Fig 4 - uploaded by Charlotte Lindqvist
Content may be subject to copyright.
Representatives of Phlomideae: A, Eremostachys laciniata ; B, Paraeremostachys phlomoides ; C, Phlomoides tuberosa ; D, Phlomoides hamosa (former Notochaete hamosa ); E, Phlomis fruticosa . — Scale bar = 20 cm (A, C), 10 cm (B, D, E). (Fig. 1A: after Rechinger, 1982 and Knorring, 1954; Fig. 1B: after Ledebour, 1830; Fig. 1C: after Konorring, 1954; Fig. 1D: after Li & Hedge, 1994; Fig. 1E: after Turpin, 1829).
Source publication
The tribe Phlomideae (Lamiaceae: Lamioideae) is divided into the three genera Phlomis, Phlomoides (incl. Pseud- eremostachys, Lamiophlomis and Notochaete), and Eremostachys (incl. Paraeremostachys), contains about 278 species and has a distribution range extending from Europe to Mongolia, China, and India. Here, we present a phylogenetic analysis b...
Contexts in source publication
Context 1
... -There are about 90 species in Phlomis char- acterized by several synapomorphies (see under Introduction; Fig. 4E; Table 1). In all topologies obtained from the analysis of single plastid markers (trnT-A, rpl32-trnL, partial trnK), com- bined plastid datasets as well as ITS sequences, the few species of Phlomis s.str. included here form a strongly supported mono- phyletic group as also found by Mathiesen & al. (2011). The generic distinctiveness ...
Context 2
... included here form a strongly supported mono- phyletic group as also found by Mathiesen & al. (2011). The generic distinctiveness of Phlomis s.str. is supported by mor- phological, anatomical, cytological, and molecular data (Azizian & Culter, 1982;Ryding, 2008;Mathiesen & al., 2011). The later- ally compressed and sickle-shaped upper corolla lip (Fig. 4E) can be regarded as synapomorphy for this genus (Table 1). The following characters occur in all species of the genus, while they are variable in the other genera of Phlomideae: basal leaves absent, margin of cauline leaves not deeply lobed (shallowly crenate to entire), indumentum of branched multinodal hairs, upper lip of the corolla ...
Context 3
... most species attributed to this genus were included in Phlomis and Eremostachys. According to Adylov & al. (1986) the most important morphological characters separat- ing Phlomoides from other taxa in Phlomideae are: flowers small in size, petals pink or purple, and corolla non-compressed dome-shaped, with uneven margins densely bearded inside ( Fig. 4C; Table 1). However, the differences in corolla size and color are far from consistent. The nutlets are more often hairy in Phlomoides than in Phlomis s.str. When present, the indumentum on the nutlets consists of short stellate hairs at the apex in Phlomis. The hairs sometimes have a small gland on one of their branches (see fig. 4F in ...
Context 4
... inside ( Fig. 4C; Table 1). However, the differences in corolla size and color are far from consistent. The nutlets are more often hairy in Phlomoides than in Phlomis s.str. When present, the indumentum on the nutlets consists of short stellate hairs at the apex in Phlomis. The hairs sometimes have a small gland on one of their branches (see fig. 4F in Ryding, 2008). The phylogenies presented here show that the genus as defined by Adylov & al. (1986) is paraphyletic, as already pointed out by Mathiesen & al. ...
Context 5
... supported group nested within Phlomoides (Figs. 1-3). Hence, our results suggest that both species of Notochaete should be included in Phlomoi- des. This conclusion is also supported by similarities in corolla shape, corolla indumentum, inflorescence structure (composed of remote spherical glomerules shared by several species of Phlo- moides, Fig. 4D) and pericarp structure (Ryding, 2008). Noto- chaete longiaristata is formally transferred to Phlomoides ...
Context 6
... ent study (Figs. 1-3) is that the genus Eremostachys (sensu Harley & al., 2004 but excluding Paraeremostachys) forms a group within Phlomoides along with three divergent species of Phlomoides. Most species of Eremostachys have robust stems, laciniate leaves, large calyces, large corollas of yellow, creamy, or white color, and bearded nutlets ( Fig. 4A; Table 1). However, the dif- ferences in the leaf and corolla characters are not consistent. The most basal group of Eremostachys based on both ITS and plastid DNA phylogenies, here indicated as the E. molucel- loides group (Figs. 1-2), as well as several other species, have dentate but undivided leaves. Some other species not related ...
Context 7
... difference in nutlet indumentum, as described by Knorring (1954) and Rechinger (1982), constitutes the most important difference between the two genera. The nutlet hairs in Eremostachys differ from those in Phlomoides in being long and all simple instead of short and usually branched (see fig. 4 in Ryding, 2008). However, there are several species of Phlomoides and at least one species of Eremostachys (E. nuda Regel) that have glabrous nutlets. According to Knorring (1954), Phlomoides vavilovii, which is nested within Eremostachys, conforms to this genus in having the nutlets bearded with long hairs. Unfortunately the data on ...
Context 8
... genus Phlomoides. The nine species with funnel-shaped calyces were kept in Eremostachys, including the widely distributed E. molucelloides. Fifteen species were transferred to the newly established genus Paraeremostachys that was considered to differ from Eremostachys s.str. in having a tubular or campanulate calyx without an expanded apical rim (Fig. 4B). The phylogenetic study presented here shows that the type of Paraeremostachys (Pa. phlomoides) is nested within the E. molucelloides group (Figs. 1-2) and clearly belongs to Eremostachys s.str. Moreover, Paraeremostachys seems to be polyphyletic as one of its members (E. thyrsiflora Benth.) is nested within the E. laciniata core ...
Citations
... ex Benth.) Spach, Nepeta L., Phlomoides Moench, Salvia L., and Scutellaria L. (Paton, 1990;Walker and Sytsma, 2007;Salmaki et al., 2012a;Hu et al., 2018Hu et al., , 2020Zhao et al., 2017Zhao et al., , 2021aZhao et al., ,b,c, 2022Chen et al., 2022a;Sun et al., 2022). During the past two decades, the circumscription of Lamiaceae and evolutionary relationships within the family have been gradually clarified based on a series of molecular phylogenetic studies (Scheen et al., 2010;Bendiksby et al., 2011;Drew and Sytsma, 2012;Li et al., 2016;Drew et al., 2017;Li and Olmstead, 2017;Zhao et al., 2021a). ...
... Phlomoides was established by Moench (1794), but it has generally been treated as a section of Phlomis L. Although Phlomoides was resurrected as a genus based on corolla shape and fruit structure (Adylov et al., 1986;Adylov, 1987;Kamelin and Machmedov, 1990a,b), the genus was not widely accepted (Ryding, 2008) until Scheen et al. (2010) confirmed it as a separate genus using molecular data. The separation of Phlomoides from Phlomis was later affirmed by studies with broader taxon sampling (Bendiksby et al., 2011;Salmaki et al., 2012a). Subsequent molecular phylogenetic studies demonstrated that at least seven genera should be transferred to Phlomoides (Scheen et al., 2010;Bendiksby et al., 2011;Mathiesen et al., 2011;Salmaki et al., 2012a;Zhao et al., 2023a,b), including Eremostachys Bunge, Lamiophlomis Kudô, Metastachydium Airy Shaw ex C.Y. Wu & H.W. Li, Notochaete Benth., Paraeremostachys Adylov et al., Pseuderemostachys Popov, and Pseudomarrubium Popov, making the redefined Phlomoides one of the largest and most heterogeneous genera of Lamiaceae, with ca. ...
... The separation of Phlomoides from Phlomis was later affirmed by studies with broader taxon sampling (Bendiksby et al., 2011;Salmaki et al., 2012a). Subsequent molecular phylogenetic studies demonstrated that at least seven genera should be transferred to Phlomoides (Scheen et al., 2010;Bendiksby et al., 2011;Mathiesen et al., 2011;Salmaki et al., 2012a;Zhao et al., 2023a,b), including Eremostachys Bunge, Lamiophlomis Kudô, Metastachydium Airy Shaw ex C.Y. Wu & H.W. Li, Notochaete Benth., Paraeremostachys Adylov et al., Pseuderemostachys Popov, and Pseudomarrubium Popov, making the redefined Phlomoides one of the largest and most heterogeneous genera of Lamiaceae, with ca. 150e170 spp. ...
Phlomoides, with 150–170 species, is the second largest and perhaps most taxonomically challenging genus within the subfamily Lamioideae (Lamiaceae). With about 60 species, China is one of three major biodiversity centers of Phlomoides. Although some Phlomoides species from China have been included in previous molecular phylogenetic studies, a robust and broad phylogeny of this lineage has yet to be completed. Moreover, given the myriad new additions to the genus, the existing infrageneric classification needs to be evaluated and revised. Here, we combine molecular and morphological data to investigate relationships within Phlomoides, with a focus on Chinese species. We observed that plastid DNA sequences can resolve relationships within Phlomoides better than nuclear ribosomal internal and external transcribed spacer regions (nrITS and nrETS). Molecular phylogenetic analyses confirm the monophyly of Phlomoides, but most previously defined infrageneric groups are not monophyletic. In addition, morphological analysis demonstrates the significant taxonomic value of eight characters to the genus. Based on our molecular phylogenetic analyses and morphological data, we establish a novel section Notochaete within Phlomoides, and propose three new combinations as well as three new synonyms. This study presents the first molecular phylogenetic analyses of Phlomoides in which taxa representative of the entire genus are included, and highlights the phylogenetic and taxonomic value of several morphological characters from species of Phlomoides from China. Our study suggests that a taxonomic revision and reclassification for the entire genus is necessary in the future.
... The tribe Phlomideae (Lamiaceae, Lamioideae) was originally established by Scheen et al. (2010) to include seven genera, Eremostachys Bunge, Lamiophlomis Kudô, Notochaete Benth., Phlomis L., Phlomoides Moench, Paraeremostachys Adylov et al. and Pseuderemostachys Popov. Subsequent phylogenetic and taxonomic studies (Bendiksby et al. 2011;Mathiesen et al. 2011;Salmaki et al. 2012b) have revised generic boundaries and Phlomoides was expanded to include Eremostachys, Lamiophlomis, Notochaete, Paraeremostachys and Pseuderemostachys. Recently, two monotypic genera, Metastachydium ...
Phlomoides is one of the largest genera of Lamiaceae with approximately 150–170 species distributed mainly in Eurasia. In this study, we describe and illustrate a new species, P. henryi, which was previously misidentified as P. bracteosa, from Yunnan Province, southwest China. Molecular phylogenetic analyses revealed that P. henryi is found within a clade in which most species lack basal leaves. In this clade, the new species is morphologically distinct from P. rotata in having an obvious stem and from the rest by having transparent to white trichomes inside the upper corolla lip. In addition, microfeatures of trichomes on the calyx and leaf epidermis can differentiate the new species from other species grouped in the same clade, and a key based on trichome morphology for these species is provided. The findings demonstrate that the use of scanning electron microscopy can reveal inconspicuous morphological affinities among morphologically similar species and play an important role in the taxonomic study of the genus Phlomoides.
... 13: Zhao (GeneBank unpublished). 14: Salmaki et al. (2012), 15: Al-Qurainy et al. (GeneBank unpublished). ...
The taxonomic classification of the recently described Lagochilus lorestanicus within the genus Lagochilus is traditionally based on its morphological characters. We present here a phylogenetic hypothesis for the genus Lagochilus (Lamiaceae, tribe Leonureae) based on plastid (trnL-trnF and matK) and nuclear (ITS) DNA sequence data, using maximum parsimony, maximum likelihood and Bayesian approaches. Our analysis indicates that while representative species from the Iranian flora are assembled in a sub-clade nested in the clade Spinosi, L. lorestanicus is retrieved in the Acanthoprasium clade in the tribe Marrubieae. The taxonomic placement of Lagochilus lorestanicus in Acanthoprasium is also corroborated by re-examination of morphological characters. Consequently, the species is transferred to the genus Acanthoprasium, which was previously restricted to Italy and France (i.e. A. frutescens) and Cyprus (i.e. A. integrifolium). The intra-specific morphological variations as well as the conservation status of Lagochilus lorestanicus are discussed.
... Phlomis is one of the largest genera in the subfamily Lamioideae (Lamiaceae), with about 90 accepted species distributed in Asia, southern Europe and northern Africa (Salmaki et al., 2012;Govaerts et al., 2018). Mathiesen et al. (2011) indicate that the Phlomis s. l. lineage has a Central Asian origin in an area around western China. ...
Phlomis lychnitis se registra por primera vez en el continente africano. Hasta el momento, sólo se conoce de Marruecos en el Alto Atlas Oriental. Se proporciona una descripción de la especie, así como los primeros datos sobre su ecología en Marruecos junto con su distribución geográfica y caracteres diagnósticos con la especie morfológicamente más cercana que comparte flores de color amarillo. También se proporcionan fotografías y una clave actualizada de las especies de Phlomis en Marruecos.
... During the past two decades, major advances have been made towards clarifying phylogenetic relationships and taxonomy of Lamioideae at various taxonomic levels (Wink & Kaufmann, 1996;Lindqvist & Albert, 2002;Scheen & Albert, 2007Scheen & al., 2008Scheen & al., , 2010Bendiksby & al., 2011Bendiksby & al., , 2014Salmaki & al., 2012Salmaki & al., , 2013Xiang & al., 2013;Chen & al., 2014;Roy & Lindqvist, 2015;Yao & al., 2016;Siadati & al., 2018). Among these studies, the most influential ones were carried out by Scheen & al. (2010) and Bendiksby & al. (2011), which reconstructed the backbone phylogeny of Lamioideae, proposed new tribes, and redefined and/or resurrected some genera. ...
... A total of 39 species of Phlomoides, covering the major distribution areas and most sections and subsections of the genus, were selected as the ingroup. Three taxa from the other genus within Phlomideae, Phlomis (Scheen & al., 2010;Bendiksby & al., 2011;Mathiesen & al., 2011;Salmaki & al., 2012;F. Zhao & al., 2021a), were sampled as the outgroup. ...
... -Total genomic DNA (gDNA) was obtained using the modified CTAB method (Doyle & Doyle, 1987) for both silica gel dried leaves and herbarium material. Polymerase chain reaction (PCR) primers, mixtures, and procedures of nrITS followed those described in Xiang & al. (2013), atpB-rbcL, psbA-trnH, and trnT-trnL followed Albaladejo & al. (2005), trnK and rpl32-trnL followed Salmaki & al. (2012), and rpl16, rps16, and trnL-trnF followed Chen & al. (2021). Amplified PCR products were sequenced on an ABI3730xl DNA Analyzer (Applied Biosystems, Foster City, California, U.S.A.) after purification with a QIAquick PCR purification Kit (BioTek, Beijing, China). ...
Distributed in Central Asia, Metastachydium (Lamiaceae) is a poorly understood and rare monotypic genus, with few collections known. The systematic position of this enigmatic genus within Lamiaceae has remained unresolved due to its poor representation in herbaria and coincident lack of available materials for molecular phylogenetic analysis. Facilitated by some recent collections, we performed Bayesian inference and maximum likelihood analyses, using an 80‐protein‐coding plastid‐gene dataset of Lamioideae, to infer the systematic placement of Metastachydium at the tribal level within Lamioideae. In addition, we used an 8‐plastid‐DNA‐region dataset as well as the nuclear ribosomal internal transcribed spacer to determine the phylogenetic affinities of Metastachydium . All phylogenetic analyses agree that Metastachydium is a member of Phlomideae and deeply nested within the genus Phlomoides , suggesting the need to expand the latter to include Metastachydium . Hence, a new combination, Phlomoides sagittata comb. nov., is proposed, and we present the first available photographs and an amended morphological description of P. sagittata . In addition, the infrageneric circumscription of Phlomoides is not supported, as most sections and subsections are not monophyletic. Hybridization and incomplete lineage sorting, following rapid diversification within Phlomoides , seem to be the source of incongruence between the nuclear and plastid tree topologies.
... Moreover, Ryding introduced Eremostachys as a monophyletic and independent genus to make Phlomoides less heterogeneous. 20 Salmaki et al. 21 conducted a molecular phylogeny study in this regard and identified several intermediate species with both Phlomoides and Eremostachys characteristics within the referred genera. Furthermore, the Phlomoides genus cannot be considered monophyletic by eliminating the Eremostachys species. ...
... Thus, Eremostachys is treated as a subclass of Phlomoides, which confirms the hypothesis proposed by the "world checklist of selected plant families" database. 21 The two principal genera of this tribe have an essential difference in their chromosome number. Phlomis and Phlomoides have x=10 and x=11 chromosomes, respectively. ...
... According to a study on the pericarp structure, the sclerenchyma region in Phlomis species is another distinguishing characteristic. 20,21,27 As previously discussed, the systematic position of Eremostachys is relatively controversial. The distinctive features of Eremostachys and Phlomoides have been addressed by various studies. ...
Phlomoides (L.) Moench belongs to the Lamiaceae family. It has recently undergone significant changes in taxonomy, with many species from Eremostachys and Phlomis added to the genus. The aforementioned species were studied in terms of morphological and phytochemical systematics. Species of Phlomoides are distinguished from Phlomis by their densely bearded upper corolla lip and nutlet. However, Eremostachys and Phlomoides have a lot in common morphologically. Plant chemosystematics present iridoids, phenylethanoids, and furanolabdanes as dominant constituents of Phlomoides species. Long-term traditional uses, such as bone fracture therapy, local analgesic, and wound healing actions, pique researchers' interest in these plants. The species and their secondary metabolites have been implicated in drug discovery by their anti-inflammatory and bone-development properties in vitro, in vivo, and clinically. A review of the taxonomic status based on phytochemical and morphological characteristics, as well as the clinical importance of the Phlomoides genus, is presented in the current study to provide a basis for further investigations.
... As one of the largest genera in Lamiaceae, Phlomoids Moench. is well known for its high local microendemism in Central Asia (Salmaki et al. 2012). ...
... According to the World Checklist (Govaerts et al. 2021) and the above circumscription, Phlomideae contains 200 records, whereas Kamelin and Makhmedov (1990) recognized about 250 species within the genera. The species are distributed from Europe to Mongolia, China, and India, with the highest number of species found in Central Asia, Afghanistan, Iran-Turanian, and Himalayan regions (Salmaki et al. 2012). ...
This paper discusses identifying Important Plant Areas (IPAs) in one of the most densely populated regions of Central Asia-the Fergana valley. The recognition of IPA sites is an attempt to introduce new ways of conserving local plant diversity with a high concentration of endemic species in Central Asia, where conservation methods of the former Soviet Union still prevail. The research revealed the current state and geography of many rare species and enriched the flora of Uzbekistan and Kyrgyzstan with several rare species. The second IPA is the transboundary territory of the Fergana valley, uniting the southern spurs of the Chatkal range and the Ungortepa-BozbuToo massif. We documented the distribution of 62 species in the IPAs under the sub-criteria of Plantlife International. Our study aimed at continuing studies on the IPAs in this region, addressing specific conservation challenges, such as conserving national endemics and endangered species that grow outside protected areas and GIS mapping of endemic species.
... Moench (1794) recognized different morphology of corollas and fruits within this genus, which then was divided into two genera. This taxonomy was accepted and developed by some researchers (Adylov et al., 1986;Adylov and Makhmedov, 1987;Kamelin and Makhmedov, 1990a;Kamelin and Makhmedov, 1990b;Scheen et al., 2010;Mathiesen et al., 2011;Salmaki et al, 2012;Xiang et al., 2014;Zhao et al., 2021). By contrast, some taxonomists recognized Phlomis and Phlomoides as two sections within the genus Phlomis (i.e., sect. ...
... Many morphological variations were observed between these two groups. For example, most Phlomis plants are perennial herbs or small shrubs, leaves are lanceolate to oblong-lanceolate, leaf venation is fan-shaped, the upper lips of corolla are laterally compressed, flattened, and sickle-shaped, and verticillasters are in a dense scapose capitulum or short spike; while most Phlomoides plants are perennial herbs with woody rhizomes and/or tubers, leaves are cordate to triangularovate, leaf venation is not fan-shaped, the upper lips of corolla are always hairy or fringed-incised, and verticillasters are axillary, lax, or dense (Mathiesen et al., 2011;Salmaki et al., 2012;Xiang et al., 2014). Anatomy, cytology, and phytochemistry analyses also revealed distinct features between Phlomis and Phlomoides. ...
... In the last two decades, phylogenetic studies based on molecular markers, including nuclear internal transcribed spacer (ITS) and chloroplast sequence (including rpl16, trnL-F, trnL intron, trnL-F intergenic spacer, rps16 intron, partial trnK, rpl32-trnL, trnT-A, and matK), supported a split of Phlomis s.l. into two monophyletic clades (Pan et al., 2009;Scheen et al., 2010;Bendiksby et al., 2011;Mathiesen et al., 2011;Salmaki et al, 2012;Xiang et al., 2014;Zhao et al., 2021). Therefore, most taxonomists accepted that Phlomis and Phlomoides should be treated as separate genera, i.e., Phlomis s.str. ...
The taxonomic terms “Phlomis” and “Phlomoides” had been used to describe two sections within the genus Phlomis belonging to the Lamiaceae family. Recently, phylogenetic analyses using molecular markers showed that Phlomis and Phlomoides formed two monophyletic clades, and thus they are generally accepted as separate genera. In this study, we assembled the complete chloroplast genome of Phlomis fruticosa, which is the first reported chloroplast genome belonging to Phlomis genus, as well as the complete chloroplast genome of Phlomoides strigosa belonging to Phlomoides genus. The results showed that the length of chloroplast genome was 151,639 bp (Phlomis fruticosa) and 152,432 bp (Phlomoides strigosa), with conserved large single copy regions, small single copy regions, and inverted repeat regions. 121 genes in Phlomis fruticosa and 120 genes in Phlomoides strigosa were annotated. The chloroplast genomes of Phlomis fruticosa, Phlomoides strigosa, and three reported Phlomoides species, as well as those of 51 species from the Lamiaceae family, which covered 12 subfamilies, were subjected to phylogenetic analyses. The Phlomis and Phlomoides species were split into two groups, which were well supported by both maximum likelihood and Bayesian inference tree analyses. Our study provided further evidence to recognize Phlomis and Phlomoides as independent genera.
... This treatment remained largely unrecognized until Adylov et al. (1986) and Adylov & Makhmedov (1987). In order to make the genus monophyletic, it was recently re-circumscribed to match molecular phylogenetic and morphological evidence (Salmaki et al. 2012b). As redefined, the genus now encompasses species previously referred to as Eremostachys Bunge (1830: 414) along with part of Phlomis Linnaeus (1753: 178), Paraeremostachys Adylov et al. (1986: 112) and other genera including Pseuderemostachys Popov (1940: 148), Notochaete Bentham (1830: 63) and Lamiophlomis Kudô (1929: 210) (Salmaki et al. 2012b). ...
... In order to make the genus monophyletic, it was recently re-circumscribed to match molecular phylogenetic and morphological evidence (Salmaki et al. 2012b). As redefined, the genus now encompasses species previously referred to as Eremostachys Bunge (1830: 414) along with part of Phlomis Linnaeus (1753: 178), Paraeremostachys Adylov et al. (1986: 112) and other genera including Pseuderemostachys Popov (1940: 148), Notochaete Bentham (1830: 63) and Lamiophlomis Kudô (1929: 210) (Salmaki et al. 2012b). In order to stabilize the nomenclature in this large genus, most of the species have been typified in several papers (Adylov & Makhmedov 1987;Kamelin & Makhmedov 1990;Sennikov & Lazkov 2010;Lazkov 2011;Salmaki et al. 2012a-b). ...
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.
... nov. and M. mimetica with data on the distribution of species of the former genus Eremostachys obtained from our own field observations and taken from literary sources (Nikitin et al. 1954;Shishkin et al. 1954;Hedge 1982;Abdullayeva et al. 1987;Hedge 1990;Salmaki et al. 2012aSalmaki et al. , 2012b. Modern taxonomy of plants under consideration, based on molecular genetic studies, suggests including all representatives of the former genus Eremostachys in the genus Phlomoides (Abdullayeva et al. 1987;Salmaki et al. 2012a). ...
... If so, the case of M. shahvarica is an example of sequential coevolution after a food shift. The molucelloides group is younger in evolutionary terms than the laciniata group (Salmaki et al. 2012b). For this reason, we consider M. shahvarica evolutionarily the youngest species in the lutko group. ...
This review considers a fascinating, from a zoogeographical viewpoint, group of closely related species: Melitaea lutko Evans, 1932, M. timandra Coutsis & van Oorschot, 2014, M. mimetica Higgins, 1940 stat. rev. and M. shahvarica sp. nov. It is a taxonomical and geographical review of these species, and data on the biology of M. shahvarica sp. nov. and nominate subspecies of M. timandra are discussed. A new species, M. shahvarica sp. nov. from Shahvar Mt. (Iran), and a new subspecies, M. timandra binaludica subsp. nov. from Kuh-e-Binalud Mts (Iran), are described. The specific structure of the group given in previous publications is critically evaluated. Hypotheses about a possible phylogenesis of the study group are provided.
