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Dicranum scoparium , examples of habit variation (moist). a – plants with straight leaves from pine heath forest; b – plants with falcate leaves from swampy forest; c – large plants with falcate leaves from high-altitude laurel forest on Madeira. 

Dicranum scoparium , examples of habit variation (moist). a – plants with straight leaves from pine heath forest; b – plants with falcate leaves from swampy forest; c – large plants with falcate leaves from high-altitude laurel forest on Madeira. 

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HEDENÄS, L. & BISANG, I. 2004. Key to European Dicranum species. - Herzogia 17: 179-197. A key to the twenty-nine European Dicranum species is presented. The features that are important for their identifi- cation are discussed. For each species, diagnostic character states and information about habitat and geographical distribution are provided. Mo...

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... scoparium Hedw. ( D. tectorum Warnst. & H. Klinggr.) D. scoparium belongs to the medium-sized species of the genus. In some respects it is one of the most variable Dicranum species (fig. 46); both phenotypes with straight and falcate leaves are frequent, and both dwarf and large male plants occur. The species has prosenchymatous upper leaf lamina cells, and has in addition (usually four) distinct lamellae on the back of the upper costa. Differences between D. scoparium, D. crassifolium , D. transsylvanicum , and poorly developed plants of D. majus are discussed under the latter species. Plants from extreme dry or Arctic environments may lack dorsal costal lamellae completely, and could therefore be confused with D. bonjeanii . However, in D. scoparium the lower costa is strong, dorsally convex, and has several stereid bands, whereas that of D. bonjeanii is weak, almost flat, and has fewer stereid bands (N YHOLM 1987, F RISVOLL & E LVEBAKK 1996). Widespread in Europe. – Found on various substrates, on soil, rock, rotten wood, tree stems etc., in both dry and humid ...

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... Dicranum Hedw. is one of the most diverse genera within the Dicranaceae, having approximately 93 accepted taxa to date (Brinda and Atwood 2023). Although numerous species are common in arcticmontane, peatland, and other boreal and arctic ecosystems, knowledge about this genus remains inadequate, and the delimitation of morphological species is perplexing due to significant variation in gametophytic characters influenced by environmental conditions (Hedenäs and Bisang 2004;Ireland 2007;Lang and Stech 2014;Ignatova et al. 2015). Recent advances in molecular phylogeny have led to a more definitive delimitation of some Dicranum species, such as those with fragile leaves (Ignatova and Fedosov 2008;Huang et al. 2023), D. fuscescens Turner complexes (Tubanova et al. 2010;Tubanova and Ignatova 2011), D. scoparium Hedw. ...
... In Shennongjia National Nature Reserve, Hubei Province, China, we discovered an intriguing species undoubtedly belonging to Dicranum, with bistratose distal laminal cells similar to those of D. crassifolium Sérgio, D. majus var. majus Turner and D. transsylvanicum Lüth (Sérgio et al. 1995;Lüth 2002;Ignatov and Ignatova 2003;Hedenäs and Bisang 2004;Hedenäs et al. 2006;Ireland 2007;Lüth 2019). However, a combination of several features, such as only one layer of guide cells in the costa, distal laminal cells with strongly protruding upper angles on the dorsal side, and a lack of lamellae on the dorsal side of the costa above the midleaf, distinguish the plants from the above-mentioned species. ...
... The Dicranum specimens from Shennongjia National Nature Reserve share morphological characteristics with D. crassifolium, D. majus var. majus and D. transsylvanicum, such as bistratose distal laminal cells and prosenchymatous laminal cells in the upper leaf portion (Sérgio et al. 1995;Lüth 2002;Ignatov and Ignatova 2003;Hedenäs and Bisang 2004;Hedenäs et al. 2006;Ireland 2007;Lüth 2019). However, D. crassifolium and D. transsylvanicum present a costa with distinct lamellae on the back (Sérgio et al. 1995;Lüth 2002), whereas this studied species only has sharp serrations on the dorsal side of the costa (see Figure 3B, F). ...
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Introduction. Dicranum Hedw. is one of the most taxonomically challenging genera of mosses, having 93 accepted species worldwide. Although recent integrative taxonomic revisions have clarified the delimitation of most species within this genus, the circumscriptions of some Dicranum species remain unsolved. During our recent survey of Dicranum in China, an intriguing species with an unknown combination of morphological traits was discovered in the Shennongjia National Nature Reserve, Central China. We carried out a study to provide morphological and phylogenetic evidence with which to determine the taxonomic status of this Chinese species. Methods. The phylogenetic evidence was obtained through the combined analysis of five loci (rps4-trnT, trnL-trnF, trnH-psbA, rps19-rpl2 and rpoB) and one nuclear region (nrITS1-5.8S-ITS2). Morphological characteristics were investigated by microscopic examination of specimens. Key results and conclusions. A new moss species, Dicranum shennongjiaense W.Z.Huang & R.L.Zhu, is described and illustrated based on molecular and morphological evidence. This new species is characterised by its falcate leaves, rectangular and elongated laminal cells with strongly protruding upper angles on the dorsal side in the upper leaf portion, irregularly bistratose laminal cells above the midleaf, one layer of guide cells in the costa, and sharp serrations on the dorsal side of the costa in the upper leaf portion. Additionally, D. baicalense Tubanova, previously known only from Russia, is newly reported for China. A key for identification of the Chinese species of Dicranum sect. Dicranum is provided.
... Until recently, reliable records for D. acutifolium from the Alps were very sparse and its occurrence was not accepted in Grims (1999) for Austria and in Meinunger and Schröder (2007) for Germany. Only recently Hedenäs and Bisang (2004) or Schlüsslmayr (2019) seriously reported its presence in the Eastern Alps and so far, only one Central European sample of the complex from Switzerland (belonging to D. brevifolium) and one from Austria (belonging to D. septentrionale) have been molecularly examined (Lang et al. 2014). ...
... In case of overlapping characteristics or discordance between the characters, the determinator had the option to assign multiple taxon names, starting with the best fit taxon name. (b) We compiled a data set of 45 morphological traits comprising the characters presented by Tubanova et al. (2010) and Lang et al. (2014) and further characters commonly used for species distinction in the genus (see e.g., Hedenäs and Bisang 2004) and analysed it using multivariate statistics. (i) We measured length and width of 5 leaves of one well-grown shoot of each specimen and estimated distal dorsal costa and upper leaf margin ornamentation and the shape of the base of these leaves. ...
... revealed only a few specimens from siliceous bedrock, suggesting that D. brevifolium is rare and that it should be prioritised in conservation. The same may apply to Scandinavia and Europe in general because D. brevifolium s.l. is normally given as basiphilous (Hedenäs and Bisang 2004;Nyholm 1987). Dicranum acutifolium seemingly occurs scattered in the limestone ranges of both Austria and Switzerland (Köckinger et al. 2021, Köckinger in litt). ...
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There is mounting evidence that reproductively isolated, but morphologically weakly differentiated species (so-called cryptic species) represent a substantial part of biological diversity, especially in bryophytes. We assessed the evolutionary history and ecological differentiation of a species pair, Dicranum brevifolium and D. septentrionale , which have overlapping ranges in the Holarctic. Despite their morphological similarity, we found similar genetic differentiation as between morphologically well-differentiated Dicranum species. Moreover, we detected gene tree discordance between plastid and nuclear markers, but neither of the two datasets resolved the two as sister species. The signal in trnL–trnF better reflects the morphological and ecological affinities and indicates a close relationship while ITS sequence data resolved the two taxa as phylogenetically distantly related. The discordance is probably unrelated to the ecological differentiation of D. septentrionale to colonise subneutral to alkaline substrates (vs. acidic in D. brevifolium ), because this ability is rare in the genus and shared with D. acutifolium . This taxon is the closest relative of D. septentrionale according to the trnL–trnF data and does not share the discordance in ITS. We furthermore demonstrate that beside D. acutifolium , both D. septentrionale and D. brevifolium occur in the Alps but D. brevifolium is most likely rarer. Based on morphological analyses including factor analysis for mixed data of 45 traits we suggest treating the latter two as near-cryptic species and we recommend verifying morphological determinations molecularly.
... The main subject of the study is D. viride (Fig. 1). This moss is a subcontinental-mountain species known from many areas in the Northern Hemisphere (central and northern Europe, the Caucasus, northern and eastern Asia, eastern parts of the North America) (Düll and Meinunger, 1989;Hedenäs and Bisang, 2004;Baisheva et al., 2013). Dicranum viride is an epiphyte that grows on the trunks of deciduous trees (mainly Fagus sp.) and facultatively as an epixylic species, very rarely found on rocks and soil (Barkman, 1958;Ellenberg et al., 1992;Köckinger et al., 2021). ...
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Functional diversity is widely recognised as the key to understanding the role of ecological mechanisms in shaping the patterns of species coexistence across different environmental gradients. Despite research in this area, there is a considerable knowledge gap on the ecological mechanisms that shape the composition of bryophyte communities, especially those at risk. One of the most endangered group of species are usually woodland specialist bryophytes, therefore, understanding the factors determining their occurrence should be a priority. One of the model species belonging to this group is Dicranum viride, a target species of international conservation importance, and of particular interest for forest managers responsible for forest ecosystems protection. We examined the effect of the tree phorophyte species on the realised niche of D. viride in the temperate forest ecosystems. The specific tree species that D. viride inhabits greatly influences its realised niche. Both phorophyte species we examined, i.e. Fagus sylvatica and Carpinus betulus, can sustain the presence of D. viride, but the key determinant is the availability of suitable microhabitats, such as specific combinations of bryophyte species within bark irregularities. Our findings indicate that even in regions where the effects of forest management are noticeable, the preservation of trees with diverse microhabitats facilitate the existence of rare moss species with D. viride as an example. As a moss with a narrow ecological amplitude, D. viride occurrence patterns can be used as a tool in monitoring studies and for improving forest management strategies aimed at harmonising wood production with the conservation of biodiversity and ecosystem functioning. As a result, this information can be significant for conservation strategies, as it highlights the importance of identifying and preserving certain tree species to protect both particular epiphyte communities and individual target species.
... E, at an altitude of 1931 m, on October 26, 2018, in Trabzon Province of Turkey. After laboratory diagnosis (Hedenäs and Bisang 2004;Ros et al. 2013;Özdemir and Batan 2017;Erata and Batan 2020), it was dried under suitable conditions and stored in the refrigerator until use so as not to spoil the substance's contents. One of the co-authors identified the plant material (Prof. ...
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Recent research shows that Dicranum species can be used to ameliorate the negative effects of honeybee bacterial diseases and that novel compounds isolated from these species may have the potential to treat bacterial diseases. This study aimed to investigate the efficacy of Dicranum polysetum Sw. against American Foulbrood using toxicity and larval model. The effectiveness of D. polysetum Sw. ethanol extract in combating AFB was investigated in vitro and in vivo. This study is important in finding an alternative treatment or prophylactic method to prevent American Foulbrood disease in honey bee colonies. Spore and vegetative forms of Paenibacillus larvae PB31B with ethanol extract of D. polysetum were tested on 2040 honey bee larvae under controlled conditions. Total phenolic and flavonoid contents of D. polysetum ethanol extracts were determined as 80.72 mg/GAE(Gallic acid equivalent) and 303.20 µg/mL, respectively. DPPH(2,2-diphenyl-1-picrylhydrazyl) radical scavenging percent inhibition value was calculated as 4.32%. In Spodoptera frugiperda (Sf9) and Lymantria dispar (LD652) cell lines, the cytotoxic activities of D. polysetum extract were below 20% at 50 µg/mL. The extract was shown to considerably decrease infection in the larvae, and the infection was clinically halted when the extract was administered during the first 24 h after spore contamination. The fact that the extract contains potent antimicrobial/antioxidant activity does not reduce larval viability and live weight, and does not interact with royal jelly is a promising development, particularly regarding its use to treat early-stage AFB infection.
... The comparison with species from this section occurring in Russia and nearby areas is presented in Table 1. Our observation on the studied material is supplemented by published data (Savicz-Ljubitzkaja & Smirnova, 1970;Gao et al., 1999;Ignatov & Ignatova, 2003;Hedenäs & Bisang, 2004;Ireland, 2007;Lang & Stech, 2014;Lüth, 2002;Noguchi & Iwatsuki, 1987;Otnyukova, 2001;Price & Maier, 2013;Sérgio at al., 1995;Tubanova & Afonina, 2016). There is also a distinction in sporophyte characters between these species: capsules of D. baicalense are arcuate, while they are straight in D. orthophyllum. ...
... Дослідження проводилися в період 2019-2022 років в межах Закарпатської області України, переважно на Вулканічному хребті Українських Карпат. Для визначення видів мохоподібних використовувалися різні визначники [MELNICHUK, 1970;HEDENÄS, BISANG, 2004]. Для з'ясування їх поширення в Україні використовували національні та регіональні бріологічні зведення й окремі публікації [MELNICHUK, 1970;ZEROV, PARTYKA, 1975;BOIKO, 2010aBOIKO, ,b, 2014BOIKO, , 2019. ...
... Dicranum scoparium is a relatively long-lived and fairly conservative plant species belonging to the Bryophyta taxon of the Dicranaceae family, and members of the wind-blown moss Dicranum are observed worldwide. Twenty-nine species are recognized in Europe, [22] and fourteen species of Dicranum have been reported from Turkey. [23] Dicranium species usually grow in upland rocky wooded areas, although it also occurs on the wet, shaded ground where competition from other plants is reduced. ...
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Bacterial diseases, such as American Foulbrood (AFB) and European Foulbrood (EFB), are known to have catastrophic effects on honey bees (if left to spread, can wipe out entire colonies), leading to severe financial losses in the beekeeping industry. The aim of this study was to evaluate the pharmacological properties of methanol extract and its fractions (ethyl acetate, n-hexane, water) derived from Dicranum scoparium Hedw., which could be utilized as a potential drug to prevent the bacterial diseases (AFB and EFB) affecting the honey bees. For this purpose, crude methanol extract and ethyl acetate/n-hexane/water fractions were prepared from the aerial part of D. scoparium , collected from Trabzon province. Bio-guided fractionation of the extract and its fractions led to the first-time isolation of five compounds. The structure of all compounds was elucidated by nuclear magnetic resonance (NMR) spectroscopy, ultraviolet (UV) spectral analysis, Fourier-transform infrared spectroscopy (FT-IR), liquid chromatography quadrupole time-of-flight mass spectroscopy (LC-QToF-MS), and by comparison of their NMR data with that of literature. The analysis of these compounds revealed significant antibacterial and sporicidal activities against bacteria causing larval diseases in honey bees. The antibacterial activity of these compounds ranged from 0.6 to 60 µg/mL against AFB and EFB causing bacteria. Therefore, the natural raw extract and fractions of D. scoparium could be used as potential therapeutic agents against bacterial agents affecting honey bees.
... Hal. is a moss that is most frequent in pine-dominated forests with rocky ground and moraine with relatively dry and acidic conditions. Its upright shoots build loose homogenous tuffs, generally surrounded by the feather moss Pleurozium schreberi, other Dicranum species and different Cladonia species (Hedenäs & Bisang, 2004). ...
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Species at their warm range margin are potentially threatened by higher temperatures, but may persist in microrefugia. Whether such microsites occur due to more suitable microclimate or due to lower biotic pressure from, for example competitive species, is still not fully resolved. We examined whether boreal bryophytes and lichens show signs of direct climate limitation, that is whether they perform better in cold and/or humid microclimates at their warm range margin. We transplanted a moss, a liverwort and a lichen to 58 boreal forest sites with different microclimates at the species' southern range margin in central Sweden. Species were grown in garden soil patches to control the effects of competitive exclusion and soil quality. We followed the transplanted species over three growing seasons (2016–2018) and modelled growth and vitality for each species as a function of subcanopy temperature, soil moisture, air humidity and forest type. In 2018, we also recorded the cover of other plants having recolonized the garden soil patches and modelled this potential future competition with the same environmental variables plus litter. Species performance increased with warmer temperatures, which was often conditional on high soil moisture, and at sites with more conifers. Soil moisture had a positive effect, especially on the moss in the last year 2018, when the growing season was exceptionally hot and dry. The lichen was mostly affected by gastropod grazing. Recolonization of other plants was also faster at warmer and moister sites. The results indicate that competition, herbivory, shading leaf litter and water scarcity might be more important than the direct effects of temperature for performance at the species' warm range margin. Synthesis. In a transplant experiment with three boreal understorey species, we did not find signs of direct temperature limitation towards the south. Forest microrefugia, that is habitats where these species could persist regional warming, may instead be sites with fewer competitors and enemies, and with sufficient moisture and more conifers in the overstorey.
... Dicranum scoparium is widely distributed in the Holarctic and is growing on various substrates, preferably on acidic soil, rotten logs, tree stems and sometimes in sympatry with other Dicranum species (Crum and Anderson, 1981;Hedenäs and Bisang, 2004;Ireland, 2007;Lang et al., 2014). Both normal-sized and dwarf males can occur simultaneously in a same cushion. ...
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Among plants, gender dimorphism occurs in about 10% of all angiosperms and more than 50% of all moss taxa, with dwarf males (DM) found exclusively in some unisexual mosses. In this study, we explore the role of male dwarfism as a reproductive strategy in the widespread acrocarpous moss Dicranum scoparium, which has facultative male dwarfism, having both dwarf males (DMs) and normal-sized males (NMs). We retrieved 119 SNP markers from transcriptomes which were used to genotype 403 samples from 11 sites at seven localities in southern Sweden. Our aims were to compare the genetic variability and genetic structure of sexually reproducing populations at different geographic levels (cushion, site, and locality) and compare in particular the relative contribution of females, dwarf males and normal-sized males to the observed genetic diversity. The numbers of DMs differed strongly between sites, but when present, they usually outnumbered both females and NMs. Low genetic differentiation was found at locality level. Genetic differentiation was strongest between cushions for females and NMs and within cushions for DMs indicating small scale structuring and sometimes inbreeding. NMs were more clonal than either DMs or females. Genetic diversity was similar between females and DMs, but lower for NMs. Two haplotypes were shared between females and DMs and one haplotype was shared between a DM and a NM. In conclusion, our results show that DMs and NMs play different roles in reproduction, inbreeding may occur at cushion level, but gene flow is high enough to prevent substantial genetic drift.
... ex Brid. (Dicranales: Dicranaceae), until recently known commonly as D. bergeri Bland., is a boreo-arctic montane circumpolar species, occurring in Asia, Europe and North America (Hill and Preston 1998;Chien and Crosby 1999;Hedenäs and Bisang 2004;Hallingbäck et al. 2006;FNA Editorial Committee 2007;Blockeel et al. 2014). It typically occupies peatlands, including treeless ombrotrophic bogs, rich fens, wooded bogs, dry pine heath forest and subalpine shrub-tundra (Birks 1977;Crum and Anderson 1981;Foster 1984;Luken 1985;Anderson et al. 1995;Belland and Vitt 1995;Nicholson and Gignac 1995;Chien and Crosby 1999;Asada 2002;Hedenäs and Bisang 2004;Locky et al. 2005;Hallingbäck et al. 2006;FNA Editorial Committee 2007;Lockhart et al. 2012), tending to be more abundant in continental than oceanic climates (Gignac et al. 1991). ...
... (Dicranales: Dicranaceae), until recently known commonly as D. bergeri Bland., is a boreo-arctic montane circumpolar species, occurring in Asia, Europe and North America (Hill and Preston 1998;Chien and Crosby 1999;Hedenäs and Bisang 2004;Hallingbäck et al. 2006;FNA Editorial Committee 2007;Blockeel et al. 2014). It typically occupies peatlands, including treeless ombrotrophic bogs, rich fens, wooded bogs, dry pine heath forest and subalpine shrub-tundra (Birks 1977;Crum and Anderson 1981;Foster 1984;Luken 1985;Anderson et al. 1995;Belland and Vitt 1995;Nicholson and Gignac 1995;Chien and Crosby 1999;Asada 2002;Hedenäs and Bisang 2004;Locky et al. 2005;Hallingbäck et al. 2006;FNA Editorial Committee 2007;Lockhart et al. 2012), tending to be more abundant in continental than oceanic climates (Gignac et al. 1991). ...
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Introduction. Dicranum undulatum has been lost from many sites due to destruction of peatlands. This study investigates its status and ecology at the last surviving location in southern Britain. Methods. The study site, Fenn’s, Whixall, Bettisfield, Wem and Cadney Mosses, is one of the largest lowland raised bogs in Britain. Distribution and abundance of Dicranum undulatum was documented with GPS-based survey and habitat and community composition via relevés. Elevation of D. undulatum and Rest Water Level (RWL) relative to the peatland surface was measured at a sample of colonies and a search for sporophytes undertaken at all colonies. Results. Dicranum undulatum was found in two areas, the vast majority at Cranberry Beds plus a small amount at Welsh Bettisfield Moss. Its community is dominated by mosses and vascular plants, most frequently Aulacomnium palustre, Erica tetralix, Eriophorum vaginatum and Vaccinium oxycoccos. Soil pH is highly acidic, similar to the upper water table. Electrical conductivity of the upper water table is low. Elevation of RWL relative to soil surface averaged −10.8 cm. Minimum and maximum elevation of shoot tips of D. undulatum relative to soil surface averaged 7.4 and 24.8 cm, respectively. Maximum height of the moss above RWL averaged 35.6 cm. No sporophytes were found. Conclusions. Strictly limited to uncut peatland, the study site supports a strong population of Dicranum undulatum. It survives at Welsh Bettisfield Moss despite it being covered by dense woodland until tree clearance in 2001. Reproduction appears limited to clonal spread via detached shoot tips.