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Systematics of the Bryophyta (mosses): From molecules to a revised classification

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... The sporomorph glossary used in this paper mainly follows Punt et al (2007) and Traverse (2007). The systematic scheme of extant plants is based on Christenhusz et al. (2011) for gymnosperms, Söderström et al. (2016) for hornworts and liverworts, Goffinet and Buck (2004) for mosses, and Schuettpelz et al. (2016) for lycophytes and ferns. The systematic scheme of fossil plants mainly follows Taylor et al. (2009). ...
... Species of extant ENCALYPTACEAE Schimper (Goffinet and Buck, 2004) are mostly distributed in high mountain or glacial related regions (Gao et al., 1996;Horton, 1978). Bryobrittonia Williams occur most commonly in sandy or silty soils along streams or rivers where the substrate is constantly moist, while Encalypta Hedwig can be found in bogs, on arid ground, or rocks. ...
... Species of extant SPHAGNACEAE Dumortier (Goffinet and Buck, 2004) are nearly cosmopolitan, but most of them are found in circumboreal regions in bogs, wetlands, or swamp forests (Boros et al., 1993;Gao, 1994). They are generally hydrophytes and eurythermic plants. ...
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The eco-group classification based on the growth-form of plants (Eco-Plant model) is widely used for extant, Cenozoic, Mesozoic, and Paleozoic palaeoenvironmental reconstructions. However, for most Mesozoic dispersed sporomorphs, the application of the Eco-Plant model is limited, because either their assignment to a specific eco-group remains uncertain or the botanical affinities to plant taxa are unclear. A new database Sporopollen (http://www.sporopollen.com) focused mainly on Mesozoic sporomorphs is created. Currently, it has collected 100,610 sporomorph pictures, 59, 498 plant pictures, 31, 922 sporomorph descriptions. At the same time, from 63, 035 references, it has collected 2, 215, 162 occurrences for both sporomorph and non-sporomorph fossils. The collected plant data include 32, 972 genera from 946 families. The collected sporomorph pictures include 5, 857 genera. With the help of the database, 861 dispersed Mesozoic sporomorph genera of Bryophytes, Pteridophytes, and Gymnosperms are reviewed by comparing the unique outline and structure/sculpture of the sporomorph wall with that of modern plants and in situ fossil plants. The results show that 474 of them can be linked to their closest parent plants and Eco-Plant model at family or order level, but 387 of them can not because of the lack of detailed ultrastructure descriptions. The use of a light microscope (LM) for determination is one of the main reasons that some dispersed sporomorphs cannot be linked precisely to their parent plants. The presented eco-groups for disperse Mesozoic sporomorphs provide the possibility to identify detailed vegetation and palaeoenvironmental change in the Mesozoic, especially in the context of climate change. A new interface (http://www.sporopollen.com/sporemesozoicsegs.php?opencode=paper1) was created based on the reviewed result to quickly link the dispersed sporomorphs to past vegetation patterns and climatic changes. Users can upload their data to the database and in return get quick results. It can automatically link all of the Mesozoic and Cenozoic sporomorphs to their possible parent plants at phylum, order, or family level. It can also automatically link all of the Triassic and Jurassic sporomorphs to the Eco-Plant model to assess the effect of humidity (EPH) and the effect of temperature (EPT). By using 30 sporomorph samples from a 10 m thick lignite bed from the Upper Triassic Haojiagou Formation (Rhaetian) as an example, the palaeovegetation and palaeoenvironment of a peat-forming wetland near the Triassic-Jurassic boundary are discussed with the help of the Eco-Plant model. The results show that the palynoflora contains both Eurasian and Gondwanan elements, and is dominated by the spores and pollen of Bennettitales, Corystospermales, Ginkgoales, and Gleicheniales. At the Triassic/Jurassic boundary (Hettangian), the palynoflora significantly changes as Cyatheales spores become the dominant elements. We analyse assemblages in terms of an Eco-Plant model, which assigns the parent plants of the palynomorphs into five groups based on humidity and four groups based on temperature, and uses multivariate statistical analyses to infer palaeoclimate and palaeoenvironmental conditions. Results suggest that the palaeoclimate of the Rhaetian was generally wet and subtropical with short seasonal drought periods. Our analysis shows that an Eco-Plant model may be a useful tool to reveal past vegetation patterns and climate changes, applicable to other Mesozoic assemblages.
... The sporomorph glossary used in this paper mainly follows Punt et al. (2007) and Traverse (2007). The systematic scheme of extant plants is based on Christenhusz et al. (2011) for gymnosperms, Söderström et al. (2016) for hornworts and liverworts, Goffinet and Buck (2004) for mosses, and Schuettpelz et al. (2016) for lycophytes and ferns. The systematic scheme of fossil plants mainly follows Taylor et al. (2009). ...
... Species of extant ENCALYPTACEAE Schimper (Goffinet and Buck, 2004) are mostly distributed in high mountain or glacial related regions (Gao et al., 1996;Horton, 1978). Bryobrittonia Williams occur most commonly in sandy or silty soils along streams or rivers where the substrate is constantly moist, while Encalypta Hedwig can be found in bogs, on arid ground, or rocks. ...
... Species of extant SPHAGNACEAE Dumortier (Goffinet and Buck, 2004) are nearly cosmopolitan, but most of them are found in circumboreal regions in bogs, wetlands, or swamp forests (Boros et al., 1993;Gao, 1994). They are generally hydrophytes and eurythermic plants. ...
Article
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The ecogroup classification based on the growth-form of plants (Eco-Plant model) is widely used for extant, Cenozoic, Mesozoic, and Paleozoic paleoenvironmental reconstructions. However, for most Mesozoic dispersed sporomorphs, the application of the Eco-Plant model is limited because either their assignment to a specific ecogroup remains uncertain or the botanical affinities to plant taxa are unclear. By comparing the unique outline and structure/sculpture of the wall of dispersed sporomorph to the sporomorph wall of modern plants and fossil plants, 861 dispersed Mesozoic sporomorph genera of Bryophytes, Pteridophytes, and Gymnosperms are reviewed. Finally, 474 of them can be linked to their closest parent plants and Eco-Plant model at family or order level. Based on the demands of the parent plants to different humidity conditions, the Eco-Plant model separates between hydrophytes, hygrophytes, mesophytes, xerophytes, and euryphytes. Additionally, due to different temperature demands a separation in megathermic, mesothermic, microthermic, and eurythermic plants is possible. In the Mesozoic, both spore-producing and pollen-producing plants are adapted to different kinds of humidity. The concept to use the spore/pollen ratio to reflect the hygrophytes/xerophytes ratio is therefore questionable. The presented ecogroups for dispersed Mesozoic sporomorphs now allow identifying at least relative plant, paleoenvironmental and paleoclimate changes in Mesozoic sedimentary records.
... Based on morphological characteristics of gametophyte, Brotherus (1925Brotherus ( , 1929 considered Sciaromiopsis to be closely related to Sciaromium for the limbate leaves, and also to Leptodictyum for elongate cell of leaf and more or less differentiated auricles. The position of Sciaromiopsis was placed in Amblystegiaceae by Brotherus (1925), and the treatment was followed by Vitt (1984), Shaw & Goffinet (2000), Buck & Goffinet (2000), Goffinet & Buck (2004), Goffinet et al. (2009), andFrey &Stech (2009). Ochyra (1986) once suggested this genus to be a member of Donrichardsiaceae, a family treated as a synonym of Amblystegiaceae subsequently (Crosby 1999). ...
... Hence, the Amblystegiaceae is difficult to delimit, serving as a catch-all family for various aquatics of uncertain affinities (Ochyra 2003). At first, Brotherus (1924Brotherus ( , 1929 considered Sciaromiopsis to be closely related to Sciaromium and Leptodictyum, two genera of Amblystegiaceae, and this opinion was accepted by many subsequent classification systems (Buck & Goffinet 2000;Frey & Stech 2009;Goffinet & Buck 2004;Goffinet et al. 2009;Shaw & Goffinet 2000;Vitt 1984). However, the classification was based mostly on old literature; the morphology of gametophytes was studied in detail after referencing the characters also important for distinguishing pleurocarpous mosses (Hedenäs 1989;Ignatov 1999). ...
Article
Sciaromiopsis comprises a single species, S. sinensis, endemic to China. The genus is currently placed in the Amblystegiaceae, solely on the basis of gametophytic characters since its sporophytes are unknown. As a semiaquatic plant, the character of a strong leaf costa and marginal borders can aid S. sinensis to adapt drought as while as aquatic conditions. In this study, the phylogenetic position of Sciaromiopsis was revisited based on morphological and molecular phylogentic evidences of three datasets, the plastid rps4 dataset with 346 mosses species, the ITS dataset with 104 moss species and 82 plastid protein-coding gene dataset with 32 moss species. According to the morphological characters of gametophyte, the stable pattern of pseudoparaphyllia, leaf cells smooth and single costa of Sciaromiopsis are consistent with key characteristics of the Brachytheciaceae, the ovate-lanceolate leaves, differentiated alar cells and rhomboidal median leaf cells are consistent with key characteristics of the Brachythecium. Maximum likelihood and Bayesian analyses of both datasets of the rps4 gene and the 82 plastid protein-coding genes clearly indicated that Sciaromiopsis is a member of Brachytheciaceae. Sciaromiopsis sinensis is either clustered in a well supported clade among species of Brachythecium, Koponeniella and Myuroclada in the rps4 tree, or strongly supported as the sister to Brachythecium rivulare in plastid 82 gene tree. Maximum likelihood analysis of the nuclear ITS locus suggested that S. sinensis is nested within Brachythecium with moderate support. Thus, S. sinensis should be considered as a member of the Brachythecium, the diagnostic character of its leaf margin bordered by 58 rows of linear cells making it an easy distinction from other species of Brachythecium.
... The specimens were identified with the help of previously published floras such as Gangulee (1969Gangulee ( -1980, Chopra (1975), Chopra & Kumar (1981), and Kumar (1980). The families are arranged according to Goffinet & Buck (2004) system of classification. ...
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The present study aims to account for the moss flora of Chail Wildlife Sanctuary, district Solan, Himachal Pradesh (HP). Frequent field visits were made in different seasons to collect moss samples. Ecological data like temperature, humidity, and habitat preferences were also recorded at the time of collection. A total of 31 moss species belonging to 22 genera and 15 families were recorded so far. The family Pottiaceae (7 spp.) was the most dominant one, followed by Brachytheciaceae (4 spp.), Polytrichaceae, Fissidentaceae, and Entodontaceae with (3 spp.) each. In the acrocarpous mosses, family Pottiaceae was highly dominant, whereas, among the pleurocarpous mosses family, Brachytheciaceae was dominant. Among the genera, Atrichum P. Beauv. (Acrocarpous), Fissidens Hedw. (Acrocarpous), and Entodon C. Muell. (Pluerocarpous) were dominantly present, represented by three species each. This study provides baseline data of moss diversity and their ecological attributes in Chail Wildlife Sanctuary, which may prove beneficial in establishing policies for future exploration of bryodiversity with proper management and conservation in the sanctuary area.
... Сфагновите мъхове се отнасят към клас Sphagnopsida, включващ един подклас Sphagnidae с два разреда: Sphagnales и Protosphagnales. В разред Sphagnales са включени всички рецентни видове, които са обединени в 4 рода: Sphagnum, Ambuchanania (1 вид), Eosphagnum (1вид) и Flatbergium (2 вида), докато разред Protosphagnales включва единствения описан досега фосилен вид (Goffinet et al. 2004(Goffinet et al. , 2008. ...
Thesis
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The thesis contains summarized data about the composition, distribution and ecology of Sphagnum-dwelling testate amoebae from Bulgaria. The materials were collected from Vitosha and Stara Planina Mountains in 2016, Rila and Pirin Mountains in 2017 and Rhodopes Mountains in 2018. A total of 266 samples (one qualitative and one quantitative) from 133 localities of 17 Sphagnum species were examined. As a result of the present study we established 148 species from 41 genera and 17 families There is a high degree of similarity between the different mountains, as the number of species varies within narrow limits (between 110 in Stara Planina and 116 species in Pirin). Most of the registered testate amoebae are typical sphagnophilic representatives belonging to the genera Assulina, Cyphoderia, Gibbocarina, Hyalosphenia, Longinebela, Nebela, Playfairina, Quadrulella and Sphenoderia. A large percent of the commonly found species are eurybionts (Argynnia dentistoma, Centropyxis aerophila, Centropyxis aculeata, Corythion dubium, Trachelocorythion pulchellum, Euglypha spp. and Trinema spp.). Among the accidentally imported as a result of meteorological conditions (wind and/or running water) are testaceans characteristic of aerophilous mosses (Awerintzewia cyclostoma and Bullinularia indica), as well as typical freshwater (Arcella spp., Difflugia spp. and Netzelia spp.) and soil (Centropyxis spp., Plagiopyxis spp. and Phryganella spp.) representatives. The analysis of the data from testate amoebae communities from different mountain shows that in general, they are characterized by extremely high similarity (82.88% – 88.70%). The small differences in the fauna of these mountains can be explained mainly by the presence of a relatively large number of rare species found in only one of the mountains (27 species in total, 18.2% of all found). When comparing different regions, the results clearly show that relatively distant areas from different mountains form well-separated clusters. A decisive role for the development of similar fauna has the presence of similar ecological conditions. 44 In the quantitative studies genera showing the largest numbers in the testacean communities in Vitosha, Stara Planina, Rila and Pirin are Hyalosphenia, Trinema, Euglypha, Assulina and Nebela. An exception is the Rhodopes Mountains, where as dominant genus is registered Archerella. In all mountains the structural parameters vary in a relatively narrow range. High values of the Shannon-Weaver, Margalef and Pielu’s indices are observed, and at the same time very low for the Simpson’s dominance index. These values show in general, that the ecological conditions in Sphagnum mosses are very favorable and the testate amoebae communities are characterized by high species diversity and high degree of species uniformity. The relationship between testate amoebae communities and environmental factors has been studied through ordinal analysis. Environmental factors with highest impact on the distribution and community structure are the type of substrate (Sphagnum moss), groundwater level, oxygen content and pH.
... As espécies de Marchantiophyta e Bryophyta foram listadas por ordem alfabética de divisões, famílias, gêneros, espécies e variedades para sete localidades, no estado de São Paulo, segundo sistemas de classificação em Goffinet & Buck (2004), para os musgos, e Crandall-Stotler & Stotler (2000), para as hepáticas. ...
Article
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O manguezal possui vegetação denominada mangue, cujas abundantes epífitas são geralmente omitidas nas abordagens sobre este ecossistema. Os resultados desse estudo incluem dados de coletas e registros na literatura sobre a ocorrência de espécies de Marchantiophyta e Bryophyta nos manguezais, em Ubatuba, São Vicente, Praia Grande, Itanhaém, Peruíbe, Iguape e Cananéia. O trabalho lista 19 famílias, com 56 gêneros e 115 espécies. A lista acrescenta 43% das 115 espécies de Marchantiophyta e Bryophyta, que ora ocorrem nos manguezais estudados do estado de São Paulo. As hepáticas predominam nas espécies de Marchantiophyta e Bryophyta destes manguezais, que são muito semelhantes às espécies encontradas na Mata Atlântica paulista.
... There is a contradictory hypothesis by Vitt (1984) which clocked the origin of the pleurocarpous members of Hypnales to be fairly recent, as the contemporaries of the early angiosperms flourishing around Cretaceous period. However, the findings of this study are supported by the molecular phylogenetic study by Goffinet and Buck (2004) claiming diplolepidous mosses to have evolved much earlier than the haplolepidous members, which apparently evolved later, with advantages favoured by the evolutionary selection. ...
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Bryophyta comprises one of the earliest lineages of land plants that had implemented remarkable innovations to their lipid metabolic systems for successful adaptation to terrestrial habitat. This study presents a comprehensive investigation of fatty acid profiles of mosses from Eastern Himalayas with an aim to trace their chemotaxonomic and evolutionary implications. Fatty acid compositions of 40 random mosses belonging to major families of Bryophyta were explored by gas chromatographic analysis. A diverse array of saturated, monounsaturated and polyunsaturated fatty acids including rare acetylenic fatty acids were detected. Hexadecanoic acid (C16:0), 9,12 (Z,Z)-octadecadienoic acid (C18:2n6) and 9,12,15 (Z,Z,Z)-octadecatrienoic acid (C18:3n3) were the predominant fatty acids in all the mosses. However, quantitative variation of C20 polyunsaturated fatty acids (PUFAs), specifically 5,8,11,14 (Z,Z,Z,Z)-eicosatetraenoic acid (C20:4n6), among the investigated mosses was the most prominent outcome. The diplolepidous members of Bryidae, especially the mosses of Hypnales, Bryales and Bartramiales contained higher amount of C20 PUFAs compared with the haplolepidous orders. Principal component analyses based on individual fatty acids and other related parameters validated C20:4n6 content and the ratio of C20:4n6/C18:2n6 as the apparent chemotaxonomic discriminants. The prevalent notion of considering 9,12,15-octadecatrien-6-ynoic acid (C18:4a) as the chemomarker of Dicranaceae has also been challenged, since the compound was detected not only in different families of Dicranales, but also in a Pottiales member, Leptodontium viticulosoides. Therefore, an ensemble of fatty acids instead of a single one can be considered as the chemical signature for taxonomic interpretation which may also be vital from an evolutionary standpoint.
... osses are small flowerless plants that typically grow in dense green clumps or mats, often in damp or shady locations (Buck and Goffinet, 2000). The individual plants are usually composed of simple leaves that are generally only one cell thick, attached to a stem that may be branched or unbranched and play only a limited role in conducting water and nutrients (Buck and Goffinet, 2000;Goffinet and Buck, 2004). Although some species have conducting tissues, these are generally poorly developed and structurally different from similar tissue found in vascular plants. ...
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(1) Background: Detailed diversity information regarding terrestrial mosses in a tropical forest ecosystem and an understanding of the drivers behind moss distribution provide crucial data for the management and conservation of forest ecosystems. Mosses are critical components of tropical forest ecosystems due to their high diversity and biomass, and they also fulfill essential ecological functions. Here, we report the first study into the relative importance of vegetation types and elevational gradient for the diversity, distribution and community structure of terrestrial moss species in southern China. (2) Methods: Five elevations spaced 200 m apart in the tropical mountain forest on the northern aspect of Tai Mo Shan were selected. The diversity, distribution, and geographical patterns of terrestrial mosses in response to altitudinal changes were examined. Differences in the biotic variables of terrestrial mosses between elevations were tested using a one-way ANOVA. Curve estimation regression models were used to describe the responses of the biotic variables to the elevation gradient. Canonical correlation analysis (CCA) was performed to identify and measure the associations among biotic variables of terrestrial mosses and abiotic environmental factors. (3) Results: Fifty-three terrestrial moss species belonging to 20 families and 31 genera were recorded along the altitudinal gradient on Tai Mo Shan. Microclimate factors including dew point and rainfall were strongly associated with the cover and thickness of the ground moss species. There were no obvious richness changes of terrestrial mosses along the elevation gradient. In total, 33 of the 51 species were tropically distributed, 14 species were found across East Asia and the tropical regions, 10 species had an East Asian pattern and 8 were temperate species. (4) Conclusions: Vegetation types significantly affect the diversity and distribution of terrestrial moss species. Although they are influenced by the East Asian and temperate climate with frequent human activities, terrestrial mosses on Tai Mo Shan are primarily tropical in nature. Forest conservation and restoration should be implemented to sustain and improve the diversity of terrestrial mosses and understory plants on Tai Mo Shan, especially at higher elevations.
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Sporoderm development in Andreaeobryum is studied and confirned to be similar to that in Andreaea in having no lamellate structure in the exine. The formation of the outer layers of the spore wall, external to the intine, proceeds by the accumulation of sporopollenin granules originating within the special wall of the sporocyte. Despite the outermost sublayer having a pattern similar to the perine in most mosses, and consequently was earlier reported as perine, it is mostly formed before the initial sporocyte walls disappear, thus questioning significant tapetal contribution. Therefore, we suggest to referring the outer sporoderm layer of Andreaeobryum to exine, whith has however is not originated by means of the tripartite lamellae, as in ferns, lycopods, hornworts, liverworts, and all other mosses excepting Andreaeobryopsida and Andreaeopsida.
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Ribosomal RNA sequences and cladistic analysis were used to infer a phylogeny for eight bryophyte taxa. Portions of the cytoplasmic large (26S-like) and small (18S-like) subunit ribosomal RNA genes were sequenced for three marchantioid liverworts (Asterella, Conocephalum, and Riccia), three mosses (Atrichum, Fissidens, and Plagiomnium), and two hornworts (Phaeoceros and Notothylas). Cladistic analysis of these data suggests that the hornworts are the sister group to the mosses, the mosses and hornworts form a clade that is sister to the tracheophytes, and the liverworts form a clade sister to the other land plants. These results differ from previous cladistic analyses based on morphology, ultrastructure, and biochemistry, wherein the mosses alone are sister group to the tracheophytes. We conclude that cladistic analysis of molecular data can provide an independent data set for the study of bryophyte phylogeny, but the differences between the molecular and morphological results are a topic for further investigation.
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The Tetraphidae is a small subclass of mosses with a nematodontous peristome that has frequently been interpreted as primitive among the true mosses. The developmental cell sequence leading to the formation of the four peristome teeth of Tetraphis pellucida is described for the first time. Comparisons are made with sequences known for other nematodontous and arthrodontous mosses. Peristome development in T. pellucida is more like that described previously for arthrodontous peristomes than to published developmental sequences for nematodontous peristomes of species in the Polytrichaceae. On the other hand, our observations confirm a basic uniformity of the earliest developmental stages in all mosses studied thus far, regardless of their systematic position.
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The systematic affinities of three monotypic genera, Kleioweisiopsis Dixon, Pleurozygodontopsis Dixon, and Trigonodictyon Dixon & P. de la Varde, as well as of the monotypic family Microtheciellaceae (Orthotrichales) are investigated. Examination of morphological characters suggests that Kleioweisiopsis should be transferred to the Ditrichaceae, Pleurozygodontopsis synonymized with Zygodon, and Trigonodictyon transferred to the Grimmiaceae. A key to the genera of the Ditrichaceae with immersed capsules is presented. Microtheciella kerrei Dixon, recently placed in its own family, is excluded from the Orthotrichales on the basis of its pleurocarpy, and placed close to the Neckeraceae in the Leucodontales.
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So far morphological approaches have produced little consensus in formulating clear definitions of the mostly southern hemispheric Lembophyllaceae and Meteoriaceae, two pleurocarpous and mainly epiphytic moss families. The generic limits proposed by various authors have resulted in an overlap of family concepts, and the systematic position of genera such as Weymouthia and Pilotrichella has been unsettled since the description of these families by Brotherus (1907) and Kindberg (1897). A molecular approach based on sequence data of two noncoding regions of the chloroplast DNA (cpDNA), trnLUAA intron and trnLUAA 3′exon-trnLGAA intergenic spacer (trn data) as well as sequence data of the internal transcribed spacer 2 (ITS2) of the nuclear ribosomal DNA (nrDNA) provides new evidence supporting the transfer of the genus Weymouthia into the Lembophyllaceae. Although the inter- and intrageneric sequence divergence is extremely low, the highly congruent maximally parsimonious trees from trn and ITS2 data sets suggest a close relationship of the Lembophyllaceae taxa sensu Tangney (1997b). The Lembophyllaceae as treated here contain five genera Camptochaete, Lembophyllum, Fifea, Fallaciella and Weymouthia. Within the Meteoriaceae Meteorium and Papillaria are retained as separate genera, while Weymouthia is excluded from the Meteoriaceae and transferred to the Lembophyllaceae. The molecular data does not align Pilotrichella with either the Meteoriaceae or the Lembophyllaceae sampled. A transfer of Pilotrichella into the Lembophyllaceae is not supported. Pulchrinodus inflatus is neither part of the genus Weymouthia nor of the Meteoriaceae or Lembophyllaceae.
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Many dung mosses (Splachnaceae) are characterized by insect-mediated spore dispersal. All of the entomophilous species are coprophilous, whereas anemophilous species are humicolous or epiphytic. The three species of the Voitioideae are coprophilous but are distinguished from other members of the family by sporangia that remain closed (cleistocarpous) and lack a peristome. Spores are released when the sporangial wall disintegrates. Phylogenetic analyses of nucleotide sequences of the trnL-trnF region and the rps4 locus (cpDNA) for 25 species of Splachnaceae suggest that this combination of characters arose twice within the Splachnaceae and that Voitia grandis is more closely related to species of Tayloria subgenus Tayloria rather to the other species of Voitia, which are nested within the genus Tetraplodon. Hence the new combination Tayloria grandis (Long) Goffinet & Shaw is made. Although the optimal trees were left unrooted, our results resolve the Voitioideae (i.e., the genus Voitia) as nested within the Splachnoideae. Tile phylogenetic significance of sporophytic characters within the family Splachnaceae is briefly discussed.