Tribe Marrubieae (Lamiaceae: Lamioideae) comprises about 91 species classified in four genera distributed from Europe to West and Central Asia and North Africa. Here, we present a phylogenetic analysis of cpDNA (rpl32-trnL, rps16, trnL intron, trnL-trnF spacer) and nuclear ribosomal (ITS) sequence data of 68 accessions representing all genera and most sections of Marrubieae. Our taxon sampling covers is dense for the genera Ballota and Marrubium, including a total of 60 of 85 currently recognized taxa (species and subspecies). Parsimony and Bayesian analyses of individual nrITS and cpDNA markers, as well as the combined plastid dataset, produced largely congruent trees. Our results confirm the monophyly of tribe Marrubieae. All analyses retrieved four well supported clades in the tribe. The phylogenetic position of the genera Acanthoprasium and Moluccella is consistent with previous studies and both are recovered as monophyletic. The most important result of the present study is the splitting of the genus Ballota, in its traditional concept, into two well-supported clades matching well with the genera Ballota and Pseudodictamnus. Moreover, Marrubium, in its current circumscription, is paraphyletic, but will become monophyletic by transferring Ballota deserti to this genus. In conclusion, we propose the recognition of five genera in tribe Marrubieae: Acanthoprasium, Ballota, Moluccella, Marrubium and the resurrected segregate Pseudodictamnus. The required new combinations are proposed.
The fruit and seed micromorphology of 22 species of Geranium, representing the eight sections of the genus represented in Iran (G. sectt. Dissecta, Geranium, and Tuberosa of subgen. Geranium; sectt. Batrachioidea, Divaricata, Lucida, Ruberta and Trilopha of subgen. Robertium), have been examined by scanning electron microscopy (SEM). Macro‐ and micromorphological characters, including fruit and seed shape, size, color, hair type and density, mericarp ornamentations, hilum position, seed coat pattern, epidermal cell shape, and anticlinal and periclinal cell walls, are presented. Two microsculpturing patterns are recognized on the mericarp surface: reticulate and pusticulate. The micromorphology of the seed coat showed four distinctive cell patterns. The seed epidermis is constructed either of polygonal, elongated polygonal, or square to rectangular cells. The polygonal type is the most common among the studied species, but the variation in testa cell characters, their size and shape, may provide further information and useful diagnostic characters at specific and infraspecific rank. The shape and color of the seeds are, however, of little systematic value. Fruit characters were found to be important for separating taxa at infrageneric rank and our results show that the species can be separated into subgenera and sections based on fruit morphology.
Cross-sections of roots, stems, petioles and peduncles were investigated in 12 species of Viola distributed mainly in northernGeneral anatomical features of Viola species are discussed. Th e following characters were found to be taxonomically informative: cross-section shapes of the aerial stem, petiole and peduncle as well as the number of vascular bundles in the aerial stem. In taxa of sect. Melanium , stem cross-sections are elliptic with two wings, but they are semi-circular with two wings in taxa of sect. Viola subsect. Rostratae . Within the latter subsection, the number of vascular bundles in the stem can be used to delimit the species V . caspia , V . reichenbachiana and V . rupestris . Calcium oxalate crystals were observed in all vegetative organs of taxa belonging to sect. Viola . All examined taxa had a secondary structure with the exception of V. somchetica . Our results show that anatomical characters are useful for delimiting species, subsections, and sections within Viola .
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