A new species of Atriplex (Amaranthaceae) from the Indian subcontinent.

A new subshrubby C4-species from the lowlands and foothills of India, Pakistan and SE Afghanistan, Atriplexpseudotatarica, is described and illustrated. Previously, it was incorrectly identified as A.crassifolia auct. non C.A.Mey. belonging to a distant C3-group of the genus. A phylogenetic analysis based on nrITS and nrETS revealed its position as sister to A.schugnanica (sect. Obionopsis). Both species share aphyllous inflorescence and smooth bract-like cover, but differ in life form, leaves, seed colour, and geographical distribution. We revised native Indian Atriplex species and excluded some of them from the flora of the country. An improved checklist of the native Atriplex species in India with their corrected synonymy and nomenclature is given, and a new diagnostic key is provided.

Biogeography and Systematics of the Genus Axyris (Amaranthaceae s.l.).

Axyris is a small genus of six species with a disjunct geographic range. Five species are present in Siberia, Central Asia, the Himalayas, and Tibet, whereas Axyris caucasica has been registered in the Central Caucasus only. Axyris species diversity is the highest in the Altai Mountains (four spp.), followed by the Tian Shan and Pamir Mountains (three spp.), and the Himalayas and Tibet (two spp.). Axyris sphaerosperma, sometimes considered endemic to Southern Siberia, in fact has a disjunct range: it is present in the lowlands of Eastern Siberia and in the Altai, Tian Shan, and Pamir Mountains. It has also been found in Mongolia and China for the first time. An updated detailed distribution of Axyris in Siberia is presented on the basis of thorough herbarium revisions. One nuclear and three plastid markers were selected for phylogenetic analysis. Divergence times were estimated using a time-calibrated Bayesian approach. Axyris shows two major clades: an Axyris amaranthoides clade and a clade including the remaining species. The latter clade consists of two subclades (A. sphaerosperma/A. caucasica and A. mira/A. prostrata + A. hybrida). The crown age for Axyris dates back to the Early Pliocene (~5.11 mya, the Zanclean). The ancestral range of Axyris covers Southern Siberia, Mongolia, NW China, and the Tian Shan/Pamir Mountains, with extensions toward Eastern Siberia, the Himalayas/Tibet, and the Caucasus. Fruit and seed characteristics of Axyris are discussed with reference to the present phylogenetic results. Closely related A. sphaerosperma and A. caucasica have the thickest seed coat among all Chenopodiaceae, and these traits have probably evolved as adaptations to extremely low winter temperatures. This reproductive peculiarity may explain the disjunct range of A. sphaerosperma, which is restricted to harsh climatic conditions.

Evolutionary relationships, biogeography and morphological characters of Glinus (Molluginaceae), with special emphasis on the genus composition in Sub-Saharan Africa.

Glinus is a small genus of Molluginaceae with 8-10 species mostly distributed in the tropics of the World. Its composition and evolutionary relationships were poorly studied. A new molecular phylogeny constructed here using nuclear (ITS) and chloroplast (rbcL, trnK-matK) markers confirmed the monophyly of the genus. Based on ITS analysis, the following well-supported lineages are present within Glinus: the G. bainesii lineage is recovered as sister to the remainder of the genus followed by G. oppositifolius. Three other clades are: G. hirtus with G. orygioides; G. radiatus and G. lotoides; the latter is represented by a sample from North America, and G. zambesiacus as sister to G. setiflorus + G. lotoides + G. dictamnoides. On the plastid gene tree, G. bainesii + G. oppositifolius form a sister clade to all other Glinus species. The next clade is formed by G. hirtus and G. orygioides followed by G. radiatus plus an American sample of G. lotoides. The next branch comprises G. setiflorus as sister to G. zambesiacus + G. lotoides + G. dictamnoides. Glinus seems to have originated from Africa around the Late Eocene or Early Miocene, with further radiations to Australia and the Americas during the Late Miocene or Late Pliocene. Compared with the previous limited character set used for the diagnostics, we have found ten new morphological and carpological traits distinguishing Glinus members. In both trees based on nuclear and plastid datasets, the major phylogenetic clades cannot be characterized by the peculiar morphological characters. Many shared character states leading to their contrasting pattern in the multivariate analysis model are interpreted as a high homoplasy in the phylogenetically distant species. We paid special attention to the composition of the genus in Sub-Saharan Africa, a region with the greatest species diversity. Our results provide new insight into the taxonomy of Glinus in this region. Glinus lotoides var. virens accepted in many previous works is a synonym of G. dictamnoides that is closely related to G. lotoides based on molecular analysis and morphological characters. The status of the American populations of G. lotoides needs further investigation due to different characters of the specimens from the Old and the New World. Many specimens previously identified as G. lotoides var. virens and as the intermediates G. lotoides × G. oppositifolius belong to G. zambesiacus sp. nov. and G. hirtus comb. nov. (≡ Mollugo hirta); the latter species is resurrected from synonymy after 200 years of unacceptance. In some African treatments, G. hirtus was known under the invalidly published name G. dahomensis. Glinus zambesiacus is distributed in the southern and eastern parts of tropical Africa, and G. hirtus previously assumed to be endemic to West Africa is indeed a species with a wide distribution across the tropical part of the continent. Glinus microphyllus previously accepted as endemic to West Tropical Africa together with other new synonyms (G. oppositifolius var. lanatus, G. herniarioides, Wycliffea rotundifolia) is considered here as G. oppositifolius var. keenaniicomb. nov. (≡ Mollugo hirta var. keenanii), a variety found across the entire distribution of G. oppositifolius (Australia, Asia, and Africa). The presence of the American G. radiatus in Africa is not confirmed, and all records of this species belong to G. hirtus. The lectotypes of some names (G. dictamnoides, G. herniarioides, Mollugo hirta, M. setiflora, Pharnaceum pentagynum, Wycliffea) as well as a neotype of G. trianthemoides are designated. A new key to the identification of all Glinus species in Sub-Saharan Africa is provided. A checklist is given of all accepted species in this region (G. bainesii, G. hirtus, G. lotoides, G. oppositifolius s.l., G. setiflorus, and G. zambesiacus) with their nomenclature, morphological description and geographical distribution.

Scorzonera sensu lato (Asteraceae, Cichorieae) - taxonomic reassessment in the light of new molecular phylogenetic and carpological analyses.

Scorzonera comprises 180-190 species and belongs to the subtribe Scorzonerinae. Its circumscription has long been the subject of debate and available molecular phylogenetic analyses affirmed the polyphyly of Scorzonera in its wide sense. We provide a re-evaluation of Scorzonera and other related genera, based on carpological (including anatomical) and extended molecular phylogenetic analyses. We present, for the first time, a comprehensive sampling, including Scorzonera in its widest sense and all other genera recognised in the Scorzonerinae. We conducted phylogenetic analyses using Maximum Parsimony, Maximum Likelihood and Bayesian analyses, based on sequences of the nuclear ribosomal ITS and of two plastid markers (partial rbcL and matK) and Maximum Parsimony for reconstructing the carpological character states at ancestral nodes. Achene characters, especially related to pericarp anatomy, such as general topography of the tissue types, disposition of the mechanical tissue and direction of its fibres, presence or absence of air cavities, provide, in certain cases, support for the phylogenetic lineages revealed. Confirming the polyphyly of Scorzonera, we propose a revised classification of the subtribe, accepting the genera Scorzonera (including four major clades: Scorzonera s. str., S. purpurea, S. albicaulis and Podospermum), Gelasia, Lipschitzia gen. nov. (for the Scorzonera divaricata clade), Pseudopodospermum, Pterachaenia (also including Scorzonera codringtonii), Ramaliella gen. nov. (for the S. polyclada clade) and Takhtajaniantha. A key to the revised genera and a characterisation of the genera and major clades are provided.