Evolution of trichomes and its systematic significance in Salvia (Mentheae; Nepetoideae; Lamiaceae)

We have investigated the trichome characteristics in representative species of Salvia and Pleudia in order to evaluate this source of morphological evidence for addressing problems regarding generic delimitation and subgeneric classification. Trichomes of 46 Salvia spp., representing three subgenera in Iran, were investigated using light microscopy and scanning electron microscopy. General trichome characteristics were constant among different populations of a certain species, but showed a degree of variability useful in the delimitation of taxa, specifically at lower taxonomic levels. Trichome characters of taxonomic interest are as follows: types of glandular hair; number of composing cells (uni‐, bi‐ or multicellular); size and thickness; branching pattern; and presence of papillae on the surface. Non‐glandular trichomes can be simple and branched. Glandular trichomes can be stalked, subsessile or sessile. Our investigation reveals the usefulness of such characters in providing fundamental taxonomic criteria for taxon delimitation in these genera at various levels, especially at the specific rank. Furthermore, the data presented here indicate the potential applicability of such characters in the determination of evolutionary trends in Salvia and allies. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 241–257.     

Trichome micromorphology of the East Asiatic genus Chelonopsis (Lamiaceae) and its systematic implications

The micromorphology of trichomes of the leaves of 17 taxa (including two varieties) of the genus Chelonopsis Miq. and of six species representing four additional genera (Bostrychanthera deflexa Benth., Colquhounia coccinea Wall. var. coccinea, Co. seguinii Vaniot. var. seguinii, Gomphostemma chinense Oliv. var. chinense, G. crinitum Wall. ex Benth. and Physostegia virginiana (L.) Benth.) was surveyed by light and scanning electron microscopy. Two basic types of trichomes can be identified: non-glandular and glandular trichomes. The non-glandular trichomes can be subdivided into two subtypes: simple unbranched and branched trichomes. Based on the cell number, simple unbranched trichomes are further divided into four shapes (unicellular, two-celled, three-celled, and more than three cells), whilst branched trichomes are separated into three shapes (biramous, stellate, and dendroid trichomes). The glandular trichomes can in turn be subdivided into four subtypes: subsessile, capitate, clavate, and sunken. Non-glandular trichomes with two cells (NGTW) and subsessile glandular trichomes (GSU) are most widespread in all taxa examined. The indumentum shows considerable variation among different sections or species. Consequently, trichome micromorphology and distribution have high taxonomic value for Chelonopsis at both infrageneric and interspecific levels. The presence of capitate glandular trichomes (GCA) provides an additional morphological character to clarify the boundaries between subgenus Chelonopsis and Aequidens Wu and Li. Within subgenus Aequidens, non-glandular trichomes with more than three cells (NGMT) and clavate glandular trichomes (GCL) are important characters for sectional division between sect. Aequidens Wu and Li and sect. Microphyllum Wu and Li. Again, three forms of three-celled trichomes can be used as a distinctive taxonomic character at specific level between C. albiflora Pax et K. Hoffm. ex Limpr., C. forrestii J. Anthony, and C. souliei (Bonati) Merr. in sect. Aequidens. This study supports Wu's delimitation of subgenus and sections and the subsequent review work by Xiang et al. Additionally, distribution of trichome types is correlated with the altitudinal distribution and habitats of some species in Chelonopsis.     

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.     

Infrasectional systematics of the genus Sideritis L. section Sideritis (Lamiaceae)

RIVERA NUÑEZ, D., OBON DE CASTRO, C., TOMAS-LORENTE, F., FERRERES, F. & TOMAS-BARBERAN, F. A., 1990. Infrasectional systematics of the genus Sideritis L. section Sideritis (Lamiaceae). A new taxonomic division of the section Sideritis is proposed on the basis of morphological, cytological and chemical characters. The following subsections art-recognized: Grandiflora, Ovata, Camarae, Linearifolia, Gymnocarpae, Stachydioides, Lacaitae, Hirsuta, Chamaedryfolia, Arborescens, Flavovirens, Leucantha, Angustifolia, Serrata and Scordioides . Possible evolutionary pathways are discussed.     

Systematics and biogeography of Klasea (Asteraceae–Cardueae) and a synopsis of the genus

Klasea , traditionally treated as a section in Serratula , is now widely accepted at the generic level. A classification of the genus is presented here, accommodating the 46 species in ten sections based on nuclear ribosomal DNA external and internal transcribed spacer sequence data and morphology. New combinations for five species and ten subspecies are published, and a new hybrid species is described. The genus ranges from the Iberian Peninsula and north Africa through southern and eastern Europe, west and central Asia to the Himalayas, and the Far East of Russia and China. The ancestral area is in west Asia, most probably eastern Anatolia and northern and western Iran. In this region, representatives of all sections are present. The largest section Klasea diversified most likely in the mountains of central Asia. A key to all Klasea species is provided.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 435–464.     

Pollen morphology of the East Asiatic genus Chelonopsis (Lamioideae: Lamiaceae) and allied genera,with reference to taxonomic implications and potential pollination ecology

Pollen grains of 15 taxa of the genus Chelonopsis (14 spp. and 1 variety) from China and Japan and 6 species of the closely related genera Bostrychanthera (1 species) and Gomphostemma (5 species) were examined by light and scanning electron microscopy. Of these, the pollen morphology of 18 taxa was studied for the first time. Pollen grains were found to be tricolpate with polar lengths of 20.8–30.0 μm and equatorial widths of 17.5–27.3 μm. The basic shape of the pollen in most taxa is subprolate or prolate-spheroidal, but spheroidal, subprolate-spheroidal, oblate-spheroidal, and prolate-subprolate grains are also found in some species. The surface is generally reticulate or suprareticulate in Chelonopsis and granulate in Bostrychanthera. In comparison with those of Chelonopsis and Bostrychanthera, the pollen grain surfaces of Gomphostemma are more diverse. In Chelonopsis, pollen is taxonomically useful at the sectional level, and some grains provide enough characters for species delimitation. The potential pollination ecology of cellular hairs on the anthers of Chelonopsis and Bostrychanthera is also briefly discussed.     

Distribution of tanshinones in the genus Salvia (family Lamiaceae) from China and its systematic significance

Salvia L.(family Lamiaceae),a large genus of over 1000 species,is widely distributed throughout tropical and temperate regions of the world.Eight-four species are native to China.However,there are many taxonomic uncertainties at the sub-generic level.Diterpenoids are a class of secondary metabolites with a large variety of structures that have been used as chemotaxonomic markers at infra-and suprageneric levels.For the sake of further chemotaxonomic understanding of Salvia,the present study investigated the...     

Molecular phylogenetics,character evolution and systematics of the genus Micranthes (Saxifragaceae)

With c. 85 species, the genus Micranthes is among the larger genera of the Saxifragaceae. It is only distantly related to the morphologically similar genus Saxifraga, in which it has frequently been included as Saxifraga section Micranthes. To study the molecular evolution of Micranthes, we analysed nuclear ribosomal (internal transcribed spacer, ITS) and plastid (trnL–trnF) DNA sequences in a comprehensive set of taxa comprising c. 75% of the species. The molecular phylogenetic tree from the combined dataset revealed eight well‐supported clades of Micranthes. These clades agree in part with previously acknowledged subsections or series of Saxifraga section Micranthes. As these eight groups can also be delineated morphologically, we suggest that they should be recognized as sections of Micranthes. New relationships were also detected for some species and species groups, e.g. section Davuricae sister to sections Intermediae and Merkianae, and M. micranthidifolia as a member of section Micranthes. Species proposed to be excluded from the genus Micranthes for morphological reasons were resolved in the molecular tree in Saxifraga. Many morphological characters surveyed were homoplasious to varying extents. Micromorphological characters support comparatively well the clades in the phylogenetic tree. An updated nomenclature and a taxonomic conspectus of sections and species of Micranthes are provided. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 47–66.     

Leaf epidermal features of Lithocarpus (Fagaceae) from China and their systematic significance

The leaves of 52 species of Lithocarpus in China were studied. The adaxial leaf epidermis was investigated by light microscopy. Epidermal cells of the adaxial surface were classified into three types on the basis of the outline of their anticlinal walls, i.e. sinuate, straight and curved. The abaxial leaf epidermis was investigated by light microscopy and scanning electron microscopy. The following types of trichome were observed: appressed parallel tuft, stellate, fused stellate, papillae, stipitate fasciculate, solitary unicellular, appressed laterally attached unicellular, curly thin‐walled unicellular, bulbous and thin‐walled peltate. The fused stellate, appressed laterally attached unicellular and curly thin‐walled unicellular trichomes were reported in Lithocarpus for the first time. The appressed parallel tuft trichome, which is recognized as a salient characteristic of Lithocarpus, was not found in 15 species. A cladistic analysis was performed on the basis of the leaf epidermal features. According to the leaf epidermal features and several morphological characteristics, 26 of the 52 species could be divided into seven groups. Similar groups can be found in Barnett's and Camus' systems. The trichomes of four genera in Fagaceae are listed and compared. Lithocarpus had 14 types of trichome, 11 of which were identical to types found in Quercus, more than in Castanopsis and Cyclobalanopsis. The evolutionary trends of trichomes in Fagaceae are discussed and a new point of view is raised. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 216–228.     

The genus Thymus (Lamiaceae) in Sicily

An update on the taxonomy and geographic distribution of the genus Thymus in Sicily is given. This study is based on literature, herbarium and field investigations. In total, five taxa belonging to Thymus sect. Serpyllum (Th.richardii subsp. nitidus, Th.longicaulis and Th.praecox subsp. parvulus) and Th. sect. Hyphodromi (Th.spinulosus and Th.paronychioides) occur in Sicily. For each one, information about taxonomy, habitat, phenology, chromosome number and distribution are provided. Furthermore, the name Thymus pedicillatus Lojac. is here typified. Finally, an analytical key for the identification of the studied taxa is proposed.     

Pollen morphology in the genus Veronica L. (Plantaginaceae) and its systematic significance

Pollen grains of 30, mainly annual, species from 134 populations of the genus Veronica (Plantaginaceae, formerly Scrophulariaceae) from the Mediterranean Region, have been studied with light and scanning electron microscopy. Three pollen types are defined based on pollen size and exine surface sculpture. In some cases within the study group pollen size can be useful in species determination. Hypothetical relationships of pollen grain size and aperture number with ploidy level, style length and corolla diameter are discussed. The ornamentation of the exine in Veronica, although generally a conservative character within the genus, gives some additional support to the most recent infrageneric classification of Veronica.     

Intraclass Evolution and Classification of the Colpodea (Ciliophora)

ABSTRACT. Using nine new taxa and statistical inferences based on morphological and molecular data, we analyze the evolution within the class Colpodea. The molecular and cladistic analyses show four well‐supported clades: platyophryids, bursariomorphids, cyrtolophosidids, and colpodids. There is a widespread occurrence of homoplasies, affecting even conspicuous morphological characteristics, e.g. the inclusion of the micronucleus in the perinuclear space of the macronucleus. The most distinct changes in the morphological classification are the lack of a basal divergence into two subclasses and the split of the cyrtolophosidids into two main clades, differing mainly by the presence vs. absence of an oral cavity. The most complex clade is that of the colpodids. We partially reconcile the morphological and molecular data using evolutionary systematics, providing a scenario in which the colpodids evolved from a Bardeliella‐like ancestor and the genus Colpoda performed an intense adaptive radiation, giving rise to three main clades: Colpodina n. subord., Grossglockneriina, and Bryophryina. Three new taxa are established: Colpodina n. subord., Tillinidae n. fam., and Ottowphryidae n. fam. Colpodean evolution and classification are far from being understood because sequences are lacking for most species and half of their diversity is possibly undescribed.     

Seed coat morphology of Aizoaceae–Sesuvioideae, Gisekiaceae and Molluginaceae and its systematic significance

Macro- and micromorphological characters of the seeds of 32 species belonging to Aizoaceae (26 species), Gisekiaceae (one species) and Molluginaceae (five species) were analysed for their taxonomic value. Seed morphology is found to be of considerable taxonomic value within the taxa investigated, although additional criteria are usually necessary for distinguishing the species. However, a subgroup of Trianthema , consisting of T. cussackiana , T. megasperma , T. pilosa , T. rhynchocalyptra and T. oxycalyptra var. oxycalyptra (Aizoaceae) shares the synapomorphy of seeds with scattered idioblast cells (papillae). There is little infrageneric variation within Sesuvium and Zaleya in contrast to Trianthema , whose seeds are considerably polymorphic. The fairly different seed structures found in Aizoaceae s.s . and Molluginaceae support their treatment as two distinct families. However, the position of Lineum (Molluginaceae) remains uncertain. Moreover, seed characters support the independence of Gisekia (Gisekiaceae) in a family of its own.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 189–206.     

Pollen morphology in the genus Coffea (Rubiaceae) and its taxonomic significance

The pollen morphology of 38 representative species and varieties within the genus Coffea L. is described. Eight pollen types, placed in two major groups, have been identified on the basis of the number of colpi, colpus characteristics, exine morphology and pollen size. Differences in pollen structures of the species studied do not correspond to present taxonomic groupings, but support a current proposal to place some of the species in the genus Paracoffea Leroy. Among the four sections in the genus, the section Eucoffea contains a high degree of pollen polymorphism.     

Systematic distribution of foliar trichome types in Crotovt (Euphorbiaceae)

Variation in foliar trichomes in Croton (Euphorbiaceae) is reviewed, and the terminology clarified by redefinitions and illustrations of terms. Evolution of trichome types appears to proceed from branched (stellate or fasciculate) hairs to simple and dendritic ones. A systematic enumeration characterizes the trichome morphology in 36 of the 40 sections, with citation of the 120 species examined.     

Systematics of the high mountain taxa of the genus Sideritis L. section Sideritis, subsection Fruticulosae Obón & D. Rivera (Lamiaceae)

Section Sideritis is an extremely diversified group which is formed mainly by species growing at low altitude. The group of high altitude taxa of this section is polyphyletic and has been taxonomically divided in different subsections in which these taxa appear normally associated with low-lying planitiary ones, the latter being the probable ancestors. The subsections comprising high altitude taxa are: subsect. Gymnocarpae , subsect. Fruticulosae; subsect. Hyssopifoliae , subsect. Luridae; subsect. Boigiae and subsect. Aranensis . Most of the high altitude taxa of section Sideritis are endemics with small distribution areas; they are incompletely known and threatened with extinction. The upper altitude limit of this section is attained at over 3000 m in Sierra Nevada (Spain) by Sideritis glacialis Boiss., which has been included in subsect. Fruticulosae Obón and D. Rivera. The taxonomy of this complex group of high altitude endemic taxa of Sideritis subsection Fruticuhsae is discussed on the basis of macro-morphological and micro-morphological characters. These are used in identification keys and for analysis of dissimilarity. The different habitats, allied species and plant communities are described. The following taxa are recognized: Sideritis glacialis subsp. glacialis, S. glacialis subsp. vestita, S. glacialis subsp. virens and S. glacialis subsp. fontquerii .     

The west Mediterranean orophilous taxa of Sideritis L. (Lamiaceae): a new species of subsection Hyssopifolia from south-eastern Spain

Orophilous taxa of Sideritis sect. Sideritis (Lamiaceae) are rare, although highly diversified in south-eastern Spain. Most of them belong to subsections Hyssopifolia and Fruticulosa and show very reduced distribution areas in the summits of the highest Betic mountains. The inaccessibility of their habitats has meant that many of them have been described only within the last twenty years. In this context, a new species Sideritis tugiensis is described in subsection Hyssopifolia , from the Oromediterranean summits of Sierra de Segura (south-eastern Spain). It is a woody, cushion-shaped plant, resembling both S. carbonellis Socorro (subsect. Hyssopifolia) and S. glacialis Boiss., s.l. (subsect. Fruticulosa) , though important morphological divergences warrant recognition at species rank. Data on morphology, ecology and chorology of the new species are reported, and affinities and differences with regard to close taxa from other subsections are presented. Evolutionary trends in the whole aggregate are briefly discussed.     

18种(1变型)鼠尾草属植物叶表皮及表皮毛微形态特征研究

鼠尾草属是唇形科中最大的属,全球有近1 000种。我国是鼠尾草属东亚分布中心,有84种,分布于全国各地,尤以西南地区最多。鼠尾草属植物具有很高的经济价值和观赏价值,但由于鼠尾草属植物分布广、种类多、形态变异大,给该属鉴定和分类学研究带来了很大的困难。此外,在采集和调查过程中发现弧隔鼠尾草亚属和荔枝草亚属的很多种类在营养生长期仅通过叶片很难鉴定。表皮毛是植物叶片上最常见的附属物,其分布特性和形态特征常作为植物鉴定和分类学研究的重要手段。该文利用光学显微镜和环境扫描电镜对18种和1变型共19份鼠尾草属植物的叶片类型、表皮细胞形状、垂周壁样式、气孔器类型和表皮毛等叶表皮微形态特征进行了观察和比较研究,为鼠尾草属的分类界定研究提供证据。结果表明:这19份鼠尾草属植物叶表皮和表皮毛形态特征在电子显微镜下表现出明显的多样性,叶表皮细胞形状为不规则形或多边形,垂周壁多数为浅波状或波状,少数为平直或弓形。气孔器为无规则型和不等细胞型,其中无规则型最为常见。样品叶表皮毛分为4个类型,盾状腺毛、头状腺毛、短非腺毛和长非腺毛。该属植物叶表皮的这些微形态特征,可为本属鉴定和分类学研究提供理论依据。