An ecosystems approach to base-rich freshwater wetlands,with special reference to fenlands

A survey of base-rich wetlands in The Netherlands is presented. The main area of their occurrence is the low-lying Holocene part of the country, until some thousand years ago a large and coherent wetland landscape: the Holland wetland. The development of various parts of the Holland wetland into marshes, fens and bogs can be understood from hydrological relations in mire basins, as recognized in the distinction of primary, secondary and tertiary mire basin stages. Presently, the remnants of the Holland wetland are separate base-rich wetlands. The succession of their vegetation reflects various abiotic conditions and human influences. Three main developmental periods are distinguished as regards these factors. The first, geological period of mire development is seen as a post-glacial relaxation, with the inertia due to the considerable mass of wetland as a stabilizing factor. Biological “grazing” influences, as an aspect of utilization by humans, converted base-rich wetlands to whole new types in the second, historical period. Presently, mass and harvesting have decreased in importance, and actual successions in terrestrializing turbaries seem to reflect rapidly changing environmental conditions. Human control could well become the most important factor in the future development of wetland nature. The present value of open fen vegetation strongly depends on the continuation of the historical harvesting. The development of wooded fen may help to increase the mass of wetland in the future. Best results in terms of biodiversity are expected when their base state is maintained through water management. The vegetation and hydrology of floating fens in terrestrializing turbaries is treated in some more detail. Various lines and phases in the succession are distinguished. Open fen vegetation at base-rich, yet nutrient-poor sites is very rich in species threatened elsewhere. The fast acidification of certain such fens is attributed to hydrological and management factors. This acidification is illustrated in the profile of a floating raft sample. At the scale of these small fens, the elemental structure comprising base-rich fen, transitional fen and bog vegetation, is not as stable as it was in the large Holland wetland. A critical role seems to be played by the supply of bases with the water influx. The changing base state is supposed to change the nutrient cycling to such an extent that it would be correct to call this trophic excitation of the ecosystem, rather than just eutrophication. Eutrophication indicates a quantitative reaction to an increased nutrient supply, the internal system being unaltered. The drainage of fens, resulting in an increased productivity of the vegetation, provides another example of excitation, to the effect that the functional system is dramatically changed internally.     

Quick Target Vegetation Recovery after Restorative Shrub Removal and Mowing in a Calcareous Fen

Removal of shrubs and trees is an important management and restoration practice to promote openness and light‐dependent vegetation in fens, especially as tree cover is increasing in previously open wetlands. The effects of woody vegetation removal on target species have been poorly documented in wetlands up to now. In this study, I investigated the effect of tree and shrub removal (especially of Juniperus communis) on the target vegetation in a partly overgrown and degraded grazed rich fen after 6 years. I also tested whether additional intensified management by mowing could promote initial recovery. Shrub removal resulted in a rapid recovery of species‐rich fen vegetation such that after 6 years brown moss cover more than tripled and target species richness doubled and became similar to values of a reference area in a favorable conservation status. Additional mowing resulted in a much higher abundance of the target rich fen vascular plants. The reasons for the success at this site may be the proximity to well‐developed rich fen vegetation, presence of cattle that dispersed diaspores, and presence of bare, colonizable substrate. Thus, it may be more beneficial to restore and expand already existing sites in a partly favorable status than to restore severely deteriorated sites. Extensive management by woody vegetation removal may be an alternative method to maintain high conservation values of open mires and other wetlands, where grazing or mowing is not necessary or feasible to meet future needs in response to overgrowth caused by global warming.     

The relation between vegetation and soil chemistry gradients in a ground water discharge fen

Abstract. The soil chemistry of a headwater valley fen is influenced by local ground water discharge that supplies base cations and alkalinity to the fen. An irrigation canal just upward of the fen is the source of this alkalinity. The ecological consequences of this artificial system are studied both on the soil and vegetation level. Rich-fen species of the alliance Caricion davallianae are connected to soil water alkalinity and soil base status. They depend directly on the alkaline ground water discharge. In addition, the local input of this water causes a gradient-rich pattern from poor to rich fen, and it is therefore concluded that it is responsible for the presence of intermediate fen vegetation too. High nutrient levels in the irrigation water have not influenced the fen until now. This case study illustrates the possibility for rich fen restoration after acidification. Irrigation with alkaline water is efficient if excess nutrients can be removed.     

The effects of groundwater discharge,mowing and eutrophication on fen vegetation evaluated over half a century

Questions: Were continued groundwater discharge and mowing regimes sufficient for vegetation preservation from 1944 to 1993? Which has a stronger effect on vegetation development; groundwater discharge or mowing? What is the role of surface water eutrophication as driver of vegetation change? Location: Het Hol, The Netherlands (ca. 92 ha, 52°13′N, 5°05′E). Methods: Hydrology was simulated for the late 1940s, early 1960s and 1987. Vegetation maps (1944, 1960, 1975 and 1993) were compared for biotope cover. Vegetation recordings in 1944 and 1987 were compared. Surface water quality was compared between 1950 and 1987. Which sites were mown was reconstructed from an interview. Effects of periodic mowing and groundwater discharge on vegetation development were tested for correlation. Results: Biotope diversity reduced significantly through decrease of semi‐aquatic and tall‐herb biotopes, and expansion of forest. The quagfen terrestrialization sere nearly disappeared from 1987 recordings, while the reed sere did well concerning abundance and species richness. Several typical (rich) fen species disappeared from recordings, while new species were mostly field margin species. Periodic mowing and discharge combined are correlated with increasing species numbers. The P‐concentration in surface water increased while N‐concentration decreased. Conclusions: Preservation of the reed sere was successful, whereas preservation of the quagfen sere was not. Periodic mowing and discharge stimulate species richness, discharge more so than periodic mowing. But slight eutrophication likely induced a shift from P‐limitation to N‐limitation, which stimulated the reed sere at the expense of the quagfen sere.     

In search of a hydrological explanation for vegetation changes along a fen gradient in the Biebrza Upper Basin (Poland)

The understanding of succession from rich fen to poorer fen types requires knowledge of changes in hydrology, water composition, peat chemistry and peat accumulation in the successional process. Water flow patterns, water levels and water chemistry, mineralisation rates and nutrient concentrations in above-ground vegetation were studied along a extreme-rich fen-moderate-rich fen gradient at Biebrza (Poland). The extreme-rich fen was a temporary groundwater discharge area, while in the moderate-rich fen groundwater flows laterally towards the river. The moderate-rich fen has a rainwater lens in spring and significant lower concentrations of calcium and higher concentrations of phosphate in the surface water. Mineralisation rates for N, P and K were higher in the moderate-rich fen. Phosphorus concentrations in plant material of the moderate-rich fen were higher than in the extreme-rich fen, but concentrations of N and K in plant material did not differ between both fen types. Water level dynamics and macro-remains of superficial peat deposits were similar in both fen types.We concluded that the differences observed in the moderate-rich and the extreme-rich fens were caused by subtile differences in the proportion of water sources at the peat surface (rainwater and calcareous groundwater, respectively). Development of an extreme-rich fen into a moderate-rich fen was ascribed to recent changes in river hydrology possibly associated with a change in management practices. The observed differences in P-availability between the fen types did not result in significantly different biomass. Moreover, biomass production in both fen types was primarily N-limited although P-availability was restricted too in the extreme-rich fen. Aulacomnium palustre, the dominant moss in the moderate-rich fen, might be favoured in competition because of its broad nutrient tolerance and its quick establishment after disturbance. It might outcompete low productive rich fen species which were shown to be N-limited in both fens. We present a conceptual model of successional pathways of rich fen vegetation in the Biebrza region.     

Experimental investigation of drought induced acidification in a rich fen soil

Intact rich fen soil cores with controlled water levels near thesurface were installed in an open greenhouse. To simulate short termsummer drought, water levels were lowered (20 cm) after two weeks inhalf of the cores (experimental cores) and remained near the surface inthe other half (blanks). After two more weeks, the water levels werebrought back to the surface in the experimental cores and remainedthere for another two weeks. In the blanks, reduction and alkalinizationof the top peat layer occurred. In the experimental soil cores oxidationand acidification started within one week after drawdown. An indicationfor a drought induced rise in soluble reactive phosphorus has beenfound. The velocity of the acidification process illustrates the dynamicnature of the hydrochemical conditions in fen soils during drought. Theprocesses controlling the acid/base status of rich fen, the effect ofdrought induced acidification on P availability and the significance forthe vegetation are discussed.     

Optimal habitat conditions for the globally threatened Aquatic Warbler Acrocephalus paludicola in eastern Poland and their implications for fen management

To identify optimal habitat for the Aquatic Warbler Acrocephalus paludicola, a flagship species of fen mires, we related numbers of singing males to habitat variables in all core breeding sites in eastern Poland. The density of male Aquatic Warblers increased with increasing ground cover by water and mosses and litter layer height, and was highest where vegetation was 60–90 cm tall. Male densities also increased with the biomass of arthropods > 10 mm length, estimated by sweep netting, and with the abundance of spiders, estimated by pan trapping. We suggest that habitat management should take into account species‐specific morphological adaptations, nest safety and arthropod productivity. Prevention of vegetation succession is a conservation priority for open fen mires. However, modern management practices to achieve this, especially mowing using tracked vehicles, should be evaluated and optimized to ensure that such practices do not adversely affect the long‐term development of moss cover and litter structure.     

Accelerated vegetation succession but no hydrological change in a boreal fen during 20 years of recent climate change

Northern mires (fens and bogs) have significant climate feedbacks and contribute to biodiversity, providing habitats to specialized biota. Many studies have found drying and degradation of bogs in response to climate change, while northern fens have received less attention. Rich fens are particularly important to biodiversity, but subject to global climate change, fen ecosystems may change via direct response of vegetation or indirectly by hydrological changes. With repeated sampling over the past 20 years, we aim to reveal trends in hydrology and vegetation in a pristine boreal fen with gradient from rich to poor fen and bog vegetation. We resampled 203 semi‐permanent plots and compared water‐table depth (WTD), pH, concentrations of mineral elements, and dissolved organic carbon (DOC), plant species occurrences, community structure, and vegetation types between 1998 and 2018. In the study area, the annual mean temperature rose by 1.0°C and precipitation by 46 mm, in 20‐year periods prior to sampling occasions. We found that wet fen vegetation decreased, while bog and poor fen vegetation increased significantly. This reflected a trend of increasing abundance of common, generalist hummock species at the expense of fen specialist species. Changes were the most pronounced in high pH plots, where Sphagnum mosses had significantly increased in plot frequency, cover, and species richness. Changes of water chemistry were mainly insignificant in concentration levels and spatial patterns. Although indications toward drier conditions were found in vegetation, WTD had not consistently increased, instead, our results revealed complex dynamics of WTD as depending on vegetation changes. Overall, we found significant trend in vegetation, conforming to common succession pattern from rich to poor fen and bog vegetation. Our results suggest that responses intrinsic to vegetation, such as increased productivity or altered species interactions, may be more significant than indirect effects via local hydrology to the ecosystem response to climate warming.     

The effect of drainage and management on peat chemistry and nutrient deficiency in the former Jegrznia-floodplain (NE-Poland)

The Jegrznia floodplain has been deprived of flooding since the late 19th century. Human influence has been limited to haymaking and cattle grazing without land reclamation. Therefore, the present vegetation reflects the impact of hydrological changes, without being much disturbed by changes in other factors. A drainage gradient on the former floodplain was examined for nutrient deficiencies and compared with an intact floodplain. For this purpose we analyzed nutrient concentrations in above ground biomass, we measured redox potentials and peat variables for a rich fen vegetation, a well-developed fen meadow as well as an impoverished fen meadow vegetation. According to P:N-ratios in above ground biomass, all vegetation types appear to be limited by phosphorus, whereas the impoverished fen meadow is also limited by potassium. These results are supported by the chemical composition of the peat, redox characteristics and the physical changes in the peat induced by drainage. It is concluded that hydrological changes in combination with an annual mowing regime resulted in a shift from no nutrient limitation to P-and/or K-limitation in the former Jegrznia floodplain. Changes in hydrology at Jegrznia stopped the input of nutrients, increased the leaching of K and turned P into unavailable Al and Fe-complexes. Continued mowing increased P-and K-deficiency.     

Effectiveness of Turf Stripping as a Measure for Restoring Species-Rich Fen Meadows in Suboptimal Hydrological Conditions

Most species‐rich fen meadows in nature reserves in The Netherlands are acidified due to weaker upwelling of base‐rich groundwater. The present study investigated whether and why turf stripping combined with superficial drainage might promote the long‐term recovery of such meadows and restore the nutrient‐poor, buffered conditions they require. In a field experiment, we analyzed changes in vegetation composition, soil parameters, and soil water chemistry in stripped plots of degraded Cirsio‐Molinietum vegetation over 12 years. After the first five years, many species from the target communities occurred in stripped plots. Both vegetation and soil data showed positive effects of turf stripping on the acid‐buffering capacity. Because sulfate concentration in the soil water decreased over time, whereas the bicarbonate concentration increased, we inferred that there was internal alkalinization driven by sulfate reduction in low‐lying stripped plots. However, the succession toward more acidophilus plant communities, in both control and stripped plots, indicated gradual acidification. This may be caused by a continuing weakening of the upward seepage of base‐rich groundwater as shown by declining calcium concentrations in the soil water. Though turf stripping exposed a nutrient‐poor soil layer with a greater acid‐buffering capacity, these positive effects might not be sufficient to combat the ongoing acidification in the long term.     

Hydro-ecological analysis of the Biebrza mire (Poland)

Vegetation composition and structure of 58 sites along gradients in the valley mire of Biebrza, Poland, are related to physical and chemical variables of groundwater and peat. The three most prominent hydrochemical processes in the valley are (a) dissolution of calcite; (b) dissolution of iron, manganese and aluminium; and (c) enrichment with nitrogen and potassium. Major factors determining these processes are vertical flow of the groundwater and river flooding.Within the rheophilous zone of the mire, calcium-richness of the shallow groundwater and base-saturation of the peat are caused by upward seepage of groundwater originating from adjacent higher grounds. This groundwater movement keeps the larger part of the mire saturated with calcium.Good correlations exist between hydrochemistry and vegetation patterns. Groundwater-fed sites support a characteristic rich fen vegetation (Caricetum limoso-diandrae) with a low biomass production. The flood-plain vegetation consists of highly-productive communities of Glycerietum maximae and Caricetum elatae. In a belt in the Upper Basin where neither flooding nor upward seepage occurs, succession, probably caused by intensified drainage, leads to a dwarf-shrub vegetation (Betuletum humilis; poor fen).     

Restoration of fen ecosystems in the Vecht River plain: cost-benefit analysis of hydrological alternatives

In the Vecht river plain, the vegetation diversity has been diminished during recent decades, due to hydrological changes and resulting eutrophication. Different options are discussed for hydrological management aimed at abating the eutrophication and at stimulating the characteristic mesotraphent succession phases of the fen vegetation. The ecological impact of the management plans are estimated with the hydro-ecological model ICHORS; this model takes into account the relations between the non-biotic environment and the presence of hydrophytes and phreatophytes. The impact of the hydrological options is assessed, after rearrangement of the predicted values of the species to values of the required fen vegetation. These results are incorporated in the management options with the water balances of the polders. Information on the costs of the options is used as a basis for a cost-benefit analysis. Technical hydrological solutions such as purification of the supplied water might contribute to the eutrophication abatement, but they are on their own unlikely to restore the ecosystems. Contrastingly, options involving the restoration of the original hydrology would probably lead to the characteristic mesotraphent fen vegetation once more, and the financial implications of this are likely to be the more favourable in the long-term.     

Key variables controlling the vegetation of a cool-temperate mire in northern Japan

Abstract. In the cool-temperate Bibi Mire, Hokkaido, Japan, valley fens and flood-plain fens have quite different vegetation. The main variables controlling the vegetation were all hydrological: mean water level, water level fluctuation and surface water flow. Chemical factors such as electrical conductivity, dissolved oxygen and related peat decomposition were less important. The pH was about neutral and has little effect. The flood-plain fen developed under fluctuating water table conditions. The dominant species are Calamagrostis langsdotffii and Carex pseudocuraica. When temporal inundation occurs in the rainy or typhoon seasons, the submergence stimulates bud germination of the stoloniferous C. pseudocuraica, which can rapidly elongate its stolons upward. Some large floating peat mats occurred in the flood-plain fen zone. On these mats some Alnus japonica saplings establish and patches of alder forest can arise. Here the water level was higher than in the peripheral alder forest zone. The valley fen is dominated by Carex lasiocarpa var. occultans and/or C. limosa. It is formed under stable water table conditions in the inundated parts of the mire -where the non-inundated wet areas are dominated by alder trees. In the area where the surface water is flowing, these two fen sedges grow in deeper water since the high oxygen content is considered to compensate the flooding stress.     

Rehabilitation of Acidified Floating Fens by Addition of Buffered Surface Water

Floating fens are species‐rich succession stages in fen areas in the Netherlands. Many of these fens are deteriorating due to acidification; Sphagnum species and Polytrichum commune build 10–25 cm thick moss carpets, and the species diversity decreases. Earlier experiments in wet ecosystems indicate that successful restoration of circum‐neutral and mesotrophic conditions requires a combination of hydrological measures and sod removal. In an acidified fen recharged by rainwater in the nature reserve Ilperveld (The Netherlands), a ditch/trench system was dug for the purpose of creating a run‐off channel for acid rainwater in wet periods and to enable circum‐neutral surface water to enter the fen in dry periods. Moreover, the sod was removed in part of the fen. Ditch/trench creation or sod cutting had no effect individually, but a combination of the two measures led to a change in the abiotic conditions (higher pH and Ca), and in turn to an increase of species‐richness and the reestablishment of a number of characteristic species. Reestablishment of rare vascular plant species and characteristic bryophytes might be a long‐term process because of incomplete recovery of site conditions and constraints in seed dispersal.     

A local rich fen fed by calcareous seepage from an artificial river water infiltration system

The habitat of low growing, herbaceous rich-fen vegetation (Caricion davallianae) is now threatened throughout Western Europe. Its conservation, restoration and even recreation receives increasing attention. In the Campine area of Belgium a rich fen, that depends entirely on an artificial river water infiltration system for its alkaline discharge, is present. The rich fen vegetation cannot be older then a century, i.e. the age of the infiltration system. This illustrates that poor-fen precursors can be transformed into rich fen by alkaline ground-water discharge. The possibilities for restoration and the difficulties with use of allochtonous water for nature conservation are discussed. Finally, the hydrochemical consequences of the alkaline discharge in the fen soil are considered: in dry summers periodical acidification of the peat soils occurs.     

Effects of stream flooding on the distribution and diversity of groundwater‐dependent vegetation in riparian areas

1. Effects of the frequency and duration of flooding on the structural and functional characteristics of riparian vegetation were studied at four sites (n = 80, 50 × 50 cm, plots) along medium‐sized naturally meandering lowland streams. Special focus was on rich fens, which – due to their high species richness – are of high priority in nature conservation. 2. Reed beds, rich fens and meadows were all regularly flooded during the 20‐year study period, with a higher frequency in reed bed areas than in rich fen and meadow areas. In rich fens, species richness was higher in low frequency flooded areas (≤3 year^?1) than in areas with a high frequency of flooding (>3 year^?1) or no flooding, whereas species richness in reed beds and meadows was unaffected by flood frequency. 3. The percentage of stress‐tolerant species was higher in low intensity flooded rich fen areas than in high intensity and non‐flooded areas, indicating that the higher species richness in low frequency flooded rich fens was caused by competitive release. We found no indication that increased productivity was associated with high flooding frequencies. 4. We conclude that the restoration of morphological features in stream channels to increase the flooding regime can be beneficial for protected vegetation within riparian areas, but also that groundwater discharge thresholds and critical levels for protected vegetation should be identified and considered when introducing stream ecosystem restoration plans.     

Multiple effects of land‐use changes impede the colonization of open water in fen ponds

Question: Dutch fen areas have become embedded in intensively used landscapes, resulting in biodiversity loss. Hence, plant species that colonize open water inducing the formation of species‐rich floating peat mats have disappeared. Despite many restoration efforts, they have not returned. Is natural succession towards floating mats impeded by site conditions, dispersal limitations or changed biotic interactions? Location: Six Dutch fen reserves: De Deelen, De Weerribben, De Wieden, Westbroek, Molenpolder and Terra Nova. Methods: In 62 fen ponds we determined plant species richness and expansion into open water. We related these to habitat quality (chemical composition of soil and surface water, pond morphology), dispersal potential (distance to remnant populations, likelihood of dispersal) and biotic interactions (presence of muskrats [Ondatra zibethicus L.] and the keystone species Stratiotes aloides). Results: Factor analysis showed that plants expanded further into open water and bank vegetation had higher species richness in areas with older ponds and lower muskrat densities. Locally, high turbidity hampered colonization. Whenever the water was clear, colonization was higher in shallow ponds, and in deep ponds only if Stratiotes was present. Species richness was negatively correlated to nutrient availability in soil and positively correlated to hydrological isolation (decreased sulphate concentrations). We also found that species richness was higher in sheltered banks. Conclusions: Multiple habitat characteristics (turbidity, water depth, nutrient and sulphate concentrations) and the influence of muskrats and Stratiotes all play a role in the lack of restoration success in Dutch fen ponds. Dispersal limitations seem to be overruled by habitat limitations, as colonization often fails even when sufficient propagule sources are present, or when connectivity is high.     

Habitat selection of the globally threatened Aquatic Warbler Acrocephalus paludicola at the western margin of its breeding range and implications for management

The globally threatened Aquatic Warbler Acrocephalus paludicola is an umbrella species for fen mires and is at risk of extinction in its westernmost breeding population due to severe habitat loss. We used boosted regression trees to model Aquatic Warbler habitat selection in order to make recommendations for effective management of the last remnant habitats. Habitat data were collected in the years 2004–2006 in all remaining breeding sites in Pomerania (eastern Germany and western Poland) as well as in recently abandoned sites. Models were validated using data from similar Aquatic Warbler habitats in Lithuania. The probability of occurrence of Aquatic Warblers in late May/early June was positively associated with low isolation from other occupied sites, less eutrophic conditions, a high proportion of area mown early in the preceding year, high availability of vegetation 60–70 cm high, high prey abundance and high habitat heterogeneity. Early summer land management is needed in the more productive sites to prevent habitat deterioration by succession to higher and denser vegetation. As this also poses a serious threat to broods, management that creates a mosaic of early and late used patches is recommended to preserve and restore productive Aquatic Warbler sites. In less productive sites, winter mowing can maintain suitable habitat conditions. Aquatic Warbler‐friendly land use supports a variety of other threatened plant and animal species typical of fens and sedge meadows and can meet the economic interests of local land users.     

Environmental conditions and fen vegetation in three lowland mires

The abiotic conditions and fen vegetation in three lowland mires were analysed. Two of these mires are in the Netherlands. They have deteriorated considerably as a result of human pressure. One mire complex is in Poland. Its hydrology is almost undisturbed. The variation in the water composition in the fens was associated with the variation in the amount of regional groundwater discharge originating from the uplands, and to a lesser extent by the infiltration of polluted water pumped into the fens.The 26 vegetation types ranged from poor fen to eutrophic reedland. Most types were typical for only one region.There were clear differences between the three regions. Both the species composition and the water quality for the vegetation types indicated that the fens in the Vecht river plain are eutrophicated, whereas the fens in Northwestern Overijssel suffer from acidification. The Biebrza valley fens are well developed and are mainly fed by fresh calcareous groundwater.Stepwise logistic regression performed on 28 fen species revealed that the hydrochemical variables explained only a part of the variance: the regional variable area explained a considerable amount of the variance for most species. In 27% of the cases it was possible to fit an optimum curve for the species response to relevant hydrochemical variables. Monotonic curves could be fitted in 64% of the cases; mostly they described the response to variables in a way that is supported by ecological literature. It was concluded that the dataset should be constructed differently to enable the calculation of generally applicable standards.     

Nutrient limitation in species-rich lowland fens

Abstract. Nitrogen, phosphorus and potassium were supplied to some Belgian fens of varying nutrient status and productivity. Plant growth in the lowest productive fen with a species-rich Caricion davallianae vegetation was strongly P-limited. N was ineffective when applied alone, but increased the effect of P-addition when applied together. Summer biomass and plant nutrient concentrations were monitored for four years, and showed partial recovery of nutrient limitation. In a more productive fen dominated by Carex lasiocarpa and in a fen meadow, nutrient limitation was less strong. N limited growth in the productive fen, and N and K were co-limiting in the fen meadow. The P-concentration in the productive fen vegetation showed a marked increase after P-fertilization, but it did not result in higher standing crop. The significance of P-limitation for the conservation of species rich low productive fens is discussed. P-limitation may be an essential feature in the conservation of low productive rich fens: because it is less mobile in the landscape than N and/or because it is an intrinsic property of this vegetation type. Plant nutrient concentrations and N:P-ratios may be used as an indication for the presence and type of nutrient limitation in the vegetation. We found N:P-ratios of 23 to 31 for a P-limited site and 8 to 15 in N-limited sites. This was in agreement with critical values from the literature: N:P > ca. 20 for P-limitation and N:P < 14 for N-limitation. Thus, this technique appears valid in the vegetation types that were studied here.