Anatomical and nutlet differentiation between <Emphasis Type="Italic">Teucrium montanum</Emphasis> and <Emphasis Type="Italic">T. polium</Emphasis> from Turkey

Teucrium montanum L. and T. polium L. are the two closest Teucrium L. species from sect. Polium (Mill.) Schreb in Turkey. In addition, they are sympatric for some part of their range in Turkey. In this study, comparative anatomical and micromorphological studies of the two species are carried out. They have been investigated by their leaf and stem anatomical features, as well as nutlet micromorphological characteristics. The results of anatomical studies show that the anatomical characters of both taxa are observed to be similar to the general features of the family Lamiaceae anatomy, except for lacking rich collenchyma at the corners. Both taxa are similar in stem anatomy and their leaves exhibit xeromorphy. However, trichome morphology on the stems and the leaves appear to have a taxonomic value in segregation of the two taxa. Light and scanning electron microscope studies on the nutlets also show that nutlet shapes, measures and surface micromorphologies are different in the two species. Whereas nutlet surfaces are bireticulate in both species, the nutlets are larger and primary sculpturing is more distinct in T. polium than in T. montanum. Moreover, the nutlets are oblong to oblong-ovoid and larger in T. polium, but ovoid and smaller in T. montanum.     

Nutlet micromorphology of Carex section Rhomboidales sensu Kükenthal (Cyperaceae) and its systematic implications

The sectional delimitation of Carex section Rhomboidales sensu Kükenthal (1909) is problematic and, here, we provide contributions to a systematic understanding of nutlet micromorphology in section Rhomboidales. Nutlet micromorphology of 87 samples representing 71 taxa was investigated using scanning electron microscopy and nine nutlet types were recognized. Although some closely related taxa show similarities, nutlet morphology in section Rhomboidales is stable at the species level and is reliable for species identification. Based on the nutlet characters, section Rhomboidales is delimited and emended as having nutlet rhombic–ovoid, trigonous; beaks conspicuous, erect, rarely curved or coiled; periclinal walls of epidermal cells straight, with zero to two silica bodies. The species with nutlets of the C. harlandii‐type and C. thibetica‐type are included, and species of the C. chinensis‐type and C. macrandrolepsis‐type are ascribed to section Mitratae and section Infossae, respectively. Carex longirostrata and C. pseudolongirostrata (= C. nodaeana), formerly ascribed to section Careyanae or section Depauperatae, are well supported as members of section Rhomboidales. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 123–143.     

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.     

Nutlets micromorphology of some <Emphasis Type="Italic">Onosoma</Emphasis> L. (Boraginaceae) species from Turkey

Morphological and micromorphological characters of the nutlets of ten Onosma species in Turkey were reported. Studied fruit surfaces show some variations. Based on the structure and patterns of epidermis cells of the nutlets, two principal types can be distinguished. In Type 1, the epidermal cells of the nutlet surface are formed in a reticulate pattern with varied sizes and shapes of the meshes. This type is the most common type in the studied species. Seven of the species observed belong to the reticulate type, i.e. O. bracteosum, O. mutabile, O. aleppicum, O. bornmuelleri, O. aucheranum, O. araraticum and O. tauricum. In Type 2, the epidermal cells of the nutlet surface are formed in an elongated type with varied sizes and shapes. Three of the observed species belong to this type, i.e. O. roussaei, O. isauricum and O. stenolobum. Significant taxonomic differences between species were determined on the surface of the nutlets.     

Nutlet micro‐morphology of the genus Microula (Boraginaceae) from the Qinghai–Tibetan Plateau,and its systematic implications

The micro‐morphology of nutlets in 30 taxa (28 species and 2 varieties) of Microula Benth. (Boraginaceae) was surveyed by light and scanning electron microscopy. A principal coordinate‐based ordination analysis (PCO) (8 morphological and 14 micro‐morphological characters) was applied to explore whether taxa clustered according to the current sectional taxonomy. The nutlet epidermis showed three major features: trichomes, papillae and lamellae. Two basic trichome types could be identified: branched and unbranched. Stellate trichomes (STT) and stipitate stellate trichomes (SST) can be considered as two subtypes of branched trichomes based on shape and branching location. Three kinds of unbranched trichomes were recognized by their shapes: slender trichomes (SLT), spear‐like trichomes (SPT) and conical trichomes (CT). Four types of papillae were revealed in this genus: central apiculate papillae (CAP), debris‐covered convex papillae (DCP), rugous papillae (RP) and tabular papillae (TP). The lamellae was classified in five types: glossy lamellae (GL), thread‐like lamellae (TL), rugous lamellae (RL), interrupted lamellae (IL) and debris‐covered lamellae (DCL). The variation in the nature of surface sculpturing, nutlet shape and size, dorsal aperture shape, and attachment scar positions proved useful diagnostic characters. Taxa belonging to the same section generally clustered together in the PCO analysis. A key based on the morphology and micro‐morphology of nutlets is provided to distinguish the sections of Microula. The presence of SLT, STT, DCP and DCL is consistent with the macro‐morphological classification and provides additional evidence for the subgeneric delimitations. The SST, SLT, CAP, DCP, RP, DCL, GL and RL are useful in discriminating species in Microula. Additionally, the distribution of some trichomes, papillae and lamellae types is correlated with the altitudinal distribution of the species.     

Nutlet sculpturing inScutellaria sect.Resinosa (Lamiaceae) and its taxonomic utility

Nutlet sculpturing of 22 taxa ofScutellaria sect.Resinosa, representing 18 of its 19 species, and the monotypic genusSalazaria was studied by scanning electron microscopy (SEM). The nutlets of sect.Resinosa, while remarkably variable among themselves, differed from hypothetically closely related sections in lacking circumferential banding and sessile glands. Species could be grouped by surface sculpturing into seemingly meaningful phenetic clusters.Scutellaria potosina var.platyphylla was sufficiently distinct micromorphically so as to be elevated to specific status asScutellaria platyphylla comb. and stat. nov. The significance of nutlet microcharacters within sect.Resinosa is discussed, especially as compared to closely related sections. The genusSalazaria had nutlet sculpturing quite different from all taxa belonging to sect.Resinosa.     

Comparative anatomical and micromorphological studies on Teucrium creticum and Teucrium orientale var. orientale (T. sect. Teucrium, Lamiaceae)

Teucrium creticum L. and T. orientale L. var. orientale are two related taxa in Teucrium sect. Teucrium . In this study, the value of anatomical and micromorphological characters for distinguishing between these two taxa is analysed. Transverse sections of the stem, as well as both transverse and surface sections of the leaves were examined anatomically. According to the data obtained, collenchyma and sclerenchyma do grow more in T. creticum than in T. orientale var. orientale . The leaves of T. creticum are hypostomatic and have a characteristic hypodermis, which is under the upper epidermis, as well as abundant spherocrystals in the upper epidermis. The leaves of T. orientale var. orientale are amphistomatic, yet no spherocristals exists in epidermal cells and the hypodermis is absent. The results obtained from studies carried out under scanning electron microscope (SEM) shows that the nutlet micromorphology varies between the two taxa. In both taxa, the nutlet surfaces are reticulate and glandular. Eglandular hairs do occur. However, in T. orientale var. orientale , 20–60 μm long eglandular hairs with scabrid surface occur only along the margins of the nutlets, whereas the ¾ dorsal sides of the nutlets in T. creticum are all covered with 90–500 μm long, smooth-surfaced hairs. Additionally, while the glandular hairs on the nutlets of T. creticum are subsessile, those on the nutlets of T. orientale var. orientale are capitate and distinctly stalked.     

FLAVONOIDS AND TAXONOMY OF THE LIMNANTHACEAE

To help clarify relationships within the Limnanthaceae, all 19 taxa were compared on the basis of flavonoids occurring in all tissues, and 14 of these taxa were additionally compared on the basis of flavonoids occurring only in the petals. Of the 46 flavonol glycosides encountered, 35 were identified as derivatives of six flavonol aglycone types: syringetin, isorhamnetin, kaempferol, laricytrin (myricetin 3'-methyl ether), quercetin and myricetin, all glycosylated with combinations of glucose and rhamnose. Varimax Factor Analysis with rotation of the flavonoid data indicated that the family probably contains 3 phyletic lines, an observation inconsistent with the conventional 2-generic interpretation of the family. Mason's sectional treatment of Limnanthes is supported by petal flavonoid results, but not by whole-plant flavonoid results, indicating that petal flavonoids more clearly reflect natural relationships in Limnanthes. Evolution of whole-plant flavonoids of Limnanthes appears to be partly linked to changes in breeding system.     

Ultrastructural features of the nutlet surface in species of the genus <Emphasis Type="Italic">Mentha</Emphasis>

The surface of nutlets and the ultrasculpture of еру pericarp were studied for the first time using a scanning electron microscopy in nine species of the genus Mentha, growing in Russia and neighboring countries. Three types of primary and four types of secondary seed surface sculptures were identified. It was found that the ultrasculptural features of the nutlet pericarp can be used in the taxonomy of the genus at the level of species and for identifying the relationship between taxa.     

Taxonomic reinstatement of Teucrium andrusi from T. paederotoides based on morphological and anatomical evidences

Teucrium andrusi, described by Post from Mardin province, Turkey, was synonimized with T. paederotoides in the flora of Turkey. According to the records in the flora of Turkey T. paederotoides was a steno‐endemic species, only known from the type locality in Gaziantep and from Mardin province. Specimens of T. paederotoides s.l. were collected from both localities and a detailed comparison of the plants from Gaziantep with those from Mardin revealed that the two populations were sufficiently distinct from each other to be recognised as separate species. Thus, T. andrusi Post is reinstated from the synonymy with T. paederotoides. Teucrium andrusi is morphologically different from T. paederotoides s.s. in terms of its general indumentum, leaf margin, bract and calyx shape, corolla colour and nutlet macro–micromorphology. It can be anatomically distinguished from T. paederotoides s.s. by stoma type, stomatal distribution on the leaf surface, crystal content in the cells and cuticula thickness. Ecological data related with the accompanying species also support species status for T. andrusi. While the two taxa are very similar in pollen and nutlet morphology, trichome density on the nutlets are different.     

Nutlet morphology in the tribe Elsholtzieae (Lamiaceae)

A comprehensive study on nutlet morphology on 46 taxa from the tribe Elsholtzieae was conducted using stereomicroscopy, light microscopy and scanning electron microscopy. The objectives were to evaluate the nutlet characteristics in a systematics context and to provide additional evidence for taxon delimitation. The nutlet shape varied from elliptical, widely elliptical, spheroidal and ovate to pyriform. The size ranged from 0.50 to 2.70 mm in length and 0.34 to 2.65 mm in width. The three types of trichomes included simple non-glandular trichomes (Collinsonia punctata), clavate non-glandular trichomes (Elsholtzia cephalantha) and peltate glandular trichomes (Mosla grosseserrata and M. chinensis). We also report the presence of a mucilaginous reaction in nine species of the genus Elsholtzia in Elsholtzieae for the first time. Four types (undulate, papillae-like projection, thorn-like pattern and reticulate) and three subtypes (reticulate, reticulate-cellular and supra-reticulate) of nutlet surface patterns in the mid-ventral region were recognized in Elsholtzieae based on the differences in cell outline, outer periclinal walls and dentations of the anticlinal walls. These nutlet micromorphological results may serve as a reference for future hypotheses on the classification of the tribe Elsholtzieae.     

Salvia siirtica sp. nov. (Lamiaceae) from Turkey

Salvia siirtica Kahraman, Celep & Do?an sp. nov.(Lamiaceae) is described and illustrated as a new species from the northern part of Siirt province in southeast Anatolia, Turkey, where it was found growing in open forests of Quercus. It is morphologically similar to S. indica L., but differs by having densely glandular pilose to villous stems and petioles, close verticillasters, smaller flowers, clearly concave calyx apices, entirely white corollas and smaller nutlets. Salvia siirtica and S. indica are also distinguished by their nutlet and pollen micromorphologies, illustrated by SEM images. The nutlet surface of S. siirtica is very finely wrinkled and its pollen has 3–4 primary lumina per 25 μm², compared to 5–8 in S. indica. Conservation status assessment, distribution map, and notes on biogeography and ecology of the new species are provided.     

Anatomical and micromorphological studies on <Emphasis Type="Italic">Teucrium</Emphasis> sect. <Emphasis Type="Italic">Isotriodon</Emphasis> (Lamiaceae) in Turkey with a taxonomic note

In this study, the anatomical features of the leaf and stem, besides the nutlet characteristics of some Teucrium sect. Isotriodon (Lamiaceae) taxa in Turkey, T. montbretii Betham subsp. montbretii, T. montbretii subsp. pamphylicum P. H. Davis, T. odontites Boiss. &; Bal., T. cavernarum P. H. Davis, T. antitauricum T. Ekim, along with an isolated population of T. montbretii (T. montbretii subsp.) were investigated. The anatomical studies revealed that the taxa share generally similar anatomical characters, such as thicker upper leaf cuticles and larger upper leaf epidermal cells compared to lower ones and diacytic to anomocytic stomata on the leaves. However, the portion of the mesophyll occupied by palisade parenchyma and the occurrence of mucilage cells in leaf epidermis shows difference among the taxa. Furthermore, the studied taxa have general stem characteristics of the Lamiaceae family, except for having poorly developed collenchyma at the corners. With the amphistomatic leaves and developed sclerenchymatic tissue in the leaf median vein, T. cavernarum is seperated from the other taxa. Trichome types on the vegetative organs and nutlet shape and sculpturing are generally the same or similar in the studied taxa, but trichomes on the nutlets are different among them. Based on nutlet characteristics and some morphological ones, it was revealed that the isolated population of T. montbretii represent a new subspecies, T. monbretii subsp. yildirimlii M.Dinç &; S.Do?u subsp. nov.     

New variation and biosystematic patterns detected by allozyme and morphological comparisons inLimnanthes sect.Reflexae (Limnanthaceae)

Sixty-one populations ofLimnanthes sect.Reflexae were surveyed for variation at 19 allozyme loci and reassessed for their morphological diversity and biosystematic relationships. Cluster analysis of allozyme data at individual population level provided evidence for a fairly good fit with the earlier systematic work on this genus but both morphological and allozyme data together defined some new clusters as well as exceptional entities. For example, only var.rosea of the fourL. douglasii varieties described byMason (1952) on the basis of geographical distribution, flower color and leaf morphology, remained as a distinct entity; the others were realigned along new geographic boundaries.L. striata appeared to have two well-developed subgroups, one of which was closely related to theL. douglasii complex. Further ecogeographic and hybridization studies are needed to fully describe these new phenetic findings in the context of phylogenetic inference. SectionInflexae was readily distinguished from sect.Reflexae by several diagnostic loci, thus supporting the validity of this division. Interspecific variation levels and population differentiation inLimnanthes were high when compared to other plant taxa, possibly indicating highly dynamic evolutionary processes for which variable habitats and extreme subdivision within species have been suggested as explanations.     

NUMERICAL TAXONOMY OF LIMNANTHACEAE

A numerical taxonomic study of the angiosperm family Limnanthaceae is presented. The 21 OTUs include all taxa recognized in a recent monograph of Limnanthes (Mason, 1952), plus Floerkea proserpinacoides, two undescribed Limnanthes species, and two artificial interspecific F₁ Limnanthes hybrids. One species (L. montana) was treated as two OTUs: measurements for one OTU were taken from field collections and for the other, from garden-grown plants. A maximum of 35 characters was used in the numerical analyses. Raw data for each character were given equal weighting by condensation. Coefficients of resemblance were established using the mean character difference (MCD) method. Tryon's clustering technique (called key communality cluster analysis) was used. Three numerical analyses of Limnanthaceae were made: one with all 35 characters (T analysis), one with 18 floral characters (F analysis), and one with 17 pre- or post-anthesis or vegetative characters (V analysis). Relative positions of OTUs in the T and F analyses are substantially different from those in a conventional taxonomic treatment of the family. The V analysis shows many differences from the T and F analyses. In all analyses the two OTUs representing L. montana were closer to other OTUs than to each other. The two hybrid OTUs showed greater resemblance to one parent than to the other and in two analyses one hybrid was not intermediate between the parents. Similarities between the T and F analyses are explained on the basis of the predominance of floral characters in both. The differences between these two analyses on the one hand and the V analysis and phylogenetic arrangement of the family on the other are attributed to parallelisms in the evolution of floral characters associated with iucreasing autogamy in each of the two phyletic lines in the family. Analyses emphasizing floral characters will place evolutionarily remotely related outcrossing OTUs together and unrelated, autogamous OTUs together. Using characters that are mostly unassociated with the breeding system the positions of OTUs in a numerical analysis are similar to their positions in a phylogenetic arrangement. The V analysis agrees best with suggestions concerning phylogenetic affinities among the taxa in the family, and provides a useful context for the generation of further evolutionary hypotheses.     

Seed-coat microsculpturing and its systematic application in Isatis (Brassicaceae) and allied genera in Iran

In order to examine the systematic application of seed-coat microsculpturing in Isatis, seed surfaces of 23 species (41 populations) in four genera of tribe Isatideae were examined using scanning electron microscopy (SEM). Eight types of basic ornamentation patterns were recognized among the studied specimens. Of these, the reticulate–areolate type was the most common and was found in the genera Isatis, Pachypterygium, Samerari and Tauscheria and 15 species (e.g., I. cappodocica, I. kotschyana and I. tinctoria). The reticulate type, the second most frequent, occurred in 7 species while other types each were represented by only one or 2 species. Although different populations of a given species show similar seed-surface sculpturing in most cases, in some polymorphic species like I. cappadocica and I. kotschyana these patterns were variable among populations. To some extent the variation corresponds to infraspecific taxa for some species, but the differences are not significant enough to be useful in the delimitation of the subspecies recognized by previous workers. Moreover, seed-coat characters do not support the separation of genera Isatis, Pachypterygium, Sameraria and Tauscheria.     

An evaluation of putative sympatric speciation within Limnanthes (Limnanthaceae)

Limnanthes floccosa ssp. floccosa and L. floccosa ssp. grandiflora are two of five subspecies within Limnanthes floccosa endemic to vernal pools in southern Oregon and northern California. Three seasons of monitoring natural populations have quantified that L. floccosa ssp. grandiflora is always found growing sympatrically with L. floccosa ssp. floccosa and that their flowering times overlap considerably. Despite their subspecific rank within the same species crossing experiments have confirmed that their F1 hybrids are sterile. An analysis of twelve microsatellite markers, with unique alleles in each taxon, also shows exceedingly low levels of gene flow between populations of the two subspecies. Due to the lack of previous phylogenetic resolution among L. floccosa subspecies, we used Illumina next generation sequencing to identify single nucleotide polymorphisms from genomic DNA libraries of L. floccosa ssp. floccosa and L. floccosa ssp. grandiflora. These data were used to identify single nucleotide polymorphisms in the chloroplast, mitochondrial, and nuclear genomes. From these variable loci, a total of 2772 bp was obtained using Sanger sequencing of ten individuals representing all subspecies of L. floccosa and an outgroup. The resulting phylogenetic reconstruction was fully resolved. Our results indicate that although L. floccosa ssp. floccosa and L. floccosa ssp. grandiflora are closely related, they are not sister taxa and therefore likely did not diverge as a result of a sympatric speciation event.