. The pattern of the embryo sac development follows a monosporic Polygonum type. Comparisons with allied groups show that Sargentodoxa shares more synapomorphies with the Lardizabalaceae than other Ranunculales. Characteristics of its reproductive morphology are consistent with the placement of Sargentodoxa as the sister group of the remaining Lardizabalaceae. It does not possess a sufficient number of apomorphic characters to justify its separation into a separate family or subfamily. It is best retained as a member of the Lardizabalaceae.     

Reproductive morphology ofMegaleranthis saniculifolia Ohwi (Ranunculaceae) and its systematic implications

To confirm the taxonomic treatment ofMegaleranthis saniculifolia Ohwi, an endemic genus and species in Korea, we compared its reproductive morphological characteristics with those ofTrollius and other genera within the Ranunculaceae. Although its external morphology might suggest thatMegaleranthis differs fromTrollius, Calathodes, and etc., we found no distinctly different features in this genus. Likewise, previous studies of their pollen structures, chromosome data, and petal morphology have indicated no differences betweenMegaleranthis andTrollius. In fact, related genera share similar characteristics, such as a tetrasporangia anther, glandular tapetum, simultaneous cytokinesis, an anatropous and bitegmic ovule, embryo sac formation of thePolygonum type, exarillate and copious albuminous seed, and several apocarps. Although the unique feature of having both tenuinucellate and crassinucellate ovules simultaneously may initially seem particular toMegaleranthis, it is present in other genera of the same family. Therefore, based on this evidence of reproductive morphology and other information, we suggest thatM. saniculifolia is closely related toTrollius, and should be included within that genus, i.e., asT. chosenensis Ohwi. Nevertheless, we have tentatively placedMegaleranthis within its own monotypic and endemic genus until definitive data become available.     

Seed coat anatomy ofCrossostylis (Rhizophoraceae): its evolutionary and systematic implications

The seed coat structure of all 13 species ofCrossostylis was studied to contribute to an understanding of species delimitation and relationships within the genus. The mature seed coat is relatively uniform and consistently constructed mainly by a well-developed exotesta and a well-developed fibrous exotegmen. The species differ in the thickness of the exotesta and exotegmen, the anatomy of exotestal cells, the presence and absence of persistent mesotesta, and so forth. On the basis of comparisons of these characters, close relationships are suggested in the species groups such as:Crossostylis banksiana andC. cominsii; C. biflora, C. raiateensis andC. multiflora; C. gandiflora, C. sebertii andC. imera; and five species in the Fiji Islands. These relationships except for those of Fijian five species are also supported by cladistics as their common characters are evaluated as synapomorphy. Species-level separation ofC. banksiana, C. pedunculata andC. raiateensis each from the closest species is doubted based on the results of seed coat structure.     

Seed morphology of modern and fossil Ampelocissus (Vitaceae) and implications for phytogeography

Seeds are useful in distinguishing among extant genera of Vitaceae and provide a good basis for interpretation of fossil remains in reconstructing the evolutionary and phytogeographic history of this putatively basal Rosid family. Seeds of Ampelocissus s.l. including Pterisanthes and Nothocissus are distinguished from those of all other vitaceous genera by long, parallel ventral infolds and a centrally positioned oval chalazal scar. Principal component analysis facilitates recognition of four Ampelocissus s.l. seed morphotypes differentiated by dorsiventral thickness, width of ventral infolds, chalazal depth, and degree of dorsal surface rugosity. While these intergrade, their end-member morphologies are distinctive and coincide well with inflorescence morphology, extant geographic distribution, and ecology. Seven fossil morphospecies are recognized. Ampelocissus parvisemina sp. n. (Paleocene of North Dakota; Eocene of Oregon) and A. auriforma Manchester (Eocene of Oregon) resemble extant Central American species; A. bravoi Berry (Eocene of Peru) is similar to one group of Old World extant species; and A. parachandleri sp. n. (Eocene of Oregon) and the three European fossil species A. chandleri (Kirchheimer) comb. n., A. lobatum (Chandler) comb. n., and A. wildei sp. n. (Eocene to Miocene) resemble another group of extant Old World Ampelocissus. All these fossils occur outside the present geographic range of the genus, reflecting warmer climates and former intercontinental links.     

Floral morphology and structure of Phyllonoma (Phyllonomaceae): systematic and evolutionary implications

Phyllonoma, a small tree genus of four species distributed from Mexico to Peru, has been placed in various families (mainly in Saxifragaceae), but now, based on molecular evidence, is placed in a distinct family Phyllonomaceae in Aquifoliales. To better understand the morphological relationships of the genus and family, I studied its floral morphology, anatomy, and vasculature using P. tenuidens. Most of the external and internal floral characteristics were described more than 120 years ago. Although some of them were confirmed, some were substantially revised, mainly those concerning the gynoecial structure. Flowers are small and basically pentamerous, consisting of five sepals, five petals, five stamens, and a gynoecium composed of two carpels usually in transversal position. Comparisons with other Aquifoliales show that Phyllonomaceae share the inferior ovary, epiphyllous inflorescence and epigynous disc nectary with East-Asian Helwingiaceae (Helwingia only), but clearly differ from Helwingiaceae in having glandular trichomes on the sepal margins and a bicarpellate, unilocular gynoecium bearing many ovules on the parietal placentae. Evidence from floral morphology and structure supports the distinctness of Phyllonomaceae and its sister-group relationship with Helwingiaceae. Its floral characteristics suggest that Phyllonomaceae have evolved by adapting to distinct biological habitats in relation to pollination and seed dispersal.     

Sporoderm characters ofTetrameranthus duckei (Annonaceae) and their systematic implications

Ultrastructural and micromorphological investigations of the sporoderm ofTetrameranthus duckei pollen confirm the interpretation of this genus as being isolated within theAnnonaceae.     

Seed morphology in New WorldAntirrhineae (Scrophulariaceae): Systematic and phylogenetic implications

Seeds of 39 species representing all native genera and sections of New WorldAntirrhineae were examined with the Scanning Electron Microscope and Light Microscope. Based on seed shape and surface ornamentation, seven morphological categories are recognized: cristate, tetracostate, foveolate, tumid tuberculate/cristate, circumalate, medusiform, and a miscellaneous category. Most sections and/or generic categories can be characterized by a single morphological type, althoughMaurandya s.l. has several distinctive seed types included within present generic boundaries. The cristate seed type is believed to be the “primitive type” from which the medusiform and tumid tuberculate/cristate types have developed. The tumid tuberculate/cristate type presumably gave rise to the circumalate, tetracostate, and foveolate patterns. The miscellaneous category (represented byLinaria andKickxia) may be phyletically remote from the other New World types. Seeds of the New WorldAntirrhineae mostly seem to be adapted for water dispersal although some adaptations for wind dispersal (e.g. wings, light weight) are evident. Free-hand sections and seed coat clearings indicate that circumalate seeds are of two different anatomical patterns; such structural diversity may be found in the other seed types.     

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.     

Phylogenetic relationships and biogeography of the desert plant genus Fagonia (Zygophyllaceae), inferred by parsimony and Bayesian model averaging

Phylogenetic relationships within Fagonia were inferred from analyses of plastid trnL intron and nuclear ribosomal ITS DNA sequences. Sampling of the genus was nearly complete, including 32 of 34 species. Phylogenetic analysis was carried out using parsimony, and Bayesian model averaging. The latter method allows model-based inference while accounting for model-selection uncertainty, and is here used for the first time in phylogenetic analyses. All species of Fagonia in the Old World, except F. cretica, form a weakly supported clade, and all Fagonia species of the New World, except F. scoparia, are well supported as sister to the Old World clade. Fagonia scoparia, from Mexico, and F. cretica, from Northern Africa, are well supported as sisters to all other Fagonia species. Vicariance-dispersal analysis, using DIVA, indicated that the occurrences of Fagonia in South America and southern Africa are due to dispersals, and also, that the ancestor of Fagonia had a distribution compatible with the boreotropics hypothesis.     

Seed and gland morphology in Euphorbia (Euphorbiaceae) with focus on their systematic and phylogenetic importance, a case study in Iranian highlands

A comprehensive study based on gland and seed micromorphology in Euphorbia (Euphorbiaceae) for species distributed in Iranian highlands is presented. A total of 86 species were studied. The gland structure was examined by direct field observations. Taxonomically important characters of glands were observed and measured: size, texture, shape, color, and horns. For species out of Iran herbarium materials were studied. Seed characteristics were examined using scanning electron microscopy (SEM) as well as dissecting light microscopy. Significant features are: seed size, seed shape, presence of caruncle, shape of caruncle, and seed surface ornamentation. A phylogenetic study using Maximum Parsimony (MP) and Bayesian Inference (BI) was performed based on sequences of nuclear DNA internal transcribed spacers (ITS) for selected species representing the main clades known in Euphorbia and with special focus on the species distributed in Iranian highlands. ITS sequences for 20 accessions representing 19 species are provided for the first time, and 48 accessions of 47 species were used from GenBank. The topologies of both analyses were congruent. The results indicate: (1) four main clades with high supports in subgen. Esula which are appropriate to be recognized at sectional rank. (2) E. larica is nested within clade A including few members of subgen. Rhizanthium and is closely related to sect. Balsamis, which is suggested here to be transferred from subgen. Esula into subgen. Rhizanthium. (3) E. osyridea of the monotypic subsect. Osyrideae is closely related to E. buhsei and to the members of sect. Esula. Tracing morphological characters on the phylogenetic tree shows that several morphological characters, such as seed ornamentation applied in previous subgeneric classification of the subgen. Esula, are homoplasious, but the gland structure and capsule surface characters are more reliable for classification purposes.     

Developmental morphology of the Asian one-leaf plant, Monophyllaea glabra (Gesneriaceae) with emphasis on inflorescence morphology

We examined the developmental morphology of the tropical Asian one-leaf plant Monophyllaea glabra, which is believed to have diverged first in the phylogenetic tree of the genus. The embryo within the seed consists of two cotyledons and a hypocotyl with no shoot or root apical meristems. The endogenous root meristem is formed nearer the hypocotyl end than in other examined Monophyllaea species. One of the cotyledons grows to form the macrocotyledon by means of the basal meristem. The groove meristem arises between the anisocotyledons, shifts toward the macrocotyledon, and is transformed to the inflorescence apex, which produces inflorescence axes in the axils of all ventral bracts of two rows, and secondary inflorescences in the axils of the lower dorsal bracts of the other two rows. The macrocotyledon may act as a ventral bract for the first inflorescence axis at the reproductive stage. This organization suggests that a common ancestor of Monophyllaea and Whytockia with decussate inflorescences diverged in one direction to become Monophyllaea and in another to become Whytockia.     

Seed morphology and its systematic implications for genus Astragalus L. sections Onobrychoidei DC., Uliginosi Gray and Ornithopodium Bunge (Fabaceae)

Forty-eight species of Astragalus L. sections Onobrychoidei DC., Uliginosi Gray and Ornithopodium Bunge (Fabaceae) in Turkey were studied in order to describe and investigate their seed morphologies and to evaluate the diagnostic value of this character using a stereoscopic microscope and a scanning electron microscope (SEM). SEM pictures of seeds were taken as a whole and in detail. The systematic implications of the findings are discussed. Six main seed types were observed. A key to identifying the different seed types is provided.     

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.     

Seed ontogeny of Chamaecrista and its systematic implications in Cassiinae (Leguminosae, Caesalpinioideae)

The genus Chamaecrista originates from the dismembering of Cassia L., which was elevated to the level of subtribe Cassiinae, giving rise to Cassia sensu stricto, Chamaecrista, and Senna. Phylogenetic analysis has suggested that Cassiinae is not monophyletic. The relationships between its genera are not clear, because they sometimes appear close to Chamaecrista and Senna and at other times close to Senna and Cassia. We describe here development of the seeds of three species of Chamaecrista and compare our data with information published in the literature for Cassiinae and non-Cassiinae legumes. The ovules appear well conserved in Cassiinae, differing only in the length of the funiculus. In terms of seed development, the shape of the endosperm haustorium is similar in Senna and Chamaecrista. Chamaecrista shares more characteristics of its mature seeds with Senna than with Cassia. Chamaecrista and Senna have variable seed shapes, the presence of pleurograms, the absence of root cap primordia, a short epicotyl, and the cotyledonary mesophyll tends to be dorsiventral. These seed characteristics place Senna close to Chamaecrista and distance it from Cassia. We also observed characteristics unique to Chamaecrista, for example the occurrence of multiple pleurograms, a partially multiplicative seed coat, and the presence of embryonic colleters.     

菝葜科种皮微结构特征及其分类学意义

在光学显微镜和扫描电镜下对菝葜科Smilacaceae3个属(菝葜属Smilax、肖菝葜属Heterosmilax和Ripogonum属)共53种5变种植物的种子形态及种皮微形态特征进行了研究。结果表明其种子形状为球形、半球形或钝三角形。在扫描电镜下种子表皮纹饰可分为7种类型,即脑纹型、粗脑纹型、网纹型、细网纹型、孔穴型、密孔穴型和细条纹型。根据种皮微形态的特征,对菝葜科内属间和属内组间的关系进行了探讨。种皮形态分析结果支持将Ripogonum属从菝葜科中分离、独立成科,支持将肖菝葜属与菝葜属合并的观点,这与孢粉学和分子证据的分析结果一致;推测肖菝葜属和菝葜属的土茯苓组sect.Coilanthus及草本组sect.Coprosmanthus的多数种类之间亲缘关系较近,菝葜组sect.China和圆锥组sect.Macranthae的大多数种类之间的亲缘关系较为密切,但种皮形态证据不支持Koyama将菝葜属分为6个组的观点。