Michael S. Ignatov

An annotated checklist of the mosses of Europe and Macaronesia

Abstract The moss flora of Europe and Macaronesia comprises 278 genera, 1292 species, 46 subspecies and 118 varieties. Of the total 1292 species, 53 are confined to Macaronesia and 21 are thought to be non-native. The checklist was derived from those for the various component countries and regions. It is based on results published up to the end of 2005. Subspecies and varieties are included; hybrids are omitted. The taxonomic hierarchy is based on one published by Goffinet & Buck in 2004. While it has been strongly influenced by results of modern molecular methods, there are still many remaining uncertainties, even at family level. Because of these uncertainties, taxonomic innovation has generally been avoided. There are four new combinations and one change of status.

ORIGIN AND EVOLUTION OF THE NORTHERN HEMISPHERE DISJUNCTION IN THE MOSS GENUS HOMALOTHECIUM (BRACHYTHECIACEAE)

Competing hypotheses that rely either on a stepping-stone dispersal via the North Atlantic or the Bering land bridges, or more recent transoceanic dispersal, have been proposed to explain the disjunct distribution of Mediterranean flora in southern Europe and western North America. These hypotheses were tested with molecular dating using a phylogeny of the moss genus Homalothecium based on ITS, atpB-rbcL, and rpl16 sequence data. The monophyly of two main lineages in Western Palearctic (Europe, central Asia and north Africa) and North America is consistent with the ancient vicariance hypothesis. The monophyly of Madeiran H. sericeum accessions supports the recognition of the Macaronesian endemic H. mandonii. A range of absolute rates of molecular evolution documented in land plants was used as probabilistic calibration prior by a Bayesian inference implementing a relaxed-clock model to derive ages for the nodes of interest. Our age estimates for the divergence of the American and Western Palearctic Homalothecium clade (5.7 Ma, IC 3.52-8.26) and the origin of H. mandonii (2.52 Myr IC 0.86-8.25) are not compatible with the ancient vicariance hypothesis. Age estimates suggests that species distributions result from rare instances of dispersal and subsequent sympatric diversification. The calibrated phylogeny indicates that Homalothecium has undergone a fast radiation during the last 4 Myr, which is consistent with the low levels of morphological divergence among sibling species.

On resurrection of the families Pseudoleskeaceae Schimp. and Pylaisiaceae Schimp. (Musci, Hypnales)

Results from our analysis of chloroplast trnL-trnF and nuclear ITS1 and ITS2 sequence data and 35 morphological characters for 134 taxa of pleurocarpous mosses provide an evidence for resurrecting two moss families, Pseudoleskeaceae and Pylaisiaceae. Both these families were described by Schimper in 1860, but soon afterwards included in Leskeaceae and Hypnaceae, respectively, and apparently never used in the 20 t h century. However, sequence-level data analysis (chloroplast encoded trnL intron and trnL-trnF spacer, nuclear encoded ITS1 & ITS2), and combined molecular and morphological analysis demonstrate the remote position of (1) the Lescuraea-group from the main part of Leskeaceae; and (2) a group of species belonging to Pylaisia, Homomallium and Hypnum (sections others than sect. Hypnum} from Hypnum sect. Hypnum (with H. cupressiforme as the type of the genus).

Phylogeny of the Brachytheciaceae (Bryophyta) based on morphology and sequence level data

Brachytheciaceae is often considered a taxonomically difficult group of mosses. For example, morphological variation has led to difficulty in generic delimitation. We used DNA sequence data (chloroplast psbT‐H and trnL‐F and nuclear ITS2) together with morphology (63 characters) to examine the relationships within this family. The combined unaligned length of the DNA sequences used in the phylogenetic analyses varied between 1277 and 1343 bp. For phylogeny reconstruction we performed direct optimization, as implemented in POY. Analyses were performed with three different gap costs and the morphological data partition was weighted both: (1) equal to gap cost, and (2) with a weight of one. The utility of sensitivity analysis has recently been cast into doubt; hence in this study it was performed only to explore the effects of weighting on homology statements and topologies and to enable more detailed comparisons between earlier studies utilizing the direct optimization method. The wide sequence length variation of non‐coding ITS2 sequences resulted in character optimizations (i.e., “alignments”) of very different lengths when various gap costs were applied. Despite this variation, the topologies of equally parsimonious trees remained fairly stable. The inclusion of several outgroups, instead of only one, was observed to increase the congruence between data sets and to slightly increase the resolution. An inversion event in the 9 bp loop region in the chloroplast psbT‐N spacer in mosses has been postulated to include only uninformative variation, thus possibly negatively impacting the phylogeny reconstruction. Despite this inversion, its variation within Brachytheciaceae was clearly congruent with information from other sources, but inclusion of these 9 bp in the analysis had only a minor effect on the phylogenetic results. In the most parsimonious topology, which was obtained with equal weighting of all data, Meteoriaceae and Brachytheciaceae were resolved as monophyletic sister groups, which had recently been suggested based on a few shared morphological characters. Our study revealed some new generic relationships within the Brachytheciaceae, which are discussed in light of the morphological characters traditionally used for generic delimitation.

And if Engler was Not Completely Wrong? Evidence for Multiple Evolutionary Origins in the Moss Flora of Macaronesia

The Macaronesian endemic flora has traditionally been interpreted as a relict of a subtropical element that spanned across Europe in the Tertiary. This hypothesis is revisited in the moss subfamily Helicodontioideae based on molecular divergence estimates derived from two independent calibration techniques either employing fossil evidence or using an Monte Carlo Markov Chain (MCMC) to sample absolute rates of nucleotide substitution from a prior distribution encompassing a wide range of rates documented across land plants. Both analyses suggest that the monotypic Madeiran endemic genus Hedenasiastrum diverged of other Helicodontioideae about 40 million years, that is, well before Macaronesian archipelagos actually emerged, in agreement with the relict hypothesis. Hedenasiastrum is characterized by a plesiomorphic morphology, which is suggestive of a complete morphological stasis over 40 million years. Macaronesian endemic Rhynchostegiella species, whose polyphyletic origin involves multiple colonization events, evolved much more recently, and yet accumulated many more morphological novelties than H. percurrens. The Macaronesian moss flora thus appears as a complex mix of ancient relicts and more recently dispersed, fast-evolving taxa.

New national and regional bryophyte records, 36

ub lis he d by M an ey P ub lis hi ng ( c) B rit is h B ry ol og ic al S oc ie ty Bryological Notes New national and regional bryophyte records, 36 L T Ellis, V A Bakalin, E Baisheva, H Bednarek-Ochyra, R Ochyra, E A Borovichev, S S Choi, B-Y Sun, P Erzberger, V E Fedosov, R Garilleti, B Albertos, P Gorski, P Hajkova, N G Hodgetts, M Ignatov, A Koczur, L E Kurbatova, M Lebouvier, A Mežaka, J Miravet, P Pawlikowski, R D Porley, J A Rossello, M S Sabovljevic, J Pantovic, A Sabovljevic, W Schroder, S Ştefănuţ, G M Suarez, M Schiavone, O T Yayintas, J Vaňa The Natural History Museum, London, UK, Botanical Garden-Institute, Vladivostok, Russia, Institute of Biology and Soil Science, Vladivostok, Russia, Institute of Biology of Ufa, Scientific Centre of Russian Academy of Sciences, Ufa, Russia, Institute of Botany, Poland, Polar-Alpine Botanical Garden-Institute, Kirovsk, Russia, Chonbuk National University, Jeonju, Korea, Berlin, Germany, M.V. Lomonosov Moscow State University, Russia, Universidad de Valencia, Burjassot, Spain, Poznan University of Life Sciences, Poland, Masaryk University, Brno, Czech Republic, Institute of Botany, Brno, Czech Republic, Earlish, Isle of Skye, UK, Main Botanical Garden, Moscow, Russia, Institute of Nature Conservation, Krakow, Poland, Komarov Botanical Institute, Russia, Universite de Rennes 1, France, University of Latvia, Riga, Latvia, Marimurtra Botanical Garden, Catalonia, Spain, University of Warsaw, Poland, Cerca dos Pomares, Portugal, Universidad de Valencia, Spain, University of Belgrade, Serbia, Ludwigsstadt, Germany, Institute of Biology of Romanian Academy, Bucharest, Romania, Facultad de Ciencias Naturales e I.M.L., Tucuman, Argentina, Canakkale Onsekiz Mart University, Biga, Canakkale, Turkey, Charles University, Praha, Czech Republic

Contribution of genosystematics to current concepts of phylogeny and classification of bryophytes

This paper is a survey of the current state of molecular studies on bryophyte phylogeny. Molecular data have greatly contributed to developing a phylogeny and classification of bryophytes. The previous traditional systems of classification based on morphological data are being significantly revised. New data of the authors are presented on phylogeny of Hypnales pleurocarpous mosses inferred from nucleotide sequence data of the nuclear DNA internal transcribed spacers ITS1-2 and the trnL-F region of the chloroplast genome.

Disentangling knots of rapid evolution: origin and diversification of the moss order Hypnales

Abstract The Hypnales are the largest order of mosses comprising approximately 4200 species. Phylogenetic reconstruction within the group has proven to be difficult due to rapid radiation at an early stage of evolution and, consequently, relationships among clades have remained poorly resolved. We compiled data from four sequence regions, namely, nuclear ITS1–5·8S–ITS2, plastid trnL–F and rps4, and mitochondrial nad5, for 122 hypnalean species and 34 species from closely related groups. Tree topologies from both Bayesian and parsimony analyses resolve the order as monophyletic. Although inferences were made from fast-evolving genes, and despite strong phylogenetic signal in the nuclear ITS1–5·8S–ITS2 data, monophyly, as well as backbone nodes within the Hypnales, remains rather poorly supported except under Bayesian inferences. Ancestral distribution based on Bayesian dispersal-vicariance analysis supports a Gondwanan origin of the Hypnales and subsequent geographical radiation in the area of the former Laurasian supercontinent. Reconstruction of historical biogeography is congruent with mainly tropical and Gondwanan distributions in the sister groups Hypnodendrales, Ptychomniales, and Hookeriales, and with the dating for the oldest pleurocarp and hypnalean fossils. We contrast groupings in the phylogenetic tree with recent classifications and other phylogenetic inferences based on molecular data, and summarise current knowledge on the evolutionary history of, and relationships among, the Hypnales.

Bryophyte flora of Altai Mountains. VIII. Brachytheciaceae

The family Brachytheciaceae in Altai is represented by 8 genera and 33 species: Brachythecium buchananii, B. cirrosum, B. collinum, B. complanatum, B. dovrense, B. erythrorrhizon ssp. asiaticum ssp. nov., B. falcatulum, B. glaciale, B. latifolium, B. mildeanum, B. oedipodium, B. ornellanum, B. plumosum, B. populeum, B. reflexum, B. rivulare, B. rotaeanum, B. rutabulum, B. salebrosum, B. starkei, B. trachypodium, B. turgidum, B. velutinum, Bryhnia novae-angliae, Cirriphyllum piliferum, Eurhynchium altaicum sp. nov., E. angustirete, E. hians, E. pulchellum, Homalothecium phili ppeanum, Myuroclada maximowiczii, Platyhypnidium riparioides, Scleropodiopsis laxiretis gen. et sp. nov. For each species descriptions, illustrations, and distribution and habitat preferences in Altai are provided.

Support from DNA data for a narrow species concept in Schistidium (Grimmiaceae, Musci)

Abstract A study of ITS1 of 28 specimens of eight species of Schistidium from well-separated populations in Russia and northwest Europe revealed that there are very big differences between species (up to 16 substitutions and 256 indels), whereas within the species differences in DNA there are very few (none to four substitutions and none to six indels). These results strongly support the narrow species concept in Schistidium. Schistidium papillosum is represented by two quite distinct genotypes and probably needs further splitting.