Nutrient dynamics of 12 Sphagnum species during establishment on a rewetted bog

• Peatland degradation through drainage and peat extraction have detrimental environmental and societal consequences. Rewetting is an option to restore lost ecosystem functions, such as carbon storage, biodiversity and nutrient sequestration. Peat mosses (Sphagnum) are the most important peat-forming species in bogs. Most Sphagnum species occur in nutrient-poor habitats; however, high growth rates have been reported in artificial nutrient-rich conditions with optimal water supply.
• Here, we demonstrate the differences in nutrient dynamics of 12 Sphagnum species during their establishment in a 1-year field experiment at a Sphagnum paludiculture area in Germany. The 12 species are categorized into three groups (slower-, medium- and fast-growing). Establishment of peat mosses is facilitated by constant supply of nutrient-rich, low pH, and low alkalinity surface water.
• Our study shows that slower-growing species (S. papillosum, S. magellancium, S. fuscum, S. rubellum, S. austinii; often forming hummocks) displayed signs of nutrient imbalance. These species accumulated higher amounts of N, P, K and Ca in their capitula, and had an elevated stem N:K quotient (>3). Additionally, this group sequestered less C and K per m² than the fast and medium-growing species (S. denticulatum, S. fallax, S. riparium, S. fimbriatum, S. squarrosum, S. palustre, S. centrale). Lower lawn thickness may have amplified negative effects of flooding in the slower-growing species.
• We conclude that nutrient dynamics and carbon/nutrient sequestration rates are species-specific. For bog restoration, generating ecosystem services or choosing suitable donor material for Sphagnum paludiculture, it is crucial to consider their compatibility with prevailing environmental conditions.

     

Sulfur cycling in a forested Sphagnum bog in northern Minnesota

The mass balance and internal cycle of sulfur within a small forested,Sphagnum bog in northern Minnesota are presented here based on a 4-year record of hydrologic inputs and outputs (precipitation, throughfall, streamflow, upland runoff) and a 3-year measurement of plant growth and sulfur uptake. Concentrations and accumulation rates of inorganic and organic sulfur species were measured in porewater. The bog is a large sink for sulfur, retaining 37% of the total sulfur input. Because of the relatively large export of organic S (21% of inputs), retention efficiency for total-S (organic S + SO ₄ ⁼ ; 37%) is less than that for SO ₄ ⁼ (58%). There is a dynamic cycle of oxidation and reduction within the bog. Annual oxidation and recycling of S is equal to total inputs in the center of the bog. Plants receive 47% of their uptake requirement from atmospheric deposition, 5% from retranslocation from foliage, and the remainder from sulfur remineralized from peat. Mineralization is most intense in the aerobic zone above the water table. Inorganic sulfur species comprise <5% of the total sulfur burden within the peat.     

High abundance of planctomycetes in anoxic layers of a Sphagnum peat bog

The depth distribution of planctomycete abundance has been examined in six different sites of the Sphagnum peat bog Bakchar, Tomsk oblast, Russia. In situ hybridization of peat with the fluorescently labeled oligonucleotide probes PLA46 and PLA886, reported to be group-specific for representatives of the phylum Planctomycetes, revealed two distinct population maxima of these bacteria in all of the profiles examined. The first population maximum was detected in the uppermost, oxic layer of the bog profile, while the second maximum was located at a depth of 30 cm below the water table level. The population sizes of planctomycetes in the uppermost layer and at a depth of 30 cm were of the same order of magnitude and comprised 0.5–1.5 × 10⁷ and 0.4?0.7 × 10⁷ cells per g^?1 of wet peat, respectively. Only 25–30% of the total number of planctomycete cells in the anoxic layer could be detected if the probe PLA886, whose target specificity is restricted to taxonomically characterized aerobic planctomycetes of the genera Gemmata, Planctomyces, Pirellula, and Isosphaera, was used alone. Other planctomycete cells in this layer were detected only with the probe PLA46, which possesses a much wider scope. This suggests the affiliation of these organisms with a yet undescribed phylogenetic subgroup within the Planctomycetes.     

High abundance of planctomycetes in anoxic layers of a Sphagnum peat bog

The depth distribution of planctomycete abundance has been examined in six different sites of the Sphagnum peat bog in Bakchar, Tomsk oblast, Russia. In situ hybridization of peat with the fluorescently labeled oligonucleotide probes PLA46 and PLA886, reported to be group-specific for representatives of the phylum Planctomycetes, revealed two distinct population maxima of these bacteria in all of the profiles examined. The first population maximum was detected in the uppermost, oxic layer of the bog profile, while the second maximum was located at a depth of 30 cm below the water table level. The population sizes of planctomycetes in the uppermost layer and at a depth of 30 cm were of the same order of magnitude and comprised 0.5-1.5 x 10(7) and 0.4-0.7 x 10(7) cells per g of wet peat, respectively. Only 25-30% of the total number of planctomycete cells in the anoxic layer could be detected if the probe PLA886, whose target specificity is restricted to taxonomically characterized aerobic planctomycetes of the genera Gemmata, Planctomyces, Pirellula, and Isosphaera, was used alone. Other planctomycete cells in this layer were detected only with the probe PLA46, which possesses a much wider scope. This suggests the affiliation of these organisms with a yet undescribed phylogenetic subgroup within the Planctomycetes.     

Effects of enhanced UV-B radiation on production-related properties of a Sphagnum fuscum dominated subarctic bog

1. The aim of the study was to investigate effects of enhanced UV-B radiation on the balance between biomass production and decay in an ombrotrophic bog which is dominated by one species of Sphagnum ( S. fuscum). This paper concerns production.
2. Enhanced UV-B radiation (simulating 15% ozone depletion under clear sky conditions) was applied by means of fluorescent tubes during two growing seasons.
3. In S. fuscum, shoot density, mass relations and length increment over time were measured and productivity was estimated. Pigment concentration, rates of dark respiration and maximum net photosynthesis were recorded.
4. Sphagnum fuscum productivity was not changed by enhanced UV-B radiation while properties determining production were highly influenced although in opposite directions.
5. Height increment was decreased by 20% in the first growing season and by 31% in the second growing season under enhanced UV-B radiation. After two growing seasons spatial shoot density was decreased by 8% by enhanced UV-B radiation. The shoots became stunted as capitulum dry mass and stem dry mass per unit length were increased by 21 and 17%, respectively, under enhanced UV-B radiation.
6. Dark respiration was significantly decreased by 31% after growth under enhanced UV-B radiation.
7. The UV-B induced change in shoot biometry together with the reduced spatial shoot density involve potential long-term effects on peat structure with possible feedback on productivity, decomposition and the strength of the system as a carbon sink.     

Summer warming and increased winter snow cover affect Sphagnum fuscum growth, structure and production in a sub-arctic bog

Sphagnum mosses form a major component of northern peatlands, which are expected to experience substantially higher increases in temperature and winter precipitation than the global average. Sphagnum may play an important role in the responses of the global carbon cycle to climate change. We investigated the responses of summer length growth, carpet structure and production in Sphagnum fuscum to experimentally induced changes in climate in a sub‐arctic bog. Thereto, we used open‐top chambers (OTCs) to create six climate scenarios including changes in summer temperatures, and changes in winter snow cover and spring temperatures. In winter, the OTCs doubled the snow thickness, resulting in 0.5–2.8°C higher average air temperatures. Spring air temperatures in OTCs increased by 1.0°C. Summer warming had a maximum effect of 0.9°C, while vapor pressure deficit was not affected. The climate manipulations had strong effects on S. fuscum. Summer warming enhanced the length increment by 42–62%, whereas bulk density decreased. This resulted in a trend (P<0.10) of enhanced biomass production. Winter snow addition enhanced dry matter production by 33%, despite the fact that the length growth and bulk density did not change significantly. The addition of spring warming to snow addition alone did not significantly enhance this effect, but we may have missed part of the early spring growth. There were no interactions between the manipulations in summer and those in winter/spring, indicating that the effects were additive. Summer warming may in the long term negatively affect productivity through the adverse effects of changes in Sphagnum structure on moisture holding and transporting capacity. Moreover, the strong length growth enhancement may affect interactions with other mosses and vascular plants. Because winter snow addition enhanced the production of S. fuscum without affecting its structure, it may increase the carbon balance of northern peatlands.     

Response of a Sphagnum bog plant community to elevated CO2 and N supply

The factors determining herbaceous canopy architecture are poorlyunderstood, especially in natural and semi-natural plant communities. Inthis study, we tested three main hypotheses: (1) the structure of herbaceouscanopies can be explained by the vertical distribution of functional groupsdefined by leaf width and the presence/absence of leaves on upright stem;(2) the degree of canopy stratification is greater in habitats that experiencelower spatial heterogeneity in the supply of light (i.e., grasslands as opposedto forest herb layers); and (3) there is significant variation among specieswithin a growth-form, with respect to their vertical position in thecanopy. We used plant foliage height distribution data from 14 grassland and 13forest herbaceous communities to test these hypotheses. A general linear mixedmodel was applied to specify the proportions of total variance in the foliageheight, accounted for by the fixed effects of plants' basicgrowth-form properties (growth-form) and community type(forest/grassland), and by the random effects of sampling site, samplingpoint, and individual species. We were also interested in the correlation ofthedegree of the stratification with various community characteristics(productivity, other canopy properties, species richness, variation ofspecies' traits) and light availability. There was some evidence ofoverall canopy stratification according toplant growth-form, since plants with leafy stem were locatedsignificantly higher. However, such a pattern of two more or less distinctlayers (grasses + upright forbs and rosette forbs) occurred withconsistency only in grasslands (greater homogeneity in light). Thebetween-species variation within a growth-form was a highlysignificant predictor of canopy vertical structure in the 27 communities. Theproportion of total observed variance, explainable throughspecies-specific effects, was comparable to that caused bybetween-site differences. The effect of community horizontal pattern wasless obvious, but still significant. The site by site analysis revealed thatthe degree to which horizontalpatchiness explained variation in vertical canopy structure was negativelyrelated to the relative importance of species-specific effects, showingthat small between-species differences lead to a more obviouswithin-community horizontal pattern, and vice versa. The upper bound ofthe degree of foliage stratification, according to growth-form, wasrelated to the variability of species light requirements and to relative (tocommunity pool size) richness, indicating that certain aspects of canopyarchitecture might be explained through community species composition anddiversity pattern.     

Cranial variables as predictors of hominine body mass

Body mass is a key variable in investigating the evolutionary biology of the hominines (Australopithecus, Paranthropus, and Homo). It is not only closely related to life-history parameters but also provides a necessary baseline for studies of encephalization or megadonty. Body mass estimates are normally based on the postcranial skeleton. However, the majority of hominid fossils are cranio-dental remains that are unassociated with postcranial material. Only rarely can postcranial material be linked with craniodentally defined hominid taxa. This study responds to this problem by evaluating body mass estimates based on 15 cranial variables to determine whether they compare in reliability with estimates determined from postcranial variables. Results establish that some cranial variables, and particularly orbital area, orbital height, and biporionic breadth, are nearly as good mass predictors for hominoids as are some of the best postcranial predictors. For the hominines in particular, orbital height is the cranial variable which produces body mass estimates that are most in line with postcranially generated estimates. Both orbital area and biporionic breadth scale differently in the hominines than they do in the other hominoids. This difference in scaling results in unusually large estimates of body mass based on these variables for the larger-sized hominines, although the three cranial variables produce equivalent predicted masses for the smaller-bodied hominines. © 1994 Wiley-Liss, Inc.     

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.     

Biomass, productivity and relative rate of photosynthesis of Sphagnum at different water levels on a South Swedish peat bog

The distribution pattern of Sphagnum species on bogs follows a hummock-hollow gradient. S . Sect. Acutifolia (that is in this study S. fuscum and S. rubellum combined) dominates hummock tops, ca 20 cm above the maximum water level with a green biomass of 50 g m⁻², S. magellancium dominates at a lower level, about 5 cm above the water level with a green biomass of 75 g m⁻² and S. cuspidatum dominates in the wettest hollows with a green biomass of about 50 g m⁻².
In situ measurements of length growth of S . Sect. Acutifolia and S. magellanicum using a ¹⁴CO₂-labelling technique during three consecutive years, revealed an unexpectedly high between-year variation in length growth of 7-23 mm yr⁻¹, and 16-22 mm yr⁻¹, respectively. Consequently the dominating producer in the transition between hummock and hollow changes from year to year, probably depending on climatic conditions.
In vitro experiments on the effects of different water levels of 2, 5, 10 and 20 cm below the moss surface, on photosynthetic activity of S . Sect. Acutifolia and S. magellanicum , measured by a second ¹⁴CO₂-technique, indicate optimal conditions for S. magellanicum at 10 cm above water level, and for S . Sect, Acutifolia at 20 cm above water level.
Differences in capillary water transport capability between the species are more important than the sensitivity of photosynthesis to water stress in explaining field patterns of productivity and distribution.     

Isolation and characterization of nitrogen-fixing bacteria of the genus Azospirillum from the soil of a Sphagnum peat bog

The presence of nitrogen-fixing bacteria of the genus Azospirillum in the soils of acidic raised Sphagnum bogs is revealed for the first time. Three Azospirillum strains, B2, B21, and B22, were isolated as a component of methane-oxidizing enrichment cultures, whereas attempts to isolate them directly from peat samples have failed. The results of comparative analysis of the nucleotide sequences of 16S rRNA genes, DNA-DNA hybridization, and the analysis of the sequences of the functional genes encoding nitrogenase and ribulose-1, 5-bisphosphate carboxylase reveal that all the newly obtained strains can be classified as Azospirillum lipoferum. Yet, unlike A. lipoferum. the isolates do not require biotin and utilize sucrose, inositol, and glycerol for growth. The cell morphology of strain B2 differs from that of the type strain and strains B21 and B22. The results obtained indicate the variability of morphological, physiological, and biochemical properties in closely related Azospirillum strains and suggest the existence of metabolic relationships between methanotrophic bacteria and the representatives of the genus Azospirillum under peat bog conditions.     

Long-term enhanced nitrogen deposition increases ecosystem respiration and carbon loss from a Sphagnum bog in the Scottish Borders

Nitrogen (N) deposition has increased in the last few decades with implications for the functioning of Sphagnum mosses, the main peat forming genus in peatlands. However, there are few in situ measurements of the carbon balance, especially where the N additions have been realistically manipulated in the field, and none with respect to the effect of N form. The aim of this study was to look at the effects of experimental N additions as oxidized or reduced N, with and without phosphorus (P) and potassium (K), on CO₂ fluxes from Sphagnum capillifolium hummocks in a long-term N addition experiment, Whim bog, in the Scottish Borders. In situ static chamber measurements were made during 2008 on 20 plots (control, and N treatments receiving 56 kg N ha⁻¹ y⁻¹ of either nitrate (NO₃⁻) or ammonium (NH₄⁺) added, with and without PK) to assess N effects on CO₂ exchange. Almost all the measured fluxes were negative, i.e. Sphagnum hummocks lost CO₂ to the atmosphere, irrespective of the treatments applied. N treatment did not have a significant effect on ecosystem respiration (ER) or net ecosystem productivity (NEP) but adding PK with N increased gross photosynthesis (P_G) significantly, compared to the other treatments. Summed monthly averages of NEP for each treatment indicated that increasing N deposition increased CO₂ loss from the system. The form of N affected the response: compared to the control, adding nitrate increased the CO₂ loss more than ammonium, both with and without PK. Nitrogen (both forms) increased the ecosystem respiration fluxes at a certain temperature, adding PK with N further enhanced the response. The positive (increasing) temperature response of ecosystem respiration with N suggests that in high N deposition areas the potential increase in ecosystem respiration, CO₂ loss, will be exacerbated with climate change.     

Photoinhibition as a control on photosynthesis and production of Sphagnum mosses

The effect of high light intensity on photosynthesis and growth of Sphagnum moss species from Alaskan arctic tundra was studied under field and laboratory conditions. Field experiments consisted of experimental shading of mosses at sites normally exposed to full ambient irradiance, and removal of the vascular plant canopy from above mosses in tundra water track habitats. Moss growth was then monitored in the experimental plots and in adjacent control areas for 50 days from late June to early August 1988. In shaded plots total moss growth was 2–3 times higher than that measured in control plots, while significant reductions in moss growth were found in canopy removal plots. The possibility that photoinhibition of photosynthesis might occur under high-light conditions and affect growth was studied under controlled laboratory conditions with mosses collected from the arctic study site, as well as from a temperate location in the Sierra Nevada, California. After 2 days of high-light treatment (800 mol photons m^–2 s^–1) in a controlled environmental chamber, moss photosynthetic capacity was significantly lowered in both arctic and temperate samples, and did not recover during the 14-day experimental period. The observed decrease in photosynthetic capacity was correlated (r ²=0.735, P<0.001) with a decrease in the ratio of variable to maximum chlorophyll fluorescence (F _v/F _m) in arctic and temperate mosses. This relationship indicates photoinhibition of photosynthesis in both arctic and temperate mosses at even moderately high light intensities. It is suggested that susceptibility to photoinhibition and failure to photoacclimate to higher light intensities in Sphagnum spp. may be related to low tissue nitrogen levels in these exclusively ombrotrophic plants. Photoinhibition of photosynthesis leading to lowered annual carbon gain in Sphagnum mosses may be an important factor affecting CO₂ flux at the ecosystem level, given the abundance of these plants in Alaskan tussock tundra.     

Phylogenetic analysis and in situ identification of bacteria community composition in an acidic Sphagnum peat bog

The Bacteria community composition in an acidic Sphagnum peat bog (pH 3.9 to 4.5) was characterized by a combination of 16S rRNA gene clone library analysis, rRNA-targeted fluorescence in situ hybridization (FISH), and cultivation. Among 84 environmental 16S rRNA gene clones, a set of only 16 cloned sequences was closely related (>or=95% similarity) to taxonomically described organisms. Main groups of clones were affiliated with the Acidobacteria (24 clones), Alphaproteobacteria (20), Verrucomicrobia (13), Actinobacteria (8), Deltaproteobacteria (4), Chloroflexi (3), and Planctomycetes (3). The proportion of cells that hybridized with oligonucleotide probes specific for members of the domains Bacteria (EUB338-mix) and Archaea (ARCH915 and ARC344) accounted for only 12 to 22% of the total cell counts. Up to 24% of the EUB338-positive cells could be assigned by FISH to specific bacterial phyla. Alphaproteobacteria and Planctomycetes were the most numerous bacterial groups (up to 1.3x10(7) and 1.1x10(7) cells g-1 peat, respectively). In contrast to conventional plating techniques, a novel biofilm-mediated enrichment approach allowed us to isolate some representatives of predominant Bacteria groups, such as Acidobacteria and Planctomycetes. This novel strategy has great potential to enable the isolation of a significant proportion of the peat bog bacterial diversity.     

Experimental climate effect on seasonal variability of polyphenol/phenoloxidase interplay along a narrow fen–bog ecological gradient in Sphagnum fallax

Extracellular phenoloxidase enzymes play an important role in the stability of soil carbon storage by contributing to the cycling of complex recalcitrant phenolic compounds. Climate warming could affect peatland functioning through an alteration of polyphenol/phenoloxidase interplay, which could lead them to becoming weaker sinks of carbon. Here, we assessed the seasonal variability of total phenolics and phenoloxidases subjected to 2–3 °C increase in air temperature using open‐top chambers. The measurements were performed along a narrow fen–bog ecological gradient over one growing season. Climate warming had a weak effect on phenoloxidases, but reduced phenolics in both fen and bog areas. Multivariate analyses revealed a split between the areas and also showed that climate warming exacerbated the seasonal variability of polyphenols, culminating in a destabilization of the carbon cycle. A negative relationship between polyphenols and phenoloxidases was recorded in controls and climate treatments suggesting an inhibitory effect of phenolics on phenoloxidases. Any significant decrease of phenolics through repeatedly elevated temperature would greatly impact the ecosystem functioning and carbon cycle through an alteration of the interaction of polyphenols with microbial communities and the production of extracellular enzymes. Our climate treatments did not have the same impact along the fen–bog gradient and suggested that not all the peatland habitats would respond similarly to climate forcing.     

遮阴对两种泥炭藓植物生长及相互作用的影响

以大泥炭藓和喙叶泥炭藓为材料,研究遮阴对其生长及植物相互作用的影响.结果表明:在单种群中,遮阴处理明显促进了大泥炭藓的高生长,但对喙叶泥炭藓的生长以及大泥炭藓生物量和分枝数未产生影响;在混合群中,喙叶泥炭藓抑制了大泥炭藓生物量和分枝数的增长,而大泥炭藓对喙叶泥炭藓的生长无影响.随遮阴胁迫的增加,邻体对喙叶泥炭藓竞争加剧,当胁迫进一步增强,邻体效应有转变为正相互作用的趋势,但邻体对大泥炭藓的效应始终为竞争,未随胁迫增加而变化.