Negative Plant–Soil Feedback and Positive Species Interaction in a Herbaceous Plant Community

Increasing evidence shows that facilitative interaction and negative plant–soil feedback are driving factors of plant population dynamics and community processes. We studied the intensity and the relative impact of negative feedback on clonal growth and seed germination of Scirpus holoschoenus, a ‘ring’ forming sedge dominant in grazed grassland, and the consequences for species coexistence. The structure of aboveground tussocks was described. A Lithium tracer assessed belowground distribution of functional roots. Seed rain and seedling emergence were compared for different positions in relation to Scirpus tussocks. Soil bioassays were used to compare growth on soil taken from inside and outside Scirpus tussocks of four coexisting species (Mentha acquatica, Pulicaria dysenterica, Scirpus holoschoenus and Dittrichia viscosa). We also compared plant performance of dominant plant species inside and outside Scirpus tussocks in the field. The ‘ring’ shaped tussocks of S. holoschoenus were generated by centrifugal rhizome development. Roots were functional and abundant under the tillers and extending outside the tussocks. The large roots mats that were present in the inner tussock zone were almost all dead. Seedling emergence and growth both showed a strong negative feedback of Scirpus in the inner tussock zone. Scirpus clonal development strongly reduced grass biomass. In the degenerated tussock zone, Pulicaria and Mentha mortality was lower, and biomass of individual plants and seed production were higher. This positive indirect interaction could be related to species-specific affinity to soil conditions generated by Scirpus, and interspecific competitive release in the degenerated tussock zone. We conclude that Scirpus negative feedback affects its seedling emergence and growth contributing to the development of the degenerated inner tussock zone. Moreover, this enhances species coexistence through facilitative interaction because the colonization of the inner tussock zone is highly species-specific.     

Mechanism of facilitation by sedge and cotton-grass tussocks on seedling establishment in a post-mined peatland

In stressful and disturbed ecosystems, seedling establishment may be facilitated by early colonizing plants. We examined the mechanism of such facilitation by tussock-forming species (Carex middendorffii and Eriophorum vaginatum), focusing on the independent and interactive effects of tussock litter and tussock mound substrate. Shading by litter on tussock mounds provides a stable but dryer substrate that may negatively affect early colonizers, owing to the co-occurrence of light deficiency and limited water availability, but positively affect late colonizers by subsequent amelioration of water availability. We used seed sowing and seedling transplant experiments with un-manipulated tussocks and manipulated shading?×?tussock mounds to examine seedling emergence, survival, and the biomass of early (Moliniopsis japonica) and late (Lobelia sessilifolia) colonizers in a post-mined peatland in northern Japan. Carex and Eriophorum tussocks facilitated seedling emergence and the growth of M. japonica and L. sessilifolia. Manipulation experiments indicated that the major positive effect was in providing stable substrates for seeds and seedlings. While the survival and growth of both colonizers were unaffected by shading alone and were negatively affected by tussock mounds alone, shading on tussock mounds decreased both the survival and growth in M. japonica but increased it in L. sessilifolia. Overall, tussock mounds with litter shading accelerated seedling establishment, especially that of late colonizers, in post-mined peatland. Our results indicate that the importance of facilitation mechanisms, for early successional plant composition that result from independent and interactive processes that co-occur as environmental conditions change.     

Mineral nutrient economy in competing species of Sphagnum mosses

Bog vegetation, which is dominated by Sphagnum mosses, depends exclusively on aerial deposition of mineral nutrients. We studied how the main mineral nutrients are distributed between intracellular and extracellular exchangeable fractions and along the vertical physiological gradient of shoot age in seven Sphagnum species occupying contrasting bog microhabitats. While the Sphagnum exchangeable cation content decreased generally in the order Ca²⁺ ≥ K⁺, Na⁺, Mg²⁺ > Al³⁺ > NH₄ ⁺, intracellular element content decreased in the order N > K > Na, Mg, P, Ca, Al. Calcium occurred mainly in the exchangeable form while Mg, Na and particularly K, Al and N occurred inside cells. Hummock species with a higher cation exchange capacity (CEC) accumulated more exchangeable Ca²⁺, while the hollow species with a lower CEC accumulated more exchangeable Na⁺, particularly in dead shoot segments. Intracellular N and P, but not metallic elements, were consistently lower in dead shoot segments, indicating the possibility of N and P reutilization from senescing segments. The greatest variation in tissue nutrient content and distribution was between species from contrasting microhabitats. The greatest variation within microhabitats was between the dissimilar species S. angustifolium and S. magellanicum. The latter species had the intracellular N content about 40% lower than other species, including even this species when grown alone. This indicates unequal competition for N, which can lead to outcompeting of S. magellanicum from mixed patches. We assume that efficient cation exchange enables Sphagnum vegetation to retain immediately the cationic nutrients from rainwater. This may represent an important mechanism of temporal extension of mineral nutrient availability to subsequent slow intracellular nutrient uptake.     

Regeneration of native vascular plants facilitated by Ischaemum aristatum var. glaucum tussocks: an experimental demonstration

Facilitation of a perennial tussock grass, Ischaemum aristatum var. glaucum, was hypothesized to be the provision of germination safe sites against inundation. We conducted field census and seed addition experiments to test this hypothesis. Facilitation effects of I. aristatum var. glaucum tussocks were obvious; in contrast to 280 seedlings/m² of ten native species observed on tussocks, seedlings hardly emerged on bare ground, even after seed addition. Although effects of moss occurrence at tussocks were not significant on the total number or species richness of emerged seedlings, significantly positive effects were observed on the seedling survival of some species, including endangered species. Conservation of facilitators will efficiently ensure the regeneration success of native vascular plants.     

Plant communities affect the species–area relationship on Carex sempervirens tussocks

The species–area relationship (SAR) has been extensively studied in a wide range of plant communities, but very few studies have directly addressed how plant communities affect the SAR and what are the underlying mechanisms. Many graminoids form distinct tussocks where many other plant species grow, but no study has investigated whether the SAR holds true for the vegetation on tussocks. In four plant communities on an abandoned subalpine pasture in the Swiss National Park, we made releves on 600 tussocks of Carex sempervirens and measured tussock basal area and other tussock traits. In all four communities, species richness on C. sempervirens tussocks was strongly positively related to tussock basal area (R²0.74), while other tussock traits explained very little (R²<0.04). Slope and intercept of the SAR on C. sempervirens tussocks differed significantly among the four communities. This was because plant communities affected richness in smaller tussocks (basal diameter <10 cm) but not that in large tussocks (basal diameter10 cm). We conclude that the SAR holds true for vegetation on C. sempervirens tussocks and changes with plant communities. Changes in the SAR on C. sempervirens tussocks are very likely because smaller tussocks are less independent of the plant communities than the larger ones, regarding disturbance or nutrients.     

Kobresia tibetica tussocks facilitate plant species inside them and increase diversity and reproduction

Many plant species grow inside tussocks of some graminoids, but the underlying mechanisms remain unclear. We address whether some species occur and flower mainly inside tussocks so that species diversity and sexual reproduction are higher inside than outside tussocks, and whether relieving biological and physical stress is the mechanism associated with the facilitative process. In a heavily grazed grassland on the eastern Tibetan Plateau, where both physical (due to high altitude) and biological conditions (due to heavy grazing) are extremely harsh, we investigated vegetation in paired plots inside and outside 150 Kobresia tibetica tussocks and measured tussock basal area (=plot area). We also measured temperatures at soil surface, 5 and 10 cm depth and the number of animals (yaks, sheep and horses) grazing inside and outside tussocks. Sixty-seven percent of the species occurred and 42% flowered more frequently inside than outside tussocks, but none less frequently. Inside tussocks 78% species flowered, but outside tussocks only 31% did. Consequently, number of species, number of flowering species and number of inflorescences were all markedly larger inside than outside tussocks. Differences in number of species, number of flowering species and number of inflorescences inside and outside tussocks increased with increasing tussock basal area. Soil temperatures were lower inside than outside tussocks, but grazing intensity was much larger outside tussocks. Therefore, tussocks of K. tibetica facilitated the species inside them likely by grazing prevention, but not by increasing warmth. This study provides evidence that plant species colonizing tussocks of graminoids can be facilitated by the tussock species, and facilitation by grazing prevention may be one mechanism causing the coexistence of the species inside tussocks.     

Multiclonal tussocks in the grass Achnatherum splendens (Trinius) Nevskia (Poaceae)

Clonal plant species often form genetically diverse populations, even when sexual reproduction in a population is rarely observed. Here we test whether the spatially discrete clusters of plants (tussocks of graminoids) formed within populations of some clonal species can likewise be multiclonal. We sampled leaves of ramets (shoots) within 20 tussocks of the grass Achnatherum splendens in the Otindag Sandland in Inner Mongolia, China, and genotyped the ramets using standard molecular protocols. The 20 tussocks were allocated to three classes: (i) small, circular, (ii) large, circular and (iii) large, irregular. Most tussocks (80%) were multiclonal and some contained at least eight different clones. Irregularly shaped tussocks contained twice as many clones as circular tussocks; neither size nor cover within a tussock affected number of clones per tussock, and the smaller clones in a tussock showed no tendency to occur on the edge or near the center of a tussock. These patterns seem more consistent with formation of multiclonal tussocks by coalescence than by colonization. Therefore, individual tussocks, especially large, irregular ones, cannot a priori be treated as genetic individuals without assessing their genetic information in, e.g., population demography, genetics and evolution studies.     

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).     

Light field heterogeneity among tussock grasses: Theoretical considerations of light harvesting and seedling establishment in tussocks and uniform tiller distributions

Although the tussock growth form of caespitose graminoids is widespread, the effect of this growth form on light interception and carbon gain of tillers has received little attention. Daily incident photosynthetic photon flux density (PFD_inc) and carbon gain in monospecific stands of tussock grasses were compared with those of a hypothetical distribution with the equivalent tiller density per total ground area, but evenly distributed rather than clumped in tussocks. This was computed for two tussock grasses Pseudoroegneria spicata (Pursh) A. Löve (bluebunch wheatgrass) and Agropyron desertorum (Fisch, ex Link) Schult. (creasted wheatgrass) at different plant densities. Daily PFD_inc and net photosynthesis (A) were greater if tillers were distributed uniformly rather than clumped in tussocks, except when the density of tussocks was so great as to approach a uniform canopy. When tussock density per ground area was low, much of the difference between tussock and uniform tiller densities in PFD_inc and A was due to shading within the tussocks; up to 50–60% of the potential carbon gain was lost in A. desertorum due to shading within tussocks. In a matrix of tussocks, the light field for establishing seedlings was very heterogeneous; potential A ranged from 7 to 96% relative to an isolated seedling. The mean of daily PFD_inc and A for seedlings in a tussock stand were nearly identical to the values in corresponding stands of uniform tiller distributions. It is hypothesized that the loss of A resulting from clumping tillers into tussocks is offset by benefits of protecting sequestered belowground resources from invasion by seedlings of competitors.     

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.

     

Avian predation at a Southern Rockhopper Penguin colony on Staten Island,Argentina

We studied predation risk in relation to nest location and subcolony size in Southern Rockhopper Penguins (Eudyptes chrysocome chrysocome) during the chick-rearing period. Striated Caracaras (Phalcoboenus australis), the main predator, preferentially attacked from tussock grasses which are found in the periphery of all subcolonies (peripheral tussocks) and often scattered within them (central tussocks). The greatest numbers of predation and attempted predation events were observed on nests in the periphery of the subcolony next to peripheral tussocks, and on those nests next to central tussocks. Central tussocks offer Striated Caracaras an additional “edge” area from which to prey, much in the same way as do the peripheral tussocks. Predation rate per individual was not correlated with subcolony size possibly due to the presence of central tussocks which, by creating an extra edge area, change the subcolony shape. There is a suggestion (P = 0.06) of increased probability of nest success with subcolony size.     

Fire temperatures and postfire plant community dynamics in Ecuadorian grass páramo

Three aspects of the páramo vegetation's response to fires were investigated: the measurement of fire temperatures, general observations of changes in plant communities following fires, and monitoring the fate of individual plants after burning.Fire temperatures were strongly influenced by the physiognomy of the vegetation, dominated by tussocks of Calamagrostis spp. Temperatures were highest amongst the upper leaves of the tussock (sometimes >500°C). The middle levels of the tussock experienced temperatures in excess of 400°C, but in the dense leaf bases temperatures were often below 65°C. On the ground between tussocks, temperatures were variable, whereas 2 cm below ground temperatures failed to reach 65°C.Plant survival depended on the intensity of the fire and the plant's position within the tussock structure. Survival was often the result of high temperature avoidance (with buds shielded by other plant parts or buried beneath the soil surface).Post-fire Calamagrostis tiller mortality rates were high and tussock regrowth was slow. Some other species appear to maintain their populations by exploiting this recovery phase for seedling establishment on tussocks.Between tussocks, changes of occupancy at the level of the individual plants were greater after fire than in control vegetation. Most transitions were random. Those which departed from random often involved gaps and were related to post-fire mortality, regrowth from below-ground parts, colonisation or, in the case of a clonal mat-forming species, to spatial rearrangement of rosettes. Recovery was slower at higher altitude. Recovery was much slower in burned plots when the upper 2 cm of soil was removed (along with buried plant parts) compared with burned plots.Qualitative observations suggest that recovery may consist of a cyclical process, mediated by the serial dominance of several species that are physiognomically important.The frequency of fires determines the amount of fuel accumulated within grass tussocks and some plants may be unable to survive repeated burning. Chance survival of species in unburned patches of vegetation and random colonisation of gaps may be important determinants of subsequent community structure.     

Vegetative structure,microclimate, and leaf growth of a páramo tussock grass species,in undisturbed,burned and grazed conditions

In neotropical alpine grasslands (páramo), the natural tussock grass vegetation is extensively grazed and occasionally burned. The low productivity of the tussock grass seems to be the reason for the disappearance of this growth form in the most frequently intervened areas. The structure, microclimate and leaf elongation rates of new emerging leaves were studied for the dominant tussock grass species Calamagrostis effusa, at an undisturbed, a moderately grazed (7 year after fire) and a heavily grazed (3.5 years after fire) site. In absence of grazing and burning, the tussocks had a high standing crop (1.07±0.09 kg DW · m^-2) and leaf area per projected tussock cover (LAI: 9.6±1.4). Two thirds of the total mass was dead and more than half of the leaves were in horizontal position. The tussock growth form protects the meristems from severe climatic conditions. At midday, the temperature was higher at meristem level than in the rest of the tussock. At this level, photosynthetic irradiance (PI) was almost extinct at 2.9±0.74% of PI above the vegetation. The red/far red ratio (R/FR) was strongly decreased. Initial leaf elongation of new born leaves was 2.3 mm · day^-1, and constant during the year; estimated net annual production was 198±73.8 g m^-2. At the moderately grazed and the heavily grazed study sites, the tussocks were smaller, greener and more erect than those at the undisturbed site. More PI reached the meristems and R/FR was higher at the base of grazed tussocks. Leaf elongation rates were lower. Most of the litter disappeared during the fires. The lower elongation rate of leaves in the grazed areas might be a response to defoliation, resulting in increased tillering and a lack growth associated with poor temperature insulation and more UV-B damage.     

Vegetation regeneration on blanket mire after mechanized peat-cutting

• 1 Blanket mire in Northern Ireland is an ecologically threatened habitat in which land use for hand peat‐cutting, forestry and agriculture has had a major influence. A recent land use change is the introduction of tractor‐powered peat‐harvesting. In this paper, the effect of machine peat‐cutting on ombrotrophic blanket mire vegetation is assessed from a regional sample of cut and uncut plots.
• 2 Principal components analysis identified water‐table depth and grazing intensity as major factors influencing the species composition of uncut mire. A key variable affecting the composition of machine‐cut mire across the drainage gradient was the number of times cut, with multiple annual cutting causing progressive decreases in acrotelm depth, catotelm bulk density and plant cover. Ericaceous species and Sphagnum spp. were particularly sensitive to cutting, with Eriophorum angustifolium and Campylopus introflexus characteristic of multiple‐cut sites.
• 3 Redundancy analysis, with number of times cut partialled out, showed that recovery time accounted for a significant amount of variance in vegetation composition. Species that significantly increased in abundance with recovery time were Sphagnum spp., Odontoschisma sphagni , Erica tetralix and Drosera rotundifolia.
• 4 Sites cut frequently, or which were grazed, recovered more slowly. Recovery from cutting was partly dependent on the post‐cutting structure of the mire surface and the species that survive cutting. The rate of recovery on sites cut once, then abandoned, is relatively rapid compared with multiple‐cut sites where species colonization is constrained by bare compacted peat.

     

The effects of spatial habitat configuration on recruitment, growth and population structure in arctic Collembola

The population density and demography of five species of arctic Collembola were studied in a naturally patchy habitat, consisting of Carex ursinae tussocks with varying degrees of isolation. Focal predictor variables were those describing the spatial configuration of tussocks, including tussock size and isolation and the amount of habitat (cover) at a 1-m² scale surrounding each tussock population. The Collembola populations were heavily influenced by environmental stochasticity in the form of winter mortality and summer drought, and the influence of patchiness on population characteristics was evaluated in this context. The five species showed very different responses to the structuring effect of the habitat, depending on life history characteristics, mobility and habitat requirements. Population density was highly variable in both time and space. Spring densities indicated larger winter mortality compared to observations from a previous study, and the snow- and ice-free season from June to August only resulted in population growth for Folsomia sexoculata. In the other species, adult mortality must have been high as there was no net population growth despite observed reproduction. The exception was Hypogastrura viatica, whose population decline was more likely to have been the result of migration out of the study area. Cover was the most important variable explaining density. No pure area or isolation effects at the tussock level were detected, even in areas with very low habitat cover. Drought was probably an important mortality factor, as July was particularly warm and dry. Due to qualitative differences in the tussocks and the matrix substrate, desiccation risk would be higher during dispersal between tussocks. We suggest that increased dispersal mortality gave the observed pattern of increased density in relation to cover, both in general and in F. quadrioculata, an opportunistic species otherwise known for rapid population growth. Onychiurus groenlandicus, which had a similar density response to cover, may also be influenced by a rescue effect sustaining densities in areas with high cover. The cover effect can be viewed as a large-scale factor which encompasses the general spatial neighbourhood of each tussock, where inter-population processes are important, as opposed to internal patch dynamics. Received: 15 March 1999 / Accepted: 22 March 2000     

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.     

Seasonal and spatial variance of seed bank species composition in an oligotrophic wet meadow

Seasonal and small-scale spatial variation of the seed bank in an oligotrophic wet meadow (Molinion) was assessed by two methods: (1) direct determination of seeds extracted from soil samples (6 cm in diameter, 10 cm depth) and (2) germination of seeds according to Ter Heerdt et al. [1996. An improved method for seed bank analysis: seedling emergence after removing the soil by sieving. Funct. Ecol. 10, 144–151]. Samples were taken three times during the year (July and November 2001, March 2002) from three distinct microhabitats (moss layer, gaps, Molinia caerulea tussocks). Using the extraction method, more seeds were found in the upper 0–3 cm soil layer than in the deeper 3–7 cm layer, significant differences in the number of seeds were found among microhabitats (with the lowest seed bank densities in Molinia tussocks). The highest number of seeds was found in the spring census, but the test was not significant (P<0.10). However, seasonal variation was found when damaged seeds were included in the analysis. Changes in damaged/undamaged and light/heavy seed ratios, and in numbers of grass/herbs/sedges seeds are also discussed. The spatial distribution of seeds is probably influenced to a great degree by abiotic factors, such as microtopography (round and heavy seeds of sedges prevail in gaps and in deeper soil layers). The species composition of the seed bank varied among microhabitats, depth, and throughout the year. No differences were found by the seed germination method.     

Linking population density and habitat structure to ecophysiological responses in semiarid Spanish steppes

We have studied the underlying factors responsible for the heterogeneous ecophysiological status of a semiarid Stipa tenacissima L. steppe in a subcatchment of SE Spain by assessing population composition and habitat structure of S. tenacissima stands. To do this, we measured and estimated 18 variables (11 biotic and seven abiotic) in 15 plots randomly distributed in the subcatchment, and then zoned this area by plot affinity using PCA. This analysis produced three sectors determined mainly by S. tenacissima cover and soil depth variables. The linear relationship fitted between S. tenacissima tussock biomass and tussock density in monospecific stands (both logarithmic) indicated a curve close to −1, suggesting that the system is close to the maximum constant yield state. Ecophysiological measurements (gas exchange, fluorescence and individual leaf area index) were taken in two periods with different water availability in a representative plot in each sector. The intraspecific competition (inferred from the density dependence of green biomass) and rock outcrops were the main factors influencing the ecophysiological status in the study area. While, in the wet season, intraspecific competition regulated water consumption in zones where S. tenacissima tussocks (monospecific stands) are dominant, during the dry season, stands in zones with extensive rock outcrops and stone cover (tussocks in “soil pockets”) had no access to non-rainfall water gains because of the adjacent bare soil, and so in these stands, gas exchange was lower and photoinhibition higher. This article stresses the importance of considering the connection between tussocks and bare-ground interspace in the functional and structural analysis of semiarid steppes.     

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.