The aim of the study is an expanded molecular phylogenetic analysis of the Holarctic taxa traditionally classified to the families Lophoziaceae, Scapaniaceae, Gymnomitriaceae and Jungermanniaceae. Also we will provide a justification of some nomenclatural changes suggested recently by Konstantinova & Vilnet (2009). The nuclear ITS1-2 and chloroplast trnL-F DNA sequences of 134 taxa (189 samples) from suborders Jungermanniineae and Cephaloziineae were analyzed. The topologies of phylogenetic trees constructed from combined sequences by maximum parsimony and Bayesian methods are congruent. The separation of family Diplophyllaceae from Scapaniaceae is not supported. The recently described family Solenostomataceae appears to be paraphyletic: the genus Nardia is in a sister relation to Gymnomitriaceae. The inclusion of the genus Isopaches in the family Anastrophyllaceae is questionable. Position of Obtusifolium and Protolophozia elongata in Scapaniaceae is not strongly supported. Phylogenetic relations within the genus Scapania are not fully resolved, but monophyly of sections Undulatae and Calcicolae is supported. Species assigned by some authors to a separate genus Scapaniella are scattered in different clades of the genus Scapania. Macrodiplophyllum imbricatum and M. plicatum can either be treated as Douinia or all these taxa included in Scapania. The distribution of species from the family Gymnomitriaceae on phylogenetic trees is congruent with the level of development or reduction of the perianth and perigynium. Taxonomically the species of Marsupella with reduced perianth are referred to Gymnomitrion, whereas the species called G. apiculatum that has a distinct perianth is transferred to Marsupella. The species status for Scapania tundrae, S. paludosa, S. crassiretis and Marsupella aquatica is supported. Genera Lophozia s.str., Schistochilopsis, Tritomaria, Protolophozia, Orthocaulis and Crossogyna are polyphyletic. The data support segregation of the recently described genera Pseudotritomaria, Heterogemma, Lophoziopsis, Pseudolophozia, Schljakovianthus, Schljakovia and Biantheridion.
Molecular phylogenic analysis of the suborder Cephaloziineae is performed, with the special attention to the family Cephaloziaceae s.l. and its implications for taxonomy are discussed. A combined alignment of nuclear ITS1-2 and chloroplast trnL-F DNA sequences of 121 species (209 samples) from the families Scapaniaceae, Anastrophyllaceae, Cephaloziaceae s.l., Cephaloziellaceae, Adelanthaceae, and Jamesoniellaceae are analyzed. The topologies of phylogenetic trees constructed by maximum parsimony, maximum likelihood and Bayesian methods show the principal congruence with previously achieved phylogenies, with exception of ambiguous position of the Cephaloziellaceae. The genus Hygrobiella does not reveal affinity to the Cephaloziaceae s.l., which supports its segregation in the monotypic family Hygrobiellaceae. The Odontoschismataceae are resolved sister to the Cephaloziaceae s.str., supporting their classification into two families expedient also from morphological evidences. The former family preliminarily includes, besides Odontoschisma and Cladopodiella, the genera Iwatsukia and Alobiellopsis from the subfamily Alobielloideae. The relationships of the subfamily Schiffnerioideae remain uncertain. The genera Cephalozia, Odontoschisma and Cladopodiella appear to be polyphyletic, but the existing section subdivisions for Cephalozia and Odontoschisma are partly supported. The genus Nowellia is found in the sister position to Cephalozia bicuspidata-complex and C. macoinii. The rest of studied Cephalozia species compose an intermingled clade with the monotypic genera Pleurocadula and Schofieldia. Thus only species of the Cephalozia bicuspidatacomplex and C. macoinii are accepted in Cephalozia, whereas other Cephalozia species and Schofieldia are transferred into the genus Pleurocadula. Infraspecific nucleotide sequence variation suggests the species status for Cephalozia affinis and Odontoschisma elongatum, whereas O. prostratum and O. sphagni appear to be conspecific. сестринским комплексу видов C. bicuspidata и C. macoinii. Остальные исследованные виды рода Cephalozia расположены в кладе с монотипными родами Pleurocadula и Schofieldia. Объем рода Cephalozia пересмотрен: только комплекс видов C. bicuspidata и C. macoinii относятся к Cephalozia, в то время как другие изученные виды Cephalozia и Schofieldia перенесены в род Pleurocadula. Исследование вариабельности нуклеотидных последовательностей подтверждает видововой статус Cephalozia affinis и Odontoschisma elongatum, в то же время O. prostratum и O. sphagni следует считать одним видом.
This study tests the phylogenetic affinity and clarifies the taxonomy of Frullania tamarisci s. l. in Russia and adjacent areas based on critical morphological reinvestigation and DNA analyses. More than 100 accessions of F. tamarisci s. l. from the collections of KPABG, VBGI and UBC were studied morphologically, and the nrITS1-5.8S-ITS2 and trnL–trnF cpDNA loci of sixteen accessions were sequenced. Molecular data were combined with accessions from GenBank and phylogenetic estimations by maximum parsimony, maximum likelihood and Bayesian approaches were performed. The tree topologies obtained were congruent and resolved the specimens previously referred to F. nisquallensis from Russian North in a clade together with an accession from Alaska. This clade is only distantly related to European F. tamarisci s. str. and temperate western North American F. nisquallensis. Taking into account the morphological differences, we describe a new species Frullania subarctica Vilnet, Borovich. & Bakalin sp. nov. The species status for F. appendiculata occurring mainly in temperate East Asia was supported by molecular evidence, based on specimens from the Primorsky Territory of Russia. Populations of F. tamarisci s. str. from the Russian North West and the Caucasus Mountains are quite similar to those from European countries both morphologically and molecularly.
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