BACKGROUND AND AIMS:Protosphagnacean mosses, whose relationships and evolutionary history are incompletely understood, were widespread during the Permian across northern regions of the Northern Hemisphere. M. F. Neuburg proposed that these Permian mosses evolved from a state of absent or weak leaf laminal cellular dimorphism (the Intia type) to marked cellular dimorphism (the Protosphagnum type). Here, we test Neuburg's hypothesis by evaluating changes in the diversity of leaf structure of protosphagnacean mosses from three fossil localities of early, middle and late Permian age. METHODS:Leaves obtained through bulk maceration of rock samples from three localities in north-eastern European Russia separated from each other by several million years were studied to document the cell differentiation patterns, with special focus on leaf apical portions. RESULTS:In the family Protosphagnaceae, the sphagnoid areolation pattern (similar to that of extant Sphagnum leaves) developed following two pathways. In the sphagnacean pathway, the sphagnoid areolation pattern derived from one initial cell via two successive unequal divisions; in the protosphagnacean pathway, the sphagnoid areolation pattern resulted from development in neighbouring cells not necessarily descended from the same initial cell. Large laminal hyaline cells became numerous only in the late Permian. CONCLUSIONS:Our results are consistent with Neuburg's hypothesis, showing a progressive increase in the frequency and diversity of leaf cellular dimorphism in protosphagnacean mosses from the early to the late Permian and indicating that a leaf developmental pathway highly similar to that of extant Sphagnum appeared in the late Permian, bolstering the hypothesis that extant Sphagnum originated from Permian protosphagnaceans.

We revised the Holarctic taxa of Ditrichaceae on the basis of both organellar DNA sequence data and morphological traits. After the removal of phylogenetically discordant elements related to Flexitrichum and Rhamphidium, the remaining representatives of the family cluster into three well-supported lineages, which are recognized here as the reinstated family Ceratodontaceae and a more restricted Ditrichaceae with two subfamilies. Rhamphidium is segregated into a newly described family, Rhamphidiaceae; Aongstroemia gayana, Dicranella vaginata and Symblepharis krausei are combined under Rhamphidium, and the genus Strombulidens is tentatively moved to Rhamphidiaceae. Ceratodontaceae includes the genera Ceratodon, Cheilothela, Trichodon, and a newly described Pseudaongstroemia, which includes a single species, known earlier as Aongstroemia fuji-alpina. The genus Trichodon is paraphyletically defined to include T. cylindricus, T. oblongus reinstated at specific rank, and a molecularly distinct lineage, which we were unable to distinguish morphologically from T. cylindricus. The monoicous genera of Ditrichaceae and the autoicous taxa of Ditrichum as previously understood are merged into a single genus, Pleuridium, which is segregated to Ditrichaceae subfam. Pleuridioideae stat. nov. A single genus is also accepted within Ditrichaceae subfam. Ditrichoideae, following the transfer of both Aongstroemia julacea s.str. and A. orientalis to Ditrichum. Ditrichum septentrionale is newly described based on material mostly from North Asia. Two ecologically specialized, putatively only locally distributed taxa are synonymized with more widespread taxa. Ditrichum plumbicola is placed into synonymy with D. lineare and similarly D. cornubicum is tentatively synonymized with D. macrorhynchum. The morphological delimitations of D. lineare, D. pusillum and D. macrorhynchum are clarified and emended.

The establishment of axenic cultures and the propagation of the rare moss Entosthodon pulchellus (H. Philip.) Brugués (Funariaceae), as well as the optimisation of its ex situ growth conditions and rapid biomass production were the goals of this study. The results obtained also provide insights into the developmental biology of this moss in in vitro conditions, particularly regarding basal media contents with or without supplements of selected plant growth regulators or sugars, i.e. its nutritional needs. The procedure for establishing axenic cultures of this rare and threatened moss species is described. The optimisation protocol of the growth and production conditions is elaborated. The spore germinability of dried herbarium samples remained rather high in the tested laboratory conditions. Suitability tests on different media types showed KNOP basal media to be the best fit for the rapid biomass production of both secondary protonemata and leafy gametophores, without the addition of sugars or plant growth regulators. However, sugar, namely sucrose, induced rapid and massive protonemal development and can be used when this developmental stage is needed.

An integrative taxonomic study based on a combined molecular and morphological analyses resulted in recognition of three species in the genus Anoectangium in Russia: A. aestivum, A. laetevirens, and A. stracheyanum.According to the obtained results, A. laetevirens is a rare East Asian species forming a sister clade to all other studied taxa of the genus.It occurs in Russia only in the Far Eastern mainland and in the Kuril Islands.Anoectangium stracheyanum is the most widespread species of the genus in the country, being known in eastern part of Asian Russia, eastwards from Baikal Lake, ranging from its southern border to the cold region of Yakutia.The clade of A. stracheyanum has two moderately supported subclades composed of northern and southern populations, which, however, are morphologically subidentical, differing only in a slightly longer leaves in plants of the southern clade.The occurrence of Anoectangium aestivum is confirmed in Russia in the Caucasus, Kola Peninsula, and the Baikal Lake area.Both A. stracheyanum and A. aestivum form supported clades, but an occasional occurrense of the DNA motifs of A. stracheyanum in some samples of A. aestivum may indicate a certain introgression.Anoectangium handelii was previously reported from Russia, but DNA barcoding found all so-called specimens belonging to depauperate plants of Gymnostomum, Hymenostylium, or Molendoa. РезюмеИнтегративное таксономическое исследование, основанное на комбинированном анализе морфологических и молекулярных данных, выявило, что род Anoectangium представлен в России тремя видами: A. aestivum, A. laetevirens и A. stracheyanum.Согласно полученным результатам, A. laetevirens, редкий восточноазиатский вид, образует кладу, сестринскую всем остальным видам рода.Он встречается в России только в материковой части юга Дальнего Востока и на Курильских островах.Anoectangium stracheyanum является самым широко распространенным видом рода в России, он известен в ее азиатской части восточнее озера Байкал, от южных границ до восточной Якутии.Клада A. stracheyanum образована двумя умеренно поддержанными кладами

DNA-barcoding revealed/confirmed the range extension of the following bryophytes: Campylostelium saxicola (Krasnodar Territory), Tortella densa (Yakutia), T. fasciculata (Crimea), T. pseudofragilis (Kamchatsky Territory), T. angustifolia (Adygea Republic), and Apopellia megaspora (Murmansk Province). РезюмеС помощью ДНК-баркодинга выявлены или подтверждены находки за пределами основного ареала следующих видов мохообразных: Campylostelium saxicola (Краснодарский край), Tortella densa (Якутия), T. fasciculata (Крым), T. pseudofragilis (Камчатский край), T. angustifolia (Адыгея), и Apopellia megaspora (Мурманская область).

A survey of the moss flora of the southernmost part of the Russian Primorsky Territory yielded several intriguing taxa, whose identity is assessed herein based on an integrative morpho-molecular approach. Bellibarbula recurva was previously known in inland Asia only from the Sino-Himalayan region and the new locality is distant from the earlier known ones to ca. 3000 km. Despite the morphological uniformity, Russian specimens are remarkably distinct in sequences of all three obtained DNA markers, approaching an American specimen in the rps4 sequence. Another probable relic, Symblepharis cf. crispifolia, appeared to be fairly common in the southern part of the Primorsky Territory, where low mountains are covered with hard-leaved forests. Russian specimens of Symblepharis cf. crispifolia var. brevipes show significant divergence from S. crispifolia s.str., which also has complex phylogenetic structure, obscuring further taxonomic implications. The description and illustrations of both taxa based on Russian specimens are provided, and the area, where both species occur, is briefly characterized; it includes numerous thermophilous species, which are rare or do not occur northwards. Our case study uncovers the problem of cryptic speciation within species distributed in temperate climate and is considered to represent relics of Arcto-Tertiary flora.

Hedwigia is a small genus that currently comprises only 12 accepted species. However, taxonomic issues within this genus remain not thoroughly resolved, and its species diversity in China is still debatable. In this study, we conducted an integrative taxonomic revision of Hedwigia in China based on both molecular and morphological evidence. We found an interesting species from Yunnan and Sichuan Province having an unknown combination of morphological characters. The molecular phylogenetic analyses based on mitochondrial nad5, chloroplast trnL-trnF and nuclear ITS markers suggest that this unknown moss represents a new species described here as H. sinica W.Z.Huang & R.L.Zhu, sp. nov. In addition, Hedwigia nemoralis Ignatova, Ignatov & Fedosov is proposed as a new synonym of H. filiformis (Michx.) P.Beauv. based on morphological and molecular evidence. Furthermore, H. czernyadjevae Ignatova, Ignatov & Fedosov, previously known only from Russia, is newly reported for China. The presence in China of H. stellata Hedenäs and H. ciliata (Hedw.) Boucher is doubtful because the voucher specimens of these two species were re-identified as H. sinica and H. emodica or H. filiformis, respectively. In conclusion, at present, four species of Hedwigia are confirmed in China.

A preliminary molecular phylogenetic analysis of Fissidens species of the F. bryoides-complex revealed one species with a relatively weak uni-to bistratose libmidium nested in a clade in which almost all the samples have stout multistratose limbidia.It is a well-defined species, relatively widely distributed in European Russia, in scattered localities eastwards to the Urals.In its nuclear ITS DNA sequences it is most similar to F. monguillonii from Altantic Europe.However, it is a much smaller plant, resembling small phenotypes of F. viridulus, and unlike F. monguillonii it has a weak uni-or bistratose limbidium.This Fissidens species is described as a new species, F. mosquensis.The clade formed by F. monguillonii and F. mosquensis further includes the East Asian F. lateralis (=F.bryoides var.lateralis), a species also characterized by a multistratose limbidium and perichaetial leaves narrower than stem leaves.Fissidens mosquensis is compared with F. bryoides and several morphotypes of F. viridulus s.l.A terrestrial form of F. rivularis from the Caucasus with a uni-to bistratose leaf margin is also discussed.Plants with this combination of traits are nested in a clade in which the other samples represent aquatic F. rivularis. РезюмеПредварительный молекулярно-филогенетический анализ видов комплекса Fissidens bryoides позволил выявить один вид с листьями, имеющими по краю сравнительно слабую 1-2-слойную кайму, который оказался в кладе, почти все представители которой имеют мощную многослойную кайму.Это хорошо морфологически отграниченный вид, довольно широко распространенный в европейской части России, от Смоленской области до Урала.По нуклеотидным последовательностям ядерного участка ITS он ближе всего к F. monguillonii, распространенному в атлантических районах Европы, но, в отличие от последнего вида, это намного более мелкое растение со слабой 1-2слойной каймой.Он описан как новый для науки вид, F. mosquensis.Клада F. monguillonii и F. mosquensis включает также восточноазиатский вид F. lateralis (=F.bryoides var.lateralis), для которого также характерны листья с многослойной каймой и более узкие по сравнению со стеблевыми перихециальные листья.Дано сравнение Fissidens mosquensis с F. bryoides и несколькими морфотипами F. viridulus s.l.Также обсуждается наземная форма F. rivularis с Кавказа, имеющая листья с не характерной для этого вида 1-2-слойной каймой, которая по данным анализа ДНК принадлежат

A recently described from Japan Fissidens neomagofukui Z. Iwats.& Tad.Suzuki differs from most species of the genus in Russia by having a reduced peristome, which teeth are triangular and not divided into two filiform branches.It also has elimbate leaves and therefore has been placed by Suzuki in subgenus Fissidens sect.Aloma, i.e. unrelated to subgenus Fissidens sect.Fissidens that includes F. bryoides and related species.Molecular phylogenetic analysis found F. neomagofukui in a sister position with specimens which are quite similar to species of F. bryoides-group, having almost complete limbidium in all parts of leaf, a typical bryoides-type peristome, and a bryoides-type costa.This species differs from other morphotypes of F. bryoides in smaller, smooth laminal cells and almost obligatory epiphytic growth on hardwood trunks; it occurs in the southern part of Primorsky Territory of Russia.Finding no appropriate existing name, we describe it as F. extremiorientalis sp.nov.Fissidens neomagofukui is confirmed in Russia only for southern part of Primorsky Territory, where it is rare and also grows exclusively on tree trunks, sometimes mixed with F. extremiorientalis. ÐåçþìåÍåäàâíî îïèñàííûé èç ßïîíèè Fissidens neomagofukui Z. Iwats.& Tad.Suzuki îòëè÷àåòñÿ îò áîëüøèíñòâà âèäîâ ðîäà ñèëüíî ðåäóöèðîâàííûì ïåðèñòîìîì ñ òðåóãîëüíûìè çóáöàìè, íå ðàçäåëåííûìè íà äâå óçêî ëèíåéíûå äîëè.Îí òàêaeå èìååò íå îêàéìëåííûå ëèñòüÿ, è ïîýòîìó Ñóçóêè ïîìåñòèë åãî â ïîäðîä Fissidens sect.Aloma, âèäû êîòîðîé íå ðîäñòâåííû òàêîâûì ïîäðîäà Fissidens sect.Fissidens, êîòîðàÿ âêëþ÷àåò F. bryoides è áëèçêèå ê íåìó âèäû.Îäíàêî, ñîãëàñíî ðåçóëüòàòàì ìîëåêóëÿðíî-ôèëîãåíåòè÷åñêîãî àíàëèçà, îí îáðàçóåò êëàäó, ñåñòðèíñêóþ îáðàçöàì, î÷åíü ïîõîaeèì íà âèäû ãðóïïû F. bryoides, ïîñêîëüêó îíè èìåþò êàéìó èç óçêèõ êëåòîê ïî÷òè ïî âñåé äëèíå è âî âñåõ ÷àñòÿõ ëèñòà, ïåðèñòîì bryoides-òèïà è aeèëêó bryoides-òèïà

The recent molecular phylogenetic study of the families Aongstroemiaceae and Dicranellaceae, which resolved the genera Aongstroemia and Dicranella as polyphyletic, indicated the need for changes in their circumscription and provided new morphological evidence to support the formal description of newly recognized lineages. Following up on these results, the present study adds another molecular marker, the highly informative trnK-psbA region, to a subset of previously analyzed taxa and presents molecular data from newly analyzed austral representatives of Dicranella and collections of Dicranella-like plants from North Asia. The molecular data are linked with morphological traits, particularly the leaf shape, tuber morphology, and capsule and peristome characters. Based on this multi-proxy evidence, we propose three new families (Dicranellopsidaceae, Rhizogemmaceae, and Ruficaulaceae) and six new genera (Bryopalisotia, Calcidicranella, Dicranellopsis, Protoaongstroemia, Rhizogemma, and Ruficaulis) to accommodate the described species according to the revealed phylogenetic affinities. Additionally, we amend the circumscriptions of the families Aongstroemiaceae and Dicranellaceae, as well as the genera Aongstroemia and Dicranella. In addition to the monotypic Protoaongstroemia that contains the newly described dicranelloid plant with a 2-3-layered distal leaf portion from Pacific Russia, P. sachalinensis, Dicranella thermalis is described for a D. heteromalla-like plant from the same region. Fourteen new combinations, including one new status change, are proposed.