Experimental warming alters the community composition,diversity, and N2 fixation activity of peat moss (Sphagnum fallax) microbiomes

Sphagnum‐dominated peatlands comprise a globally important pool of soil carbon (C) and are vulnerable to climate change. While peat mosses of the genus Sphagnum are known to harbor diverse microbial communities that mediate C and nitrogen (N) cycling in peatlands, the effects of climate change on Sphagnum microbiome composition and functioning are largely unknown. We investigated the impacts of experimental whole‐ecosystem warming on the Sphagnum moss microbiome, focusing on N₂ fixing microorganisms (diazotrophs). To characterize the microbiome response to warming, we performed next‐generation sequencing of small subunit (SSU) rRNA and nitrogenase (nifH) gene amplicons and quantified rates of N₂ fixation activity in Sphagnum fallax individuals sampled from experimental enclosures over 2 years in a northern Minnesota, USA bog. The taxonomic diversity of overall microbial communities and diazotroph communities, as well as N₂ fixation rates, decreased with warming (p < 0.05). Following warming, diazotrophs shifted from a mixed community of Nostocales (Cyanobacteria) and Rhizobiales (Alphaproteobacteria) to predominance of Nostocales. Microbiome community composition differed between years, with some diazotroph populations persisting while others declined in relative abundance in warmed plots in the second year. Our results demonstrate that warming substantially alters the community composition, diversity, and N₂ fixation activity of peat moss microbiomes, which may ultimately impact host fitness, ecosystem productivity, and C storage potential in peatlands.     

Sphagnum in peatlands of Australasia: Their distribution,utilisation and management

In comparison to the northern hemisphere, Sphagnum peatlands are an unusual andinfrequent component of the Australasianlandscape. Most peatlands in Australasiaare primarily composed of eitherRestionaceous or Cyperaceous peats. Sphagnum peatlands in Australia and PapuaNew Guinea/Irian Jaya (now West Papua) arelargely located in montane and alpineenvironments, but also occur down to sealevel in New Zealand and as moss patches onsome subantarctic islands. Fire is a majordeterminant of the characteristics ofpeatlands in Australasia. Peatlandmanagement in Australasia is hindered bythe need for increased understanding ofpeatland processes to enable a sustainablebalance of conservation of a small resourcewith localised utilisation. Themanagement focus in Australasia has largelybeen on ensuring ecologically sustainable Sphagnum moss harvesting, withlimited peat mining. We have found thatgeneral recovery of Sphagnum after moss harvesting canbe enhanced by harvesting larger peatlands,and by leaving one-third of the acrotelm toregenerate. The largest upland peat swampin mainland Australia, Wingecarribee Swamp,suffered a major collapse in 1998 followingpeat mining. Environmental and managementconsequences of this collapse have majorramifications for rehabilitation options. Sphagnum peatlands in Australasia arelikely to be adversely affected bydrainage, burning, grazing, trampling,global warming and peat mining.     

Vascular plant‐mediated controls on atmospheric carbon assimilation and peat carbon decomposition under climate change

Climate change can alter peatland plant community composition by promoting the growth of vascular plants. How such vegetation change affects peatland carbon dynamics remains, however, unclear. In order to assess the effect of vegetation change on carbon uptake and release, we performed a vascular plant‐removal experiment in two Sphagnum‐dominated peatlands that represent contrasting stages of natural vegetation succession along a climatic gradient. Periodic measurements of net ecosystem CO₂ exchange revealed that vascular plants play a crucial role in assuring the potential for net carbon uptake, particularly with a warmer climate. The presence of vascular plants, however, also increased ecosystem respiration, and by using the seasonal variation of respired CO₂ radiocarbon (bomb‐¹⁴C) signature we demonstrate an enhanced heterotrophic decomposition of peat carbon due to rhizosphere priming. The observed rhizosphere priming of peat carbon decomposition was matched by more advanced humification of dissolved organic matter, which remained apparent beyond the plant growing season. Our results underline the relevance of rhizosphere priming in peatlands, especially when assessing the future carbon sink function of peatlands undergoing a shift in vegetation community composition in association with climate change.     

Abundant Trimethylornithine Lipids and Specific Gene Sequences Are Indicative of Planctomycete Importance at the Oxic/Anoxic Interface in Sphagnum-Dominated Northern Wetlands

Northern wetlands make up a substantial terrestrial carbon sink and are often dominated by decay-resistant Sphagnum mosses. Recent studies have shown that planctomycetes appear to be involved in degradation of Sphagnum-derived debris. Novel trimethylornithine (TMO) lipids have recently been characterized as abundant lipids in various Sphagnum wetland planctomycete isolates, but their occurrence in the environment has not yet been confirmed. We applied a combined intact polar lipid (IPL) and molecular analysis of peat cores collected from two northern wetlands (Saxnäs Mosse [Sweden] and Obukhovskoye [Russia]) in order to investigate the preferred niche and abundance of TMO-producing planctomycetes. TMOs were present throughout the profiles of Sphagnum bogs, but their concentration peaked at the oxic/anoxic interface, which coincided with a maximum abundance of planctomycete-specific 16S rRNA gene sequences. The sequences detected at the oxic/anoxic interface were affiliated with the Isosphaera group, while sequences present in the anoxic peat layers were related to an uncultured planctomycete group. Pyrosequencing-based analysis identified Planctomycetes as the major bacterial group at the oxic/anoxic interface at the Obukhovskoye peat (54% of total 16S rRNA gene sequence reads), followed by Acidobacteria (19% reads), while in the Saxnäs Mosse peat, Acidobacteria were dominant (46%), and Planctomycetes contributed to 6% of the total reads. The detection of abundant TMO lipids in planctomycetes isolated from peat bogs and the lack of TMO production by cultures of acidobacteria suggest that planctomycetes are the producers of TMOs in peat bogs. The higher accumulation of TMOs at the oxic/anoxic interface and the change in the planctomycete community with depth suggest that these IPLs could be synthesized as a response to changing redox conditions at the oxic/anoxic interface.     

Simulated Small Scale Disturbances Increase Decomposition Rates and Facilitates Invasive Species Encroachment in a High Elevation Tropical Andean Peatland

Tropical alpine peatlands are important carbon reservoirs and are a critical component of local hydrological cycles. In high elevation peatlands slow decomposition rates result from a nutrient‐poor substrate resistant to decay. The responses of páramo peatland ecosystems to increased nutrient additions and physical disturbance due to agricultural activities are unknown. Here, we conducted a two‐year fertilization and physical disturbance experiment in a Sphagnum—dominated peatland in the Central Andes of Colombia. We hypothesized that fertilization and physical disturbance will diminish the ability of the peat to store organic matter by increasing decomposition and that vascular plants will displace Sphagnum as the dominant plant group. We simulated cattle activity by adding manure as a fertilizer and physical disturbance as a proxy for cattle trampling. Species composition varied in proportion to the intensity of disturbance. Sphagnum cover was reduced under any disturbance treatment. Non‐native grasses usually found in cattle pastures invaded treatments with fertilizer additions or physical disturbance. Overall aboveground plant biomass doubled in fertilized treatments, suggesting that plant biomass production was nutrient limited. Decomposition rates tripled in disturbed treatments as compared to controls. This reduces the ability of the peatland to store organic matter. Andean peatlands are prized ecological assets; however, our results show that the El Morro páramo peatland experienced increased decomposition rates over short time periods after small‐scale disturbances. This created profound consequences for the ecological services offered by these peatlands.     

Acidobacteria in microbial communities of the bog and tundra lichens

Bacterial communities of the lichens from a Sphagnum bog (Karelia) and tundra (Vorkuta oblast) were investigated. Members of the phylum Acidobacteria were numerous in the thallus of living and decaying lichens (3.8 × 10⁸ cells/g), constituting 6 to 32% of the total bacterial number. Pure cultures of acidobacteria were isolated from the samples of living and decaying lichen thallus. Ten of them were identified and classified as members of subgroup 1 of the Acidobacteria. The hydrolytic activity of two strains isolated from the living and decomposing zones of the thallus was investigated. They were capable of growth on xylan, starch, pectin, laminarin, and lichen extract. Acidobacteria were shown to be a stable and numerous component of microbial communities of the bog and tundra lichens.     

Restoration of Sphagnum and restiad peatlands in Australia and New Zealand reveals similar approaches

Peatlands in Australia and New Zealand are composed mainly of Restionaceous and Cyperaceous peats, although Sphagnum peat is common in wetter climates (Mean Annual Precipitation > 1,000 mm) and at higher altitudes (>1,000 m). Experimental trials in two contrasting peatland types—fire‐damaged Sphagnum peatlands in the Australian Alps and cutover restiad bogs in lowland New Zealand—revealed similar approaches to peatland restoration. Hydrological restoration and rehydration of drying peats involved blocking drainage ditches to raise water tables or, additionally in burnt Sphagnum peatlands, peat‐trenching, and the use of sterilized straw bales to form semipermanent “dam walls” and barriers to spread and slow surface water movement. Recovery to the predisturbance vegetation community was most successful once protective microclimates had been established, either artificially or naturally. Specifically, horizontally laid shadecloth resulted in Sphagnum cristatum regeneration rates and biomass production 3–4 times that of unshaded vegetation (Australia), and early successional nurse shrubs facilitated establishment of Sporadanthus ferrugineus (New Zealand) within 2–3 years. On severely burnt or cutover sites, a patch dynamic approach using transplants of Sphagnum or creation of restiad peat “islands” markedly improved vegetation recovery. In New Zealand, this approach has been scaled up to whole mine‐site restoration, in which the newly vegetated islands provide habitat and seed sources for plants and invertebrates to spread onto surrounding areas. Although a vegetation cover can be established relatively rapidly in both peatland types, restoration of invertebrate communities, ecosystem processes, and peat hydrological function and accumulation may take many decades.     

Microbial community composition and methanotroph diversity of a subarctic wetland in Russia

This study assessed the microbial diversity, activity, and composition of methane-oxidizing communities of a subarctic wetland in Russia with mosaic cover of Sphagnum mosses and lichens of the genera Cladonia and Cetraria. Potential methane-oxidizing activity of peat sampled from lichen-dominated wetland sites was higher than that in the sites dominated by Sphagnum mosses. In peat from lichen-dominated sites, major bacterial groups identified by high-throughput sequencing of the 16S rRNA genes were the Acidobacteria (35.4–41.2% of total 16S rRNA gene reads), Alphaproteobacteria (19.1–24.2%), Gammaproteobacteria (7.9–11.1%), Actinobacteria (5.5–13.2%), Planctomycetes (7.2–9.5%), and Verrucomicrobia (5.1–9.5%). The distinctive feature of this community was high proportion of Subdivision 2 Acidobacteria, which are not characteristic for boreal Sphagnum peat bogs. Methanotrophic community composition was determined by molecular analysis of the pmoA gene encoding particulate methane monooxygenase. Most (~80%) of all pmoA gene fragments revealed in peat from lichen-dominated sites belonged to the phylogenetic lineage represented by a microaerobic spiral-shaped methanotroph, “Candidatus Methylospira mobilis”. Members of the genus Methylocystis, which are typical inhabitants of boreal Sphagnum peat bogs, represented only a minor group of indigenous methanotrophs. The specific feature of a methanotrophic community in peat from lichen-dominated sites was the presence of uncultivated USCα (Upland Soil Cluster alpha) methanotrophs, which are typical for acidic upland soils showing atmospheric methane oxidation. The methanotrophic community composition in lichen-dominated sites of a tundra wetland, therefore, was markedly different from that in boreal Sphagnum peat bogs.     

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.     

Rapid loss of an ecosystem engineer: Sphagnum decline in an experimentally warmed bog

Sphagnum mosses are keystone components of peatland ecosystems. They facilitate the accumulation of carbon in peat deposits, but climate change is predicted to expose peatland ecosystem to sustained and unprecedented warming leading to a significant release of carbon to the atmosphere. Sphagnum responses to climate change, and their interaction with other components of the ecosystem, will determine the future trajectory of carbon fluxes in peatlands. We measured the growth and productivity of Sphagnum in an ombrotrophic bog in northern Minnesota, where ten 12.8‐m‐diameter plots were exposed to a range of whole‐ecosystem (air and soil) warming treatments (+0 to +9°C) in ambient or elevated (+500 ppm) CO₂. The experiment is unique in its spatial and temporal scale, a focus on response surface analysis encompassing the range of elevated temperature predicted to occur this century, and consideration of an effect of co‐occurring CO₂ altering the temperature response surface. In the second year of warming, dry matter increment of Sphagnum increased with modest warming to a maximum at 5°C above ambient and decreased with additional warming. Sphagnum cover declined from close to 100% of the ground area to <50% in the warmest enclosures. After three years of warming, annual Sphagnum productivity declined linearly with increasing temperature (13–29 g C/m² per °C warming) due to widespread desiccation and loss of Sphagnum. Productivity was less in elevated CO₂ enclosures, which we attribute to increased shading by shrubs. Sphagnum desiccation and growth responses were associated with the effects of warming on hydrology. The rapid decline of the Sphagnum community with sustained warming, which appears to be irreversible, can be expected to have many follow‐on consequences to the structure and function of this and similar ecosystems, with significant feedbacks to the global carbon cycle and climate change.     

Relationships between bryophyte production and substrate properties in restored milled peatlands

The relationship between the small‐scale distribution pattern of bryophyte biomass on restored milled peatlands and substrate properties (e.g. moisture, pH, nutrients, and their ratios) was studied. Substrate properties may determine the species composition of bryophyte communities that have developed in such areas. Two experimental sites were established in northern Estonia where the moss‐layer‐transfer technique had been used for the revegetation of abandoned peatfields for almost a decade before sampling. Diaspores of Sphagnum species common on bogs were distributed in these sites. After 7 years one site was mainly dominated by Sphagnum whereas true mosses (Polytrichum strictum, Aulacomnium palustre, and Pleurozium schreberi) were abundant in the other site. Three moss groups were distinguished: Sphagnum, P. strictum, and other mosses based on cluster analysis. The biomass of Sphagnum was related to peat moisture and potassium content. For P. strictum the N/K ratio was important, and the production of A. palustre grew with the increase in the N/P ratio of peat. It was concluded that peat properties played an important role in the formation and development of bryophyte communities on revegetated peatfields on a small scale (<0.1 ha).     

Wide distribution of Phycisphaera-like planctomycetes from WD2101 soil group in peatlands and genome analysis of the first cultivated representative

Phycisphaera-like WD2101 ‘soil group’ is one of the as-yet-uncultivated phylogenetic clades within the phylum Planctomycetes. Members of this clade are commonly detected in various terrestrial habitats. This study shows that WD2101 represented one of the major planctomycete groups in 10 boreal peatlands, comprising up to 76% and 36% of all Planctomycetes-affiliated 16S rRNA gene reads in raised bogs and eutrophic fens respectively. These types of peatlands displayed clearly distinct intra-group diversity of WD2101-affiliated planctomycetes. The first isolate of this enigmatic planctomycete group, strain M1803, was obtained from a humic lake surrounded by Sphagnum peat bogs. Strain M1803 displayed 89.2% 16S rRNA gene similarity to Tepidisphaera mucosa and was represented by motile cocci that divided by binary fission and grew under micro-oxic conditions. The complete 7.19 Mb genome of strain M1803 contained an array of genes encoding Planctomycetal type bacterial microcompartment organelle likely involved in l -rhamnose metabolism, suggesting participation of M1803-like planctomycetes in polysaccharide degradation in peatlands. The corresponding cellular microcompartments were revealed in ultrathin cell sections. Strain M1803 was classified as a novel genus and species, Humisphaera borealis gen. nov., sp. nov., affiliated with the formerly recognized WD2101 ‘soil group’.     

长白山哈泥泥炭地丘间生境泥炭藓孢子的寿命

长寿有性繁殖体对于植物种群的长存具有重要意义,迄今,泥炭地苔藓植物孢子长寿性研究还很少。在长白山哈泥泥炭地钻取丘间表层泥炭样品,测定泥炭腐殖化度和烧失量,逐层提取和培养泥炭藓孢子,研究埋藏时间对孢子萌发的影响。结果表明,丘间泥炭藓孢子埋藏环境中,随着埋深的增加即埋藏年限的增加,泥炭腐殖化度和烧失量总体上分别呈现增加和递减的趋势,而地层泥炭藓孢子萌发率呈现直线递减的规律,但在埋藏近150余年后孢子萌发率仍可达40%。研究进一步证明泥炭藓具有长期持久孢子库,根据推算,泥炭地丘间埋藏环境中,泥炭藓孢子最大寿命可超过400a。     

Biogeography and comparative ecology of testate amoebae inhabiting Sphagnum-dominated peatlands in the Great Lakes and Rocky Mountain regions of North America

Testate amoebae (Protozoa: Rhizopoda) are common inhabitants of peatlands. Strong relationships between community composition and substrate moisture in Sphagnum‐dominated peatlands have made them particularly useful as hydrological proxies in environmental and palaeoenvironmental research. However, stability of these relationships in geographical space is important for widespread applicability. In this study, we compared testate amoeba communities inhabiting Sphagnum‐dominated peatlands of the Great Lakes and Rocky Mountain regions of North America. Our primary objectives were to describe patterns of community composition in the two regions, develop hypotheses to explain differences, and determine if taxa occupy similar ecological niches with respect to substrate moisture in both places. Our results indicated that testate amoeba communities are relatively different in the two regions, and these differences are probably caused by differences in climate and peatland trophic status, although other factors may also play a role. However, many taxa do occur in both regions and these taxa had comparable moisture preferences in each region, suggesting that the ecological niches of taxa with respect to substrate moisture are similar even within communities of relatively different composition.     

Comparison of carbon and nitrogen accumulation rate between bog and fen phases in a pristine peatland with the fen-bog transition

Long-term carbon and nitrogen dynamics in peatlands are affected by both vegetation production and decomposition processes. Here, we examined the carbon accumulation rate (CAR), nitrogen accumulation rate (NAR) and δ¹³C, δ¹⁵N of plant residuals in a peat core dated back to ~8500 cal year BP in a temperate peatland in Northeast China. Impacted by the tephra during 1160 and 789 cal year BP and climate change, the peatland changed from a fen dominated by vascular plants to a bog dominated by Sphagnum mosses. We used the Clymo model to quantify peat addition rate and decay constant for acrotelm and catotelm layers during both bog and fen phases. Our studied peatland was dominated by Sphagnum fuscum during the bog phase (789 to −59 cal year BP) and lower accumulation rates in the acrotelm layer was found during this phase, suggesting the dominant role of volcanic eruption in the CAR of the peat core. Both mean CAR and NAR were higher during the bog phase than during the fen phase in our study, consistent with the results of the only one similar study in the literature. Because the input rate of organic matter was considered to be lower during the bog phase, the decomposition process must have been much lower during the bog phase than during the fen phase and potentially controlled CAR and NAR. During the fen phase, CAR was also lower under higher temperature and summer insolation, conditions beneficial for decomposition. δ¹⁵N of Sphagnum hinted that nitrogen fixation had a positive effect on nitrogen accumulation, particular in recent decades. Our study suggested that decomposition is more important for carbon and nitrogen sequestration than production in peatlands in most conditions and if future climate changes or human disturbance increase decomposition rate, carbon sequestration in peatlands will be jeopardized.     

Consistent centennial-scale change in European sub-Arctic peatland vegetation toward Sphagnum dominance—Implications for carbon sink capacity

Climate warming is leading to permafrost thaw in northern peatlands, and current predictions suggest that thawing will drive greater surface wetness and an increase in methane emissions. Hydrology largely drives peatland vegetation composition, which is a key element in peatland functioning and thus in carbon dynamics. These processes are expected to change. Peatland carbon accumulation is determined by the balance between plant production and peat decomposition. But both processes are expected to accelerate in northern peatlands due to warming, leading to uncertainty in future peatland carbon budgets. Here, we compile a dataset of vegetation changes and apparent carbon accumulation data reconstructed from 33 peat cores collected from 16 sub-arctic peatlands in Fennoscandia and European Russia. The data cover the past two millennia that has undergone prominent changes in climate and a notable increase in annual temperatures toward present times. We show a pattern where European sub-Arctic peatland microhabitats have undergone a habitat change where currently drier habitats dominated by Sphagnum mosses replaced wetter sedge-dominated vegetation and these new habitats have remained relatively stable over the recent decades. Our results suggest an alternative future pathway where sub-arctic peatlands may at least partly sustain dry vegetation and enhance the carbon sink capacity of northern peatlands.     

A Race for Space? How Sphagnum fuscum stabilizes vegetation composition during long‐term climate manipulations

Strong climate warming is predicted at higher latitudes this century, with potentially major consequences for productivity and carbon sequestration. Although northern peatlands contain one‐third of the world's soil organic carbon, little is known about the long‐term responses to experimental climate change of vascular plant communities in these Sphagnum‐dominated ecosystems. We aimed to see how long‐term experimental climate manipulations, relevant to different predicted future climate scenarios, affect total vascular plant abundance and species composition when the community is dominated by mosses. During 8 years, we investigated how the vascular plant community of a Sphagnum fuscum‐dominated subarctic peat bog responded to six experimental climate regimes, including factorial combinations of summer as well as spring warming and a thicker snow cover. Vascular plant species composition in our peat bog was more stable than is typically observed in (sub)arctic experiments: neither changes in total vascular plant abundance, nor in individual species abundances, Shannon's diversity or evenness were found in response to the climate manipulations. For three key species (Empetrum hermaphroditum, Betula nana and S. fuscum) we also measured whether the treatments had a sustained effect on plant length growth responses and how these responses interacted. Contrasting with the stability at the community level, both key shrubs and the peatmoss showed sustained positive growth responses at the plant level to the climate treatments. However, a higher percentage of moss‐encroached E. hermaphroditum shoots and a lack of change in B. nana net shrub height indicated encroachment by S. fuscum, resulting in long‐term stability of the vascular community composition: in a warmer world, vascular species of subarctic peat bogs appear to just keep pace with growing Sphagnum in their race for space. Our findings contribute to general ecological theory by demonstrating that community resistance to environmental changes does not necessarily mean inertia in vegetation response.