Energy and moisture considerations on cutover peatlands: surface microtopography, mulch cover and Sphagnum regeneration

This study examined (i) the effect of artificially created microtopography and straw mulch on the soil moisture and (ii) energy balance and the establishment of a Sphagnum cover on a cutover peatland. Straw mulch caused rainfall interception approaching 2 mm per event. Although interception represented 44% of the total rainfall over the measurement period, water that evaporated from the mulch used energy that would otherwise have been used to evaporate soil water. Thus, the net effect of interception by mulch was negligible. The soil heat flux below the mulch was only 13% of the bare soil value and was decoupled from the daily net radiation. Net radiation over the bare soil was 15% greater than over the mulch. However, because of the greater heat flux into the bare peat, the energy available for sensible and latent heat fluxes was similar between the mulch covered and bare peat. Average evaporation from mulch and bare soil was estimated to be 2.6 and 3.1 mm d⁻¹, respectively. Soil water tension 1 cm below the surface remained above −100 cm (mb) all season (100% of the time) when a mulch was used, compared to only 30% of the time in the bare soil. Correspondingly, the water table was sustained above the 40 cm depth, 60% of time in the mulch covered site, compared to only 40% of the time in the bare peat site. Negative relief elements of the microtopography were wetter and cooler than positive relief elements. However, when under a mulch, the negative relief elements provided no additional benefit, in terms of temperature or soil moisture amelioration. The control site with a mulch cover was equivalent or better than negative relief elements with a mulch cover. Taking into account the poorer performance of positive relief elements, even when mulch covered, the creation of surface microtopography reduced the overall moisture content of the site. Sphagnum established and spread only when the diaspores were protected with a straw mulch. All microtopography types tested had no effect on the establishment of Sphagnum mosses when the microtopography treatments, including positive and negative relief elements, were treated as a whole, although being in a depression helped Sphagnum establishment.     

On the Use of Shallow Basins to Restore Cutover Peatlands: Plant Establishment

Since the early 1990s, restoration techniques have been developed for milled and cutover peatlands in eastern Canada. These techniques are based on the active reintroduction of peatland plant diaspores, blocking drainage, and the use of straw mulch to improve surface conditions. This study examines the effectiveness of using shallow (20 cm deep) basins of various widths to improve the success of current peatland restoration techniques. It comprises three different experiments, each spanning three or four growing seasons and combining both small‐scale manual and large‐scale mechanized plant reintroductions. Cover data recorded in two of the experiments were regressed against a series of environmental factors to determine how Sphagnum establishment success was related to abiotic variables such as moisture, water ponding, surface roughness, and mulch cover. Results of these experiments demonstrate that shallow basins were generally effective at promoting Sphagnum establishment and that this effect extends beyond the positive impact that basins have on hydrological conditions. Basins of various widths were equally successful. Soil‐moisture content (linear positive effect) and duration and severity of flooding events (quadratic effect) were shown to be determinant of plant recovery. Other factors such as the density of straw cover (positive effect) and surface roughness (negative effect) were also instrumental in explaining local variation in Sphagnum cover. Plant cover after three and four growing seasons averaged 20–25% in mechanical reintroductions and 40–60% in manual reintroductions, demonstrating the overall effectiveness of the restoration techniques used.     

North American approach to the restoration of Sphagnum dominated peatlands

Sphagnum dominated peatlands do not rehabilitate well after being cutover (mined) for peat and some action needs to be taken in order to restore these sites within a human generation. Peatland restoration is recent and has seen significant advances in the 1990s. A new approach addressing the North American context has been developed and is presentedin this paper. The short-term goal of this approach is to establish a plant cover composed of peat bog species and to restore a water regime characteristic of peatland ecosystems. The long-term objective is to return the cutover areas to functional peat accumulating ecosystems. The approach developed for peatland restoration in North America involves the following steps: 1)field preparation, 2) diaspore collection, 3) diaspore introduction, 4) diaspore protection, and 5) fertilization. Field preparation aims at providing suitable hydrological conditions for diaspores through creation of microtopography and water retention basins, re-shaping cutover fields and blocking ditches. It is site specific because it depends largely onlocal conditions. The second step is the collection of the top 10 centimetres of the living vegetation in a natural bog as a source of diaspores. It is recommended to use a ratio of surface collected to surface restored between 1: 10 and 1: 15 in order to minimize the impact on natural bogs and to insure rapid plant establishment in less than four years. Diaspores are then spread as a thin layer on the bare peat surfaces to be restored. It has been demonstrated that too scant or too thick a layer decreases plant establishment success. Diaspores are then covered by a straw mulch applied at a rate of 3 000 kg ha^-1 which provides improved water availabilityand temperature conditions. Finally, phosphorus fertilization favours more rapid substrate colonization by vascular plants, which have been shown to help stabilize the bare peat surface and act as nurse plants to the Sphagnum mosses.     

Experimental restoration of a fen plant community after peat mining

Question: Which restoration measures (introduction of donor diaspore material, application of straw mulch, alteration of residual peat depths) contribute to the establishment of a fen plant community on minerotrophic surfaces after peat mining? Location: Rivière‐du‐Loup peatland, southern Québec, Canada at 100 m a.s.1. Methods: The effectiveness of introducing fen plants with the application of donor diaspore material was tested. The donor diaspore material, containing seeds, rhizomes, moss fragments, and other plant propagules, was collected from two different types of natural fens. We tested whether the application of straw mulch would increase fen species cover and biodiversity compared to control plots without straw mulch. Terrace levels of different peat depths (15 cm, 40 cm, and 56 cm) were created to test the effects of different environmental site conditions on the success of re‐vegetation. Results: Applying donor seed bank from natural fens was found to significantly increase fen plant cover and richness after the two growing seasons. Straw mulch proved to significantly increase fen plant richness. The intermediate terrace level (40 cm) had the highest fen plant establishment. Compared to reference sites, the low terrace level (15 cm) was richer in base cations, whereas the high terrace level (56 cm) was much drier. Conclusions: The application of donor diaspore material was demonstrated as an effective technique for establishing vascular fen plants. Further re wetting measures are considered necessary at the restoration site to create a fen ecosystem rather than simply restoring some fen species.     

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.     

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.     

不同覆盖施肥措施对黄土旱塬冬小麦土壤水分的影响

于2007年9月—2008年7月在位于黄土高原渭北旱塬的王东沟试验区进行冬小麦不同覆盖施肥措施（包括不施肥、农民习惯施肥、推荐施肥、推荐施肥+有机肥、推荐施肥+垄上覆膜、推荐施肥+垄上覆膜+沟内覆草、推荐施肥+全区覆草7个处理）田间试验,并采用水分中子仪定期观测土壤含水量,研究黄土高原旱塬区不同栽培措施下土壤水分的变化特征.结果表明：在干旱季节（春季）,推荐施肥+垄上覆膜+沟内覆草措施有利于贮存更多的土壤水分,其土壤储水量约比最低值（推荐施肥+有机肥）高48.2 mm,并可将土壤水分保持到冬小麦需水的关键期,而且推荐施肥+垄上覆膜措施仅次于推荐施肥+垄上覆膜+沟内覆草,表明这两种措施能够在田间蓄积较多天然降水,有利于黄土高原旱区雨养农业的发展.     

Sensitivity of C Sequestration in Reintroduced Sphagnum to Water-Level Variation in a Cutaway Peatland

The reintroduction of Sphagnum fragments has been found to be a promising method for restoring mire vegetation in a cutaway peatland. Although it is known that moisture controls Sphagnum photosynthesis, information concerning the sensitivity of carbon dynamics on water‐level variation is still scarce. In a 4‐year field experiment, we studied the carbon dynamics of reintroduced Sphagnum angustifolium material in a restored (rewetted) cutaway peatland. Cutaway peatland restored by Sphagnum reintroduction showed high sensitivity to variation in water level. Water level controlled both photosynthesis and respiration. Gross photosynthesis (P_G) had a unimodal response to water‐level variation with optimum level at ?12 cm. The range of water level for high P_G (above 60% of the maximum light‐saturated P_G) was between 22 and 1 cm below soil surface. Water level had a dual effect on total respiration. When the water level was below soil surface, peat respiration increased rapidly along the lowering water level until the respiration rate started to slow down at approximately ?30 cm. Contrary to peat respiration, the response of Sphagnum respiration to water‐level variation resembled that of photosynthesis with an optimum at ?12 cm. In optimal conditions, Sphagnum reintroduction turned the cutaway site from carbon source to a sink of 23 g C/m² per season (mid‐May to the end of September). In dry conditions, lowered photosynthesis together with the higher peat respiration led to a net loss of 56 g C/m². Although the water level above the optimum amplitude restricted CO₂ fixation, a decrease in peat respiration led to a positive CO₂ balance of 9 g C/m².     

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).     

Range change evolution of peat mosses (Sphagnum) within and between climate zones

Peat mosses (Sphagnum) hold exceptional importance in the control of global carbon fluxes and climate because of the vast stores of carbon bound up in partially decomposed biomass (peat). This study tests the hypothesis that the early diversification of Sphagnum was in the Northern Hemisphere, with subsequent range expansions to tropical latitudes and the Southern Hemisphere. A phylogenetic analysis of 192 accessions representing the moss class Sphagnopsida based on four plastid loci was conducted in conjunction with biogeographic analyses using BioGeoBEARS to investigate the tempo and mode of geographic range evolution. Analyses support the hypothesis that the major intrageneric clades of peat‐forming species accounting for >90% of peat moss diversity originated and diversified at northern latitudes. The genus underwent multiple range expansions into tropical and Southern Hemisphere regions. Range evolution in peat mosses was most common within latitudinal zones, attesting to the relative difficulty of successfully invading new climate zones. Allopolyploidy in Sphagnum (inferred from microsatellite heterozygosity) does not appear to be biased with regard to geographic region nor intrageneric clade. The inference that Sphagnum diversified in cool‐or cold‐climate regions and repeatedly expanded its range into tropical regions makes the genus an excellent model for studying morphological, physiological, and genomic traits associated with adaptation to warming climates.     

Release of inorganic N,P and K in peat soils; effect of temperature,water chemistry and water level

In the Netherlands, fens that are fed by polluted river water are often eutrophic, whereas fens fed by calcium-rich groundwater often are mesotrophic. Differences in trophic status can not always be attributed to differences in the nutrient load of the water. In this paper we try to determine if the inflow of river water in fens, in fact, accelerates the soil nutrient release, thereby creating more eutrophic conditions (‘internal eutrophication’). For this purpose, we compared nutrient release rates (N, P and K) in soil cores fromSphagnum peat andCarex peat saturated with different media, that were artificially created to mimic the three basic water sources: polluted river water, unpolluted calcium-rich groundwater and rainwater. In addition, we studied the effect of temperature and water level on nutrient release rates. The experiments proved thatSphagnum peat released much more P and ammonium thanCarex peat. The strong site effect proved consistent throughout the water chemistry treatments, which indicates that soil quality may be the most important agent determining nutrient release rates. Nevertheless, it was established that water chemistry and water level are of significant influence on nutrient release rates in peat soils. In particular, river water stimulated P release by the peat, most notably in theSphagnum peat. P-release in both soils was only minor when the soils were incubated in clean Ca-rich groundwater. It is suggested that P release is strongly associated with soil chemical processes, and that high P release rates after incubation in river water are due to the high sulphate content of the water. The net release from the soil of ammonium, potassium and phosphate increased with increasing temperature. A freezing treatment significantly increased nutrient availability. The results of the experiments are examined in the context of hydrologic management strategies for the conservation of fens in agricultural landscapes.     

Influence of temperature on the δ13C values and distribution of methanotroph‐related hopanoids in Sphagnum‐dominated peat bogs

Methane emissions from peat bogs are mitigated by methanotrophs, which live in symbiosis with peat moss (e.g. Sphagnum). Here, we investigate the influence of temperature and resultant changes in methane fluxes on Sphagnum and methanotroph‐related biomarkers, evaluating their potential as proxies in ancient bogs. A pulse‐chase experiment using ¹³C‐labelled methane in the field clearly showed label uptake in diploptene, a biomarker for methanotrophs, demonstrating in situ methanotrophic activity in Sphagnum under natural conditions. Peat cores containing live Sphagnum were incubated at 5, 10, 15, 20 and 25°C for two months, causing differences in net methane fluxes. The natural δ¹³C values of diploptene extracted from Sphagnum showed a strong correlation with temperature and methane production. The δ¹³C values ranged from ?34‰ at 5°C to ?41‰ at 25°C. These results are best explained by enhanced expression of the methanotrophic enzymatic isotope effect at higher methane concentrations. Hence, δ¹³C values of diploptene, or its diagenetic products, potentially provide a useful tool to assess methanotrophic activity in past environments. Increased methane fluxes towards Sphagnum did not affect δ¹³C values of bulk Sphagnum and its specific marker, the C₂₃ n‐alkane. The concentration of methanotroph‐specific bacteriohopanepolyols (BHPs), aminobacteriohopanetetrol (aminotetrol, characteristic for type II and to a lesser extent type I methanotrophs) and aminobacteriohopanepentol (aminopentol, a marker for type I methanotrophs) showed a non‐linear response to increased methane fluxes, with relatively high abundances at 25°C compared to those at 20°C or below. Aminotetrol was more abundant than aminopentol, in contrast to similar abundances of aminotetrol and aminopentol in fresh Sphagnum. This probably indicates that type II methanotrophs became prevalent under the experimental conditions relative to type I methanotrophs. Even though BHP concentrations may not directly reflect bacterial activity, they may provide insight into the presence of different types of methanotrophs.     

Fine‐scale five‐year vegetation change in boreal bog vegetation

Abstract. Within an ombrogenous part of N. Kisselbergmosen, Rødenes, SE Norway, fine‐scale changes in species abundance, successional trends relative to the main gradients (as represented by DCA axes), and co‐ordinated change within pairs of the bottom layer species are studied. Data sets were sampled twice with a five‐year interval, and included species abundance and cover of mud bottom, naked peat and litter in 436 sample plots (16 cm× 16 cm), and species abundance in 6976 subplots (4 cm× 4 cm). Depth from the surface of subplots to the water table was estimated in 1991. Most summers and growing seasons were somewhat drier than normal in the 5‐yr period. The area covered by mud‐bottom, naked peat and litter increased significantly, as did the frequencies of the dwarf shrubs Calluna vulgaris and Andromeda polifolia in hummocks and upper lawn. Sample plots were significantly displaced downward the peat productivity gradient (DCA 2), reflecting the reduced cover of many bottom layer species, including all Sphagnum spp. Significant coordinated changes in cover of bottom layer species are described. The changes observed in hummocks support the existence of a local regeneration cycle, as suggested by other researchers. Some of the vegetation changes seem parallel to those reported from areas with a higher nitrogen deposition, but it is not likely that nitrogen deposition alone is the major cause of the observed changes. Between‐year variation in population size and climatic fluctuations may as well explain the observed changes.     

Polytrichum Strictum as a Solution to Frost Heaving in Disturbed Ecosystems: A Case Study with Milled Peatlands

Substrate instability is a common problem in many disturbed ecosystems. In the case of milled harvested peatlands, the pioneer moss Polytrichum strictum is commonly found; it is well adapted to tolerate the harsh microclimatic conditions and peat instability of these sites. A field experiment was used to determine the effectiveness of P. strictum against frost heaving, a major type of disturbance on bare peat. Wooden dowels and fir trees (Abies balsamea) placed in a P. strictum carpet experienced almost no frost heaving, whereas heaving was severe on bare peat. Reintroduced P. strictum fragments thinly spread on bare peat reduced but did not eliminate frost heaving. Straw mulch (a protective cover often required in peatland restoration) effectively reduced heaving in the fall, but was less effective in the spring because it had partially decomposed. The P. strictum carpet, P. strictum fragments, and straw mulch reduced frost heaving by reducing the number of freeze–thaw cycles, by slowing the rate of ground thaw in the spring, and by reducing the unfrozen water content of the peat during the spring thaw. Different species of Polytrichum mosses should be considered for the restoration or regeneration of disturbed ecosystems where soil stability is problematic.     

Sphagnum growth under N saturation: interactive effects of water level and P or K fertilization

• Sphagnum biomass is a promising material that could be used as a substitute for peat in growing media and can be sustainably produced by converting existing drainage‐based peatland agriculture into wet, climate‐friendly agriculture (paludiculture). Our study focuses on yield maximization of Sphagnum as a crop.
• We tested the effects of three water level regimes and of phosphorus or potassium fertilization on the growth of four Sphagnum species (S. papillosum, S. palustre, S. fimbriatum, S. fallax). To simulate field conditions in Central and Western Europe we carried out a glasshouse experiment under nitrogen‐saturated conditions.
• A constant high water table (remaining at 2 cm below capitulum during growth) led to highest productivity for all tested species. Water table fluctuations between 2 and 9 cm below capitulum during growth and a water level 2 cm below capitulum at the start but falling relatively during plant growth led to significantly lower productivity. Fertilization had no effect on Sphagnum growth under conditions with high atmospheric deposition such as in NW Germany (38 kg N, 0.3 kg P, 7.6 kg K·ha^?1·year^?1).
• Large‐scale maximization of Sphagnum yields requires precise water management, with water tables just below the capitula and rising with Sphagnum growth. The nutrient load in large areas of Central and Western Europe from atmospheric deposition and irrigation water is high but, with an optimal water supply, does not hamper Sphagnum growth, at least not of regional provenances of Sphagnum.

     

Microclimatic buffering and resource‐based habitat in a glacial relict butterfly: significance for conservation under climate change

In contrast to several organisms that have already shown range shifts to the north as a response to climate change, southern populations of relict species are trapped in isolated altitudinal habitats. Therefore, there is a growing interest to better understand their habitat use, with particular attention to the thermal aspects and associated significance for their habitat management. We address this issue by a study of larval habitat use relative to vegetation structure and microclimate in a glacial relict butterfly of peat bog ecosystems, using a functional, resource‐based habitat approach. We analysed caterpillar presence and density relative to vegetation composition (reflecting gradients of humidity, temperature, and natural succession of the peat bog) and to the availability and quality of thermal refuges for caterpillars (i.e., structures provided by Sphagnum hummocks). We also tested caterpillar survival rates under different temperature and humidity treatments. We found that (1) Boloria aquilonaris was a specialist butterfly of early successional stages with very humid zones of peat bog, (2) the lack of Sphagnum hummocks reduced larval habitat suitability, and hence the population density, and (3) a reduction of the thermal buffering ability of Sphagnum hummocks was observed in less humid, degraded parts, or late‐successional stages of peat bog. A larval rearing experiment showed a significant impact of temperature on caterpillar survival; survival being higher at lower temperature. Our field and laboratory results support the idea that the thermal environment exploited by caterpillars should be considered as a functional resource and included in a population‐specific habitat definition. Appropriate management of the peat bog habitat of this glacial relict species should not exclusively focus on the larval and adult feeding resources, but also on the quality of thermal refuges provided by Sphagnum hummocks in humid zones of the peat bog, especially in the current critical context of climate warming.     

Development of microsatellites for the peat moss Sphagnum capillifolium using ISSR cloning

Sphagnum mosses are major components of peat bogs but populations of many species are under threat due to habitat fragmentation resulting from the cutting of peat for fuel. We have used an intersimple sequence repeat (ISSR)‐based cloning method to develop nine polymorphic nuclear microsatellites for the peat moss species Sphagnum capillifolium. Between three and seven alleles per locus were detected in a sample of 48 haploid gametophytes and levels of gene diversity ranged from 0.5391 to 0.7960. These represent the first microsatellite markers developed for this important genus and most also exhibited cross‐species amplification across a range of common Sphagnum species.     

Significant nonsymbiotic nitrogen fixation in Patagonian ombrotrophic bogs

Nitrogen (N) nutrition in pristine peatlands relies on the natural input of inorganic N through atmospheric deposition or biological dinitrogen (N₂) fixation. However, N₂ fixation and its significance for N cycling, plant productivity, and peat buildup are mostly associated with the presence of Sphagnum mosses. Here, we report high nonsymbiotic N₂‐fixation rates in two pristine Patagonian bogs with diversified vegetation and natural N deposition. Nonsymbiotic N₂ fixation was measured in samples from 0 to 10, 10 to 20, and 40 to 50 cm depth using the ¹⁵N₂ assay as well as the acetylene reduction assay (ARA). The ARA considerably underestimated N₂ fixation and can thus not be recommended for peatland studies. Based on the ¹⁵N₂ assay, high nonsymbiotic N₂‐fixation rates of 0.3–1.4 μmol N₂ g^?1 day^?1 were found down to 50 cm under micro‐oxic conditions (2 vol.%) in samples from plots covered by Sphagnum magellanicum or by vascular cushion plants, latter characterized by dense and deep aerenchyma roots. Peat N concentrations point to greater potential of nonsymbiotic N₂ fixation under cushion plants, likely because of the availability of easily decomposable organic compounds and oxic conditions in the rhizosphere. In the Sphagnum plots, high N₂ fixation below 10 cm depth rather reflects the potential during dry periods or low water level when oxygen penetrates the top peat layer and triggers peat mineralization. Natural abundance of the ¹⁵N isotope of live Sphagnum (5.6 δ‰) from 0 to 10 cm points to solely N uptake from atmospheric deposition and nonsymbiotic N₂ fixation. A mean ¹⁵N signature of ?0.7 δ‰ of peat from the cushion plant plots indicates additional N supply from N mineralization. Our findings suggest that nonsymbiotic N₂ fixation overcomes N deficiency in different vegetation communities and has great significance for N cycling and peat accumulation in pristine peatlands.     

Species diversity,niche metrics and species associations in harvested and undisturbed bogs

Abstract. Competition is considered an important force in structuring plant communities and in governing niche relations, but communities recovering from disturbance, may be less governed by species interactions and less orderly organized. To address this issue, we studied species richness, abundance and patterns of association between plant species at three spatial scales (1 m², 1/25 m², 1/625 m²) in two ombrotrophic mires in east-central Sweden. One was at a secondary successional stage following peat extraction 50 yr ago and the other was undisturbed. Peat extraction leads to a change in hydrology which is slowly restored by the formation of new peat. Niche breadth and niche overlap along the gradient of height above the water table were calculated for the five common Sphagnum species occurring at both mires in an attempt to better understand differences in species co-occurrence at each mire. Species cover differed between the mires, and the number of species per plot was higher in the undisturbed community at all scales, suggesting that the degree of species intermingling was greater than at the harvested site. At all scales, the number of non-random associations was higher, and niche overlap lower among ecologically similar species (e.g. hollow Sphagnum species) in the undisturbed mire. These differences indicate that random events are important in colonization, and that biotic interactions between neighbours later result in a higher degree of non-randomness. In addition, we surveyed a number of abandoned peat pit sites to test the effect of disturbance for species composition at a regional scale. Ombrotrophic peat pits contained several Sphagnum species normally associated with minerotrophic mires, and species of wooded mires occurred frequently in peat pits, making them more species-rich than undisturbed bogs. There were also Sphagnum species new to, or rare in, this part of Sweden which indicates effective long-distance dispersal. Even 50 yr after peat extraction had ceased, the vegetation had not recovered to its original composition.