Sphagnum growth and ecophysiology during mire succession

Sphagnum mosses are widespread in areas where mires exist and constitute a globally important carbon sink. Their ecophysiology is known to be related to the water level, but very little is currently known about the successional trend in Sphagnum. We hypothesized that moss species follow the known vascular plant growth strategy along the successional gradient (i.e., decrease in production and maximal photosynthesis while succession proceeds). To address this hypothesis, we studied links between the growth and related ecophysiological processes of Sphagnum mosses from a time-since-initiation chronosequence of five wetlands. We quantified the rates of increase in biomass and length of different Sphagnum species in relation to their CO₂ assimilation rates, their photosynthetic light reaction efficiencies, and their physiological states, as measured by the chlorophyll fluorescence method. In agreement with our hypothesis, increase in biomass and CO₂ exchange rate of Sphagnum mosses decreased along the successional gradient, following the tactics of more intensely studied vascular plants. Mosses at the young and old ends of the chronosequence showed indications of downregulation, measured as a low ratio between variable and maximum fluorescence (F _v/F _m). Our study divided the species into three groups; pioneer species, hollow species, and ombrotrophic hummock formers. The pioneer species S. fimbriatum is a ruderal plant that occurred at the first sites along the chronosequence, which were characterized by low stress but high disturbance. Hollow species are competitive plants that occurred at sites with low stress and low disturbance (i.e., in the wet depressions in the middle and at the old end of the chronosequence). Ombrotrophic hummock species are stress-tolerant plants that occurred at sites with high stress and low disturbance (i.e., at the old end of the chronosequence). The three groups along the mire successional gradient appeared to be somewhat analogous to the three primary strategies suggested by Grime.     

Initiation of microtopography in revegetated cutover peatlands

Question: How many years are required for a gradient of microtopography to be initiated in revegetated cutover peatlands and become similar to natural bogs? Location: Newly formed Sphagnum carpets on cutover peatlands that revegetated spontaneously after site abandonment (in Estonia), or following active restoration (in Canada) and on undisturbed natural bogs nearby. Methods: Moss surface height was measured along linear transects above a local reference level (the lowest point for a given transect). Heights of at least 20 cm were associated with hummocks. Frequency distributions of surface height and principal component analyses (separately for Canada and Estonia) were conducted to follow the evolution of microtopography in revegetated sites and their similarity with those of natural peatlands. In Canada, regressions were also performed to estimate the time required for the microtopography in revegetated cutover peatlands to become similar to that found in natural bogs. Results: Only 10–30 yr were needed for microstructures comparable to those in natural bogs to develop on restored peatlands where Sphagnum diaspores have been reintroduced. However, this process may take more than a century in cutover peatlands left to revegetate spontaneously. Conclusions: In cutover peatlands with spontaneous revegetation, hummock–hollow formation starts on bare peat which lacks both plant propagules and viable seed banks, and the initiation of microstructures is probably more akin to the process that occurs naturally. Nonetheless, hummock–hollow microtopography resembling that found in natural bogs without pools appeared, in all of the examined cutover peatlands, over periods that are short in terms of peatland development time‐scales. Active peatland restoration could effectively reduce the time required for initiation of microtopography by about 70 yr.     

Photosynthetic traits of Sphagnum and feather moss species in undrained,drained and rewetted boreal spruce swamp forests

In restored peatlands, recovery of carbon assimilation by peat‐forming plants is a prerequisite for the recovery of ecosystem functioning. Restoration by rewetting may affect moss photosynthesis and respiration directly and/or through species successional turnover. To quantify the importance of the direct effects and the effects mediated by species change in boreal spruce swamp forests, we used a dual approach: (i) we measured successional changes in moss communities at 36 sites (nine undrained, nine drained, 18 rewetted) and (ii) photosynthetic properties of the dominant Sphagnum and feather mosses at nine of these sites (three undrained, three drained, three rewetted). Drainage and rewetting affected moss carbon assimilation mainly through species successional turnover. The species differed along a light‐adaptation gradient, which separated shade‐adapted feather mosses from Sphagnum mosses and Sphagnum girgensohnii from other Sphagna, and a productivity and moisture gradient, which separated Sphagnum riparium and Sphagnum girgensohnii from the less productive S. angustifolium, S. magellanicum and S. russowii. Undrained and drained sites harbored conservative, low‐production species: hummock‐Sphagna and feather mosses, respectively. Ditch creation and rewetting produced niches for species with opportunistic strategies and high carbon assimilation. The direct effects also caused higher photosynthetic productivity in ditches and in rewetted sites than in undrained and drained main sites.     

Facilitation vs. competition: Does interspecific interaction affect drought responses in Sphagnum?

The stress-gradient hypothesis (SGH) predicts that the relative importance of competition decreases and facilitation increases with an increase in abiotic stress. In peatlands, Sphagnum faces the threat of drought and differentiates into hummock species (drought-tolerant) and hollow species. Whether interspecific interaction affects the influence of drought on bryophyte composition in peatlands is unknown. We established an experiment by simulating drought and building bryophyte communities with two hummock species (S. palustre and S. capillifolium) and one hollow species (S. fallax). In all three species, drought decreased biomass production, height increment and side-shoot production. Sphagnum stores water in the hyaline cells, and leaf hyaline cell percentage (HCP) in the two hummock species increased with drought while no effect was found in S. fallax, suggesting that adjusting HCP is not an effective response to drought for the hollow species. Morphological traits and carbon and nitrogen contents in hummock species responded more to drought than in the hollow species, indicating a rapid response in phenotypic plasticity is an important strategy to resist drought in the hummock species. The presence of neighboring Sphagnum species, rather than drought, decreased carbon content for all three species. All three bryophytes showed interaction between drought and neighbor in two or more plant traits. Our study, however, did not support SGH, and there were no changes from competition under wet to facilitation under dry treatments in any of the six species combinations. On the contrary, when S. fallax was the target species, a change from facilitation under wet to competition under dry treatments was observed. The results suggest that hummock species can facilitate hollow species in wet environments but they could suppress hollow species under drought conditions by competing for water resources. Both drought and strong competition are the probable reasons why hollow species rarely grow in hummocks.     

Testate Amoeba (Arcellinida,Euglyphida) Ecology along a Poor‐Rich Gradient in Fens of Western Poland

We investigated the relationships between testate amoebae (Arcellinida, Euglyphida), vegetation and water chemistry along environmental gradients in minerotrophic peatlands (fens) in western Poland. We hypothesized that: a) hydrochemistry significantly influences structure of testate amoeba communities, and b) testate amoeba communities are more closely correlated with the hydrochemical variables (environment) than with the vegetation data. Testate amoeba communities and vegetation from 71 sample plots were investigated together with the hydro‐chemistry and hydrology based on 16 environmental variables and vegetation composition. Testate amoeba communities revealed a distinctive poor‐rich gradient in analysed fens. Mineral‐rich habitats, which were dominated by brown mosses, were preferred by a higher number of taxa than acidic habitats, which were dominated by Sphagnum. We recorded a total of 107 testate amoebae taxa. The average species richness of testate amoebae for brown mosses was higher (20) than for Sphagnum (13). We found that testate amoebae communities were similarly correlated with vascular plants, mosses and environmental parameters. Results of direct ordination demonstrate that hydrology, pH, Mg²⁺ and sodium remain the most important environmental control for the entire data set. CCA showed that in case of brown mosses hydrology, sodium and oxygen affect testate amoeba communities significantly whereas in Sphagnum only sodium emerge as most significant determining testate amoeba assemblages. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Biochemical determinants of litter quality in 15 species of <Emphasis Type="Italic">Sphagnum</Emphasis>

Background and aims

Sphagnum mosses are ecosystem engineers that create and maintain boreal peatlands. With unique biochemistry, waterlogging and acidifying capacities, they build up meters-thick layers of peat, reducing competition and impeding decomposition. We quantify within-genus differences in biochemical composition to make inferences about decay rates, related to hummock–hollow and fen–bog gradients and to phylogeny.

Methods

We sampled litter from 15 Sphagnum species, abundant over the whole northern hemisphere. We used regression and Principal Components Analysis (PCA) to evaluate general relationships between litter quality parameters and decay rates measured under laboratory and field conditions.

Results

Both concentrations of the polysaccharide sphagnan and the soluble phenolics were positively correlated with intrinsic decay resistance, however, so were the previously understudied lignin-like phenolics. More resistant litter had more of all the important metabolites; consequently, PC1 scores were related to lab mass loss (R²?=?0.57). There was no such relationship with field mass loss, which is also affected by the environment. PCA also revealed that metabolites clearly group Sphagnum sections (subgenera).

Conclusions

We suggest that the commonly stated growth-decomposition trade-off is largely due to litter quality. We show a strong phylogenetic control on Sphagnum metabolites, but their effects on decay are affected by nutrient availability in the habitat.

     

Community and species-performance patterns along an alpine poor-rich mire gradient

Abstract. Vegetation, water table depth and water chemistry of 16 peatlands in the southern Alps, Italy, were analysed in 115 sample plots. A poor-rich gradient could be detected along the first axis of a partial Detrended Canonical Correspondence Analysis. Both vegetation and hydrochemistry vary gradually along the gradient. Vascular plants have much broader niches along the gradient than bryophytes. Mosses (except Sphangnum) and hepatics have narrower niches than Sphagnum. The various species of Sphagnum segregate more clearly from each other along the moisture gradient than along the poor-rich one. The positions of species optima along the latter gradient largely reflect the ecological preferences of mire plants in peatlands with respect to nutrient status.     

Fine-Scale Horizontal and Vertical Micro-distribution Patterns of Testate Amoebae Along a Narrow Fen/Bog Gradient

The ecology of peatland testate amoebae is well studied along broad gradient from very wet (pool) to dry (hummock) micro-sites where testate amoebae are often found to respond primarily to the depth to water table (DWT). Much less is known on their responses to finer-scale gradients, and nothing is known of their possible response to phenolic compounds, which play a key role in carbon storage in peatlands. We studied the vertical (0–3, 3–6, and 6–9 cm sampling depths) micro-distribution patterns of testate amoebae in the same microhabitat (Sphagnum fallax lawn) along a narrow ecological gradient between a poor fen with an almost flat and homogeneous Sphagnum carpet (fen) and a “young bog” (bog) with more marked micro-topography and mosaic of poor fen and bog vegetation. We analyzed the relationships between the testate amoeba data and three sets of variables (1) “chemical” (pH, Eh potential, and conductivity), (2) “physical” (water temperature, altitude, i.e., Sphagnum mat micro-topography, and DWT), and (3) phenolic compounds in/from Sphagnum (water-soluble and primarily bound phenolics) as well as the habitat (fen/bog) and the sampling depth. Testate amoeba Shannon H′ diversity, equitability J of communities, and total density peaked in lower parts of Sphagnum, but the patterns differed between the fen and bog micro-sites. Redundancy analyses revealed that testate amoeba communities differed significantly in relation to Eh, conductivity, water temperature, altitude, water-soluble phenolics, habitat, and sampling depth, but not to DWT, pH, or primarily bound phenolics. The sensitivity of testate amoebae to weak environmental gradients makes them particularly good integrators of micro-environmental variations and has implications for their use in paleoecology and environmental monitoring. The correlation between testate amoeba communities and the concentration of water-soluble phenolic suggests direct (e.g., physiological) and/or indirect (e.g., through impact on prey organisms) effects on testate amoebae, which requires further research.     

Desiccation tolerance of Sphagnum revisited: a puzzle resolved

As ecosystem engineers, Sphagnum mosses control their surroundings through water retention, acidification and peat accumulation. Because water retention avoids desiccation, sphagna are generally intolerant to drought; however, the literature on Sphagnum desiccation tolerance (DT) provides puzzling results, indicating the inducible nature of their DT. To test this, various Sphagnum species and other mesic bryophytes were hardened to drought by (i) slow drying; (ii) ABA application and (iii) chilling or frost. DT tolerance was assessed as recovery of chlorophyll fluorescence parameters after severe desiccation. We monitored the seasonal course of DT in bog bryophytes. Under laboratory conditions, following initial de‐hardening, untreated Sphagnum shoots lacked DT; however, DT was induced by all hardening treatments except chilling, notably by slow drying, and in Sphagnum species of the section Cuspidata. In the field, sphagna in hollows and lawns developed DT several times during the growing season, responding to reduced precipitation and a lowered water table. Hummock and aquatic species developed DT only in late autumn, probably as a response to frost. Sphagnum protonemata failed to develop DT; hence, desiccation may limit Sphagnum establishment in drier habitats with suitable substrate chemistry. Desiccation avoiders among sphagna form compact hummocks or live submerged; thus, they do not develop DT in the field, lacking the initial desiccation experience, which is frequent in hollow and lawn habitats. We confirmed the morpho‐physiological trade‐off: in contrast to typical hollow sphagna, hummock species invest more resources in water retention (desiccation avoidance), while they have a lower ability to develop physiological DT.     

Fire history and vegetation recovery in two raised bogs at the Baltic Sea

Questions: What were the bog fire patterns and frequencies in two boreal peatlands during the last 5000 years? What is the nature and time‐scale of post‐fire vegetation successions? Were fire events related to climate? Location: Männikjärve bog, central east Estonia; Kontolanrahka bog, southwest Finland. Methods: Macroscopic charcoal, plant macrofossils and radiocarbon dating were examined. Redundancy analysis was used in the assessments. Results: During the last 5000 years, both of the above peatlands have experienced several fire events. A typical pre‐fire vegetation community consisted of dry hummock Sphagnum spp., often accompanied by Calluna vulgaris. Only the most severe occasional fires resulted in a dramatic change in the vegetation composition. In these cases, a wet shift occurred, where the pre‐fire hummock community was replaced by a wet hollow community. Calluna vulgaris was found to be a key species in both pre‐ and post‐fire vegetation dynamics. The recovery time of dry microtopes following severe combustion and the subsequent hydrological change could take up to 350 years. Even after a long‐lasting wet phase, the post‐fire disturbance succession led towards a dry hummock community. Conclusions: Fire succession appeared to be cyclic, starting as and developing towards a dry hummock community. Fires have been a regular phenomenon in boreal bogs, even in regions with rather low human impact. The fire history records did not indicate any direct link to the regional long‐term climate.     

A climatic threshold triggers the die‐off of peat mosses during an extreme heat wave

Heat waves, which are projected to be more frequent and intense in a warmer climate, could become a serious threat to plants that rely on water surplus availability, such as bryophytes. Here, I take the advantage of the European summer 2003 climate anomaly to assess the impact of an extreme heat wave on peat mosses of the genus Sphagnum, a group of bryophytes forming the bulk of living and dead biomass in peatlands. With this aim, 20 selected bogs in the Italian Alps were checked for Sphagnum survival in the years following the heat wave. Over the study area, the period May–September 2003 was characterized by higher mean monthly air temperature (13.5 °C) and lower mean monthly precipitation (87 mm) compared with normal climatic conditions (11.5 °C and 117 mm, respectively) so that the heat wave coincided with a drought spell. As a consequence of the unusual water stress, I documented an increased mortality of peat mosses forming high hummocks. In particular, at habitat scale, the distribution of desiccated peat mosses was restricted to the hummock face receiving the greatest amount of solar irradiation. However, at regional scale, the present study identified a climatic threshold, simply defined by the ratio of precipitation to temperature (P : T), which triggered an irreversible desiccation of peat mosses when mean monthly P : T dropped below 6.5 (mm : °C) during May–September 2003. The absence of any sign of recovery after 4 years since the drought must be seen as a harbinger of the deleterious effects of extreme heat waves on organisms not adapted to cope with abrupt climate anomaly.     

Field Simulation of Global Change: Transplanting Northern Bog Mesocosms Southward

A large proportion of northern peatlands consists of Sphagnum-dominated ombrotrophic bogs. In these bogs, peat mosses (Sphagnum) and vascular plants occur in an apparent stable equilibrium, thereby sustaining the carbon sink function of the bog ecosystem. How global warming and increased nitrogen (N) deposition will affect the species composition in bog vegetation is still unclear. We performed a transplantation experiment in which mesocosms with intact vegetation were transplanted southward from north Sweden to north-east Germany along a transect of four bog sites, in which both temperature and N deposition increased. In addition, we monitored undisturbed vegetation in control plots at the four sites of the latitudinal gradient. Four growing seasons after transplantation, ericaceous dwarf shrubs had become much more abundant when transplanted to the warmest site which also had highest N deposition. As a result ericoid aboveground biomass in the transplanted mesocosms increased most at the southernmost site, this site also had highest ericoid biomass in the undisturbed vegetation. The two dominant Sphagnum species showed opposing responses when transplanted southward; Sphagnum balticum height increment decreased, whereas S. fuscum height increment increased when transplanted southward. Sphagnum production did not differ significantly among the transplanted mesocosms, but was lowest in the southernmost control plots. The dwarf shrub expansion and increased N concentrations in plant tissues we observed, point in the direction of a positive feedback toward vascular plant-dominance suppressing peat-forming Sphagnum in the long term. However, our data also indicate that precipitation and phosphorus availability influence the competitive balance between Sphagnum, dwarf shrubs and graminoids.     

Mechanisms involved in the re-establishment of Sphagnum-dominated vegetation in rewetted bog remnants

Restoration of peat bog vegetation inhighly degraded peatlands is generallyattempted by improving the hydrology ofthese areas. The present paper discussesand explains various restoration strategiesrelating to peat quality, water chemistryand hydrology. In some cases, (shallow)inundation of bog remnants leads to a rapidredevelopment of (floating) Sphagnumvegetation, usually when poorly humifiedSphagnum peat is still present. Afterinundation, the peat either swells up tothe newly created water table or becomesbuoyant, in both cases creating a favorablesubstrate for Sphagnum mosses. Bulkdensity and methane production rate play animportant role in the buoyancy of floatingpeat, methane providing buoyancy to thesubstrates. The presence of (slightly)calcareous groundwater in the peat base mayenhance the development of floating raftsby stimulating decomposition processes.Alternatively, the growth of submerged Sphagnum species can also lead to thedevelopment of floating rafts. This dependson the penetration of light into the waterlayer and the availability of carbondioxide in the water layer.Many bog remnants, however, only havestrongly humified peat, which does notfavor the redevelopment of Sphagnumcarpets after deep inundation. On the otherhand, most peat moss species appear to dovery well on surface soaked black peat,which is why shallow inundation (< 0.3 m)is to be preferred in such cases.Compartmentalization of the terrain willprobably be necessary to ensure a more orless constant water table.An important prerequisite for thesuccessful restoration of bog remnants isthe development of a hydrologicallyself-regulating acrotelm. Key speciesinvolved in this development are Sphagnum magellanicum, Sphagnumpapillosum and Sphagnum rubellum.These typical hummock and lawn species areusually very slow colonizers compared tohollow species such as Sphagnumcuspidatum and Sphagnum fallax.Introduction of key species in carpetsdominated by hollow species or on baresubstrates appears to be very successful,indicating that the main constraint iscolonization.     

Climate drivers alter nitrogen availability in surface peat and decouple N2 fixation from CH4 oxidation in the Sphagnum moss microbiome

Peat mosses (Sphagnum spp.) are keystone species in boreal peatlands, where they dominate net primary productivity and facilitate the accumulation of carbon in thick peat deposits. Sphagnum mosses harbor a diverse assemblage of microbial partners, including N₂-fixing (diazotrophic) and CH₄-oxidizing (methanotrophic) taxa that support ecosystem function by regulating transformations of carbon and nitrogen. Here, we investigate the response of the Sphagnum phytobiome (plant + constituent microbiome + environment) to a gradient of experimental warming (+0°C to +9°C) and elevated CO₂ (+500 ppm) in an ombrotrophic peatland in northern Minnesota (USA). By tracking changes in carbon (CH₄, CO₂) and nitrogen (NH₄-N) cycling from the belowground environment up to Sphagnum and its associated microbiome, we identified a series of cascading impacts to the Sphagnum phytobiome triggered by warming and elevated CO₂. Under ambient CO₂, warming increased plant-available NH₄-N in surface peat, excess N accumulated in Sphagnum tissue, and N₂ fixation activity decreased. Elevated CO₂ offset the effects of warming, disrupting the accumulation of N in peat and Sphagnum tissue. Methane concentrations in porewater increased with warming irrespective of CO₂ treatment, resulting in a ~10× rise in methanotrophic activity within Sphagnum from the +9°C enclosures. Warming's divergent impacts on diazotrophy and methanotrophy caused these processes to become decoupled at warmer temperatures, as evidenced by declining rates of methane-induced N₂ fixation and significant losses of keystone microbial taxa. In addition to changes in the Sphagnum microbiome, we observed ~94% mortality of Sphagnum between the +0°C and +9°C treatments, possibly due to the interactive effects of warming on N-availability and competition from vascular plant species. Collectively, these results highlight the vulnerability of the Sphagnum phytobiome to rising temperatures and atmospheric CO₂ concentrations, with significant implications for carbon and nitrogen cycling in boreal peatlands.     

Sphagnum divinum (sp. nov.) and S. medium Limpr. and their relationship to S. magellanicum Brid.

Sphagnum magellanicum has been viewed as being a predominantly circumpolar species in the northern hemisphere, but it occurs in the southern hemisphere and was originally described from the southern parts of Chile. It is an ecologically important species in mire ecosystems and has been extensively used as a model to study processes of growth, carbon sequestration and peat decomposition. Molecular and experimental studies have, however, revealed genetic structure within S. magellanicum, and morphological differences associated with these genetic groups. Here we describe Sphagnum divinum in Sphagnum subgenus Sphagnum (Sphagnaceae, Bryophyta) as a new species, based on molecular and morphological evidence. Sphagnum medium is reinstated as a distinct species and is epitypified. Consequently, a new species concept of S. magellanicum is presented including an epitypification. Important morphological characters to separate these three species in the field and under the microscope are presented. Ecology and distribution differ among the species; S. divinium has a wide habitat range including mire margin, forested peatlands and moist heaths, and a circumpolar distribution around the northern hemisphere. Sphagnum medium seems to be more restricted to ombrotrophic mire expanse habitats and shows an amphi-Atlantic distribution in the northern hemisphere. Sphagnum magellanicum has a very broad ecological niche in peatlands and is found in most mire habitats in Tierra del Fuego on the southern tip of South America.     

Litter Decomposition in Temperate Peatland Ecosystems: The Effect of Substrate and Site

Abstract The large accumulation of organic matter in peatlands is primarily caused by slow rates of litter decomposition. We determined rates of decomposition of major peat-forming litters of vascular plants and mosses at five sites: a poor fen in New Hampshire and a bog hummock, a poor fen, a beaver pond margin and a beaver pond in Ontario. We used the litterbag technique, retrieving triplicate litterbags six or seven times over 3–5 years, and found that simple exponential decay and continuous-quality non-linear regression models could adequately characterize the decomposition in most cases. Within each site, the rate of decomposition at the surface was generally Typha latifolia leaves = Chamaedaphne calyculata leaves = Carex leaves > Chamaedaphne calyculata stems > hummock Sphagnum = lawn/hollow Sphagnum, with exponential decay constant (k) values generally ranging from 0.05 to 0.37 and continuous-quality model initial quality (q ₀ ) values ranging from 1.0 (arbitrarily set for Typha leaves) to 0.7 (Sphagnum). In general, surface decay rates were slowest at the bog hummock site, which had the lowest water table, and in the beaver pond, which was inundated, and fastest at the fens. The continuous-quality model site decomposition parameter (u ₀ ) ranged from 0.80 to 0.17. Analysis of original litter samples for carbon, nitrogen and proximate fractions revealed a relatively poor explanation of decomposition rates, as defined by k and q ₀ , compared to most well-drained ecosystems. Three litters, roots of sedge and a shrub and Typha leaves, were placed at depths of 10, 30 and 60 cm at the sites. Decomposition rates decreased with depth at each site, with k means of 0.15, 0.08 and 0.05 y⁻¹ at 10, 30 and 60 cm, respectively, and u ₀ of 0.25, 0.13 and 0.07. These differences are primarily related to the position of the water table at each site and to a lesser extent the cooler temperatures in the lower layers of the peat. The distinction between bog and fen was less important than the position of the water table. These results show that we can characterize decomposition rates of surface litter in northern peatlands, but given the large primary productivity below-ground in these ecosystems, and the differential rates of decomposition with depth, subsurface input and decomposition of organic matter is an important and relatively uncertain attribute.     

Old Lineages in a New Ecosystem: Diversification of Arcellinid Amoebae (Amoebozoa) and Peatland Mosses

Arcellinid testate amoebae (Amoebozoa) form a group of free-living microbial eukaryotes with one of the oldest fossil records known, yet several aspects of their evolutionary history remain poorly understood. Arcellinids occur in a range of terrestrial, freshwater and even brackish habitats; however, many arcellinid morphospecies such as Hyalosphenia papilio are particularly abundant in Sphagnum-dominated peatlands, a relatively new ecosystem that appeared during the diversification of Sphagnum species in the Miocene (5–20 Myr ago). Here, we reconstruct divergence times in arcellinid testate amoebae after selecting several fossils for clock calibrations and then infer whether or not arcellinids followed a pattern of diversification that parallels the pattern described for Sphagnum. We found that the diversification of core arcellinids occurred during the Phanerozoic, which is congruent with most arcellinid fossils but not with the oldest known amoebozoan fossil (i.e. at ca. 662 or ca. 750 Myr). Overall, Sphagnum and the Hyalospheniidae exhibit different patterns of diversification. However, an extensive molecular phylogenetic analysis of distinct clades within H. papilio species complex demonstrated a correlation between the recent diversification of H. papilio, the recent diversification of Sphagnum mosses, and the establishment of peatlands.     

Environmental Controls of Cryptogam Composition and Diversity in Anthropogenic and Natural Peatland Ecosystems of Chilean Patagonia

Peatlands exhibit highly characteristic ecological traits and are unique complex ecosystems. Nevertheless, knowledge about southern South American peatlands is very limited. In this study, we analyzed species composition of bryophytes and lichens of Southern Hemisphere peatlands, specifically from eight peatlands of Isla Grande de Chiloé (Chiloé Island) in southern Chile (42°–43°S and 75°–73°W). Two kinds of Sphagnum peatlands were studied: natural and anthropogenic peatlands. Our results indicate the existence of clear environmental gradients affecting the structure of bryo-lichenic communities in the Sphagnum peatlands of Chiloé. Canonical correspondence analysis suggests that variation in bryophyte and lichen species composition mainly follows ombrotrophic–minerotrophic and lithotrophic-thalassotrophic gradients. Surface-water chemistry is the most significant factor accounting for changes in floristic composition among our study sites. In contrast to our expectations, bog origin (natural or anthropic) was not the most significant factor accounting for changes in floristic composition among peatlands. Other elements, such as the water source supplying peatlands or the influence of sea spray, were more relevant in the bryo-lichenic flora species occurrence in the peatlands of Chiloé. We also observed clear differences in ecological niches among species in general additive model response curves. Therefore, our results show that despite the origin, the ecology of peatlands follows common rules with peatlands from the Northern Hemisphere.     

Dry/wet cycling reduces spore germination and viability in six peatland bryophytes

• Dry/wet cycling driven by water level fluctuation in wetlands may strongly influence the destiny of seeds. However, how dry/wet cycling affects spore survival and germinability in peatland bryophytes is poorly understood.
• Six peatland bryophytes, three hummock- and three hollow-dwelling Sphagnum species, were chosen as study species. We tested the effects of dry (60% air RH)/wet (waterlogging) cycle frequency (once per 12, 8 or 4 days for low, medium or high, respectively) and ratio (3:1, 1:1 or 1:3 dry:wet time per cycle) on spore germinability, viability, dormancy percentage and protonema development.
• Dry/wet cycling significantly reduced spore germination percentage and viability and slowed protonema development in all Sphagnum species, being more pronounced with higher dry/wet cycling frequencies. The hummock species S. capillifolium and S. fuscum had higher spore germination percentage after the continuous dry treatment, while the hollow species S. angustifolium, S. squarrosum and S. subsecundum showed the opposite response, compared to the continuously wet treatment. Except for S. squarrosum, spore viability was higher after the dry than after the wet treatment. Spore viability and dormancy percentage were higher after a dry/wet ratio of 1:3 than after ratios of 3:1 and 1:1.
• Our study shows that both germinability and viability of bryophyte spores are reduced by dry/wet cycling (especially when frequent) in peatlands. This emphasizes the need to ensure constant water levels and low frequencies of water level fluctuation, which are relevant in connection with wetland restoration, to promote Sphagnum spore survival and establishment in peatlands after disturbances.

     

Sphagnum distribution patterns along environmental gradients in Bulgaria

Abstract

The distribution patterns of 18 Sphagnum species along base-richness and altitudinal gradients were studied in Bulgarian treeless wetlands which are noteworthy because of the edge-of-range occurrence of many mire species including Sphagnum. Of 483 spring and mire sites studied, 202 samples contained some Sphagnum species. The most common species were S. subsecundum (n=85), S. platyphyllum (46), S. contortum (41), S. teres (40) and S. capillifolium (26). The significance of Sphagnum responses to environmental gradients was tested by comparing generalized additive models against the null model. Many Sphagnum species displayed a significant response to the altitudinal gradient. Several species were clearly linked to low or to high altitudes, but the realized niche of other species was wide with respect to altitude. Most species significantly responded to water pH, both above and below the timberline. The same result was obtained for water conductivity below the timberline, whereas only a few species had a significant response to conductivity above the timberline. The highest water conductivity under which Sphagnum species occurred was 280 μS cm^?1. Sphagnum contortum was the species occupying the mires with the highest mineral content. On the contrary, Sphagnum warnstorfii, one of the most calcitolerant species in many regions of Europe, often occurred in extremely mineral-poor mires above the timberline. Some other Sphagnum species growing in mineral-rich mires below the timberline, also inhabited extremely mineral-poor mires above the timberline. This could be explained by adaptation to local conditions during long-term isolation on mineral-poor bedrock or by changed competition pressure.