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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.

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Nutlets of 20 taxa of Phlomoides, representing most of the currently recognized sections were examined by scanning electron microscopy. The basic shape of nutlets in most taxa studied is broadly ovate, but ovate, triangular and oblong ones can also be found in few species. Regarding the sculpturing pattern of nutlet surface, four basic types can be distinguished: reticulate, scalariform, ruminate 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 them to each other. For example, sect. Filipendula show four different types of sculpturing. Therefore, 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.
 

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Nutlets of 20 taxa of the tribe Marrubieae were examined by scanning electron microscopy (SEM) and detailed descriptions of nutlet micromorphological features for all examined taxa are provided. The nutlets exhibited variation in size, shape, color and surface sculpturing. The nutlets shape of most species studied is ovate, but rounded, broad ovate, elliptic, lanceolate, triangular and oblong can also be found in a few species. Six basic types of the sculpturing pattern of nutlet surface can be distinguished: reticulate, foveolate, scalariform, ruminate, pusticulate and colliculate-granulate. The most common type of nutlet sculpturing among the studied species is reticulate, but the variation in size and shape of their composing cells provided useful diagnostic characters. Our investigation revealed that the type of sculpturing was more useful in taxon delimitation among Marrubium species and allies at the species rank.

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Stachys sect. Satureoides includes four species characterized by being annual herbs having oblanceolate leaves, urceolate calyx in fruit and dense hairs at mouth and are distributed from Eastern and Southern Turkey, Syria, Northern Iraq to Western Iran. The main aim of present study is to determine the phylogenetic placement of S. sect. Satureoides based on Stachys melampyroides, as the only representative of this section in Iran. The present molecular study was conducted based on nuclear ribosomal ITS marker, using Maximum Parsimony and Bayesian Inference analyses. In total, 86 accessions (including nine outgroups and 77 ingroups) were included. Our results demonstrated that sect. Satureoides was placed in Olisia clade which is one of the most heterogenous and taxonomically challenging clades of Stachys as a paraphyletic genus. Moreover, not only sect. Satureoides was not found to be phylogenetically close to sect. Neurocalyx, as had been suggested by previous morphological studies, but also it was placed as a sister group of sect Fragilicaulis which was characterized by chasmophilus species with fragile stems and yellow flowers. Our results revealed that sect. Neurocalyx could be the closest group to the Satureoides+Fragilicaulis clade. In addition, based on the nrITS phylogenetic tree, annual species of Stachys showed affinity to perennial Stachys species. Hence, it seems that “Annual habit” have been evolved among their perennial ancestors, independently.