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The genus Phlomoides (Lamiaceae, tribe Phlomideae) has been recently re-circumscribed to include members of Eremostachys and a few other genera. As a consequence, several new combinations and new names have been suggested in this genus recently. Here, typification of 14 names in Eremostachys, viz. E. adpressa, E. alberti, E. boissieriana, E. czuiliensis, E. eriocalyx, E. gymnocalyx, E. hissarica, E. iliensis, E. laciniata var. labiosiformis, E. sarawschanica, E. speciosa var. schugnanica, E. subspicata, E. trautvetteriana and E. zenaidae is presented. Moreover, two new combinations (Phlomoides calophyta and P. transjordanica) and two new synonyms (P. kermanica and P. semnanensis) are proposed.

Stachydeae, comprising c. 470 species, are one of the most diverse and taxonomically puzzling groups in Lamioideae. In the present study, the phylogenetic relationships in the Eurystachys clade (a phylogenetic name for all genera attributed to Stachydeae except Melittis) were reconstructed utilizing nuclear ribosomal DNA sequences (nrETS, 5S-NTS) from 148 accessions in 12 genera. Our phylogenetic results recovered Stachys as paraphyletic with numerous traditionally recognized genera nested in it. A broadly defined Eurystachys clade, however, was monophyletic. Unlike previous studies, the present study was able to resolve the group into 12 well-supported clades, named here as (1) Eriostomum, (2) Stachys, (3) Prasium, (4) Setifolia, (5) Distantes, (6) Burgsdorfia, (7) Hesiodia, (8) Empedoclia, (9) Sideritis, (10) Marrubiastrum, (11) Swainsoniana and (12) Olisia. These 12 clades were formally named in a phylogenetic nomenclature for the Eurystachys clade. Several infrageneric units were retrieved as monophyletic, namely Sideritis sections Burgsdorfia, Empedoclia and Hesiodia, Sideritis subgenus Marrubiastrum and Stachys sections Eriostomum (including Stachys section Mucronata) and Setifolia. The findings of this study also provide the basis for a future formal classification, with two options: (1) splitting of the Eurystachys clade into 12 monophyletic genera, all of them based on pre-existing genus names and redefined to encompass additional taxa, but without clear morphological apomorphies; or (2) lumping of all segregates into a broadly defined Stachys, including widely recognized and well-defined segregates such as Prasium and Sideritis.

Background: A robust molecular phylogeny is fundamental for developing a stable classification and providing a solid framework to understand patterns of diversification, historical biogeography, and character evolution. As the sixth largest angiosperm family, Lamiaceae, or the mint family, consitutes a major source of aromatic oil, wood, ornamentals, and culinary and medicinal herbs, making it an exceptionally important group ecologically, ethnobotanically, and floristically. The lack of a reliable phylogenetic framework for this family has thus far hindered broad-scale biogeographic studies and our comprehension of diversification. Although significant progress has been made towards clarifying Lamiaceae relationships during the past three decades, the resolution of a phylogenetic backbone at the tribal level has remained one of the greatest challenges due to limited availability of genetic data. Results: We performed phylogenetic analyses of Lamiaceae to infer relationships at the tribal level using 79 protein-coding plastid genes from 175 accessions representing 170 taxa, 79 genera, and all 12 subfamilies. Both maximum likelihood and Bayesian analyses yielded a more robust phylogenetic hypothesis relative to previous studies and supported the monophyly of all 12 subfamilies, and a classification for 22 tribes, three of which are newly recognized in this study. As a consequence, we propose an updated phylogenetically informed tribal classification for Lamiaceae that is supplemented with a detailed summary of taxonomic history, generic and species diversity, morphology, synapomorphies, and distribution for each subfamily and tribe. Conclusions: Increased taxon sampling conjoined with phylogenetic analyses based on plastome sequences has provided robust support at both deep and shallow nodes and offers new insights into the phylogenetic relationships among tribes and subfamilies of Lamiaceae. This robust phylogenetic backbone of Lamiaceae will serve as a framework for future studies on mint classification, biogeography, character evolution, and diversification. Graphical abstract: [Figure not available: see fulltext.]. © 2021, The Author(s). Ten Thousand Talent Plans for Young Top-notch Talents of Yunnan Province: YNWR-QNBJ-2018-279National Science Foundation, NSF: TUBITAK-BIDEB 2219, DEB-1655611RG19-172019FI009 This study was in part supported by Yunnan Fundamental Research Projects (Grant No. 2019FI009) and “Ten Thousand Talents Program of Yunnan” (Grant No. YNWR-QNBJ-2018-279) awarded to CLX, the CAS “Light of West China” program to CLX and YPC, the Australian Biological Resources Study National Taxonomy Research Grant Program (Grant No. RG19-17) awarded to TCW, CLX, and BL, NSF DEB-1655611 awarded to BTD, and the Postdoctoral Research Program (TUBITAK-BIDEB 2219) awarded to FC.

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.

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.

Despite a number of recent molecular phylogenetic studies on Phlomoides, in terms of trichome morphology the genus is still among the most poorly studied taxa in the family Lamiaceae. In order to test the utility of trichome characters for delimitation of sections, subsection and species of Phlomoides, we examined trichomes of 64 species representing all recognized sections and subsections using stereomicroscopy and scaning electron microscopy. Two basic types of trichomes could be identified: non‐glandular and glandular. Both trichome types can be simple or branched. The glandular trichomes were sessile, short stalked or long stalked. Different kinds of branched trichomes were observed in most species of P. sect. Phlomoides, i.e. symmetrically stellate, stellate with a central long branch, bi‐ or trifurcate. The species of P. sect. Filipendula were mostly covered by simple trichomes. Moreover, variation in trichome characters appears to have particular value, not only for the classification at sectional or subsectional rank, but also for delimitation of species within each section. For example, all studied species of P. subsect. Fulgentes are characterized by various kinds of stellate trichomes, while the trichome variability in P. subsect. Tetragonae was sufficiently high for species discrimination. An ancestral character state reconstruction was performed in order to investigate the evolution of trichome types and it revealed the following evolutionary trends in trichome characters of Phlomoides: 1) branched trichomes are primitive in Phlomoides as compared to simple ones, 2) long simple non‐glandular trichomes are derived as compared to short simple ones and 3) the presence of stalked glandular trichomes is advanced as compared to subsessile or sessile ones.

Trichomes of 60 taxa of tribe Marrubieae were investigated using light and scanning electron microscopy (SEM) in order to assess their importance in generic delimitation. In general, two main types of trichomes were observed: glandular (capitate, peltate or branched) and non-glandular (simple or branched) trichomes. The results revealed trichome micromorphology as useful for taxon delimitation within Marrubieae at various ranks, especially at the generic level in accordance with molecular investigations. Among the characters investigated, regular (non-peltate) glandular trichomes with capitate heads (stalked or sessile) and short simple trichomes were common among all studied species. Peltate glandular trichomes were also common except for Acanthoprasium. Long simple non-glandular trichomes were apomorphic and were found in all species of Ballota. In some species of Pseudodictamnus the glandular trichomes were branched. This condition is likely derived, and might be useful in subgeneric classification. Additional character state assessments in Marrubieae in terms of trichome morphology include: long trichomes are apomorphic versus the short ones, branched trichomes are apomorphic versus the simple ones, smooth surfaces of trichomes are apomorphic versus papillate surfaces, and peltate glandular trichomes are apomorphic with respect to the non-peltate ones.

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.

Trichomes of 32 species of the genus Phlomoides representing its various natural groups were examined using scanning electron (SEM) and light microscopy (LM). Trichomes varied among different species of this genus but were identical among different populations of a certain species, and therefore provide valuable characteristics in delimitation of sections and species. The indumentum might bear the following trichomes alone or a mixture of them: glandular, non-glandular, branched, simple, papillate, and/or smooth on surface. Two basic types of trichomes can be identified: non-glandular and glandular. Both trichome types can be simple or divided. According to their size, simple non-glandular trichomes can in turn be subdivided into three subtypes: short, long and extremely long (longer than 3 mm). Variation in trichome characters appears to have particular value, not only in classification at sectional rank, but also in separating different species of Phlomoides from their relatives. For example, most species of P. sect. Filipendula are characterized by dense extremely long or lanate trichomes. Moreover, P. tuberosa is unique among all studied species by having branched non-glandular trichomes papillate on surface. This result indicates the presence of branched trichomes with papillate surface as characteristic features of indumentum in the type species of this section. The following evolutionary trends are suggested here regarding trichome characters for Phlomoides : (1) multi-nodal branched trichomes are primitive against branched trichomes, (2) extremely long trichomes are primitive against the long or short simple trichomes, and (3) loss of glandular trichomes is advanced against their presence.

Although tribe Stachydeae (Lamiaceae) is considered monophyletic, relationships within the tribe are still poorly understood. The complexity of Stachydeae includes paraphyletic genera, considerable morphological plasticity, a range of ploidy levels, and presumably frequent natural hybridization. We performed parsimony and Bayesian phylogenetic analyses of nuclear (ribosomal ITS) and plastid (trnL intron, trnL-trnF spacer, rps16 intron) DNA sequence data from a taxonomically and geographically broad sampling of the tribe to identify major evolutionary lineages and to test taxonomic hypotheses within this largest of all lamioid tribes. We included 143 accessions corresponding to 121 species, representing both Old and New World species, and all 12 recognized genera of tribe Stachydeae. Both nuclear and plastid data corroborate monophyly of the tribe, with Melittis as sister to all remaining Stachydeae. For the latter well-supported clade, we suggest the phylogenetic name Eurystachys. Within Eurystachys, although monophyly is supported by both nuclear and plastid data for several named and unnamed groups, the majority of recognized taxa appear to be para- or polyphyletic. The taxon compositions of most subclades are congruent between the plastid and nuclear tree topologies, whereas their relative phylogenetic placements are often not. This level of plastid-nuclear incongruence suggests considerable impact of hybridization in the evolution of Stachydeae.

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 (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.

The new species Phlomoides binaludensis (Phlomideae, Lamioideae, Lamiaceae) from Binalud Mountains in the Khorasan province in NE Iran, is here described and illustrated. It belongs to Phlomoides section Filipendula and resembles P. labiosiformis and P. laciniata in general morphology, but differs primarily in corolla and calyx size, corolla color and stem indumentum. Notes on the ecology and conservation status of the new species are also given.