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The evolutionary trend of the morphological characters of the genus Stachys in Iran, on the basis of nrITS se-quence data, was investigated. A total of 28 nrITS sequences, representing 26 species of Stachys as well as Betonica of-ficinalis and Melittis mellisophyllum, were obtained from GenBank. Patterns of character evolution were assessed for 10 vegetative and reproductive characters with emphasis on taxonomic treatments of Stachys performed earlier to infer its history and interpret processes of change. The most important characters included habitat, growth form, leaf morp-hology, inflorescence, calyx and corolla as well as trichome morphology. Phylogenetic tree, based on the nrITS dataset, were constructed by Bayesian analysis. Parsimony mapping was performed in Mesquite v. 1.12. The present analysis revealed that the state of symmetrical calyx, the presence of basal leaves as well as the presence of glandular trichomes were not in agreement with the results of molecular data, which indicated the artificial nature of these characters in pr-evious classification. The presence of simple trichomes as well as the growth forms were consistent with phylogeny ba-sed on nrITS sequences. Evolutionary trend of morphological characters demonstrated that the sect. Eriostomum and sect. Fragilicaulis were the most primitive and the most advanced sections in Iran, respectively.

We have investigated the phylogenetic position of Stachys tibetica by use of both plastid (rps16 intron, trnL-F region and matK) and nuclear ribosomal (ITS) DNA sequence data and both parsimony and Bayesian phylogenetic methods. All data corroborate that S. tibetica is extraneous in Stachys (tribe Stachydeae) and suggest a phylogenetic affiliation in a different lamioid tribe, Lamieae. The molecular data suggest that the species is most closely related to the genus Stachyopsis, while most of the morphological data suggest an intermediate position between Stachyopsis and Eriophyton. Due to difficulties in distinguishing Stachyopsis and Eriophyton, and the possible embedding of the former in the latter genus, we prefer to amalgamate the two closely related genera and refer S. tibetica to the expanded Eriophyton. As a consequence, five new combinations are made: Eriophyton lamiflorum (Rupr.) Brauchler, Eriophyton maleolens (Rech.f.) Salmaki, Eriophyton marrubioides (Regel) Ryding, Eriophyton oblongatum (Schrenk) Bendiksby and Eriophyton tibeticum (Vatke) Ryding.

Stachys is a large and taxonomically complex genus of Lamiaceae (Lamioideae: Stachydeae). On the basis of morphological examination and field investigation, we revised the genus Stachys in Iran. We examined about 3500 specimens from various herbaria and produced a key for identification of all taxa found in Iran. The following are provided for each taxon: a description along with the complete synonymy, notes on nomenclature and relationships within the genus, geographical distribution, habitat, IUCN conservation status assessment and selected materials examined. The following taxa are reported here as new for Iran: S. atherocalyx, S. kotschyi, S. melampyroides and S. recta subsp. subcrenata. Several taxa are placed in synonymy for the first time here: S. persica is a synonym of S. alpina, S. koelzii is a synonym of S. aucheri, S. ballotiformis is a synonym of S. kurdica subsp. kurdica, and S. setifera subsp. daenensis and S. setifera subsp. iranica are synonyms of S. setifera. Two taxa are reduced to the rank of subspecies: S. kurdica subsp. asterocalyx and S. pilifera subsp. ixodes. According to the present revision, the genus contains 32 species, nine subspecies and two hybrids in Iran with 17 endemic taxa. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170, 573–617.

Stachys (Lamiaceae: Lamioideae) is a species-rich, widespread, and taxonomically complex genus. A comparative anatomical study of the petioles and leaf lamina of 34 Stachys taxa representing 12 sections of the genus distributed in Iran was carried out to evaluate interspecific relationships and anatomical features that may be useful in species identification and subgeneric classification. The general leaf anatomy of Stachys species presented here corroborates earlier studies in Lamiaceae and on a few studied species in the genus. Leaf anatomy provides valuable characters that are useful in subgeneric classification as well as species discrimination in Stachys. The most important diagnostic characters are as follows: the shape of transverse section, length of ventral and dorsiventral axis, number of median bundles in the petiole, number of cell layers of palisade and spongy parenchyma, type and thickness of collenchyma as well as trichome type. Based on the present study and in accordance with previous works, some large sections such as Ambleia, Aucheriana, and Fragilicaulis appear to be natural and need minor or no changes in delimitation, while circumscription of sect. Olisia should be revised. Isolated positions of some monotypic or oligotypic sections, such as Pontostachys, Thamnostachys, and Trinerves, as well as Zietenia, are supported by unique leaf anatomical features.

Trichomes of 37 taxa of the genus Stachys and one species of Sideritis (S. montana) were examined using light and scanning electron microscopy. The indumentum shows considerable variability among different species, but is constant among different populations of one species, and therefore, affords valuable characters in delimitation of sections and species. The characters of taxonomic interest were presence of glandular and non-glandular trichomes, thickness of the cell walls, number of cells (unicellular or multi-cellular), presence of branched (dendroid) trichomes, presence of vermiform trichomes, orientation of trichomes in relation to the epidermal surface, curviness of trichomes, and presence of papillae on trichome surface. Two basic types of trichomes can be distinguished: glandular and non-glandular trichomes. The glandular trichomes can in turn be subdivided into subtypes: stalked, subsessile, or sessile. The stalks of the glandular trichomes can be uni- or multi-cellular. Simple unbranched and branched trichomes constitute two subtypes of non-glandular trichomes. Our data do not provide any support for separation of Sideritis from Stachys. The following evolutionary trends are suggested here for Stachys: vermiform trichomes with stellate base are primitive against vermiform trichomes with tuberculate base, long vermiform trichomes are primitive against the short simple trichomes, appressed trichomes are advanced against spreading ones, and loss of glandular trichomes is advanced against their presence. Overall, trichome micromorphology is more useful in separation of species within sections rather than characterizing large natural groups known as sections, except for few cases.

Fresh leaves of 150 individuals from 15 populations representing five putative species of Stachys sect. Fragilicaulis, subsect. Fragiles distributed in the Zagros mountains in western Iran were analyzed for isoenzyme variation. Four enzyme systems (peroxidase, esterase, superoxide dismutase and ascorbate peroxidase) have been studied and a total of 32 bands representing seven putative loci have been analyzed by UPGMA using the Euclidean distances. UPGMA analysis resulted in the recognition of three distinct clusters among the considered populations. The average value of Shannon diversity index (H′) estimated for each population ranged from 0.05 to 2.17. The mean number of bands per isoenzyme ranges from 1.60 to 2.50. The value of Euclidean distance ranges from 0.85 to 3.79. The analysis of isoenzyme variation patterns suggests recognizing following species in Stachys sect. Fragilicaulis subsect. Fragiles: Stachys benthamiana (including Stachys megalodonta), Stachys kurdica (including Stachys ballotiformis) and Stachys asterocalyx. These data also suggest paying more attention to geographical distribution of taxa in Stachys for their taxonomic delimitation. This study provides another strong support for application of electrophoretic analysis of isoenzymes in taxonomy at species level.

Nutlets of 31 taxa of Stachys (29 species and two subspecies), representing nine of the currently recognized sections and one species of the closely related genus Sideritis (S. montana) distributed in Iran were examined by light and scanning electron microscopy. The basic shape of nutlets in most taxa studied is ovate, but rounded; broad ovate, oval, triangular and oblong can also be found in few species. Regarding sculpturing pattern of nutlet surface seven basic types can be distinguished: reticulate, colliculate, scalariform, verrucate, ruminate, foveate and rugose. The reticulate type is the most common among the studied species, but the variation in alignment, size and shape of its composing cells provide further evidence that are useful as diagnostic characteristics. However, the type of sculpturing is more useful for separating species within the sections, rather than correlating the related ones to each other. For example in sect. Eriostemum, five species known in Iran, show four different types of sculpturing. However, nutlet microsculpturing is not useful in separating large natural groups like sections in this genus. It seems also that contrary to other genera of Lamiaceae, nutlet characters are of low phylogenetic value in this genus, especially due to high variation even among closely related species. (© 2008 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim) Mikromorphology der Nusschen und ihre systematische Bedeutung fur die Gattung Stachys L. (Lamiaceae) in Iran Die Nusschen von 31 Taxa aus der Gattung Stachys (29 Spezies, zwei Subspezies) aus neun der gegenwartig bekannten Sektionen und eine Art aus der eng verwandten Gattung Siderites (S. montana) aus Iran wurden mittels Licht- und Elektronenmikroskopie untersucht. Die Grundform der Nusschen der hier untersuchten Taxa ist ovat aber abgerundet, bei einigen Arten wurden aber auch breit-ovate, ovale, dreieckige oder langliche Formen gefunden. Als Skulptur-Muster fur die Oberflache der Nusschen konnen sieben Grundtypen ausgeschieden werden: retikulat, kollikulat, sklariform, verrucat, ruminat, foveat und rugos. Am haufigsten bei den hier untersuchten Arten ist der retikulate Typ, aber die Variabilitat in Anordnung, Grose und Form der Zellen erfordern weitere Beweise, dass sie als diagnostisches Merkmal geeignet sind. Die Art der Skulptur ist jedoch besser zur Trennung der Arten in den Sektionen geeignet, besser als die verwandten Arten untereinander zu vergleichen. So zum Beispiel zeigen die funf iranischen Arten der Sekt. Eriostemum vier unterschiedliche Skulptur-Typen. Die Mikroskulptur ist indes nicht zur Trennung solch groser Gruppen wie die Sektionen in dieser Gattung, geeignet. So scheint es, dass, im Gegensatz zu anderen Genera der Lamiaceae, die Nusschen in dieser Gattung von geringem phylogenetischem Wert sind, ganz besonders in Bezug auf die hohe Variationsbreite zwischen verwandten Arten.

Pollen grains of 30 taxa of the genus Stachys (29 spp. and one subsp.), representing 9 of the currently recognized sections and 1 species of the closely related genus Sideritis (Si. montana) distributed in Iran were examined by light and scanning electron microscopy. Twenty-eight taxa are studied for the first time under aspects of pollen morphology. The basic shape of the pollen grains in most taxa studied is prolate-spheroidal, but subprolate, spheroidal and oblate-spheroidal pollen grains can also be found in few species. The grains are usually tricolpate (the amb triangular), but also tetracolpate (the amb circular to more or less square) in some species (S. iberica, S. atherocalyx and Si. montana). The surface is microreticulate (the frequent type), reticulate, perforate, foveolate-psilate or foveolate. The lumina are separated by smooth or sinuate muri which make them polygonal, more or less rounded and elongate. Major pollen morphological features of the taxa studied are compared and discussed on the basis of taxonomical concepts. In some cases, these characters are useful in delimitation of formerly introduced sections while they mostly provide further characters in separating related species from each other. For example, all members of S. sect. Aucheriana are characterized by elongated lumina. Based on the oblate-spheroidal shape of its pollen as well as tetracolpate aperture type, the results of the present study confirm sect. Pontostachys as including S. angustifolia, S. iberica, S. sparsipilosa as well as S. atherocalyx. Our results also suggest that although some species like S. fruticolosa and S. lavandulifolia are morphologically well characterized, they cannot be separated from other species of Stachys based on pollen morphology.

The main purpose of this study was to test the hypothesis that Stachys persepolitana is extraneous in Stachys (tribe Stachydeae, subfam. Lamioideae, Lamiaceae) and instead belongs to the genus Lamium (Lamieae, Lamioideae). We investigated the phylogenetic position of S. persepolitana using plastid (rps16 intron, trnL-F and matK regions) and nuclear (nrITS) DNA sequence data with both parsimony and Bayesian phylogenetic approaches. Plastid and nuclear data strongly support that S. persepolitana is extraneous in Stachys and belongs in fact to the genus Lamium. Morphological characters also corroborate its placement in Lamium. Most of the morphological features used to distinguish S. persepolitana from the rest of Stachys fit with Lamium. Anterior pair of stamens bending outward after pollination, anthers hairy and mericarps triquetrous are the most important characters correlating S. persepolitana to Lamium.