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The occurrence of species in both polar regions (bipolarity) is a common phenomenon in the Antarctic flora. Considering the high morphological variation in polar regions due to extreme conditions, the use of molecular tools is indispensable for testing whether Arctic and Antarctic populations indeed belong to the same species. However, few phylogeographic studies of bipolar bryophytes have been conducted so far, especially when comparing molecular and morphological variation. Here, we assess the bipolarity and intraspecific variation of Roaldia revoluta, a strictly bipolar species of pleurocarpous mosses. Phylogenetic analyses based on ITS sequences clearly resolve R. revoluta as monophyletic and confirm its bipolar distribution pattern. Low intraspecific molecular variation in the markers ITS/26S and rpl16 was observed, and most specimens from both polar regions belong to a single haplotype, making it difficult to infer the origin and dispersal routes between both polar regions of R. revoluta. Morphometric analysis furthermore suggests that there are no significant morphological differences among populations from both polar regions and that morphological variation is mainly influenced by local environmental conditions. Our data do not unequivocally support the recent separation of the former intraspecific taxon Hypnum revolutum var. dolomiticum at the species level as Roaldia dolomitica.

Increasing evidence indicates that wide distributed bryophyte taxa with homogeneous morphology may represent separate evolutionary lineages. The evolutionary histories of these cryptic lineages may be related to historical factors, such as the climatic oscillations in the Quaternary. Thus, the post-glacial demographic signatures paired with cryptic speciation may result in complex phylogeographic patterns. This research has two aims: to determine whether the widespread moss Racomitrium lanuginosum represents cryptic molecular taxa across the Northern Hemisphere and to infer the effects of Quaternary glaciations on spatial genetic diversity. We used the internal transcribed spacer (ITS) marker to resolve the phylogeographic history of the species and single nucleotide polymorphisms (genotyping-by-sequencing) to infer the genetic structure and demographic history. Finally, we assessed the historical changes in the distribution range using species distribution models. Racomitrium lanuginosum comprises distinct molecular lineages sympatrically distributed in the Northern Hemisphere. We also uncovered long-distance dispersal from eastern North America to Scandinavia and potential in situ survival in northern Scandinavia. Due to the genetic signatures, the Alaska Peninsula could be considered a glacial refugium. The species experienced post-glacial expansion northwards in the Northern Hemisphere, mainly from the Alaska Peninsula. Our results exemplify the complex phylogeographic history in cold environments and contribute to recognizing evolutionary patterns in the Northern Hemisphere.

The Macaronesian endemic pleurocarpous moss species Andoa berthelotiana occurs in the Madeira, Azores and Canary Islands archipelagos. It has a checkered taxonomic history with placements in different genera or, as monospecific genus Andoa, in different families, most frequently in the Hypnaceae. Earlier molecular phylogenetic analyses indicated a close relationship of Andoa with Ctenidium, Hyocomium and Myurium in a clade corresponding to Myuriaceae, which was supported in the present study based on a larger sampling and markers from all three genomes (chloroplast trnL-trnF, nuclear ribosomal ITS and mitochondrial nad5). Haplotype networks of the nad5 and ITS sequences as well as AFLP fingerprinting data revealed the existence of two intraspecific lineages in Andoa berthelotiana, one occurring in Madeira and the Canary Islands and the other in the Azores. The trnL-trnF haplotype network contradicts this geographic pattern, however, the position of the two mutations differentiating the trnL-trnF haplotypes suggests that they may not be phylogenetically informative. A detailed morphological analysis of plants from the Azores and Madeira indicates the existence of distinguishing characters that correspond with the two molecular lineages. However, the considerable morphological overlap observed would hamper the identification of specimens if both lineages were formally described as separate taxa. We therefore suggest treating the two lineages of Andoa as semi-cryptic species.

Trindade is a Brazilian oceanic island approximately 1,100 km from the American mainland. The Cyathea Sm. plants from Trindade were originally considered an endemic species, Cyathea copelandii Kuhn & Luerssen, but later included in Cyathea delgadii Sternb. Based on molecular data (trnL-trnF, trnG-trnR and rbcL-accD), morphometric analysis of leaf characters, scanning electron microscopy of indusia and spores, and habitat suitability modeling, the present study confirms the status of C. copelandii as a synonym of C. delgadii. Intraspecific molecular variation suggests phylogeographic structuring in C. delgadii and differentiation between the coastal Atlantic Forest domain and the Cerrado domain (savanna) in the interior of Brazil. The Trindade populations are more closely related to the Atlantic Forest populations, indicating a colonization of the island from the latter area.

Molecular phylogenetic reconstructions of Rhabdoweisiaceae based on plastid (trnS-rps4 and trnL-F) and mitochondrial (nad5) DNA sequences and a comprehensive taxon sampling were performed, including 15 out of 16 genera currently attributed to the family, taxa recently found belonging to Rhabdoweisiaceae and expanded outgroups from related families of Dicranidae. The suggested revised delimitation of the family comprises Camptodontium (previously referred to Dicranaceae), three subaquatic species currently classified in Blindia (Seligeriaceae), and Glyphomitrium (previously referred to Ptychomitriaceae, Erpodiaceae or its own family) and Eucamptodon perichaetialis (the type species of the genus, previously referred to Dicnemonaceae). The emended delimitation of Rhabdoweisiaceae thus adds to the already high morphological heterogeneity resulting from the divergent evolution of specialized traits in different lineages of epiphytes and saxicolous hygrophytes. Taxonomic inferences from the phylogenetic analyses include segregation of three new genera, Pseudoblindia (for species currently recognized as Kiaeria falcata, Blindia robusta, B. lewinskyae and B. inundata), Ripariella (for Kiaeria riparia) and Brideliella (for Oncophorus wahlenbergii, O. demetrii and Symblepharis lindigii). Hollia is reinstated for the dicnemonoid Australasian species previously classified in Eucamptodon. Additionally, we propose to merge (1) Oreoweisia, Pocsiella and Pseudohyophila with Cynodontium, and (2) Kiaeria s.s. with Arctoa. Furthermore, we propose to limit Oncophorus to O. virens and O. integerrimus and expand the circumscription of Symblepharis to accommodate O. crispifolius, O. dendrophilus, O. elongatus and O. raui. The generic status and inclusion in Rhabdoweisiaceae are confirmed for Cnestrum and the recently described Notocynodontium, and the exclusion of Amphidium, Dichodontium, Holodontium and Hymenoloma from the family is supported.

Introduction. Half a century since the creation of the International Association of Bryologists, we carried out a review to identify outstanding challenges and future perspectives in bryology. Specifically, we have identified 50 fundamental questions that are critical in advancing the discipline. Methods. We have adapted a deep-rooted methodology of horizon scanning to identify key research foci. An initial pool of 258 questions was prepared by a multidisciplinary and international working group of 32 bryologists. A series of online surveys completed by a broader community of researchers in bryology, followed by quality-control steps implemented by the working group, were used to create a list of top-priority questions. This final list was restricted to 50 questions with a broad conceptual scope and answerable through realistic research approaches. Key results. The top list of 50 fundamental questions was organised into four general topics: Bryophyte Biodiversity and Biogeography; Bryophyte Ecology, Physiology and Reproductive Biology; Bryophyte Conservation and Management; and Bryophyte Evolution and Systematics. These topics included 9, 19, 14 and 8 questions, respectively. Conclusions. Although many of the research challenges identified are not newly conceived, our horizon-scanning exercise has established a significant foundation for future bryological research. We suggest analytical and conceptual strategies and novel developments for potential use in advancing the research agenda for bryology., J. Patiño was funded by the Spanish Ministerio de Ciencia e Innovación (MICINN) through the Ramón y Cajal Program (RYC-2016-20506) and supported by the Fundación BBVA project (PR19_ECO_0046) and the MICINN project (PID2019-110538GA-I00). S. McDaniel was supported by grants from the NSF (DEB 1542609, 1541005, 1541506). J. Martínez-Abaigar was supported by the grant PGC2018-093824-B-C42, funded by MCIN/AEI/10.13039/501100011033 and by the European Regional Development Fund. B. Goffinet was supported by NSF grant DEB-1753811.

An updated list of the taxa of the moss family Pottiaceae recorded for El Salvador is presented, based on literature reports and own investigations. At present, 20 genera of Pottiaceae are known from El Salvador, of which four (Barbula, Scopelophila, Trichostomum, Weissia) are reported as new for the country. Eight species and two varieties are reported as new for El Salvador: Barbula orizabensis, Bryoerythrophyllum ferruginascens, B. inaequalifolium, Chionoloma tenuirostre, Didymodon acutus var. icmadophilus, Hyophila nymaniana, Leptodontium viticulosoides var. viticulosoides, Scopelophila cataractae, Trichostomum brachydontium, and Weissia jamaicensis, increasing the total number of species to 35. Information on the distribution, main substrate types, and approximate elevational range in Mesoamerica as well as the general distribution are provided for all taxa.

Species of the moss genus Schlotheimia are often difficult to identify morphologically, because it is necessary to study sporophytic characters and fertile plants are quite rare. In this paper, we ...

In Antarctica, the genus Bartramia has been restricted to a single polymorphic species, B. patens. Its status as a separate species or a subspecies of the Northern Hemisphere B. ithyphylla was debated. In the present paper, we combine analyses of chloroplast (trnS–rps4–trnT–trnL–trnF region) and nuclear ITS sequences with a reinvestigation of morphological characteristics to infer the identity of Antarctic Bartramia. Phylogenetic and Automatic Barcode Gap Discovery (ABGD) species delimitation analyses indicate that the species diversity of Bartramia in Antarctica has been underestimated, since two species were identified, both belonging to Bartramia sect. Pyridium. Of these, B. subsymmetrica is a new record of the species for Antarctica, as it has previously only been recorded from Livingston Island, South Shetlands. The other species is B. patens, which is separated from B. ithyphylla by newly inferred morphological characteristics and is a sister species to the latter in the molecular phylogenetic analyses. Consequently, we consider B. ithyphylla to be a Northern Hemisphere instead of a bipolar species. The suggested conspecificity of both taxa into one species in the ABGD analysis is considered to result from overlumping by this species delimitation method. The delimitation of the three species of section Bartramia (B. halleriana, B. mossmaniana and B. pomiformis) and the circumscription of the genus Bartramia are discussed.

Research on bryophyte biology has made exciting advances during the last 10 to 15 years since the publications of Goffinet & Shaw (2008) and Frey & Stech (2009) that summarized the knowledge of the field. New fossils provided insights into past bryophyte diversity and integrative taxonomic approaches combine the ever increasing molecular data with thorough assessments of morphology and anatomy. Patterns of speciation, diversity at population level and geographic distributions are becoming better understood, and the interactions of bryophytes with their biotic and abiotic environment are increasingly being revealed. Nevertheless, important knowledge gaps remain, and anthropogenic threats such as habitat alterations and global climate change on bryophyte diversity increase the urgency of further research.

Botany is one of the oldest sciences done south of parallel 60 °S, although few professional botanists have dedicated themselves to investigating the Antarctic bryoflora. After the publications of liverwort and moss floras in 2000 and 2008, respectively, new species were described. Currently, the Antarctic bryoflora comprises 28 liverwort and 116 moss species. Furthermore, Antarctic bryology has entered a new phase characterized by the use of molecular tools, in particular DNA sequencing. Although the molecular studies of Antarctic bryophytes have focused exclusively on mosses, molecular data (fingerprinting data and/or DNA sequences) have already been published for 36 % of the Antarctic moss species. In this paper we review the current state of Antarctic bryological research, focusing on molecular studies and conservation, and discuss future questions of Antarctic bryology in the light of global challenges.

Cover of 50th anniversary of IAB

The ‘Rand flora’ is a biogeographical disjunction which refers to plant lineages occurring at the margins of the African continent and neighbouring oceanic archipelagos. Here, we tested whether the phylogeographical pattern of Exormotheca pustulosa Mitt. was the result of vicariance induced by past climatic changes or the outcome of a series of recent long-distance dispersal events. Two chloroplast markers (rps4-trnF region and psbA-trnH spacer) and one nuclear marker (ITS2) were analysed. Phylogenetic and phylogeographical relationships were inferred as well as divergence time estimates and ancestral areas. Exormotheca possibly originated in Eastern Africa during the Late Oligocene/Early Miocene while Exormotheca putulosa diversified during the Late Miocene. Three main E. pustulosa groups were found: the northern Macaronesia/Western Mediterranean, the South Africa/Saint Helena and the Cape Verde groups. The major splits among these groups occurred during the Late Miocene/Pliocene; diversification was recent, dating back to the Pleistocene. Climate-driven vicariance and subsequent long-distance dispersal events may have shaped the current disjunct distribution of E. pustulosa that corresponds to the Rand Flora pattern. Colonization of Macaronesia seems to have occurred twice by two independent lineages. The evolutionary history of E. pustulosa populations of Cape Verde warrants further study., This work was supported by Fundação para a Ciência e Tecnologia, Portugal (Project PTDC/AGR-FOR/3427/2014). We thank the LISU, E and PRE herbaria for providing material for the development of this work. We also thank Vera Nunes for her support in the laboratory. J.P. was funded by the Ministerio de Economía y Competitividad through the Ramón y Cajal Program (RYC-2016-20506), and Marie Sklodowska-Curie COFUND, Researchers’ Night and Individual Fellowships Global (MSCA grant agreement No. 747238, ‘UNISLAND’). We thank the anonymous reviewers for their helpful comments.

Molecular phylogenetic reconstructions support the monophyly of Pylaisiaceae. The family includes Calliergonella Loeske, Homomallium (Schimp.) Loeske, Pseudohygrohypnum Kanda, Pylaisia Schimp., 2 f...

The diversity of bryophyte and lichen collections in 9 of the oldest preserved herbaria (dating from ca 1542 to 1577) was compared, including the first reports of bryophytes and lichens from the ‘En Tibi’ herbarium (possibly 1542–1544) and the herbarium of Leonhard Rauwolf (1560–1563). Bryophytes and lichens formed only a minority in each herbarium compared to the numbers of vascular plant specimens; numbers ranged from representatives of 21 genera in the herbarium of Ulisse Aldrovandi to the single genus Conocephalum in the Rauwolf herbarium. The focus was on large, handsome species of bryophytes and macrolichens, apart from small amounts of additional species collected as ‘by-catch’ in mixed collections. All herbaria together included 34 genera of bryophytes (36 species and 10 specimens identified to genus level) and 13 genera of lichens (24 species and 4 specimens identified to genus level). The diversity of mosses was higher than that of liverworts, and pleurocarpous mosses were more diverse than acro...

We studied suprageneric relationships and patterns of morphological character evolution within the moss family Leucobryaceae, based on phylogenetic analyses of markers from all three genomes and representatives from 11 out of 14 genera. Three main lineages are strongly supported within the Leucobryaceae, which display different morphologies of the leaf costa, two dicranoid clades (Campylopus clade and Dicranodontium clade) and a leucobryoid clade. Character evolution analyses resolved the leucobryoid morphology as a synapomorphy for the leucobryoid clade and provided data to evaluate generic circumscriptions within the family. Patterns of molecular variation, species richness, geographical distribution, ecological amplitude and variation in costa structure seem to covary: The two lineages with the most modified morphologies (deviating from the basic dicranoid type; Campylopus and leucobryoid clades) are also the most molecularly variable, species diverse, and occupy the broadest distributions and widest variety of habitats. Most sampled genera have clear morpho-molecular circumscriptions, except Campylopodiella, Dicranodontium and Leucobryum. For the latter, two alternative topologies were resolved (Leucobryum monophyletic or the Asian genera Cladopodanthus and Schistomitrium nested within Leucobryum).

The delimitation of the invasive moss species Campylopus introflexus from its closest relative, Campylopus pilifer, has been long debated based on morphology. Previous molecular phylogenetic reconstructions based on the nuclear ribosomal internal transcribed spacers (ITS) 1 and 2 showed that C. pilifer is split into an Old World and a New World lineage, but remained partly inconclusive concerning the relationships between these two clades and C. introflexus. Analyses of an extended ITS dataset displayed statistically supported incongruence between ITS1 and ITS2. ITS1 separates the New World clade of C. pilifer from a clade comprising C. introflexus and the Old World C. pilifer. Ancestral state reconstruction showed that this topology is morphologically supported by differences in the height of the dorsal costal lamellae in leaf cross‐section (despite some overlap). ITS2, in contrast, supports the current morphological species concept, i.e., separating C. introflexus from C. pilifer, which is morphologically supported by the orientation of the hyaline hair point at leaf apex as well as costal lamellae height. Re‐analysis of published and newly generated plastid atpB‐rbcL spacer sequences supported the three ITS lineages. Ecological niche modeling proved a useful approach and showed that all three molecular lineages occupy distinct environmental spaces that are similar, but undoubtedly not equivalent. In line with the ITS1 topology, the C. pilifer lineage from the New World occupies the most distinct environmental niche, whereas the niches of Old World C. pilifer and C. introflexus are very similar. Taking the inferences from ecological niche comparisons, phylogenetics, and morphology together, we conclude that all three molecular lineages represent different taxa that should be recognized as independent species, viz. C. introflexus, C. pilifer (Old World clade), and the reinstated C. lamellatus Mont. (New World clade).

The development of evidence-based international strategies for the conservation and management of Arctic ecosystems in the face of climate change is hindered by critical knowledge gaps in Arctic floristic diversity and evolution. Particularly poorly studied are the bryophytes, which dominate the vegetation across vast areas of the Arctic and consequently play an important role in global biogeochemical cycles. Currently, much of what is known about Arctic floristic evolution is based on studies of vascular plants. Bryophytes, however, possess a number of features, such as poikilohydry, totipotency, several reproductive strategies, and the ability to disperse through microscopic diaspores, that may cause their responses to Arctic environments to differ from those of the vascular plants. Here we discuss several priority areas identified in the Arctic Council’s “Arctic Biodiversity Assessment” that are necessary to illuminate patterns of Arctic bryophyte evolution and diversity, including dispersal, glacial refugia, local adaptation, and ecological interactions with bryophyte-associated microbiomes. A survey of digitally available herbarium data archived in the largest online aggregate, GBIF, across the Arctic to boreal zones indicates that sampling coverage of mosses is heterogeneous and relatively sparse in the Arctic sensu stricto. A coordinated international effort across the Arctic will be necessary to address knowledge gaps in Arctic bryophyte diversity and evolution in the context of ongoing climate change. This article is open access. This work is licensed under a Creative Commons Attribution 4.0 International License (CC BY 4.0). http://creativecommons.org/licenses/by/4.0/deed.en_GB.

The extreme conditions of life in Antarctica pose challenges to many organisms. Bryophytes comprise the main vegetation of the continent and they endure many adversities, thriving in an environment hostile to most living organisms. Mosses are known to exhibit phenotypic plasticity, due to genotypic heterogeneity, environmental influences, or a combination of both. We investigated morphological diversity and its possible relationship with genetic diversity in the Antarctic moss, Roaldia revoluta, and also if the geographic distribution of morphotypes was related with environmental variables. We obtained 49 samples from Antarctica, including King George Island (K), James Ross Island (J), and several locations in the Antarctic Peninsula (P), representing most of the habitats where this species is found. A principal component analysis (PCA) was performed using quantitative measures of gametophytic morphological characters. For molecular analyses, we used DNA sequences of five molecular markers obtained from 29 specimens. Canonical correspondence analysis (CCA) was carried out to compare morphological diversity with environmental variables. Our data support the existence of two morphotypes of R. revoluta, morphotype A (apiculate and ovate leaves), more frequent in K, and morphotype B (acuminate and more oblong leaves), more common in J. Both morphotypes were present in P. The distinct morphotypes could not be related with genetic heterogeneity, since no sequence differences were found in any of the selected markers, supporting the morphological distinction being explained by environmental factors. The results from CCA supported a relationship between the observed morphological variation and the local environmental characteristics of wind speed and minimum temperature.

An integrative taxonomic approach, including molecular phylogenetic reconstructions based on plastid rps4-trnF and nuclear ITS sequences, statistical analysis of morphological-anatomical characters, and classical taxonomy, indicates that the reduction of 13 Amphidium species to three in a recent morphological revision represents a case of too extensive lumping. Instead, six Amphidium species can be distinguished based on molecular and morphological data, the widespread Amphidium lapponicum, A. mougeotii, and A. tortuosum, as well as the Macaronesian endemic A. curvipes, the North American endemic A. californicum, and a newly discovered species from Central Asia (southern Siberia and northern Mongolia), A. asiaticum sp. nov. Diagnostic morphological characters for all six species are discussed. The present data confirm that species diversity of Amphidium is highest in the Holarctic, where all six species occur.

The systematic position of the tropical moss genus Octoblepharum remains under discussion, and in addition, species circumscriptions and relationships of its 18 accepted species remain poorly known. Here we analysed 21 Octoblepharum samples representing four species (O. albidum, O. arthrocormoides, O. cocuiense, O. pulvinatum), within a dataset of haplolepideous mosses and markers from all three genomes. Octoblepharum is resolved as monophyletic and sister to the Calymperaceae, and this clade as sister to the Hypodontiaceae. Based on the molecular results and morphological characters, we suggest Octoblepharum to be placed in its own family, the Octoblepharaceae. The analysed Octoblepharum species are molecularly clearly distinguishable. The two included specimens of the rare and little known Asian species O. arthrocormoides were originally labelled as the common and widespread O. albidum, and their recognition may represent the first records of the species after its description. Octoblepharum albidum show...

Two species of Campylopus, C. fragiliformis and C. pilifer, were previously reported from Trindade Island, a Brazilian island situated in the South Atlantic Ocean around 1100 km off the coast of mainland Brazil. Phylogenetic inferences from sequences of the nuclear ribosomal internal transcribed spacers and three plastid markers are consistent with the recognition of two Campylopus species on Trindade. However, the island populations of C. pilifer are nested among those of C. introflexus and those of C. fragiliformis among those of C. occultus. A re-investigation of morphological characters confirms that the populations from Trindade Island belong to these species. Based on phylogenetic relationships, both C. introflexus and C. occultus probably reached Trindade from continental South America. The colonization of Trindade by C. introflexus in particular might have been facilitated by the severe human impact on the original vegetation throughout the last five centuries.

Hybridization, polyploidization, and reticulate evolution are major mechanisms that shape the current fern diversity. Based on molecular phylogenetic analyses of chloroplast DNA ( psbA-trnH , trnG-trnR ) and low-copy nuclear DNA ( gapCp ) sequences and flow cytometric analysis, we attempt to clarify the origin of putative Nephrolepis hybrids and cultivars. Our results confirm the presence of several hybrids, among which Nephrolepis ‘bostoniensis’ and Nephrolepis x hippocrepicis . Two widespread species, Nephrolepis biserrata and Nephrolepis cordifolia are revealed as the central species in the formation of these hybrids, with Nephrolepis brownii probably occasionally contributing as well. The C-values range indicates that all analysed Nephrolepis taxa are diploid except for Nephrolepis hirsutula , which is probably a tetraploid. All hybrids and cultivars are found to be diploid or homoploid hybrids. Additionally, our results suggest that Nephrolepis davallioides might also be a hybrid, possibly of multiple origins.

A morpho-molecular study was undertaken to solve the taxonomic identity of Riccia plants from the Canary Islands. These plants were assumed to belong to the South African endemic Riccia section Pilifer, but could not be assigned to a particular species in that section. In the interim they were named R. lamellosa (section Riccia), since R. lamellosa is the only European species with conspicuous white ventral scales. Molecular phylogenetic reconstructions based on trnL-F and ITS2 sequences confirmed that the respective Riccia plants belong to section Pilifer. The respective clade is clearly separated from the clades of R. lamellosa and R. elongata, the latter being morphologically most similar within section Pilifer. Based on the combined molecular and morphological evidence the Riccia plants from Canary Islands are described as a new species, R. boumanii, which represents the first species of section Pilifer outside South Africa. Based on revised herbarium specimens, R. boumanii occurs on five islands of t...

Bryophytes have been included in rather few studies of elevational gradients and studies on epiphytic bryophytes on different slopes of a single mountain have not been conducted in Indonesia. This research aims to compare species richness pattern, species composition, and zonation of epiphytic bryophytes on tree trunks on two different slopes of Mt Gede, West Java, Indonesia. We established an elevational transect on the North-North-East (Cibodas) and the South-South-West (Selabintana) slopes of the mountain. The transects ranged from 1500 to 2700 m each and sampling was carried out at 200 m intervals. The Cibodas slope is richer than the Selabintana slope, with 160 species (71 mosses and 89 liverworts) compared to 149 species (57 mosses and 92 liverworts), respectively. Species richness pattern on both slopes showed quadratic responses for mosses and cubic responses for liverworts. Pairwise elevation comparison between slopes showed that species composition is significantly different except for the highest plot (2700 m). Pairwise elevation comparison within the two transects revealed significant species composition changes at different elevations, indicating the presence of an elevational zonation of epiphytic bryophytes along the slopes. Based on overall bryophyte species composition, five elevational zones were recognized on each slope.

The Antarctic microbial community is an important component of the ecosystem in Antarctica. Fungi are among the major Antarctic microorganisms, and several taxa have already been isolated from various substrates including vascular plants, bryophytes, and lichens. Compared with tropical and temperate ecosystems, the diversity of vascular plants in Antarctica is relatively low, such that basal plants, lichens and only two native angiosperms, Colobanthus quitensis (Caryophyllaceae) and Deschampsia antarctica (Poaceae), are dominant. Considering the extreme conditions in Antarctica, angiosperms, bryophytes and lichens, which have adapted to these conditions, represent hotspot microhabitats for fungal diversity. The fungal assemblages associated with these organisms include various taxa categorised as cosmopolitan cold-adapted and endemic species. In the present chapter, we focus on the richness, diversity, and ecology of cryptic fungal assemblages associated with bryophytes, vascular plants and lichens of Antarctica.

© 2018 Biersma, Jackson, Stech, Griffiths, Linse and Convey. From glacial reconstructions it is clear that Antarctic terrestrial life must have been extremely limited throughout Quaternary glacial periods. In contrast, recent biological studies provide clear evidence for long-term in situ persistence throughout glacial times within most extant Antarctic faunal and several microbial groups. However, even now, the evolutionary history of the Antarctic flora-despite playing major role in Antarctic ecosystems-remains poorly studied. We assessed the diversity, richness and relative age divergences within Schistidium (Grimmiaceae, Bryophyta), the most species-rich plant genus in the Antarctic, as well as the plant genus containing most Antarctic endemic species. We applied phylogenetic and molecular dating methods based on nuclear ribosomal Internal Transcribed Spacer sequences, including all known Antarctic Schistidium species with available sample material. We additionally investigated the continent-wide genetic diversity within the most common Antarctic representative of the genus-the endemic species Schistidium antarctici-and performed preliminary phylogeographic analyses of the bipolar species Schistidium rivulare. Most previously described Antarctic Schistidium species were genetically distinct, confirming their specific status. Interspecific divergences of all species took place at least ~1 Mya, suggesting a likely in situ persistence in Antarctica for (at least) all endemic Schistidium species. The widespread endemic species, Schistidium antarctici, diverged from other Antarctic congeners in the late Miocene, thereby revealing the oldest extant plant species currently known in Antarctica, and providing increasing support for the hypothesis of vegetation survival through multiple glacial periods. Within S. antarctici we identified several distinct clades dividing the eastern Antarctic Peninsula and Scotia Arc islands from the western Antarctic Peninsula and all continental locations. This suggests that the mountainous spine on the Antarctic Peninsula forms a strong barrier to gene flow in this species, while increased genetic diversity in the northern Maritime Antarctic indicates likely glacial refugia in this area. This study provides an important first step toward assessing the diversity and evolutionary history of the most speciose moss genus in the Antarctic. The multi-million year presence of several endemic species contributes to studies on their adaptive potential to survive climate change over both historical and contemporary timescales.

The cosmopolitan genus Porella L. is the largest genus of Porellaceae with about 50–60 species (Schuster, 1980). Identification of Porella species can be difficult, due to high morphological plasti...

How many species of Isothecium(Lembophyllaceae, Bryophyta) are there in Macaronesia? A survey usingintegrative taxonomy

Because of their morphological plasticity and broad geographic distribution, the taxonomy of Dicranum is difficult. The circumscription of the different species included in the Dicranum acutifolium complex is poorly understood and taxonomic confusions are frequent. The present study extends earlier internal transcribed spacer (ITS)-based phylogenetic reconstructions of the D. acutifolium complex by analysing five additional chloroplast markers (trnT– rps4, trnL–F, psbA–trnH, rps19–rpl2, rpoB) together with ITS from a larger taxon sample. The phylogenetic analyses delimit Dicranum acutifolium (Lindb. & Arnell) C.E.O.Jensen, D. bardunovii Tubanova & Ignatova, D. brevifolium (Lindb.) Lindb., and D. septentrionale Tubanova & Ignatova, which together form the D. acutifolium complex, and confirm that D. pseudoacutifolium Otnyukova is synonymous with D. flexicaule Brid. Previously Dicranum septentrionale was known from across Russia, but it also occurs in Scandinavia, where it was probably overlooked due to the morphological resemblance to D. brevifolium. The problem of mixed collections for identification is exemplified by the holotype of D. bardunovii, which also contains individuals of the morphologically most similar D. acutifolium, according to the molecular data. Morphometric analyses support the differentiation of the D. acutifolium complex. Furthermore, ordination analyses point to a continuous range of variation among species within the D. acutifolium complex, especially due to the larger variation in D. septentrionale.

Dicranum is a large (ca. 90 species) and taxonomically complex moss genus. Circumscriptions and relationships of many Dicranum species remain ambiguous due to the absence of a worldwide revision and comprehensive phylogenetic analyses. In this study, we address species circumscriptions and relationships of presumed close allies within Dicranum sect. Dicranum. Molecular phylogenetic reconstructions based on five chloroplast regions and nrITS suggest a close relationship between D. bonjeanii, D. howellii, D. nipponense, D. japonicum, D. cf. lorifolium, and D. scoparium, which can be regarded as the D. scoparium species complex. In contrast, D. majus and D. polysetum, as well as D. fuscescens and D. spadiceum (former varieties of D. scoparium), are separated from the complex. Molecular data are generally congruent with the morphological species concept, but the circumscriptions of D. bonjeanii, D. japonicum, D. cf. lorifolium, and D. scoparium need further study. Most analyzed D. scoparium specimens from across its Holarctic distribution are contained in one clade (D. scoparium s. s.), but a number of North American specimens are resolved as closely related to D. japonicum and D. cf. lorifolium. Costa cross sections and characters of the leaf apex (shape, serrulation of margins) are most relevant for identifying the studied Dicranum species morphologically.

The identification of bryophytes from the Arctic is often difficult due to deviating morphologies under the extreme environmental conditions. This is especially true for species-rich and taxonomically complex genera, such as the moss genus Dicranum. DNA barcoding is expected to improve the identification of Arctic bryophyte species, but the optimal combination of barcoding markers for mosses in general, especially for delimiting closely related, is still under discussion. In this paper, we test the discrimination capacity of six potential barcode markers (rps4-trnT UGU, trnL UAA-trnF GAA, trnH GUG-psbA, rps19-rpl2, rpoB, and nrITS1-5.8S-ITS2) based on phylogenetic reconstructions of 30 Dicranum samples from Spitsbergen (Svalbard, Norway), and reference samples from all ten Dicranum species confirmed for the Svalbard archipelago and six additional Arctic Dicranum species. All 16 species (possibly except D. fuscescens) were distinguishable with bootstrap support >70 % based on the combined sequence data, but none of the individual markers could delimit all included species. All Svalbard collections could be readily assigned to five species, D. acutifolium, D. elongatum, D. laevidens, D. majus, and D. spadiceum, respectively. It is concluded that DNA barcoding improves species identification of Arctic Dicranum plants, but that a combination of several markers is necessary in order to obtain reliable identification results, with the single loci ITS1, trnL-F and rps4-trnT being the most promising regions.