Anatomy, palynology and nutlet micromorphology of Turkish endemic Teucrium sandrasicum (Lamiaceae)

In this study, the anatomical features of the leaf and stem, besides the pollen and nutlet characteristics of Teucrium sandrasicum are investigated. T. sandrasicum, belonging to sect. Teucrium, is an endemic perennial herb growing on serpentine around Muğla province. The anatomical studies on T. sandrasicum revealed that the stem shares the general characteristics of the Labiatae family. The leaves clearly exhibit xeromorphy due to features such as the distribution of stomata on the lower surface (hipostomatic), the occurrence of guard cells below the epidermis (xeromorphic type), inrolled margins, thick cuticle layer, thick outer epidermal cell wall, a high density of trichomes and thick palisade layer of the mesophyll. The anatomical studies showed that the upper epidermal cells of the leaf include many spherocrystals. The pollen grains are prolate, medium in size, 3-colpate with verrucate ornamentation. The nutlets are ellipsoid with a reticulate-verrucate surface. The results have proven that T. sandrasicum is different from the other species of the sect. Teucrium because of the branched trichomes on the stem and the lack of eglandular trichomes on the nutlets.     

Trichome micromorphology in Turkish species of Ziziphora (Lamiaceae)

Ziziphora L. is represented by 5 species and 2 subspecies in the flora of Turkey: Z. clinopodioides, Z. capitata, Z. persica, Z. tenuior, Z. taurica subsp. taurica, Z. taurica subsp. cleonioides. It is difficult to distinguish between some Ziziphora taxa because of their morphological similarities. In this study, the leaf and calyx trichomes of Ziziphora taxa in Turkey were studied in order to assess anatomical variations that may serve as distinguishing characters. Their micromorphological features were surveyed by scanning electron microscopy (SEM) and light microscopy (LM). Trichomes on leaves and calyx can be divided into two general types: non‐glandular trichomes and glandular (secretory) trichomes. The non‐ glandular trichomes are simple, acicular or curved with cuticular micropapillae. They usually consist of one or more additional cells. The glandular trichomes are divided into two types: peltate and capitate and Ziziphora taxa can easily be distinguished by presence/absence, density and types of glandular trichomes on leaves and calyx. The peltate trichomes consist of 12 or 18 secretory head cells in a single disc; four or six central cells surrounded by eight or twelve peripheral ones. Peltate trichomes are absent on the adaxial leaf surface of Z. capitata and Z. persica. Two types of capitate trichomes are present in Ziziphora. The capitate trichomes are only absent on the calyx surface of Z. persica. In addition, the trichome micromorphology provides some support for separating the two subspecies of Z. taurica. In conclusion, Ziziphora taxa can easily be distinguished by cell number, cell shape presence/absence and density of the glandular trichomes on leaves and calyx.     

Cypsela micromorphology and anatomy in Cirsium sect. Epitrachys (Asteraceae,Carduoideae) and its taxonomic implications

Cirsium Miller, one of the largest genera of the Asteraceae (Carduoideae), contains more than 250 taxa. Turkey is one of its diversity centers and according to recent studies this genus is represented by 80 taxa. In the present study, cypsela morphologies, micromorphologies and anatomical structures of 17 C. sect. Epitrachys taxa distributed in Turkey are characterized in detail by using stereomicroscope, light microscope (LM) and scanning electron microscope (SEM). Cypselas are found to be obovate to narrowly oblong, or rarely oblanceolate. Based on SEM characters, six cypselas surface ornamentation types are identified, pericarp anatomical structures are described, as well as the structure of the testa. Crystal formations were observed in three taxa and secretory materials in the mesocarp were evaluated. Data obtained from this study are compared with previously published results and it is concluded that fruit micromorphological and anatomical characters are useful for the delimitation of taxa in Cirsium and may be useful for supraspecific classification.     

Morphological studies on the nutlet of Turkish Cyclotrichium Manden. & Scheng. (Lamiaceae)

Morphological characters of the nutlets of Cyclotrichium Manden. & Scheng. in Turkey were examined using scanning electron microscopy (SEM). The genus Cyclotrichium is represented by six species in Turkey, of which two are endemic for Turkey. Nutlet morphology of the species was studied using stereoscopic microscopy and SEM, and the features were described and illustrated. Nutlet morphology of the examined specimens exhibits some variation in size, shape and sculpture. Nutlets range from 1.1 to 2.0 mm in length and 0.5 to 1.0 mm in width and are elliptic, oblong to ovoid and trigonous in shape. Nutlet sculpture can be divided into two main types: distinctly or weakly reticulate pattern in C. niveum, C. origanifolium, C. longiflorum and rounded cell arrangement in the remaining species. Micromorphological characters could be useful in solving taxonomic problems of Cyclotrichium.     

Morphology, anatomy, and taxonomic position of Pagameopsis (Rubiaceae-Rubioideae)

Morphological and anatomical features (including wood anatomy and pollen morphology) of the small neotropical genusPagameopsis (Rubiaceae-Rubioideae) are discussed and illustrated. The fused ovaries, fenestrate corolla tube, basally attached anther filaments, and absence of raphides are especially noteworthy.Pagameopsis is definitely not a member of the Psychotrieae because of significant differences in wood anatomy and gynoecial and fruit structure. A close affinity with Gaertnereae seems doubtful for similar reasons. The taxonomic affinities ofPagameopsis remain obscure. The genus shows similarities with several taxa of the Rubioideae, such as Coccocypseleae, Morindeae, Hedyotideae, andMetabolos. The absence of raphides, on the other hand, makes a position in the Rubioideae doubtful.     

Pollen morphology of some endemic Turkish Centaurea L. (Asteraceae, section Phaloletis) and their taxonomic value

In this study, the detailed pollen morphological structures of some endemic Turkish species of Centaurea amaena Boiss., C. antalyense H. Duman & A. Duran, C. aphrodisea Boiss., C. hierapolitana Boiss., C. luschaniana Heimerl, C. lycia Boiss., C. tossiensis Freyn. Et Sint., and C. wagenitzii Hub.-Mor. (Asteraceae, section Phaloletis) were studied under light microscopy (LM) and scanning electron microscopy (SEM) for the first time. LM and SEM investigations showed that the pollen grains of eight taxa are more or less spheroidal-subprolate, the amb triangular and tricolporate. The exine sculpture is tectate, microechinate-scabrate in the pollen of Centaurea taxa. Spinules are less dense in Centaurea amaena, C. antalyense, C. hierapolitana, and C. lycia, but they are more dense in C. aphrodisea, C. luschaniana, C. tossiensis, and C. wagenitzii. Spinule dimensions are different from each other. The exine has one layer of columellae beneath the spines. We determined all taxa that have the Helianthoid type. Exine sctructure and sculpture as well as spine density and dimensions in Asteraceae are the most reliable characteristics for discriminating taxa.     

Functional implications of the staminal lever mechanism in Salvia cyclostegia (Lamiaceae)

Background and Aims

Flower morphology and inflorescence architecture affect pollinator foraging behaviour and thereby influence the process of pollination and the reproductive success of plants. This study explored possible ecological functions of the lever-like stamens and the floral design in Salvia cyclostegia.

Methods

Flower construction was experimentally manipulated by removing either the lower lever arms or the upper fertile thecae of the two stamens from a flower. The two types of manipulated individuals were intermixed with the control ones and randomly distributed in the population.

Key Results

Removing the sterile lower lever arms significantly reduced handling time per flower of the main pollinator, Bombus personatus. Interestingly, this manipulation did not increase the number of flowers probed per plant visit, but instead reduced it, i.e. shortened the visit sequence of the bumble-bees. Both loss of staminal lever function by removing lower lever arms and exclusion of self pollen by removing upper fertile thecae significantly reduced seed set per flower and seed set per plant. Both the manipulations interacted significantly with inflorescence size for the effect on female reproductive output.

Conclusions

Though the intact flowers demand a long handling time for pollinators, the reversible staminal lever is of advantage by promoting dispersal of pollen and thus the male function. The particular floral design in S. cyclostegia contributes to the floral constancy of B. personatus bumble-bees, with the lower lever arms acting as an optical cue for foraging cognition.     

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.     

Nutlet micromorphology of Iranian Nepeta (Lamiaceae) species

To evaluate nutlets characteristics for systematic relationships, a comprehensive morphological and micro‐morphological study of the nutlets of 16 Iranian taxa of Nepeta (Lamiaceae) was conducted using scanning electron microscopy (SEM). Differences in surface ornamentation, size, shape and color were observed between the species. The studied taxa were categorized in two basic types based on surface ornamentation: non‐sculptured (smooth‐type) and sculptured nutlets. The non‐sculptured nutlets type could be further divided into four sub‐types, including smooth, granulate, undulate‐cellular and reticulate. In the sculptured nutlets, three subtypes of tuberculate, tuberculate‐cellular and verrucate were recognized. The shape of nutlets were described as oblong, ovoid‐oblong, oblong‐linear, to elliptic‐oblong and their size range are 1.2–2.0 mm in length and 0.5–2.0 mm in width. Based on micrographs, the areole form and location were defined as bi‐lobed, straight, basal, sub‐basal or lateral. Nutlet micromorphological characteristics such as surface ornamentation can be useful for classification and identification of e.g. medicinal species of Nepeta in Iran.     

Salvia (Lamiaceae) is not monophyletic: implications for the systematics, radiation, and ecological specializations of Salvia and tribe Mentheae

Salvia, with over 900 species from both the Old and New World, is the largest genus in the Lamiaceae. Unlike most members of the subfamily Nepetoideae to which it belongs, only two stamens are expressed in Salvia. Although the structure of these stamens is remarkably variable across the genus, generally each stamen has an elongate connective and divergent anther thecae, which form a lever mechanism important in pollination. In a preliminary investigation of infrageneric relationships within Salvia, the monophyly of the genus and its relationship to other members of the tribe Mentheae were investigated using the chloroplast DNA regions rbcL and trnL-F. Significant conclusions drawn from the data include: Salvia is not monophyletic, Rosmarinus and Perovskia together are sister to an Old World clade of Salvia, the section Audibertia is sister to subgenus Calosphace or the monotypic Asian genus Dorystaechas, and the New World members of section Heterosphace are sister to section Salviastrum. Owing to the non-monophyly of Salvia, relationships at the next clearly monophyletic level, tribe Mentheae, were investigated.     

The taxonomic value of leaf anatomy and trichome morphology of the genus Cyclotrichium (Lamiaceae) in Turkey

The genus Cyclotrichium (Boiss.) Manden. & Scheng. is represented by six species in Turkey: C. glabrescens, C. leucotrichum, C. longiflorum, C. niveum, C. origanifolium and C. stamineum. They are aromatic perennial subshrubs used as spices or herbal teas in traditional Turkish medicine. The leaf anatomy and tomentum morphology of leaves and calyces of Cyclotrichium species in Turkey was investigated by scanning electron microscopy (SEM) and light microscopy (LM). The investigated species have equifacial (C. niveum, C. origanifolium) or bifacial leaves (C. glabressens, C. leucotrichum, C. longiflorum, C. stamineum). All species have peltate and capitate glandular, and simple (all species) or branched (C. niveum) eglandular trichomes and diacytic stomata. Peltate trichomes consist of a basal cell embedded in the epidermis, a stalk cell, and a broad 12 (–13)‐celled secretory head arranged in two concentric circles. The capitate trichomes observed in Cyclotrichium can be grouped into five types, differing in structure and size. They consist of either a pear‐shaped or globose unicellular head and uni‐or bicellular stalk, or a bicellular head and bicellular stalk. The density of peltate, capitate and eglandular trichomes differs between species. Peltate trichomes are densely spaced only on the calyx and on the leaf surface of C. niveum and C. origanifolium and on the abaxial leaf surface of C. longiflorum and C. stamineum. The significance of trichome architecture for taxonomy in Cyclotrichium and Lamiaceae in general is discussed.     

The anatomical and micromorphological properties of three endemic and medicinal Salvia species (Lamiaceae) in Erzincan (Turkey)

In this study, we comparatively investigated three endemic Salvia species spreading in Erzincan (Turkey) in terms of anatomy and micromorphology. For anatomical investigation, cross sections taken from stems and leaves of the species were examined under light microscope. For micromorphological investigation, epidermal surface and nutlet structure were examined using scanning electron microscopy. The S. euphratica and S. divaricata species were examined anatomically and micromorphologically for the first time, and S. hypargea was examined micromorphologically for the first time. In anatomical examinations, it was seen that stem and leaf structures of the species were similar. In micromorphological analyses, it has been seen that hairiness of the nutlet surface and nutlet ornamentations (verrucate and rugose type) created a difference between the species.     

Anatomy,trichome morphology and palynology of Salvia chrysophylla Stapf (Lamiaceae)

The anatomy, palynology, morphology and distribution of the trichomes on the aerial parts of Salvia chrysophylla Stapf, an endemic species in Turkey, were studied in order to understand the usefulness of these characteristics for systematic purposes. Some anatomical characters such as (1–)2–24-rowed pith rays in roots, dorsiventral leaves, obviously larger upper epidermal cells, and two to three large vascular bundles in the center and two to four small subsidiary bundles in the wings of petiole provide information of taxonomical significance. Three main types of trichomes were observed on the stem, inflorescence axis, leaf and calyx surfaces of S. chrysophylla. They are peltate, capitate glandular and non-glandular. Capitate glandular and non-glandular trichomes were further subdivided into several kinds. Glandular trichomes are present in abundance on the inflorescence axis and calyx, but non-glandular ones were mainly situated on the leaf and stem. Scanning Electron Microscopy (SEM) studies on the pollen grains have revealed that they are oblate-spheroidal and their exine ornamentation is bireticulate-perforate.     

Comparative leaf micromorphology and anatomy of the dragon tree group of <Emphasis Type="Italic">Dracaena</Emphasis> (Asparagaceae) and their taxonomic implications

Micromorphological features of the leaf epidermis and the inner structure of leaf tissues of eight arborescent taxa of the genus Dracaena were analysed using light and scanning electron microscopy. The plants are xeromorphic or mesomorphic. Their leaves are isobilateral and amphistomatic, and the stomata are anomocytic and tetracytic. The mesophyll in all the species is divided into an outer chlorenchyma and a central region with colourless water-storage cells, chlorophyll cells and vascular bundles. Water-storage cells have wall bands and reticulate thickenings on the walls. The article describes and illustrates several new quantitative and qualitative leaf characters of the dragon tree group. Our findings can be used to identify the dragon tree group leaves, while the shape of epidermal cells and stomata types may be useful in the identification and classification of fragments of fossil leaves. We conclude that D. ombet and D. schizantha are not two distinct species, but should be treated as subspecies of D. ombet. Leaf characters, especially stomata depth on adaxial epidermis, height of adaxial epidermal cells and the presence and thickness of hypodermal fibre bundles markedly differ between geographical groups: Macaronesian species (D. draco and D. tamaranae), the species found in East Africa and Arabian Peninsula (D. ombet subsp. ombet, D. ombet subsp. schizantha, D. serrulata and D. cinnabari) and Southeast Asian species (D. kaweesakii and D. jayniana).     

Floral micromorphology and microsporogenesis of the gynodioecious herb Glechoma longituba (Lamiaceae)

Gynodioecy, the phenomenon of having both hermaphrodite and female (i.e. male‐sterile) individuals within the same population, is an important intermediate step in the evolution of separate sexes in flowering plants. In this study, we investigated the floral micromorphology and microsporogenesis of the gynodioecious herb Glechoma longituba from four natural populations in Korea. The floral micromorphological characters of the different sex morphs were examined and compared using scanning electron microscopy (SEM), and the ultrastructure of microspores during microsporogenesis was studied. We also examined the development of anthers and pollen grains in the three sexual morphs (i.e. hermaphrodites, females, and gynomonoecious, i.e. individuals with a mixture of female and hermaphroditic flowers) by embryological investigation. The major difference in anther development between the three phenotypes was the early disintegration of the tapetal cells in the anthers of female flowers. While mature fertile pollen grains were found in both hermaphrodite and gynomonoecious phenotypes, females did not produce any pollen grains. In addition, both fertile and sterile pollen grains in gynomonoecious phenotypes were frequently observed. The results of the present study indicate that floral micromorphological characters were not distinct between sexual morphs of G. longituba, except for the structure of the inner cell surfaces of the anther. The observed tapetum abnormalities and degeneration of pollen grains in both gynomonoecious phenotypes and females may be the consequence of inbreeding depression in hermaphrodites.