Taxonomic importance of seed macro‐ and micro‐morphology in Abelmoschus (Malvaceae)

Seed morphology of Abelmoschus is known to be variable, but patterns of variation have never been critically studied. We studied seed macro‐ and micro‐morphological characters, including seed shape/size, seed coat pattern and trichome density/structure in multiple samples to evaluate the taxonomic significance of seed characters. Among the studied characters, seed shape and trichome structure were found to have major taxonomic importance and proved to be valuable characters for separating taxa. Two main seed types were present: seeds with deciduous trichomes and seeds with persistent trichomes. These characters offer significant evidence to the distinctness of certain species (A. esculentus, A. moschatus subsp. moschatus, A. moschatus subsp. tuberosus, A. crinitus and A. angulosus). Further, our results indicate that A. moschatus subsp. tuberosus should be maintained as a separate subspecies while A. manihot subsp. tetraphyllus var. pungens may be merged in A. angulosus. No significant intraspecific variation was observed, except in A. esculentus. We conclude that seed coat sculpturing and seed trichomes do indeed provide stable and diagnostic characters for many morphologically closely related taxa of Abelmoschus and that LM/SEM techniques can be useful in solving systematic problems and management of Abelmoschus genetic resources.     

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.     

中国东北牻牛儿苗科(Geraniaceae)种子形态特征及其分类学意义

本文对中国东北牛儿苗科(Geraniaceae)二属(太阳花属Erodium, 老鹳草属(Geranium)计14种(species)1变种(Variety)和1变型(form)植物的种皮亚显微结构(SEM)及种子形态进行了研究.结果表明, 太阳花属与老鹳草属植物无论从种子形状, 长宽比, 还是种皮初级雕纹及次级雕纹形态都表现出明显差异。在老鹤草属下种及种下等级的鉴定和划分上, 种皮微形态及种子形态特征提供了大量有价值的信息。     

Seed micro‐morphology and its systematic significance in Gypsophila (Caryophyllaceae) and allied genera

In order to examine the systematic application of seed‐coat micro‐morphology in Gypsophila and allied genera, seed surfaces of 30 species and two varieties representing four genera of tribe Caryophylleae were examined with scanning electron microscope. The seeds of examined species range between 0.5–2.1 mm in length and 0.3–2.4 mm in diameter. The exomorphology of the seed coat shows two distinctive cell patterns. The epidermis is constructed either of elongated polygonal or of broad polygonal cells. The elongated type is the most common among the studied species, but the variation in alignment of testa cells, their size and shape as well as the density of protuberances may provide further information and useful diagnostic characters at generic and specific rank. The testa cells in Gypsophila and Saponaria are shallowly undulate, deeply undulate, lobed and armed at anticlinal walls. Deeply undulate anticlinal walls were observed in both Gypsophila and Ankyropetalum and a few species of Saponaria. Non‐ or indistinctly grooved anticlinal walls is the more common type in Allochrusa. Seed‐coat characters support the separation of Gypsophila and Saponaria to some extent but disagree with recognition of Ankyropetalum as a genus separate from Gypsophila.     

Micromorphology of fruits and seeds of Iranian Geranium (Geraniaceae), and its systematic significance

The fruit and seed micromorphology of 22 species of Geranium, representing the eight sections of the genus represented in Iran (G. sectt. Dissecta, Geranium, and Tuberosa of subgen. Geranium; sectt. Batrachioidea, Divaricata, Lucida, Ruberta and Trilopha of subgen. Robertium), have been examined by scanning electron microscopy (SEM). Macro‐ and micromorphological characters, including fruit and seed shape, size, color, hair type and density, mericarp ornamentations, hilum position, seed coat pattern, epidermal cell shape, and anticlinal and periclinal cell walls, are presented. Two microsculpturing patterns are recognized on the mericarp surface: reticulate and pusticulate. The micromorphology of the seed coat showed four distinctive cell patterns. The seed epidermis is constructed either of polygonal, elongated polygonal, or square to rectangular cells. The polygonal type is the most common among the studied species, but the variation in testa cell characters, their size and shape, may provide further information and useful diagnostic characters at specific and infraspecific rank. The shape and color of the seeds are, however, of little systematic value. Fruit characters were found to be important for separating taxa at infrageneric rank and our results show that the species can be separated into subgenera and sections based on fruit morphology.     

Seed morphology in the tribe Chloraeeae (Orchidaceae): combining traditional and geometric morphometrics

Previous work on orchid seeds has shown that characters associated with the seed coat may be useful for classification and phylogeny at a suprageneric level. The seed morphology of several species of the tribe Chloraeeae was analysed using traditional morphometrics, and the seed shape was studied, for the first time, using tools of geometric morphometrics. Seed characters were evaluated by their discriminative power and the information they may provide in a phylogenetic context. By contrast with previous findings, seed shape resulted in a continuum among the taxa studied, and in only a few cases could genera or groups of species be discriminated on the basis of shape. However, seed size, expressed as centroid size, was a variable character and informative at a phylogenetic level. Traditional measures of seed coat, mainly those of seed coat cells, were also helpful for the discrimination of genera and species, agreeing with previous statements about their utility in taxonomy and phylogeny. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 171–183.     

SEED COAT MORPHOLOGY IN CORDYLANTHUS (SCROPHULARIACEAE) AND ITS TAXONOMIC SIGNIFICANCE

A study of seed coat sculpturing in Cordylanthus (Scrophulariaceae—Rhinantheae) using the scanning electron microscope (SEM) shows that seed surface patterns are characteristic and constant for a given species or a group of species. Seeds of 23 species were examined and classified into four types (irregularly crested, deeply reticulate, shallowly reticulate, and irregularly striate) based on differences in the reticulated seed coat. Anatomical studies of sections with the light microscope show that the reticular patterns result from enlargement of the epidermal cells of the integument, followed by formation of characteristic wall patterns through lignification. Seed characters furnish useful data for formulating the taxonomy of Cordylanthus both on the sectional and subsectional level and for the delimitation of certain species. Seed coat morphology offers evidence for including the genus Dicranostegia in Cordylanthus and for maintaining C. hispidus and C. palmatus as separate species. The distinctness of two seed coat types within section Cordylanthus suggests two major phylogenetic lines within this section.     

Triterpene saponins from Fagonia scabra Forssk and other Fagonia species

This article reports on the structure of triterpenoids saponins (1–49) isolated from Fagonia species. In addition, it describes one known flavonoid (50) with luteolin as aglycone and seven known triterpenoid saponins (26, 51–56) isolated from the aerial parts of Fagonia scabra Forssk. Compounds 50–56, were isolated for the first time from the genus Fagonia, while compounds 50–53, 55–56 have never been encountered in the Zygophyllaceae family. The chemotaxonomic relationship between F. scabra and other Fagonia species was also discussed.     

Ultra-sculpturing of seed morphotypes in selected species of genus Salvia L. and their taxonomic significance

• The taxonomic importance of macromorphological and micromorphological seed characteristics was investigated using SEM of ten species of the Salvia genus from Pakistan. The aim was to identify diagnostic seed ultrastructural features that could aid in species delimitation, correct identification and phylogenetic position. The ultrastructure of Salvia varies greatly, and a wide range of unique micromorphological features have been observed.
• Seed micromorphological features were explored by SEM, including seed shape, colour, texture, cell outline, surface sculpturing, epidermal cell arrangement, anticlinal, and periclinal wall pattern.
• Seed shapes were categorized as obovate, spherical, spheroid, broadly elliptic, elliptic and oblong, mostly with a terminal hilum. Seed colours were black, light brown, dark brown, brown and yellow. Exo-morphological characters, i.e. epidermal cell arrangements, included irregular, wavy pentagonal-hexagonal, regular pentagonal-hexagonal. Cluster analysis was used to assess similar and distinct species within Salvia with a feasible explanation. Taxonomic keys were made based on micromorphological qualitative features that help to delimit species and identify them quickly within the Salvia genus.
• Seed morphology of ten Salvia species was described and investigated, and the diagnostic significance of features evaluated using SEM. This study analysed seed features, especially at the species level, which might provide much new taxonomic information. The results revealed that, in seed morphology, using SEM can help with taxon identification, especially at the genus and species levels.

     

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.     

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.     

Numerical analyses of seed morphology and its taxonomic significance in the tribe Nigelleae (Ranunculaceae)

We examined the seed morphology of Nigella, Garidella and Komaroffia to aid systematics and taxonomy. The seeds of 21 taxa in 20 species from Mediterranean and West Asian countries were studied using both zoom stereo and scanning electron microscopy (SEM). Principal coordinate analysis (PCoA) and unweighted average linkage analysis (UPGMA) were performed to assess the contribution of seed characters to the taxonomy of the tribe. Representative pictures from zoom stereo microscopy and SEM studies and measurement of seeds are presented. Most studied species showed unique seed surface sculpturing features, and seed shape, size and surface sculpturing separated the studied taxa into five main groups. However, the variation in seed shape and color were found to be in conflict with the generic taxonomy while macromorphological characters were found more congruent with the sectional taxonomy than with the generic taxonomy. Five main types of seed ornamentations – rugulose + striate, granulate, smooth, rugulose and mucronulate – were identified. Patterns in seed micromorphology strongly supported the sectional level taxonomy of Nigella. Two of the three sections of the genus were found to be well differentiated in terms of seed characters. Furthermore, seed morphology alone provide good support to Garidella as a distinct genus, but shows less taxonomic value for Komaroffia.     

十大功劳属植物种子形态研究

利用光镜(LM)和扫描电镜(SEM)对十大功劳属24种植物的种子形态进行观察。结果发现十大功劳属植物种子颜色为黑色,长3.4～6.7mm,宽1.7～3.5mm,属大型种子,形状多样;外种皮纹饰为网纹状,网眼形状、大小及网脊粗细、凸凹在不同类群间不完全相同,种内变异稳定,据此可将十大功劳属植物种皮纹饰划分为9种类型。研究结果表明:种子形态在十大功劳属内具有重要的系统学意义,但不支持Ahrendt将十大功劳属下分为2个groups,4个sections,14个subsections的观点。     

<Emphasis Type="Italic">Anacamptis</Emphasis> versus <Emphasis Type="Italic">Orchis</Emphasis> (Orchidaceae): seed micromorphology and its taxonomic significance

Seed micromorphology of 24 taxa of the genera Anacamptis and Orchis was examined under light and scanning electron microscopy. Seed qualitative characters appear very useful at the supraspecific level in the subtribe Orchidinae. Based on our observations, the sculpturing of the periclinal walls of the medial testa cells, the seed shape and several features of the anticlinal walls of the apical cells showed variability between the studied taxa, with special relevance of the first character. According to this factor, we found one type of seed for Anacamptis and two for Orchis. The taxonomic value of seed coat characters is compared with the last taxonomic proposals for the genera Anacamptis and Orchis.     

Fruit and seed morphology of the tribe Orobancheae (Orobanchaceae) genera in Turkey and its taxonomic significance

Seed and fruit morphology of four genera of Orobanchaceae: Cistanche, Diphelypaea, Orobanche and Phelipanche, native to Turkey, were studied using light and scanning electron microscopy. In this study, 100 accessions belonging to 39 species were investigated. The examined fruits had loculicidal capsules, generally dehiscing with 2 slits, and the seeds were small and with reticulate ornamentation. The basic shape of the seeds in most of the studied taxa was oblong, ellipsoid to ovoid, globose or subglobose. The results indicate that characters such as seed size, shape and color are useless for identification of the taxa at the species level. However, the anticlinal wall pattern and periclinal wall ornamentation proved useful for identifying the different genera. Four morphological types could be distinguished: smooth, perforate, microfibrillar and granulate. Type I corresponds to the genus Cistanche and Orobanche anatolica, whereas types II, III and IV were found in Diphelypaea, Orobanche and Phelipanche, respectively. The utility of seed characteristics is discussed in terms of inter‐ and intra‐generic taxonomy.     

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: an addition to the taxonomy of Tephrosia (Leguminosae, Papilionoideae, Millettieae) from South America

Seed morphology was studied using scanning electron microscopy in 13 species, one subspecies and one forma of Tephrosia Pers. (Leguminosae, Papilionoideae, Millettieae) occurring in South America. Macromorphological and micromorphological characters were examined, including seed form, colour and size, testa pattern, reticulum anticlinal wall, boundaries of anticlinal wall and hilum form. Crested and simple-reticulate testa patterns were predominant. A foveolate pattern, multifoveolate pattern and subgrooved pattern are all recorded for the first time in this genus. The macromorphological characters displayed continuous variation in shape and size and are thus are not significant for species separation. The data obtained in this study together with data from the literature provide additional characters to help classify the genus. A key to the taxa under study is presented.     

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.     

勾儿茶属和小勾儿茶属(鼠李科)的果实及种子形态研究及其系统学意义

在扫描电镜和解剖镜下研究鼠李科(Rhamnaceae)勾儿茶属(Berchemia)和小勾儿茶属(Berchemiella)共17种植物的果实和种子形态.结果表明：小勾儿茶属的核果1室,具1枚种子,勾儿茶属大多数种的核果2室,每室具1枚种子,但多叶勾儿茶(B．polyphylla)的核果1室较大,具1枚种子,另一室较小而没有种子,该种可能是连接勾儿茶属和小勾儿茶属的中间类群.这两个属的种子形状通常为不规则长椭圆形,种皮纹饰可划分为光滑或几乎光滑、具不明显或稀疏的条纹以及具明显的条纹或沟这三种类型.种皮纹饰的差异对这两个属属下种类的划分具有一定的意义.讨论了与前人研究结果不同之处和可能的原因.