Bryophytes are an important component of plant diversity, may be found from sea level to mountaintops, and are particularly conspicuous on the Azores islands. These plants rely on environmental water, which acquires intercepting rain and dew (liquid water) and uses fog (water vapor), and transports both externally, by capillary forces, and internally, in different cells (specialized or not). This study characterizes and quantifies the ability of six liverworts and eight mosses to retain water, through different pathways, and to lose water by evaporation. Twelve replicates of each species were collected in Azorean native vegetation during the summer of 2016. The absolute water content (AWC) was obtained through measurements of specimens saturated, without free water, and completely dry. Most of the 14-target species showed an ectohydric behavior pattern retaining more than 60% of water through gametophyte surface. The AWC value ranged from 646% in Polytrichum commune to 5584% in Sphagnum subnitens. The water loss by direct evaporation showed, for most of species, an exponential decay curve along time. Understanding how much native bryophytes, acquire, store, and release water into the system contributes not only to the knowledge of native vegetation resilience but also to potential impacts on the availability and quality of water—a major ecosystem service performed by bryophytes.

Two peat moss species, frequent both in the mainland Portugal and in the Azores archipelago, were evaluated for essential oil composition and establishment under in vitro culture. Sphagnum auriculatum and Sphagnum subnitens essential oils were isolated by hydrodistillation and analysed by Gas Chromatography (GC) and Gas Chromatography-Mass Spectrometry (GC-MS). The essential oil of S. auriculatum was dominated by an as yet unidentified sesquiterpene, whereas zierene was the main component of S. subnitens essential oil. The in vitro cultures were successfully established for future studies of their chemical profile. The components present in essential oils obtained from S. auriculatum and S. subnitens together with morphological traits could be used to support the taxonomy of this plant group.

Peat mosses are a key functional group in peatla nds, driving biogeochemical cycles, habitat development and changes in species composition. They are generally intolerant of calcium and magnesium bicarbonate, but some species are ada pted to mineral-rich fens. A previous study found a coincidence between genetic variation and the ability to tolerate high pH/calcium levels in Sphagnum warnstorfii . Here we compare its microsatellite variation with that of two rarer cal- cium-tolerant species ( Sphagnum subnitens , S. contortum ), using a novel data set from Eurasia. Because physiological experiments indicate that S. warnstorfii can tolerate high magnesium lev- els, we included also samples from dolomite and serpentinite. Genetic diversity of S. warnstorfii was higher than that of other species. The Bayesian analysis in program Structure resulted in two population groups of S. warnstorfii . One group coincided with dolomite (Italy, Austria, Estonia) and moderately magnesium-rich (but calcium-poor) r ocks (serpentinite, metadolerite, cordierite- bearing migmatite on the Bohemian Massif), wh ile the second one coincided with magnesium- poor bedrock across Eurasia. The principal coor dinate analysis revealed a cline between popula - tions from magnesium-rich and magnesium-poor bedrocks, with populations from dolomite and serpentinite forming one extreme. Populations from magnesium-poor bedrock located far from any dolomite or serpentinite formed the opposite ex treme of the cline. We demonstrate for the first time that magnesium toxicity may drive bryophyte microevolution, as has repeatedly been shown for vascular plants, including ferns.

As builders and major components of peatlands, Sphagnopsida (peat mosses) are very important organisms for ecosystems and world’s climate. Nowadays many Sphagnum species as well as their habitats are largely protected, while their scientific and economic relevance remains considerable. Advanced methods of in vitro cultivation provide the potential to work in a sustainable way with peat mosses and address aspects of basic research as well as biotechnological and economical topics like biomonitoring or the production of renewable substrates for horticulture (Sphagnum farming). Here, we describe the establishment of axenic in vitro cultures of the five peat moss species Sphagnum fimbriatum Wils. and Hook., Sphagnum magellanicum Brid., Sphagnum palustre L., Sphagnum rubellum Wils. and Sphagnum subnitens Russ. and Warnst. with specific focus on large-scale cultivation of S. palustre in bioreactors. Axenic, clonal cultures were established to produce high quantities of biomass under standardized laboratory conditions. For advanced production of S. palustre we tested different cultivation techniques, growth media and inocula, and analyzed the effects of tissue disruption. While cultivation on solid medium is suitable for long term storage, submerse cultivation in liquid medium yielded highest amounts of biomass. By addition of sucrose and ammonium nitrate we were able to increase the biomass by around 10- to 30-fold within 4 weeks. The morphology of in vitro-cultivated gametophores showed similar phenotypic characteristics compared to material from the field. Thus the tested culture techniques are suitable to produce S. palustre material for basic and applied research.Electronic supplementary materialThe online version of this article (doi:10.1007/s11240-014-0658-2) contains supplementary material, which is available to authorized users.

Abstract QuestionHow did frequency and local abundance of key bryophytes in Dutch fens change between 1940 and 2000?LocationThe Netherlands.MethodsA total of 1939 vegetation relevés, objectively assigned to the class Parvocaricetea and dating from the period 1940–1999 inclusive, were extracted from a national Netherlands database. From this relevé set, two additional data sets were derived; these were syntaxonomically homogeneous at the association level. One contained 418 Scorpidio‐Caricetum diandrae relevés and the other 956 Pallavicinio‐Sphagnetum relevés. The following analyses were done for these three data sets: frequency and local abundance change of 13 bryophyte species over three 20‐yr periods were described using logistic regression and the nonparametric Jonckheere–Terpstra test, respectively.ResultsAcidophytic, late‐successional species, notably Sphagnum palustre and Polytrichum uliginosum, have replaced relatively early‐successional species such as brown mosses, Sphagnum subnitens and Sphagnum squarrosum during the studied period. The results were mostly consistent across the three data sets. Local abundance patterns generally paralleled those of frequency, although cover trends were often less robust.ConclusionsDrastic changes have taken place in the bryophyte layer of Dutch fens between 1940 and 2000. Our study has shown that acidophytic, late‐successional bryophytes have replaced relatively early‐successional brown moss and Sphagnum species. Both hydrological changes and nutrient shifts (in particular phosphorus enrichment and ammonium toxicity) appear to have been responsible for the observed rapid shifts in the bryophyte layer of Dutch fens.

Sphagnum subnitens var. nitidum newly added to the Korean Peninsula

The monoicous peatmoss Sphagnum subnitens has a tripartite distribution that includes disjunct population systems in Europe (including the Azores), northwestern North America and New Zealand. Regional genetic diversity was highest in European S.subnitens but in northwestern North America, a single microsatellite-based multilocus haploid genotype was detected across 16 sites ranging from Coos County, Oregon, to Kavalga Island in the Western Aleutians (a distance of some 4115 km). Two multilocus haploid genotypes were detected across 14 sites on South Island, New Zealand. The microsatellite-based regional genetic diversity detected in New Zealand and North American S.subnitens is the lowest reported for any Sphagnum. The low genetic diversity detected in both of these regions most likely resulted from a founder event associated with vegetative propagation and complete selfing, with one founding haploid plant in northwest North America and two in New Zealand. Thus, one plant appears to have contributed 100% of the gene pool for the population systems of S.subnitens occurring in northwest North America, and this is arguably the most genetically uniform group of plants having a widespread distribution yet detected. Although having a distribution spanning 12.5° of latitude and 56° of longitude, there was no evidence of any genetic diversification in S. subnitens in northwest North America. No genetic structure was detected among the three regions, and it appears that European plants of S.subnitens provided the source for New Zealand and northwest North American populations.

Mosses of the planned national park “Tulos” were studied in 1997, 2004–2006. In total 150 moss species and 1 variety were recorded. Comments on ecology and distribution of all the species, and label data for rare mosses are given. Sixty six moss species and 1 variety are recorded as new to the planned national park “Tulos”, and 11 species as new to Kuitozero-Leksozero floristic district. Three species (Neckera pennata, Pseudotaxiphyllum elegans and Sphagnum auriculatum) are listed in the Red Data Book of Karelian Republic, and 4 species (Neckera pennata, Polytrichastrum pallidisetum, Sphagnum subnitens, S. inundatum) are in the Red Data Book of Finland.

Summary This study illustrates the results of a floristic and phytosociological study conducted in a small portion (375 ha) of the Merse valley. Following a brief description of the geomorphologic and climatic aspects, the bryological and vascular flora is presented, obtained through floristic field studies and bibliographic data. The bryological flora includes 25 moss species including Sphagnum subnitens and S. capillifolium, which are of interest from a conservationist viewpoint, and 5 liverworts; the vascular flora comprises 252 species (belonging to 68 families and 186 genera); there are only three endemic entities but many species considered as rare or of interest, including Anagallis minima, Blechnum spicant, Carex viridula, Dryopteris dilatata, Erythronium dens-canis, Juncus bulbosus, Osmunda regalis and Radiola linoides, thus highlighting the phytogeographical relevance of this biotope. Chorological analysis highlighted the dominance of eurosiberian and boreal elements. The principal vegetation types sampled can be ascribed to the following alliances: Cicendio filiformis-Solenopsion laurentiae (Isoetetalia, Isoeto-Nanojuncetea), Osmundo-Alnion and Alnion incanae (Populetalia albae, Querco-Fagetea), Erythronio dens-canis-Quercion petraeae and Teucrio siculi-Quercion cerridis (Quercetalia-pubescen- ti-petraeae, Querco-Fagetea), Sarothamnion scoparii (Cytisetalia scopario-striati, Cytisetea scopario-striati). There are three habitats of Community interest: alluvial forests with Alnus glutinosa (code 91EO), woods of Castanea sativa (code 9260) and rare short-lived small coenoses of Isoeto-Nanojuncetea (code 3130).

Abstract Question: Why is bryophyte succession in eutrophicated fens faster than in natural fens?Location: Mineral‐rich fens in The Netherlands and NW Europe.Methods: Literature review on the ecology of four bryophyte species in various successional types as observed in Dutch fens.Results: Bryophyte succession in eutrophicated fens from the brown moss Calliergonella cuspidata to Sphagnum squarrosum is much faster than in natural fens with species shifts from Scorpidium scorpioides to Sphagnum subnitens. Under P‐poor conditions, the brown moss stage is stabilized as long as mineral‐rich water is supplied. This is because S. scorpioides is tolerant of rainwater, is a strong competitor and can counteract acidification to some extent while S. subnitens is intolerant to groundwater and has low growth rates and low acidification capacity. In contrast, the Sphagnum stage is stable after rapid succession from rich‐fen mosses under P‐rich conditions. Calliergonella cuspidata has suboptimal growth in rainwater, possibly due to ammonium toxicity, while the high growth rates of S. squarrosum in nutrient‐rich and highly acidic groundwater allow early establishment and rapid expansion.Conclusions: If measures to improve fen base status occur in environments of increased nutrient (P) availability, the management may not lead to the desired restoration of brown moss stages, but instead to rapid acidification by S. squarrosum.

Question: Why is bryophyte succession in eutrophicated fens faster than in natural fens? Location: Mineral-rich fens in The Netherlands and NW Europe. Methods: Literature review on the ecology of four bryophyte species in various successional types as observed in Dutch fens. Results: Bryophyte succession in eutrophicated fens from the brown moss Calliergonella cuspidata to Sphagnum squarrosum is much faster than in natural fens with species shifts from Scorpidium scorpioides to Sphagnum subnitens. Under P-poor conditions, the brown moss stage is stabilized as long as mineral-rich water is supplied. This is because S. scorpioides is tolerant of rainwater, is a strong competitor and can counteract acidification to some extent while S. subnitens is intolerant to groundwater and has low growth rates and low acidification capacity. In contrast, the Sphagnum stage is stable after rapid succession from rich-fen mosses under P-rich conditions. Calliergonella cuspidata has suboptimal growth in rainwater, possibly due to ammonium toxicity, while the high growth rates of S. squarrosum in nutrient-rich and highly acidic groundwater allow early establishment and rapid expansion. Conclusions: If measures to improve fen base status occur in environments of increased nutrient (P) availability, the management may not lead to the desired restoration of brown moss stages, but instead to rapid acidification by S. squarrosum.

Over half of the world's peat originated from Sphagnum, representing 10–15% of the terrestrial carbon stock. However, information regarding the release and exudation of organic carbon by living Sphagnum plants into the surface peat is scarce. In this study, we examined the contribution of recent Sphagnum subnitens (Russ. and Warnst.) photosynthate carbon to the peatland dissolved organic carbon (DOC) pool. This was done using a 13CO2 pulse-chase experimental approach during the growing season. Despite the importance of Sphagnum in long-term carbon accumulation, results showed that the Sphagnum community rapidly contributes recently synthesized carbon to the peatland DOC pool. We estimate that by 4 h up to 4% of the total DOC in peat leachate was derived from 13CO2 pulse labelling at ambient CO2 concentrations. Nonetheless, a huge 64% of the 13C initially assimilated by photosynthesis was retained in Sphagnum subnitens for 23 days after labelling, consistent with the role of Sphagnum in peatland carbon accumulation. The majority of 13C loss as respired CO2 came within the few days post 13CO2 labelling, suggesting that it was derived from plant respiration of photosynthates.

Summary Sphagnum subnitens Russ. & Warnst, and Sphagnum subsecundum Nees are reported from a new site in Tuscany. The two species were found in a recently described Betula pendula stand, located in the Belagaio forest, Colline Metallifere, southern Tuscany. Fifty-two bryological taxa (19 liverworts and 33 mosses) were recorded in this area. The forest of Belagaio is the southernmost site known for Sphagnum subnitens and Sphagnum subsecundum in Tuscany. Plagiomnium ellipticum is another new species for Tuscany.

Detailed floristic records from 12 mires on three Suffolk fens were gathered in 1959 and 1991. The data were analysed using tabulation and ordination to elucidate change at the plant community and individual species levels. Substantial changes at both levels were observed. The respective roles of three factors, namely lack of traditional management, changes in water regime and increased fertility, in accounting for these changes, have been explored. At community level, those mires which showed the greatest alteration were those subject to hydrological disturbance combined with dereliction. In terms of the British National Vegetation Classification they changed from short sedge mires allied to M13 Schoeno-Juncetum towards highly fertile tall herb fens with many ruderals and allied to S26 Phragmites australis-Urtica dioica fen. Partial dewatering led to M27 Filipendula ulmaria-Angelica sylvestris tall herb fen with few non-fen ruderals. Mires where ground water influx was reduced or eliminated but where traditional management was sustained showed less community change, with more of the principle community dominants being maintained. Here M13 Scheono-Juncetum mires developed into dry variants of M24 Molinia caerulea-Cirsium dissectum fen meadows. Mires with undisturbed hydrology but which became derelict were transformed from M13 Schoeno-Juncetum into hydrophytic Cladium-dominated communities of the S25 Phragmites australis-Eupatorium cannabinum tall herb fens. Many fen species were retained or recruited, while the colonization of ruderals, scrub and non-fen species that was noted in the dewatered mires did not occur. The dewatered mires showed the greatest alteration at species level. In unmanaged areas, all of the fen associates and bryophytes were lost and replaced by non-fen species and ruderal tall herbs. Where traditional management was maintained, most of the small species characteristic of wet fens and semi-aquatic conditions were still lost, although recruitment of ruderals and non-fen plants was less evident, being restricted to species of mesotrophic meadows. Retained species tended to be common fen plants or non-fen species. Mires which were derelict, but where the natural hydrology persisted, showed the least change in associated flora. Nevertheless, many of the smaller herbs were still eliminated by competition in the now dense tall-herb fen, indicating that traditional management was required as well as hydrological integrity for the survival of most wet-fen species. However, some wet-fen species were retained or recruited, indicating management to be subordinate to hydrology in their maintenance. They include Caltha palustris, Valeriana dioica, Lotus uliginosus and the bryophytes Campylium stellatum, Aneura pinguis, Sphagnum subnitens and Riccardia multifida. All mires showed a decline in conservation importance as measured by indices of botanical 'quality'. The mires which showed least decline (15-64% reduction in Rarity Weighted Principal Fen Species Score) were those which retained their hydrological integrity, regardless of management status. One mire, only partially dewatered but still managed, declined by nearly 70%. Although there were problems with the data, tentative comments are made on the relative importance of the three environmental factors to the maintenance of mire systems. It is concluded that maintenance of stable hydrologies is more important than extensive management in the long-term conservation of these sites.

1 Four bryophyte species, Scorpidium scorpioides (characteristic of mineral-rich, nutrient-poor fens), Sphagnum subnitens (mineral-poor, nutrient-poor), Calliergonella cuspidata (mineral-rich, nutrient-rich) and Sphagnum squarrosum (mineral-poor, nutrient-rich), were cultivated to study whether their mutual replacement was regulated by abiotic conditions or whether interspecific interactions played a major role. 2 Water type may contribute to the replacement of some rich-fen species by Sphagnum spp. Growth of Calliergonella cuspidata was best in (mineral-rich) groundwater and reduced in (mineral-poor) rain water, whereas growth of Sphagnum subnitens was much better in rain water than in groundwater. Scorpidium scorpioides and Sphagnum squarrosum, however, performed well in both water types. 3 Nutrient enrichment may also explain species replacements to some extent. As expected, Sphagnum squarrosum was clearly stimulated by nutrient supply, while S. subnitens did not respond. However, S. scorpioides showed increased rather than decreased growth rates and C. cuspidata was not affected by nutrient supply. 4 Interspecific interactions played a role in all replacement series. Generally, mixed culture stimulated growth of the species whose natural environment was simulated and/or reduced growth of the other species, suggesting that competitive interactions are involved. However, interactions between S. scorpioides and S. subnitens appear to counteract the observed succession. 5 The different species responses may lead to different rates of succession in nutrientpoor ('slow succession') and nutrient-rich fens ('rapid succession').

The acidification capacity of wetland bryophytes as influenced by simulated clean and polluted rain

Abstract In Dutch rich fens, mid-successional stages dominated by Sphagnum subnitens are rapidly being replaced by other vegetation, dominated by S. fallax. In both species growth was strongly reduced in more mineral-rich ground water and simulated river water, compared to growth in rain water.Both species were positively affected by added phosphorus supply, whereas added supply of nitrogen did not alter growth rates.Nutrient supply seemed to compensate for the intolerance to high mineral levels in S. fallax: nutrient addition stimulated growth in length in ground water and simulated river water. Such compensation was not found in S. subnitens. Growth in mixed culture positively affected Sphagnum subnitens and negatively affected its successor S. fallax. The success of Sphagnum fallax in early stages of succession in more eutrophic fens may to some extent be explained by its tolerance to high mineral levels with increased nutrient supply.

The blue-grey sheen of dried specimens of Sphagnum subnitens is attributed to thin film interference in the walls of the hyaline cells, this being produced by reflection from upper and lower film surfaces. The reflectance is large in air and small in water, so that the colour is visible only in dried specimens. S. rubellurn lacks the blue-grey sheen because only a small part of the total area of the hyaline cells is thin enough to generate interference colour.

Abstract Sphagnum subnitens, S. squarrosum, and Drepanocladus revolvens are recorded from New Zealand for the first time, all three occurring in mires. Both Sphagnum species are apparently rare but Drepanocladus revolvens is widespread and probably overlooked.