Hydrology of Dutch Cirsio‐Molinietum meadows: Prospects for restoration

Abstract. Fen meadows (Cirsio dissecti‐Molinietum) are seriously threatened by desiccation, acidification and eutro‐phication. In The Netherlands several projects were launched to restore damaged fen meadows. This review describes how successes and failures of these restoration projects depend on hydrological systems. Six hydrological systems have been distinguished, which all provide the site conditions required by this community. Nowadays, the best developed fen meadows are found in the higher Pleistocene landscape of The Netherlands, where they depend on base‐rich groundwater discharging from local or large groundwater systems. Fen meadows of the lower Holocene landscape usually occur in man‐made surface water systems. Almost all stands have been severely deteriorated. Restoration of fen meadows in the Pleistocene landscape is promising when the hydrology is only slightly disturbed or when hydrological measures are taken in combination with sod cutting. Restoration prospects of fen meadows in the Holocene landscape are low. Until now a complete regeneration of Cirsio‐Molinietum meadows has not been realized. Restoration measures failed to restore high pH values in the top soil. It is hypothesized that viable seeds of many target species lack in the soil seed bank. In addition, the dispersal capacities of these species seem to be limited.     

Sod cutting and soil biota effects on seedling performance

Sod cutting (i.e. top soil removal) is a restoration management option for enhancing seedling establishment and for lowering the nutrient concentration in eutrophicated soils of nutrient-poor species-rich grasslands. Removal of the upper soil changes not only abiotic soil properties but may also affect the resident soil community. We investigated the effects of sod cutting on the establishment and performance of two endangered plant species (Cirsium dissectum and Succisa pratensis) while simultaneously manipulating the interaction between seedlings and soil biota. In intact grassland and sod-cut areas at two localities, seedlings were grown in plastic tubes. Half of the tubes had a filter that excluded roots but allowed entry of fungal hyphae and soil microorganisms. The other tubes were closed (i.e. no contact with the surrounding soil). In a greenhouse experiment we studied the effect of soil solutions (with or without fungal tissue) from three grasslands and three sod-cut areas on seedling growth. Sod cutting had a positive net effect on seedling growth for S. pratensis. Access to (mycorrhizal) fungi and other soil biota resulted in a negative impact on seedling growth of both plant species, both in grassland and sod-cut areas. The greenhouse experiment confirmed that the soil biota in these meadows reduced seedling growth. Although sod cutting did not mitigate negative plant-soil feedback, it enhanced seedling growth, presumably by decreasing competition for light. Sod cutting is therefore very useful when seedling establishment needs to be stimulated.     

Long‐term influence of sod cutting depth on the restoration of degraded wet heaths

Temperate heaths have an unfavorable conservation status in most European biogeographical regions. Increasing nitrogen levels promote competitive grass species such as Molinia caerulea, which is a main threat to heathland conservation in Europe. This article investigates the long‐term influence of sod cutting and the resulting changes in soil properties on the heath composition, integrity, and structure. In 15 nature reserves across the northern half of Belgium, we used (1) a large number of plots (203); (2) a broad range of sod cut depths (2–40 cm), and (3) a temporal dimension that describes how long the effects of sod cutting persist (census up to 19 years after sod cutting). Multivariate analyses were used in order to explore the influence of sod cut depth and time after sod cutting on the soil and vegetation properties. There was a positive relationship between sod cut depth and soil pH and water level, and a negative relationship with Al³⁺, NH₄⁺, and total organic matter (TOM). However, only a limited number of typical (target) species appeared after sod cutting, and then only weakly. Most of the time they remained a minor component of the restored vegetation. Moreover, M. caerulea reappeared and its cover significantly increased during the years following sod cutting. Although we were able to show that sod cut depth has a differential effect on soil properties and vegetation recovery, it also appeared that sod cutting does not restore wet heaths in the long term when applied in regions with high nitrogen deposition.     

The restoration of species-rich heathland communities in the Netherlands

The wet heathland communities of the Ericetum tetralicis and the Cirsio-Molinietum have declined in the Netherlands due to acidification, eutrophication and lowering of the water table. To investigate the prospects of restoration of both communities, the effects of sod cutting and hydrological measures on vegetation and soil chemistry were studied in two nature reserves where these plant communities occurred decades ago. The combination of sod cutting and hydrological measures has restored several rare, groundwater dependent heathland communities. Sod cutting has restored the Ericetum tetralicis, but not the Cirsio-Molinietum. This might be due to the absence of viable seeds of characteristic species of the Cirsio-Molinietum and/or the absence of optimal site conditions, especially high phosphorus concentrations in the top soil. The high phosphorus concentrations might be a consequence of high mineralization rates and/or prolonged inundation with iron-poor water and the decreased flux of iron-rich groundwater into the topsoil. Restoration of the Cirsio-Molinietum only seems possible when sod cutting is carried out together with hydrological measures that counter prolonged inundation and reinforce the discharge of base and iron-rich groundwater.     

Dynamics of species richness and composition in a limestone grassland restored after tree cutting

Abstract. After abandonment many limestone grasslands have been overgrown by trees and shrubs; as a result, species-rich communities with many regionally rare and endangered species are vanishing. Some studies suggest that, in cases where grassland is being restored, the species composition and rate of change is strongly dependent on the initial conditions, i.e. the earlier presence of grassland species and the opportunity for colonization of new sites by grassland species. These hypotheses were tested in a five-year restoration experiment after the clearing of a 35-yr-old secondary pine wood developed on abandoned grassland. Tree cutting induced rapid changes in the floristic composition and species cover. The number of grassland species from the class Festuco-Brometea increased significantly in the restored grassland, but their cover was much lower than in the old grassland. Canonical Correspondence Analysis showed significant differences in species composition between the grassland restored in former wood gaps and that developed in former closed wood. In wood gap sites the cover of species from the class Molinio-Arrhenatheretea and tufted perennials was much higher, whereas the cover of Festuco-Brometea species was lower. Significantly more shrubs, woodland species, ruderal and nitrophilous species as well as annual and biennial species occurred in the former closed wood site. It was found that richness and composition of the restored grassland depended strongly on the community composition before tree cutting, as well as on the presence of grassland species in the neighbourhood. Periodical tree cutting enables the maintenance of a temporal-spatial mosaic of scrub-grassland communities in isolated habitats and the preservation of local species diversity.     

Long-term effects on the nitrogen budget of a short-rotation grey alder (Alnus incana (L.) Moench) forest on abandoned agricultural land

Short-rotation energy forestry is one of the potential ways for management of abandoned agricultural areas. It helps sequestrate carbon and mitigate human-induced climate changes. Owing to symbiotic dinitrogen (N₂) fixation by actinomycetes and the soil fertilizing capacity and fast biomass growth of grey alders, the latter can be suitable species for short-rotation forestry. In our study of a young grey alder stand (Alnus incana (L.) Moench) on abandoned arable land in Estonia we tested the following hypotheses: (1) afforestation of abandoned agricultural land by grey alder significantly affects the soil nitrogen (N) status already during the first rotation period; (2) input of symbiotic fixation covers an essential part of the plant annual N demand of the stand; (3) despite a considerable N input into the ecosystem of a young alder stand, there will occur no significant environmental hazards (N leaching or N₂O emissions). The first two hypotheses can be accepted: there was a significant increase in N and C content in the topsoil (from 0.11 to 0.14%, and from 1.4 to 1.7%, respectively), and N fixation (151.5 kg N ha⁻¹ yr⁻¹) covered about 74% of the annual N demand of the stand. The third hypothesis met support as well: N₂O emissions (0.5 kg N ha⁻¹ yr⁻¹) were low, while most of the annual gaseous N losses were in the form of N₂ (73.8 kg N ha⁻¹ yr⁻¹). Annual average NO₃-N leaching was 15 kg N ha⁻¹ yr⁻¹ but the N that leached from topsoil accumulated in deeper soil layers. The soil acidifying effect of alders was clearly evident; during the 14-year period soil acidity increased 1.3 units in the upper 0-10 cm topsoil layer.     

Dynamics of gaseous nitrogen and carbon fluxes in riparian alder forests

We studied greenhouse gas (GHG) fluxes in two differently loaded riparian Alnus incana-dominated forests in agricultural landscapes of southern Estonia: a 33-year-old stand in Porijõgi, in which the uphill agricultural activities had been abandoned since the middle of the 1990s, and a 50-year-old stand in Viiratsi, which still receives polluted lateral flow from uphill fields fertilized with pig slurry. In Porijõgi, closed-chamber based sampling lasted from October 2001 to October 2009, whereas in Viiratsi the sampling period was from November 2003 to October 2009. Both temporal and spatial variations in all GHG gas fluxes were remarkable. Local differences in GHG fluxes between micro-sites (“Edge”, “Dry” and “Wet” in Porijõgi, and “Wet”, “Slope” and “Dry” in Viiratsi) were sometimes greater than those between sites. Median values of GHG fluxes from both sites over the whole study period and all microsites did not differ significantly, being 45 and 42 mg CO₂-C m⁻² h⁻¹, 8 and 0.5 μg CH₄-C m⁻² h⁻¹, 1.0 and 2.1 mg N₂-N m⁻² h⁻¹, and 5 and 9 μg N₂O-N m⁻² h⁻¹, in Porijõgi and Viiratsi, respectively. The N₂:N₂O ratio in Viiratsi (40-1200) was lower than in Porijõgi (10-7600). The median values-based estimation of the Global Warming Potential of CH₄ and N₂O was 19 and 185 kg CO₂ equivalents (eq) ha⁻¹ yr⁻¹ in Porijõgi and −14 and 336 kg CO₂ eq ha⁻¹ yr⁻¹ in Viiratsi, respectively. A significant Spearman rank correlation was found between the mean monthly air temperature and CO₂, CH₄ and N₂ fluxes in Porijõgi, and N₂O flux in Viiratsi, and between the monthly precipitation and CH₄ fluxes in both study sites. Higher groundwater level significantly increases CH₄ emission and decreases CO₂ and N₂O emission, whereas higher soil temperature significantly increases N₂O, CH₄ and N₂ emission values. In Porijõgi, GHG emissions did not display any discernable trend, whereas in Viiratsi a significant increase in CO₂, N₂, and N₂O emissions has been found. This may be a result of the age of the grey alder stand, but may also be caused by the long-term nutrient load of this riparian alder stand, which indicates a need for the management of similar heavily loaded riparian alder stands.     

Restoration of brook valley meadows in the Netherlands

Until recently, restoration measures in Dutch brook valley meadows consisted of re-introducing traditional management techniques, such as mowing without fertilisation and low-intensity grazing. In the Netherlands, additional measures, such as rewetting and sod cutting, are now carried out on a large scale to combat negative influences of drainage and acidifying influences by atmospheric deposition. An analysis of successful and unsuccessful projects shows that restoration of brook valley meadows is most successful if traditional management techniques are applied in recently abandoned fields that had not been drained or fertilised. Large-scale topsoil removal in former agricultural fields that had been used intensively for several decades is often unsuccessful since seed banks are depleted, while hydrological conditions and seed dispersal mechanisms are sub-optimal. In areas with an organic topsoil, long-term drainage had often led to irreversible changes in chemical and physical properties of the soil. Successful sites were all characterised by a regular discharge of calcareous groundwater provided by local or regional hydrological systems, and, where not very long ago, populations of target species existed. On mineral soils, in particular, sod removal in established nature reserves was a successful measure to increase the number of endangered fen meadow species. It is argued that attempts to restore species-rich meadows should be avoided on former agricultural fields, where pedological processes have led to almost irreversible changes in the soil profile and where soil seed banks have been completely depleted. From a soil conservation point of view, such areas should be exploited as eutrophic wetlands that are regularly flooded.     

Are Sulfurous Soil Amendments (S0, Fe(II)SO4, Fe(III)SO4) an Effective Tool in the Restoration of Heathland and Acidic Grassland after Four Decades of Rock Phosphate Fertilization?

This paper deals with the complex issue of reversing long‐term improvements of fertility in soils derived from heathlands and acidic grasslands using sulfur‐based amendments. The experiment was conducted on a former heathland and acid grassland in the U.K. that was heavily fertilized and limed with rock phosphate, chalk, and marl. The experimental work had three aims. First, to determine whether sulfurous soil amendments are able to lower pH to a level suitable for heathland and acidic grassland re‐creation (approximately 3 pH units). Second, to determine what effect the soil amendments have on the available pool of some basic cations and some potentially toxic acidic cations that may affect the plant community. Third, to determine whether the addition of Fe to the soil system would sequester PO₄^? ions that might be liberated from rock phosphate by the experimental treatments. The application of S⁰ and Fe^(II)SO₄^? to the soil was able to reduce pH. However, only the highest S⁰ treatment (2,000 kg/ha S) lowered pH sufficiently for heathland restoration purposes but effectively so. Where pH was lowered, basic cations were lost from the exchangeable pool and replaced by acidic cations. Where Fe was added to the soil, there was no evidence of PO₄^? sequestration from soil test data (Olsen P), but sequestration was apparent because of lower foliar P in the grass sward. The ability of the forb Rumex acetosella to apparently detoxify Al³⁺, prevalent in acidified soils, appeared to give it a competitive advantage over other less tolerant species. We would anticipate further changes in plant community structure through time, driven by Al³⁺ toxicity, leading to the competitive exclusion of less tolerant species. This, we suggest, is a key abiotic driver in the restoration of biotic (acidic plant) communities.     

Nitrogen removal in subsurface water by narrow buffer strips in the intensive farming landscape of the Po River watershed, Italy

In many countries buffer strips have become an important management tool widely accepted for controlling the diffuse pollution and supporting the development of more sustainable agriculture. However, there is the need to investigate their role in intensive farming systems where a realistic and shareable proposal to realize buffer strips can only foresee the use of a limited space. We evaluated the nitrogen buffering capacities of two narrow riparian strips (5-8 m) along irrigation ditches located in a typical flat agricultural watershed of the alluvial plain of the River Po (Northern Italy). Subsurface water level and nutrient concentrations were monitored along transects of piezometers installed from crop fields to ditches in two different areas. Spatial and temporal variation in water chemistry and hydrology were investigated to individuate the main processes (biological or physical) leading to groundwater nitrate depletion related to fertilization, pluviometric regime and seasonal variation. The results obtained indicate an elevated nitrate removal efficiency in both riparian areas. Compared to the high mean concentrations measured at the exit of the crop fields (10-90 mg l⁻¹ N-NO₃⁻), nitrate levels within riparian sites can be very low, completely disappearing below the ditches. The patterns of some chemical species (O₂, SO₄²⁻ and HCO₃⁻) and the potential denitrification rates suggest that denitrification plays a predominant role in the N-NO₃⁻ depletion observed in the first few meters of the herbaceous strip. The key factors in the system are the elevated groundwater residence time and the effect of the evapotranspiration. The water uptake by woody vegetation affects the subsurface water to flow through the riparian zone and, at the same time, it contributes to completely remove the nitrate from the groundwater.Our findings also suggest the double role of riparian vegetation both in ecohydrological and biological terms. In fact the water uptake by trees affects the subsurface flow pattern and contributes to completely remove the nitrate in the riparian zone.     

Stimulating denitrification in a stream mesocosm with elemental sulfur as an electron donor

The heavy use of fertilizers in agricultural lands can result in significant nitrate (NO₃⁻) loadings to the aquatic environment. We hypothesized that biological denitrification in agricultural ditches and streams could be enhanced by adding elemental sulfur (S^o) to the sediment layer, where it could act as a biofilm support and electron donor. Using a bench-scale stream mesocosm with a bed of S^o granules, we explored NO₃⁻ removal fluxes as a function of the effluent NO₃⁻ concentrations. With effluent NO₃⁻ ranging from 0.5 mg N L⁻¹ to 4.1 mg N L⁻¹, NO₃⁻ removal fluxes ranged from 228 mg N m⁻² d⁻¹ to 708 mg N m⁻² d⁻¹. This is as much as 100 times higher than for agricultural drainage streams. Sulfate (SO₄²⁻) production was high due to aerobic sulfur oxidation. Molecular studies demonstrated that the S^o amendment selected for Thiobacillus species, and that no special inoculum was required for establishing a S^o-based autotrophic denitrifying community. Modeling studies suggested that denitrification was diffusion limited, and advective flow through the bed would greatly enhance NO₃⁻ removal fluxes. Our results indicate that amendment with S^o is an effective means to stimulate denitrification in a stream environment. To minimize SO₄²⁻ production, it may be better to place S^o deeper in the sediment layer.     

The effects of sod cutting and additional liming on potential net nitrification in heathland soils

The effects of sod cutting, a common restoration measure to remove excess nutrients from grass-dominated heathlands, on nitrification were studied in dry and wet Dutch heathlands and in incubation experiments. In the field, soil ammonium and nitrate concentrations were measured after treatment by sod cutting, with or without additional liming. Potential net nitrification was measured by incubating soil samples of all treatments with extra ammonium in a climate chamber at pH 6. Potential net nitrification of heaths dominated by Molinia caerulea was significantly higher than that of dwarf-shrub dominated heaths. Sod cutting of the former areas significantly decreased potential net nitrification, whereas in the latter areas no differences were found. Liming of sod-cut soils greatly increased potential net nitrification and the accumulation of ammonium in the soil up to toxic concentrations could be prevented. Our results show that the combination of sod cutting and liming would create suitable soil conditions for the germination and establishment of endangered plant species of dry and wet heathlands. The success of restoration projects of these areas can thus be increased.     

Restoration management of abandoned chalk grassland in the Netherlands

After abandonment of chalk grassland in the Netherlands, Brachypodium pinnatum had become very dominant and this resulted in a dramatic decrease of plant diversity. Restoration of these abandoned sites is important, because of its former high diversity with many nationally endangered species. To restore the chalk grassland vegetation, the impact of the introduction of different experimental cutting regimes was investigated during three years. The thick litter layer and the dominance of Brachypodium was strongly reduced after introduction of all cutting regimes. In all cutting treatments forb phytomass increased considerably, especially in the twice-a-year cutting vegetation, where the forb phytomass became as high as that of Brachypodium. Phanerogamic species diversity was stimulated in all cutting regimes. Especially the number of short-lived forbs. with a persistent seed bank, increased markedly in the twice-a-year cutting treatment. It is concluded that cutting the vegetation twice is adequate to reduce the negative effects of the thick litter layer and the abundant growth of Brachypodium. It creates an appropriate starting point for more detinite management, whether mowing or grazing. However, the restoration of a more complete species assortment strongly depends on the re-invasion possibilities of these species.     

Groundwater input affecting plant distribution by controlling ammonium and iron availability

Question: How does groundwater input affect plant distribution in Alnus glutinosa (black alder) carrs? Location: Alder carrs along the river Meuse, SE Netherlands. Methods: Three types of site, characterized by groundwater flow, were sampled in 17 A. glutinosa carrs. Vegetation and abiotic data (soil and water chemistry) were collected and analysed using a Canonical Correspondence Analysis. Based on the results, a laboratory experiment tested the effect of groundwater input (Ca²⁺) on pore water chemistry (NH₄⁺ availability). Results: Environmental factors indicating groundwater input (Ca²⁺ and Fe²⁺), correlating with the NH⁺₄ concentration in the pore water, best explained the variation in plant distribution. NH₄⁺ availability was determined by Ca²⁺ input via the groundwater and subsequent competition for exchange sites in the sediment. As a result, nutrient‐poor seepage locations fully fed by groundwater were dominated by small iron resistant plants such as Caltha palustris and Equisetum fluviatile. More nutrient‐rich locations, fed by a combination of groundwater and surface water, allowed the growth of taller iron resistant plant species such as Carex paniculata. Nutrient‐rich locations with stagnating surface water were hardly fed by groundwater, allowing the occurrence of fast growing and less iron tolerant wetland grasses such as Glyceria fluitans and G. maxima. Conclusion: Groundwater input affects plant composition in A. glutinosa carrs along the river Meuse by determining nutrient availability (ammonium) and concentrations of toxic iron.     

Diversity models applied to a chalk grassland

In a permanent plot experiment started in 1971, the effects of several management regimes on diversity of a Dutch chalk grassland have been investigated and the results have been compared to existing models predicting general trends in diversity. Treatments included grazing, mowing, and leaving untouched; in the mown plots, the effects of fertilizing and sod cutting were also studied. Grazing resulted in the highest diversity, leaving untouched in the lowest (ca. 42 and 15 spp. per m², respectively). Within the mown plots, fertilizing decreased diversity. The effects of sod cutting disappeared after some years. The results conform best with the response surface model of Huston (1979) relating diversity to frequency of population reduction and rate of competitive displacement, but a slight modification of the shape of the surface is suggested.Nomenclature follows van der Meijden et al., 1983.The authors are very grateful to Dr J. Miles (Banchory, U.K.) for stimulating remarks and to Prof. Dr. M. J. A. Werger, Dr R. K. Peet and two reviewers for many valuable comments and suggestions.     

Population dynamics of Gentiana pneumonanthe and Rhynchospora fusca during wet heathland restoration

Abstract. The population dynamics and reproductive strategies of two rare wet heathland species, Gentiana pneumonanthe and Rhynchospora, were studied in experimental permanent plots in a wet heathland near Bremen, NW Germany, to assess how effective sod cutting and mowing are in promoting these species. In one experiment, small plots (0.25 m² - 4 m²) were sod-cut or mown; in the second, one large plot 30 mx 50m) was sod-cut. The development of the vegetation and the number of shoots of the two target species were recorded annually. Sod cutting lead to the highest shoot numbers of Gentiana in the long run, whilst mowing was more effective at the beginning of the experiment. Seedlings and adult shoots slowly became more and more abundant after sod cutting. In contrast, Rhynchospora soon formed closed stands after sod-cutting, first through seed germination and then through fast clonal growth of the established individuals. Moist soil or short inundations promoted the germination and seedling establishment of Gentiana, whereas drought negatively affected seedling recruitment. Long periods of inundation severely reduced the population at first, but high numbers of seedlings were found in the following growing season. For the disturbance-dependent Gentiana and Rhynchospora, the availability of gaps within the vegetation is of crucial importance. To promote existing populations, we suggest small-scale sod cuttings which create gaps without disturbing existing flowering individuals too much. For degenerated stands of wet heathland we recommend large-scale sod cutting to activate the seed bank. Additionally, seed introduction may be helpful to encourage the development of a wet heathland with characteristic floristic composition.     

Integrated stream and wetland restoration: A watershed approach to improved water quality on the landscape

Water quality in Upper Sandy Creek, a headwater stream for the Cape Fear River in the North Carolina Piedmont, is impaired due to high N and P concentrations, sediment load, and coliform bacteria. The creek and floodplain ecosystem had become dysfunctional due to the effects of altered storm water delivery following urban watershed development where the impervious surface reached nearly 30% in some sub-watersheds. At Duke University, an 8-ha Stream and Wetland Assessment Management Park (SWAMP) was created in the lower portion of the watershed to assess the cumulative effect of restoring multiple portions of stream and former adjacent wetlands, with specific goals of quantifying water quality improvements. To accomplish these goals, a three-phase stream/riparian floodplain restoration (600 m), storm water reservoir/wetland complex (1.6 ha) along with a surface flow treatment wetland (0.5 ha) was ecologically designed to increase the stream wetland connection, and restore groundwater wetland hydrology. The multi-phased restoration of Sandy Creek and adjacent wetlands resulted in functioning riparian hydrology, which reduced downstream water pulses, nutrients, coliform bacteria, sediment, and stream erosion. Storm water event nutrient budgets indicated a substantial attenuation of N and P within the SWAMP project. Most notably, (NO₂⁻ + NO₃⁻)-N loads were reduced by 64% and P loads were reduced by 28%. Sediment retention in the stormwater reservoir and riparian wetlands showed accretion rates of 1.8 cm year⁻¹ and 1.1 cm year⁻¹, respectively. Sediment retention totaled nearly 500 MT year⁻¹.     

Effects of removal of organic matter on the productivity of heathlands

Abstract. Effects of milling, mowing and sod cutting on productivity in a Deschampsia grass heath and a Molinia-Deschampsia grass heath were studied from 1977 to 1986. The sum of above-ground biomass, litter and accumulated humified matter (TOM) in both types was ca. 70 ton/ha and the primary productivity 3–5 ton ha^-1yr^-1. Mulching did not affect the annual production. Sod cutting reduced the productivity to 1–2 ton ha^-1yr^-1; on the Molinia-Deschampsia site this reduction lasted for at least a decade, while the Deschampsia heath started to recover from sod cutting soon and increased in productivity again. Annual mowing both with and without removal of hay reduced the production as well, particularly on the Molinia-Deschampsia heath. Lower production was either the result of phosphorus depletion - Molinia-Deschampsia site - or nitrogen depletion - Deschampsia site. In the mowing treatments there was also a depletion of carbohydrate reserves.     

Nodulation potential of soils from red alder stands covering a wide age range

Red alder (Alnus rubra Bong.) stands in the Pacific Northwest are the common first stage in succession following disturbance. These stands are highly productive and contribute a large amount of N to the soils as a result of their N₂-fixing symbiosis with Frankia. As these alder stands age, the soils not only increase in total N, but concentrations of NO^– ₃ increase and pH decreases as a result of nitrification. The objective of this study was to determine how the nodulation capacity of Frankia varies as red alder stands age and if differences in nodulation capacity are related to changes in soil properties. Nodulation capacity was determined by a red alder seedling bioassay for soils from red alder stands in the Oregon coast range covering a wide range of ages. Six chronosequences were sampled, each containing a young, an intermediate, and an older alder stand. Soil total N, total C, NO^– ₃, NH⁺ ₄, and pH were measured on the same soil samples. These factors as well as alder stand characteristics were compared with nodulation capacity in an attempt to identify soil characteristics typical in developing alder stands that most strongly affect nodulation capacity. Soil pH and NO^– ₃ concentration were highly correlated with nodulation capacity and with each other. Cluster analysis of the sites using these two variables identified two groups with distinctly different nodulation capacities. The cluster with the higher nodulation capacity was lower in NO^– ₃ and higher in pH than the other cluster, which included the majority of sites. There was substantial overlap in the age ranges for the two clusters and there was no significant correlation between age and nodulation capacity. Thus nodulation capacity appears to be most closely related to soil properties than to stand age.