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Phylogenetic and biosystematic relationships in four highly disjunct polyploid complexes in the subgenera Ceterach and Phyllitis in Asplenium (Aspleniaceae) |
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| Authors: | Istvan Pinter Freek Bakker John Barrett Cymon Cox Mary Gibby Sally Henderson Mary Morgan-Richards Fred Rumsey Stephen Russell Steve Trewick Harald Schneider Johannes Vogel |
| Institution: | aDepartment of Genetics, Eötvös Loránd University, Budapest, Hungary bWageningen University Branch, National Herbarium Netherlands, The Netherlands cDepartment of Genetics, University of Cambridge, UK dDepartment of Biology, Duke University, Durham, NC, U.S.A. eRoyal Botanic Garden Edinburgh, Scotland fDepartment of Botany, The Natural History Museum, London, UK gAlbrecht-von-Haller Institut für Pflanzenwissenschaften, Universität Göttingen, Germany |
| Abstract: | Phylogenetic studies using DNA sequences of two chloroplast regions, rbcL and trnL-F, demonstrate that the proposed genus Ceterach is a small clade within the large genus Asplenium, and sister to the Phyllitis clade. The Ceterach clade is characterised by irregular anastomosing veins and often densely scaled leaf blades. Its taxonomic status as a group nested within Asplenium is confirmed, and it is accepted here as a subgenus with seven species. The Ceterach clade comprises four lineages that correspond to disjunct polyploid complexes: the A. aureum clade forming a polyploid complex (4×, 6×, 8×) in Macaronesia, the A. ceterach clade forming a polyploid complex (2×, 4×, 6×) in the Mediterranean Basin, the A. paucivenosum clade (4×, 6×) in central Asia, and the A. dalhousiae clade (2×) with a disjunct distribution in the Himalaya, Yemen and Eritrea, and southwestern North America. Asplenium paucivenosum is sister to all other members of the Ceterach clade, whereas A. dalhousiae is sister to the A. aureum clade that includes tetraploid A. aureum, hexaploid A. lolegnamense, and octoploid A. parvifolium. Asplenium ceterach and its variations – including the hexaploid A. ceterach subsp. mediterraneum subsp. nov. first described below – form a monophyletic unit, sister to a clade consisting of A. aureum and A. dalhousiae. Asplenium cordatum from Africa and A. haugthonii from the isolated atlantic island of St. Helena are not members of the Ceterach clade, which suggests that leaf blades with dense indumenta have evolved at least twice within asplenioid ferns. The allotetraploid species A. hybridum has the chloroplast DNA from A. ceterach, and therefore the latter species is the maternal ancestor of the former. The other parent of this hybrid species is A. sagittatum that is nested within the sister clade of Ceterach, the Phyllitis clade comprising A. sagittatum and A. scolopendrium. The findings suggest that the current distribution of Ceterach is either the result of long-distance dispersal or represents fragmented relicts of a previously more widely distributed species. |
| Keywords: | biogeography long-distance dispersal oceanic islands radiations molecular phylogeny plant taxonomy |
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