New insight into the taxonomy of Cephaloziellaceae (Marchantiophyta): the family of the smallest higher plants on Earth.

An analysis of the phylogeny of Cephaloziellaceae was carried out based on trees constructed for previously and newly obtained sequences of five genes: nuclear ITS1-2 and chloroplast trnL-F, trnG, rbcL, and psbA. Phylogenetic trees inferred from different genes are congruent for the main details; however, the position of several taxa is variable. As a result, a new phylogenetic system of the family was proposed. The narrow genus concept seems to be more appropriate for the family. Cephaloziella spinicaulis is segregated into the new genus Douiniella, the generic status for Prionolobus and Metacephalozia is confirmed, and the dubious generic status of Kymatocalyx is substantiated. The generic independence of Cylindrocolea from Cephaloziella s. str. is confirmed. The small amount of data hinders the description of two more genera from Cephaloziella s.l.

Liverworts of Magadan: Life on the Edge of Beringia.

Magadan Province, located on the southwestern edge of Greater Beringia, combines various floristic elements in its flora, including Beringian and circum-boreal species. The geographic position and habitat diversity of Magadan Province predicts its liverwort flora is one of the richest hemiarctic floras in Asia. The distribution of species throughout the region is uneven, and while its southern part shows close connections with the suboceanic floras of Kamchatka, the middle and central parts are clearly related to the floras of Chukotka, where the Beringian land bridge directly lies in the past. The wide distribution of basic rocks here leads to the existence of a significant complex of basiphylous taxa. A total of 214 taxa (including 205 species and nine varieties) are reported in the present paper. The study of liverworts of Magadan Province should be continued and several new additions to the floral list are highly likely.

The Type Specimens of Plectocolea and Solenostoma (Marchantiophyta) in Some Japanese and European Herbaria.

Solenostomataceae are among the most diverse and, at the same time, least described liverwort families in Pacific Asia. Many species therein are known only from type descriptions, which are sometimes incomplete. The present account is based on a study of 81 type specimens belonging to 61 species known mostly in East Asia, although also in other regions; it provides line-art figures and photomicrographs of all the studied taxa. One lectotype and one epitype are designated, and six new combinations are proposed. as Additionally, the indication of a necessity of an epitype for Plectocolea setulosa Herzog is justified.

The Liverwort and Hornwort Flora of Jeju Island, Republic of Korea: A Volcanic Island with a Unique Mixture of Subtropical, Temperate, Boreal, and Arctomontane Taxa.

Jeju Island, due to its position at the southern tip of the Korean Peninsula in Northeast Asia, is a on the unique enclave of the many southern elements in the area and features a mixture of subtropical, temperate, boreal, and arctomontane taxa. Among the arctomontane species recorded in this study was Anthelia juratzkana; among the temperate species was Dactyloradula brunnea, and subtropical species were Cavicularia densa, Pallavicinia subciliata, Wiesnerella denudata, and Megaceros flagellaris. A valuable species as first recorded for the Jeju Island is Cryptocoleopsis imbricata. The distribution patterns of these species suggest that the flora of Jeju Island is a meeting place between boreal and subtropical floras. We recorded 222 taxa belonging to 45 families, 80 genera, 209 species, 9 subspecies, and 4 varieties. Among these, 86 species are reported as new to the flora of Jeju Island. A checklist based on a study of 1697 specimens is also provided.

Liverwort and Hornwort Flora of Hoàng Liên National Park and the Adjacent Areas (North Vietnam, Indochina).

The study of the flora located in the central part of the Hoàng Liên SÆ¡n Range in the northern region of Indochina has revealed 279 species of liverwort and hornwort, 26 of which are newly reported for the flora of Vietnam. The uniqueness and peculiarity of the studied flora are explained by the significant altitudinal range in the area treated and its position in the contact zone of the Sikang-Yunnan floristic province of the East Asian Floristic Region with the Indochina Floristic Region. The checklist includes data on the distribution of each species in the studied region, habitats, and accompanying taxa. The high disunity of the regional floras of the southern tip of the East Asian region compared to the lesser disunity of the regional floras in the north of the East Asian region is shown. In general, the studied flora possess Sino-Himalayan mountain subtropical characteristics with the large participation of tropical elements.

Marsupellabrasiliensis sp. nov. (Gymnomitriaceae, Marchantiophyta) from Brazil - the distribution of sect. Stolonicaulon in Neotropics is now confirmed.

The specimen previously identified as Marsupellamicrophylla from Brazil is reassessed and described as a new species, M.brasiliensis. The new species is characterized by paroicous inflorescence, bispiral elaters, scale-like, commonly unlobed leaves and very small leaf cells. Descriptions and drawings are provided along with a corresponding discussion of the morphological peculiarity of the new species. Marsupellabrasiliensis belongs to sect. Stolonicaulon, and the distribution of Marsupellasect.Stolonicaulon in the New World is confirmed. The infrageneric position of M.microphylla remains unresolved, and whether it belongs to the same section is still unclear.

Konstantinovia Is Not Monotypic, and a New Attempt to Determine Relationships in Cephaloziellaceae-Scapaniaceae Superclade (Marchantiophyta).

The exploration of liverworts on Bering Island (the westernmost Aleutians) has revealed plants assigned to the recently described and previously monotypic Konstantinovia, previously known only from Yunnan Province of China, and belonging to the bigeneric Obtusifoliaceae. The collected plants are described here as Konstantinovia beringii sp. nov. The known localities of two species of Konstantinovia are separated by more than 6000 km, while the presence of the genus on the Commander Islands is probably a relict. Phylogenetic examination of both collected specimens and new material from other related families resulted in the construction of a fairly well-supported phylogenetic tree for the entire Cephaloziellaceae s.l. + Scapaniaceae s.l. clade. The constructed trees have confirmed the previously stated assumption that it is necessary to segregate one more family within this superclade, described here as Oleolophoziaceae fam. nov.

Hidden Diversity within Tetralophozia filiformis (Marchantiophyta, Anastrophyllaceae) in East Asia.

Tetralophozia filiformis s.l. is known from a number of localities mostly in amphi-oceanic areas in Northern Hemisphere, including Atlantic Europe, amphi-Pacific Asia, South Siberia, and western North America. The newly obtained collections of this 'species' show strong variation in morphology of the taxon across amphi-Pacific Asia although connected by some 'intergrading' modifications. This implies the genetic diversity within this unit earlier recognized as a single taxon. Authors used molecular-genetic, morphological, and chorological methods to understand if the geographically correlated morphological variation also correlates with genetic differences and if it is possible to distinguish some additional taxa within the series of specimens originating from the various areas in amphi-Pacific Asia. It was found that Tetralophozia filiformis is a complex of at least three morphologically similar species, including one long forgotten name (Chandonanthus pusillus) that should be reinstated as separate species and one taxon (Tetralophozia sibirica) that should be described as new. Tetralophozia filiformis and Chandonanthus pusillus are lectotypified, and the new combination is provided for the latter. The three accepted taxa distinctly differ one from another in distribution patterns, preferable climate characteristics, and genetic distances, besides minor differentiations in morphology. The main morphological distinguishing features are the leaf cell size, height of undivided part in leaf lamina, and leaf dentation characteristics. Taking into account the robust correlation between the climate-based and molecular-genetic-based clusters, one more (fourth) taxon could be probably segregated from Tetralophozia filiformis.

The Taxonomically Richest Liverwort Hemiboreal Flora in Eurasia Is in the South Kurils.

The long coexistence of various floral elements, landscape diversity, and island isolation led to the formation of the richest Eurasian hemiboreal liverwort flora in the southern Kurils. This land that covers less than 5000 square kilometres and houses 242 species and two varieties of liverworts and hornworts. The flora 'core' is represented by hemiboreal East Asian and boreal circumpolar taxa. Other elements that have noticeable input in the flora formation are cool-temperate East Asian hypoarctomontane circumpolar and arctomontane. The distribution of some species is restricted to the thermal pools near active or dormant volcanoes or volcanic ash deposits; such species generally provide specificity to the flora. Despite the territorial proximity, the climate of each considered island is characterized by features that, in the vast majority of cases, distinguish it from the climate of the neighbouring island. The last circumstance may inspire the difference in the liverwort taxonomic composition of each of the islands. The comparison of the taxonomic composition of district floras in the Amphi-Pacific hemiarctic, boreal, and cool-temperate Asia revealed four main focal centres: East Kamchatka Peninsula and Sakhalin Island, the southern Sikhote-Alin and the East Manchurian Mountains, the mountains of the southern part of the Korean Peninsula, and the South Kurils plus northern Hokkaido. The remaining floras involved in the comparison occupy an intermediate position between these four centres.

Stolonicaulon: A Section-Puzzle within Marsupella (Gymnomitriaceae, Marchantiophyta).

Marsupella sect. Stolonicaulon is not speciose and is a commonly neglected section within the genus, which currently includes three species with somewhat similar morphologies (wiry shoots with distanced leaves) and distributions in the mountains of tropical and subtropical regions (SE (Southeast) Asia, the Venezuelan Andes, and the high mountains of SE Brazil). After studying materials that were found to be dissimilar to the "traditional" Marsupella that were collected in the last decade by the authors of this article, it was found that these plants belong to three new-for-science species, and all of these species should be included in Marsupella sect. Stolonicaulon. The newly described species have expanded the boundaries of morphological variability, not only for the section itself, but also for the genus based on two findings: (1) the leaves of Marsupella sect. Stolonicaulon can be either appressed and entire or spaced and deeply divided (thus, the plants could occasionally be similar to Cephaloziella or Anastrophyllum); (2) some species of the section possess regular underleaf production. The first discovery of regular underleaves in Marsupella, as noted in two of the three newly described taxa, is the main morphological novelty described in this paper. The development of regular underleaves is a presumable relict character that brings Marsupella closer to Nardia, which was recently transferred to the Gymnomitriaceae and occupies an isolated position within its own subfamily, Nardioideae.

Calypogeia (Calypogeiaceae, Marchantiophyta) in Pacific Asia: Updates from Molecular Revision with Particular Attention to the Genus in North Indochina.

Calypogeia is a genus in Pacific Asia that is difficult to classify taxonomically. These difficulties arise from (1) considering the presence of oil bodies as anatomical characters for taxonomic differentiation, (2) the wide occurrence of sibling, semicryptic and geographical vicariant taxa and (3) the inevitable need to organize new datasets for molecular genetic revision of the genus. The present study uses an integrative approach, including molecular genetic, morphological, chorological and ecological methods, to understand the taxonomy of the genus in Amphi-Pacific Asia. As a result, a set of new-to-science taxa was revealed, and the suite of morphological features necessary for reliable discrimination of the taxa was revised. These results are based on the study of a large set of 'fresh' collections suitable for molecular analysis and morphological comparison and include data on oil bodies. The most basal branch in Calypogeia s.l. is segregated into a new genus, Asperifolia. Descriptions of the new taxa and the key to Calypogeia in Vietnam are provided.

The systematic position of puzzling Sino-Himalayan Lophocoleasikkimensis (Lophocoleaceae, Marchantiophyta) is identified.

Lophocoleasikkimensis, a little-known Sino-Himalayan species, was collected in North Vietnam and its taxonomic position was identified by molecular genetic techniques. The species is characterized by generally narrowly pointed leaves, which are not seen in other representatives of Lophocoleaceae. We found that it belongs to the recently described genus Cryptolophocolea, although it is clearly morphologically dissimilar to other members of the genus. We propose a corresponding nomenclature combination: Cryptolophocoleasikkimensis comb. nov. This species is the only one in its genus with a predominantly Sino-Himalayan distribution; the vast majority of congeners are distributed in the Southern Hemisphere (mostly in Australasia). Reports of this species in Vietnam further confirm the close phytogeographic relationships of the flora of northern Indochina with those of the Sino-Himalayas and suggest that this species is found in other parts of the Hoang Lien Range and the southern Hengduan Range.

Integrating continental mainland and islands in temperate East Asia: liverworts and hornworts of the Korean Peninsula.

The liverwort and hornwort flora of the Korean Peninsula possesses some unique traits arising from the geographic position of the Peninsula, where the mainland flora meets insular flora. This flora is still not exhaustively studied, due not only to political reasons, but also because much less attention has been paid than to adjacent lands by hepaticologists. A checklist presented is based on a study of ca. 15,500 specimens collected by the authors and a review of relevant literature. This study provides the checklist of liverworts and hornworts known from Korea and the geographical distribution of each species within the peninsula. The liverworts and hornworts in Korean flora include 346 taxa (326 species, 16 subspecies and four varieties) in 112 genera and 50 families. Since 2007, 75 taxa of liverworts and four taxa of hornworts are reported as new to the Korean Peninsula, with a number of the new records arising following application of new taxonomic concepts that have become apparent over the last few decades. While compiling the checklist, 42 species, previously reported to Korea, are excluded from the Korean liverwort flora.

Revision of Gymnomitriaceae (Marchantiophyta) in the Korean Peninsula.

This paper provides a revision of Gymnomitrion and Marsupella in the Korean Peninsula based on a study of the collections housed in the herbaria of Jeonbuk National University (JNU) and the Botanical Garden-Institute in Vladivostok (VBGI). In total, 12 species were recorded (six in Gymnomitrion and seven in Marsupella), including four taxa whose identity was not confirmed with the available materials and suspected to be recorded wrongly. Each confirmed species is annotated by morphological descriptions based on available Korean material, data on ecology, distribution, specimens examined as well as illustrations.

Liverwort flora of Ayan - a gained link between subarctic and hemiboreal floras in West Okhotiya (Pacific Russia).

The liverwort flora of Ayan was first investigated one hundred and fifty years after the first exploration of vascular plants. A number of factors has determined the relatively high taxonomic diversity of liverworts in this hemiarctic flora of small-sized area: 118 species and one subspecies were revealed. These data are new not only for the studied area, but also for the huge land adjacent to the western coast of the Sea of Okhotsk. The liverwort flora possesses the domination of taxa common in the hemiarctic, although with a lot of taxa more common in boreal as well as arctic-alpine environments. The presence of Mega-Beringian and calciphilous taxa is the peculiar trait of the studied liverwort flora. Based on detrended correspondence analysis (DCA), Ayan liverwort flora shows relationships to the continental mainland floras situated both in North-East Asian hemiarctic and hemiboreal East Asia and is, therefore, the link between both. The flora of Ayan surroundings is one of the newly-filled 'blank spots' in the possible floral exchange way between Arctic Northeast Asia and mountainous floras of temperate East Asia.

The complete mitochondrial genome of Douinia plicata (Lindb.) Konstant. et. Vilnet (Scapaniaceae, Jungermanniales).

Douinia plicata (Lindb.) Konstant. & Vilnet is the endemic species in Northeast Asia. Here, we reported complete mitochondrial genome of D. plicata. It is 144,206 bp long and includes 72 genes (42 protein-coding genes, three rRNAs, and 27 tRNAs). The overall GC content is 45.1%. Intergeneic variations against S. amplicata, which is slightly higher than intraspecific variations of S. ampliata and W. denudata. Phylogenetic trees show D. plicatum is clustered with three Scapania mitochondrial genomes with high supportive values, which is congruent with previous studies.

The complete mitochondrial genome of Scapania ampliata Steph., 1897 (Scapaniaceae, Jungermanniales).

Scapania ampliata Steph. is the endemic species in East Asia. To investigate intraspecific variations on mitochondrial genomes of S. ampliata, we completed mitochondrial genome of S. ampliata isolated in Korea. It is 143,664 bp long and contains 73 genes (41 protein-coding genes, three rRNAs, 28 tRNAs, and one pseudogene). 823 single nucleotide polymorphisms (SNPs; 0.057%) and 2,242 insertions and deletions were identified between two S. ampliata mitochondrial genomes, which is large number of intraspecific variations in comparison to the other cases of Bryophyte mitochondrial genomes. Phylogenetic trees show that S. ampliata is clustered with those of two Scapania species with high supportive values.

Blepharostoma trichophyllum S.L. (Marchantiophyta): The Complex of Sibling Species and Hybrids.

Blepharostoma trichophyllum was found to be a species collectiva formed by several strongly genetically different species. The taxonomic diversity in the group is the possible result of radiation in early stages; then, these taxa likely survived for a long time in similar environmental conditions, which resulted in stasis. Presently, the existing taxa are similar one to another and may be morphologically distinguished with difficulties. The most taxonomically valuable morphological characteristics include oil bodies and cells in the leaf segment features. The most diverse genotypes (the vast majority of which are treated here as distinct species) were found in amphi-Pacific Asia, which may reflect the evolutionary history of the genus or may be the consequence of more profound sampling in the macro-region in comparison with other parts of the Holarctic.

The complete chloroplast genome of Scapania ampliata Steph., 1897 (Scapaniaceae, Jungermanniales).

We completed chloroplast genome of Scapania ampliata Steph., presenting distinct morphological features including yellowish brown, one-celled gemmae, and decurrent dorsal lobe. It is 118,026 bp long and has four subregions: 80,850 bp of large single copy (LSC) and 19,436 bp of small single copy (SSC) regions are separated by 8,870 bp of inverted repeat (IR) regions including 130 genes (86 protein-coding genes, 8 ribosomal RNAs, and 36 transfer RNAs). The overall guanine cytosine (GC) content is 34.0% and those in the LSC, SSC, and IR regions are 31.9%, 31.0%, and 46.3%, respectively. Phylogenetic trees show S. ampliata is clustered with Scapania ciliata.

The complete chloroplast genome of Wiesnerella denudata (Mitt.) Steph. (Wiesnerellaceae, Marchantiophyta).

We completed chloroplast genome of Wiesnerella denudata (Mitt.) Steph, only one species of the monotypic Wiesnerella genus and family Wiesnerellaceae Inoue. It is 122,500 bp and has four subregions: 82,143 bp of large single copy (LSC) and 20,009 bp of small single copy (SSC) regions are separated by 10,174 bp of inverted repeat (IR) regions including 132 genes (88 protein-coding genes, eight rRNAs, and 36 tRNAs). The overall GC content is 28.8% and those in the LSC, SSC, and IR regions are 26.4, 24.6, and 42.8%, respectively. Phylogenetic trees show incongruencies of phylogenetic relationship of W. denudata, requiring additional research.