Systematic implication of seed micromorphology in Arenaria (Caryophyllaceae) and allied genera

Variations in seed micromorphology of 64 taxa of Arenaria and allied genera were examined using scanning electron microscopy (SEM) in order to evaluate their diagnostic value for systematic studies in the genus and providing additional evidence on delimitation of natural groups. Significant diversity was found in seed coat morphology both among and within genera. Size, color, and shape of seeds appear to be of low importance at species level but provide useful evidence at generic rank. Cerastium and Stellaria are well distinguished from Arenaria by serrate seed margins and solid columellae on testa cells. The sculpturing pattern renders most characters potentially applicable in taxonomy and phylogeny assessment in Arenaria. The studied taxa show four basic types of sculpturing, viz. colliculate, rugose, papillate, and smooth. The colliculate type which is characteristic for the core group of Arenaria (A. subg. Arenaria) can be further divided into five subtypes. The smooth seed surface with indistinct cell boundaries represents a synapomorphy for A. subg. Leiosperma, whereas the winged seeds along with rugose surface characterize subg. Odontostemma. Otherwise seed micromorphology rarely provides adequate evidence for separation of natural groups, especially at sectional rank. We hypothesize that the seeds are highly polymorphic in certain groups of taxa showing recent adaptive radiation, such as the annual species of Arenaria, and do not provide strong support for natural groups within the genus.     

Seed morphology under SEM and light microscopy in Kansas Juncus (Juncaceae)

Seed morphology was studied in 15 species of four subgenera ofJuncus occurring in Kansas, to determine if seeds provide traits useful in assessing systematic relations within the genus. In this study seed size and shape were of limited value, while surface ornamentation of the hard inner seed coat provided encouraging results. SubgenusPoiophylli showed little variation in surface ornamentation among taxa; similar ornamentation was observed in subgenusGenuini. SubgeneraGraminifolii andSeptati were separately distinct with the taxa in theSeptati forming a continuum of variation.     

Morphological variation in pollen grains of <Emphasis Type="Italic">Mucuna</Emphasis> (Leguminosae): new biogeographic and evolutionary patterns

Mucuna comprises 105 species with an overall pantropical distribution and is divided into three subgenera: M. subg. Mucuna, M. subg. Stizolobium and M. subg. Macrocarpa. Although phylogenetic studies have supported the occurrence of three main clades, evolutionary relationships among them are not fully resolved. The objective of this study was to examine pollen grain morphology from representatives of all three subgenera and map these onto the phylogenetic trees generated by analysis of other characters. Pollen grain surface, form, size, and aperture number were compared. A Bayesian inference tree using matK sequences was constructed. The results indicate that the representatives of M. subg. Macrocarpa have the smallest pollen grains in the genus (a synapomorphic character here identified for this subgenus) and that species of subgenus Mucuna (those with umbelliform inflorescences) have the largest pollen grains. Additional morphological diversity of the pollen grain surface was noted: reticulate and/or micro-reticulate (in all three subgenera), perforate, gemmate or verrucose (only in M. subg. Mucuna). For all studied taxa, the pollen grains are triaperturate, except for two species of M. subg. Mucuna, which have tetraperturate pollen. The phylogenetic tree obtained using the matK marker resolved M. subg. Stizolobium as the earliest diverging lineage in Mucuna. Based on this phylogeny, a reticulate ornamentation pattern of the pollen surface may represent the ancestral state for the genus, while the larger pollen size and the foraminate, gemmate, and verrucose ornamentations are derived characteristics within the genus. These putative derived ornamentations have been observed only in neotropical species.     

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 micromorphology of South American species of Hybanthus (Violaceae)

The seed‐coat of several species of Hybanthus subg. Ionidium from South America were studied by SEM (scanning electronic microscopy). Three different levels of sculpture were observed on the seed coat. The micro‐morphological patterns showed great variation between the species and should be considered as additional characters in future taxonomic treatments. The presence of a conspicuous elaiosome was observed; this structure is well known in the genus Viola and is most likely related to seed dispersal mediated by ants.     

Comparative seed morphology and character evolution in the genus Lysimachia (Myrsinaceae) and related taxa

Summary We investigated seed morphology in 34 species of the genus Lysimachia and in 14 species and two subspecies of six additional genera (Anagallis, Ardisiandra, Asterolinon, Glaux, Pelletiera, Trientalis), which have been shown to be closely related to, or are placed within Lysimachia in previous molecular studies. We studied seed shape, seed coat structure, and seed coat surface patterns. Three major types of seed shape were identified: (1) sectoroid, (2) polyhedral, and (3) coarsely rugose with concave hilar area. In addition, seeds may be keeled or winged. The outer layer of the seed coat is either sponge-like and adhering only loosely to the inner seed coat or it is thin and tightly adhering to the underlying tissue. Seed surface patterns can be divided into six main types: (1) reticulate, (2) tuberculate, (3) vesiculose, (4) colliculate, (5) undulate, or (6) poroid-alveolate. Seed surface patterns are mostly congruent with molecular phylogenetic relationships. A reticulate surface pattern is diagnostic of, e.g. Lysimachia subgenera Palladia and Hawaiian Lysimachiopsis. Mapping seed characters onto a recent phylogenetic tree, reveals that they provide potentially synapomorphic character states for various subclades of Lysimachia. Salient examples include a rugose seed shape, which turns out to be synapomorphic for the clade comprising the genus Pelletiera plus Asterolinon linum-stellatum and a sponge-like outer seed coat layer, which characterizes a clade with Lysimachia vulgaris, L. thyrsiflora, and L. terrestris, with an analogue that apparently evolved in parallel in Trientalis europaea. We also discuss possible habitat factors that may have favored the independent evolution of particular seed types such as winged seeds in various lineages.     

Genetic relatedness and taxonomy in closely related species of Hedysarum (Fabaceae)

A multidisciplinary study, engaging morphological, carpological and molecular data, has been performed to investigate the genetic relatedness and taxonomic boundaries of the close species Hedysarum gmelinii, H. setigerum and H. chaiyrakanicum (Fabaceae) with overlapped distribution areas in southern Siberia. The diagnostic features of these legume species are analyzed and discussed, including their macro- and micromorphological characteristics, seed coat ornamentation and inter-simple sequence repeat (ISSR) profiles. The morphometric features, pod and seed microsculpture traits of H. chaiyrakanicum and the ISSR patterns of the three species have been determined for the first time. Sprout, leaf, calyx, corolla, and stem rachis measurements, leaflet indumentum type and ISSR patterns significantly discriminate H. chaiyrakanicum from the other two species, whereas plant height, lengths of stem and leaf, and length and width of leaflet show opposite ranges of variation for H. gmelinii and H. setigerum though none of them is reliable in species identification. Ornamentation of seed coat and ISSR patterns does not differ significantly in the species. Therefore, our study supports the separate taxonomic treatment of H. chaiyrakanicum and the subordination of the cryptic species H. setigerum within H. gmelinii.     

Systematic relevance of seed coat anatomy in the European heathers (Ericeae, Ericaceae)

The anatomy of the seed coat of the European species of tribe Ericeae (Calluna, Daboecia and Erica) of the Ericaceae family was studied, and the taxonomic importance of their characters was analyzed. The seed coat is mostly formed by a one-cell layer with thick, pitted inner walls and thin outer walls that collapse at maturity over the inner walls. The cell junctions are either raised with anticlinal walls up to four times the height of the periclinal walls or are not raised with similar values for the height of both the anticlinal and periclinal walls. Three main cell junction types were found and described. The thickness of the inner walls is variable, but there is a large overlap among the results for different species. Calluna vulgaris is the only species with no pits, and E. multiflora has a pitted pattern on its inner walls, which is distinctive from the rest of the species. Our main results agree with the external seed morphology, and valuable new data were obtained for certain groups such as the E. cinerea-E. terminalis or the E. scoparia complex. The similarities that are found in seed coat characters are not in accordance with the classical taxonomic delimitation of infrageneric groups within Erica.     

Anatomy of Brazilian Eriocaulaceae: correlation with taxonomy and habitat using multivariate analyses

In this study we present a survey on vegetative anatomy in species of Actinocephalus, Blastocaulon, Eriocaulon, Leiothrix, Paepalanthus, Philodice, Syngonanthus, and Tonina (Eriocaulaceae). Multivariate analyses were used to correlate anatomical characters to taxa and the habitats where the species occur. Root and stem anatomical characters seem to be more affected by environmental factors where these species occur, and seem of little value for delimiting major taxonomic groups within the family. Other characters in the leaves, such as epidermis with thickened wall cells, compartmented substomatal chambers, mesophyll with hypodermis, compact chlorenchyma, collenchymatous bundle sheath extensions, and numerous vascular bundles, were shown to be important for defining species clusters in Leiothrix, Syngonanthus, and Paepalanthus subg. Platycaulon. Similarly, loosely aggregated chlorenchyma caused Blastocaulon, Eriocaulon, Philodice, Syngonanthus sect. Carpocephalus, S. sect. Syngonanthus, and Tonina, genera from humid environments, to cluster. Scape characters appear to be more informative in discriminating groups. This situation probably reflects lower selection pressures determining anatomical characters of this organ.     

国产打碗花属植物种子形态及其分类学意义

本文借助于又目解剖镜和扫描电电镜对我国打碗花属５种植物的种子进行了观察。结果表明：１、该属植物的种子可分为两类,一类为旋花型,较小,表面具疣状突起,种皮纹饰呈网状,包括打碗花、毛打碗花及旋花;另一类为藤水苗型较大,无疣状突起,种皮纹饰呈咀烂状,包括藤长苗和肾叶打碗花。２、疠状突起由数个相邻种皮细胞胞间壁强烈突出而成,呈山峰状,相连或单独存在。３、该属植物种子形态存在明显区别,可作为分种的依据。     

SYSTEMATIC SIGNIFICANCE OF SEED-SURFACE FEATURES IN ORTHOCARPUS (SCROPHULARIACEAE—SUBTRIBE CASTILLEJINAE)

Scanning electron microscope and light microscope examination of seed-coat features of 26 species of Orthocarpus have allowed recognition of many species-level differences (summarized in a key) and of three seed-coat types that parallel taxonomic subgroups but support realignments at generic and infrageneric levels. Type 1 seeds (subg. Orthocarpus, sect. Orthocarpus) have a lateral hilum, sculptured inner tangential seed-coat walls, and a tightly fitting outer seed coat. They are very similar to seeds of Cordylanthus. Seeds of Types 2 and 3 have a terminal hilum and membranous inner tangential cell walls. Type 2 seeds (subg. Orthocarpus, sects. Castillejoides and Cordylanthoides, with one exception) have a net-like, loosely fitting outer seed coat that shows close relationship to seeds of Castilleja. Inner tangential walls of Type 2 seeds normally rupture. Type 3 seeds (subg. Triphysaria, with two exceptions) have a tightly fitting outer seed coat and inner tangential walls are always retained. Seed features support evidence from floral morphology and chromosome numbers that Orthocarpus as currently recognized is not a monophyletic lineage.     

Seed Coat Microsculpturing Is Related to Genomic Components in Wild Brassica juncea and Sinapis arvensis

It has been reported that wild Brassica and related species are widely distributed across Xinjiang, China, and there has been an argument for species identification. Seed coat microsculpturing (SCM) is known to be an excellent character for taxonomic and evolutionary studies. By identifying collections from Xinjiang, China, and combining SCM pattern, flow cytometry, and genome-specific DNA markers as well as sexual compatibility with known species, this study aimed to detect potential relationships between SCM and genomic types in wild Brassica and related species. Three wild collections were found to be tetraploid with a SCM reticulate pattern similar to B. juncea, and containing A and B genome-specific loci, indicating relatively high sexual compatibility with B. juncea. The others were diploid, carrying S-genome-specific DNA markers, and having relatively high sexual compatibility with Sinapis arvensis. Moreover, their SCM was in a rugose pattern similar to that of S. arvensis. It was suggested that SCM, as a morphological characteristic, can reflect genomic type, and be used to distinguish B-genome species such as B. juncea from the related S. arvensis. The relationship between SCM and genomic type can support taxonomic studies of the wild Brassica species and related species.     

凤仙花属棒凤仙亚属(凤仙花科)植物花粉形态的分类学意义

棒凤仙亚属代表着凤仙花属最先分化出的一支,以多年生草本,多于5朵花的总状花序,侧生萼片4枚,3沟花粉,心皮四室且每室具1枚种子,果实棒锤状,种子椭圆形以及三沟花粉其极面观为三角形而区别于其它种类。该亚属主要分布于中国南部,少数种类辐射分布至华中和中南半岛地区。已有报道证实花粉的形态特征对凤仙花属植物的属下分类具有重要意义,但对棒凤仙亚属的种类少有涉及。为了进一步探讨该亚属花粉的形态特征及其分类学价值,该研究应用扫描电镜技术对国产棒凤仙亚属植物的14种以及凤仙花亚属的窄萼凤仙花的花粉进行了观察。结果表明：棒凤仙亚属的花粉以单粒存在,具3条萌发沟,辐射对称,极面观为三角形或三角状圆形,花粉粒外壁具均匀的网状纹饰,网脊平滑或具波状边缘,网眼中具不同密度的颗粒状突起。棒凤仙亚属的花粉形态与水角属的花粉形态相似,表现出了其原始性;其花粉形态特征与花部形态性状等相关性差,对该亚属的组级分类意义较小,但对于该亚属的种间界定具有重要意义;而其花粉的极面观形状、赤道面观形状以及网状纹饰等变异相对稳定,对于界定该亚属具有重要的分类学价值。     

Karyological studies of Iranian <Emphasis Type="Italic">Allium</Emphasis> L. (Amaryllidaceae) species with focus on sect. <Emphasis Type="Italic">Acanthoprason</Emphasis>. 1. Mitotic chromosomes

Eighteen species and subspecies (34 accessions) of Allium sect. Acanthoprason and 11 species (17 accessions) belonging to other subgenera and sections of Allium were karyologically investigated and include first reports for 12 species. The examined plants of 47 accessions were diploid, three accessions of two species were tetraploid, and in the A. bisotunense accession, we found a mix of di- and triploid individuals. B chromosomes were found in 10 accessions. A basic chromosome number of x = 8 was confirmed for all investigated members of subg. Melanocrommyum and subg. Allium, and x = 9 for Allium tripedale of subg. Nectaroscordum. Idiograms were drawn for each accession, and metaphase images are presented illustrating observed chromosomal variations. Also, karyotype features and asymmetry parameters were calculated for all accessions. Chromosomal aberrations, e.g. aneuploid cells or loss of whole or parts of chromosome arms, were rarely observed. In general, the karyotypes showed low variation in inter- and intrachromosomal asymmetry especially inside of the taxonomic groups, though satellited chromosomes were good markers for subgenera and even specific for two studied sections of subg. Allium. Six different types of satellites were recognized, two of them were newly described: Type P was prevalent in subg. Melanocrommyum, and type O in sect. Codonoprasum. Statistical analyses were performed on five karyological parameters to test correct relationships and also to test previous grouping hypotheses. Although our data confirm distinct karyological characters for the subgenera investigated, the remarkable morphological diversity inside of subg. Melanocrommyum is not mirrored by striking karyological differences.     

A Contribution to the Taxonomy of the Genus Xiphinema Cobb, 1913

In a comparative study of different populations of Xiphinema, the significance of the female gonad structure in species determination is stressed, and monodelphic, pseudomonodelphic, and didelphic forms in the genus are defined. X. chambersi is redescribed and illustrated, X. ensiculiferum and X. krugi are redefined, and X. ensiculiferoides n.sp. and X. orthotenum n.sp. are described. Proposed new synonymies are: X. itanhaense = X. brasiliense; X. bulgariensis and X. conurum = X. italiae; X. ifacolum = X. basiri; X. macrostylum = X. ensiculiferum; X. truncatum = X. elongatum; and X. vulgare = X. setariae. X. yapoense is considered species inquirenda and X. obtusum a nomen dubium. The following eight subgenera of the genus Xiphinema are proposed: Radiphinema n. subg., Krugiphinema n. subg., Xiphinema n. subg., Elongiphinema n. subg., Halliphinema n. subg., Basiphinema n. subg., Rotundiphinema n. subg., and Diversiphinema n. subg. A key to the subgenera of Xiphinema is presented, plus a list of 50 species in the genus, their synonyms, and 10 species inquirendae.     

Systematic significance of seed morphology in the genus Veronica (Plantaginaceae), with special reference to the Egyptian taxa

Seed morphology of 29 taxa including 9 subgenera belonging to the genus Veronica was compared using scanning electron microscopy to assess their diagnostic value for systematic studies. Subgenus Beccabunga is the largest in this study represented by nine taxa. Seed surface often varies from ridged reticulate or verrucose/granulate reticulate to rugose-reticulate. Seed color and size have limited taxonomic significance as their variation is uninformative. Thereagainst, seed shape and seed coat ornamentation present informative characters that can be used efficiently in distinguishing the studied taxa. Our seed features support the DNA sequence data in showing close relationships between V. biloba and V. campylopoda, and between V. fruticans and V. fruticulosa. The present study indicates thatV. anagallis-aquatica, V. anagalloides, V. comosa, and V. catenata have more or less the same seed features; consequently, these four former species are most likely representing a single species as reported by earlier studies. Furthermore, V. polita and V. persica are very similar in terms of seed characteristics. Our results justify placement of V. peregina within subgenus Beccabunga and support the monophyly of the subgenus Veronica.     

Phylogenetic analysis of <Emphasis Type="Italic">IRIS</Emphasis> L. from China on chloroplast <Emphasis Type="Italic">TRN</Emphasis>L-F sequences

Phylogenetic relationships among the six Iris subgenera were reconstructed by chloroplast trnL-F sequences data using maximum likelihood. The entire matrix of aligned bases analyzed includes 1043 characters and the length of all sequences varied from 747 bp to 893 bp, and mutation sites accounted for 18.79% of the total length. The cluster analysis results accorded well with the subgeneric classification of Chinese Iris species. Results suggested rhizomes and sepals lacking ornament are ancestral characters, and subg. Xyridion and subg. Limniris are more primordial than another four subgenera. Subgenus Nepalensis and subg. Xyridion were resolved as monophyletic.     

罗布麻与大花罗布麻种了微结构特征

罗布麻（Ａｐｏｃｙｎｕｍ ｖｅｎｔｒｍ Ｌ．）和大花罗布麻（Ｐｏａｃｙｎｕｍ ｈｅｎｄｅｒｓｏｎｉｉ（Ｈｏｏｋ．ｆ．）Ｗｏｏｄｓｏｎ）分别属于夹竹桃科罗布麻属（Ａｐｏｃｙｎｕｍ Ｌ．）和白麻属（Ｐｏａｃｙｎｕｍ Ｂａｉｌｌ．）。其外部形态区分明显,但种子微结构特征却颇为相似。种子形状、颜色相同,大小相近;种子表面基本纹饰均为网穴状,次级纹饰均为颗粒状。     

Three new species of Lactarius (Russulaceae,Russulales) from Northeast China

Three new species of Lactarius are described from Northeast China, based on morphological comparisons and analyses on ITS nrDNA sequences. Samples of these species were mostly collected from deciduous or mixed forests with oaks. Lactarius albidocinereus, a member of L. subg. Plinthogalus is very similar to European L. azonites in the morphology and the sequence data. It differs from L. azonites by more ellipsoid spores with thinner ornamentation and shorter terminal cells in the pileipellis. Lactarius dilutisalmoneus belongs to L. sect. Zonarii of L. subg. Lactarius due to the pale salmoneus subzonate pileus with glabrous margin, unchanging latex and spore ornamentation composed of isolated elements. In comparison with its relatives, i.e. European L. zonarioides and North American L. olympianus, this new species has smaller spores with more prominent ornamentation. Lactarius olivaceorimosellus, a member of L. subg. Russularia, has areolate-rimulose pileus and lacks rosettes in the pileus and stipe cortex. Apart from the concentrically cracked pileus and absence of rosettes, the species can be easily distinguished by the olivaceo-brownish tinge of the pileus and spores with more dispersed elements of the ornamentation. Of the three species, at least L. albidocinereus represents a link with the Lactarius mycota of subtropical China.