Vegetation development in dune slacks: the role of persistent seed banks

Abstract. The soil seed bank composition was determined at four sites in the dune slack ‘Koegelwieck’ on the Dutch Wadden Sea island of Terschelling. At three different sites in the slack, where sod-cutting experiments down to the mineral sand had been carried out, the established vegetation and seed bank were assessed after 5, 9 and 39 yr of undisturbed development, respectively. In addition, a fourth site in the slack was investigated, where vegetation development had proceeded for 80 yr since plant colonization of bare soil and where nowadays a vegetation dominated by Calamagrostis epigejos and Salix repens occurs. Together these four sites can be regarded as a chronosequence of dune slack formation. Clear time sequences were detected in the seed bank data. Many late successional species showed a significant increase in the number of seeds during the succession. Some of the early successional basiphilous pioneer species such as Anagallis minima, Centaurium littorale, Littorella uniflora, Radiola linoides and Samolus valerandi, showed either a decrease during the time of succession or an optimum in the vegetation while remaining present in the seed bank in low but detectable numbers. They could, therefore, play a role in re-establishment of the vegetation after sod-cutting. One of the target species, Schoenus nigricans, established within a few years after removal of the sod. However, no seeds of this species have been detected in the soil below either of the successional stages. Based on the species disappearance from the established vegetation and based on the independent data of Thompson et al. (1997) an estimation of seed longevity could be made for several Red List species of wet dune slacks.     

Changes in soil and vegetation during dune slack succession

Many rare plant species occur in Dutch wet dune slacks, particularly in the Junco baltici‐Schoenetum nigricantis. For nature management it is important to understand the processes controlling the presence of these basiphilous early successional communities, which is why we investigated vegetation and soil development during succession in coastal dune slacks. We compared 12 chronosequential stages of 0, 2, 4, 9, 10, 13, 25, 30, 43, 60, 70 and 85 yr in five different dune slack systems. In four of these locations turf had earlier been removed in order to restore the basiphilous pioneer stage. The main variation in the vegetation is related to the acidification/soil enrichment gradient and the salinity/maritime gradient. During succession, organic matter accumulates and acidification takes place. Maritime influence can buffer the soil and postpone the succession of basiphilous pioneer communities for many years. A significant correlation with age was found for 18 variables. Multiple regressions predicted changes in the vegetation (dependent variables: biomass, cover of Salix repens, Calamagrostis epigejos and Schoenus nigricans) as a function of acidification, organic matter accumulation, increase in available P and presence of Na in the soil. We conclude that natural ageing of the vegetation and the associated process of accumulation of biomass drive succession in this hydrosere. The underlying soil processes are acidification and organic matter accumulation. During succession dominance shifts from S. nigricans to S. repens or C. epigejos. Maintenance of the pioneer character of the habitat is only possible by local intervention or by natural or man‐induced dune forming. The effect of sod‐stripping depends on the environmental conditions and, in case of acidification, success is limited. Succession can be postponed by mowing.     

Preference of wet dune species for waterlogged conditions can be explained by adaptations and specific recruitment requirements

The preference of wetland angiosperms for waterlogged soils has been explained by several hypotheses: (1) wetland species are adapted to waterlogging and sensitive to drought; (2) wetland species are tolerant to drought, but inferior competitors at drier conditions; (3) wetland species have narrow moisture optima for recruitment. We tested these hypotheses by the application of permanent and fluctuating water levels to experimental wet dune plant communities with four species that frequently occur in wet dune slacks (Carex flacca, Molinia caerulea, Samolus valerandi and Schoenus nigricans) and one competitive species of drier conditions (Calamagrostis epigejos). After 3 years, fluctuating water levels had led to lower total biomass production than permanent water levels, indicating that switching from aerated to anoxic soil conditions involved physiological costs. The collective biomass of wetland species was highest at permanently waterlogged conditions with interspecific variation in the biomass optima. At the nutrient-poor growth conditions of this long-term experiment, biomass of Calamagrostis was independent of water level treatment, thus the hypothesized competitive superiority of this species at drier conditions could not explain the biomass responses of the wetland species in this study. Instead, this is evidence for an ecological preference of adult wet dune plants for waterlogged to moist conditions. Recruitment of most wet dune slack species occurred in a narrower range of water levels than adult growth, indicating that recruitment requirements also pose a limitation to the distribution of these wet dune species. Incorporation of recruitment into nature management support models may improve their predictions.     

Effects of Ca- and Fe-rich Seepage on P Availability and Plant Performance in Calcareous Dune Soils

Ca- and Fe-rich seepage in wet dune slacks often sustains oligotrophic, species rich vegetation as a result of reduced P availability. While this effect has been attributed to pH buffering, we tested whether Ca- and Fe-rich seepage also immobilised P in calcareous soils with a strong pH buffer. Two oligotrophic species (Carex flacca and Schoenus nigricans) and two eutrophic species (Calamagrostis epigejos and Molinia caerulea) were planted in experimental sods. After 4 months supply with water of seepage or infiltration quality, with or without the addition of P, soil P fractions and a number of plant physiological responses were measured. A field validation was performed in a flow-through lake in calcareous dunes where the seepage flow had been restored recently. The readily available water soluble P fraction (Pw) was reduced by more than 80% by seepage, both in the greenhouse experiment and in the field, but the P Olsen-inorganic fraction was unaffected. All four test species had elevated N: P ratios in aboveground tissues when treated with seepage, indicating that seepage water had indeed reduced P availability to plants. Formation of dauciform roots by Carex flacca was diminished by P addition to less than 25% of treatments without P addition, indicating sensitivity to P availability, while seepage on average halved production of these root structures. Dauciform root formation by Schoenus nigricans was unaffected by the P addition and the hydrological treatment. Biomass of the test species in the experiment as well as vegetation biomass and relative abundance of oligotrophic species in the field were unrelated to seepage patterns, suggesting that compensatory mechanisms enabled the plants to sustain biomass production within the time frame of this experiment. In conclusion, Ca- and Fe-rich seepage can lower P availability in soils with a strong pH buffer. In the long term, this may create favourable conditions for species that have low P requirements or efficient P uptake. In the short term, however, existing vegetation seems to be resilient to changes in P supply.     

Accumulation rates of soil organic matter in wet dune slacks on the Dutch Wadden Sea islands

Background and aims

A long-term monitoring program (ranging from 16 to 77 years) on the Dutch Wadden Sea Islands provided well-documented examples of vegetation succession in wet dune slacks. We used this opportunity to study soil organic matter (SOM) accumulation in relation to vegetation succession. The aim of this paper is to identify the factors which regulate accumulation rates of SOM in wet dune slacks.

Methods

We used several soil chronosequences using data from the monitoring program together with data from a long-term research activity and more recent measurements. We used several soil chronosequences using data from the monitoring program together with data from a long-term research (up to 150 years) and more recent measurements. Field measurements included pH, soil organic matter, above ground standing crop and water levels. Water level regimes (inundation duration and mean minimum water level), were simulated using a hydrological model. Capable of simulating inundation duration and water-level fluctuations, this model used field measurements collected over more than 5 years, as well as precipitation and evapotranspiration data collected over a period of 25 years.

Results

Sampling two synchronic chronosequences showed that SOM accumulations increased linearly during the first 50–60 years and then levelled off. Sampling various diachronic chronosequences over time showed a wide variation in accumulation rates. Slacks with low productive species, such as Littorella uniflora, showed low accumulation rates (0.02–0.08 kg/m²/year), and persisted even over a period of more than 90 years. In contrast, slacks dominated by high productive species, such as Phragmites australis, showed ten times higher accumulation rates (0.17–0.26 kg/m²/year) over a similar time period and comparable annual inundation periods (176–240 days). A multiple linear regression showed that variation in SOM accumulation rates was best explained by above-ground biomass of the vegetation.

Conclusions

We conclude that the rate of SOM accumulation in wet dune slacks is primarily controlled by plant above-ground biomass. Both above-ground biomass and SOM accumulation can remain very low over a long period of time when dune slacks are flooded during most of the year and plants with adaptive traits are able to maintain vegetation succession at a pioneer stage.     

Plant responses to rising water tables and nutrient management in calcareous dune slacks

Plant species of oligotrophic wet dune slacks have dramatically decreased as a result of desiccation and eutrophication. The aim of this study was to test in a field experiment the effects of restoration management in oligotrophic, wet dune slacks (groundwater level rise in combination with topsoil removal or mowing) on abiotic variables and on survival and biomass of four plant species. The effect of groundwater level rise on abiotic variables strongly differed between mown sampling locations and those with topsoil removal. At locations with a mowing treatment, a large rise in water tables led to increased N availability and higher reduced iron concentrations than at other locations. Such effects were absent at locations with recent topsoil removal. No effect of groundwater level rise on P-availability was found. Topsoil removal on average lowered N availability by 13%, P availability by 65% and Fe²⁺ by 56%. All phytometer species survived better in mown dune slacks than in dune slacks that had received topsoil removal. Survival of all species was negatively related to groundwater level rise. On the short term local extinction risks of small populations may be enhanced by rewetting and topsoil removal. On the long-term, however, such measures are crucial to maintain vegetation of oligotrophic wet dune slacks in a degraded dune landscape.     

Wet dune slacks: decline and new opportunities

For a number of infiltrated coastal dune areas it is discussed to what extent artificial infiltration for the public water supply affects the quality of soil, groundwater and vegetation around pools and ponds, and what its effect is on the vegetation. Further, the results of investigations into the quality of vegetation, soil and water of a number of non-infiltrated, less affected dune areas are presented. The emphasis is on changes in groundwater flow pattern and on changes in the chemical composition of groundwater on the vegetation of wet dune slacks. Finally, recommendations for the management of wet dune slacks are presented. It can be concluded that the introduction of nutrients through infiltration causes an abundance of nitrophilous herbaceous vegetation along the banks of all infiltration ponds and most dune pools. Of the three investigated macro-nutrients, nitrate, potassium and phosphate, the latter shows the most significant correlation with the composition, cover and biomass of the vegetation. The moist biotopes of non-infiltrated dunes have largely disappeared because of desiccation, mainly as a consequence of water withdrawal, afforestation and coastal erosion. Relatively unaffected dune slacks can be found in the dunes on the Dutch Wadden Sea islands and a small number of dune areas on the mainland. In most areas, however, a serious decline in many rare species has been observed during the past twenty years because of eutrophic and acid precipitation, often in combination with disturbances of the groundwater regime.     

Dune slack restoration in Dutch mainland coastal dunes

The 45,000 ha Dutch dunes are relatively well protected but they suffer from severe desiccation. During the last decade a lot of knowledge has been obtained about the steering processes of eco-hydrology in dunes resulting in well-founded schemes to restore wet dune slack communities. In two case studies a large-scale restoration of the landscape was necessary. Since 1987 in the North-Holland Dune Reserve the groundwater catchment has been reduced (from 8 million m³ to 2 million m³ per year) and the resulting development monitored. In another case, large-scale intervention on 35 ha of the Amsterdam Water Supply Dunes in 1995 resulted in the restoration of active sand drift with opportunities for the establishment of pioneer vegetation. In small parts of the area, the surface has stabilised as a result of vegetation growth, but elsewhere drifting sand has spread and covered previously stable surfaces. Many wet to moist dune slacks developed and results relating to geomorphology, hydrology and vegetation are presented. Finally, in 1996 yet another project of 50 ha was carried out in the Meijendel dunes, in which infiltration ponds were removed to allow vegetation succession to develop from scratch. Results of these three case studies will be presented. If only hydrological restoration is carried out, sod-cutting was found to be necessary to restore the whole series of vegetation types of dune slacks. In two cases an area was denuded. The seed bank and surrounding slacks seem to play an important role in the success of restoration, this is especially true for the pioneer vegetation types. Future monitoring will prove whether critical (Red List) plant species will return.     

Oxidation of elemental-S in coastal-dune sands and soils

Summary S-oxidation was studied in samples of (a) coastal sands lacking vegetation; (b) sands from beneath isolated stands ofAmmophila arenaria andHippophaë rhamnoides; and (c) dune soils obtained from beneath vegetation growing on mature dunes. S-oxidation in samples taken from dune environments was compared with the process in a fertile garden soil.Elemental-S was oxidized to SO ₄ ^2– in all samples, with S₂O ₃ ^2– being formed as intermediates. S-oxidation was most pronounced in the dune soil, followed by the garden soil,Ammophila arenaria andH. rhamnoides rhizospheres and finally the non-vegetated sand. The rate of S-oxidation thus generally increased with increasing C and N content, increasing vegetation cover and decreasing soil-sand pH.Maximum S-oxidation occurred at 30–37°C, but some of the intermediates appeared even at 45°C, presumably indicating abiotic S-oxidation at high temperatures. S-oxidation decreased the pH of the two soils studied, but did not markedly acidify the unvegetated or rhizosphere sands.     

Dynamics of seasonal bicarbonate supply in a dune slack: effects on organic matter, nitrogen pool and vegetation succession

The seasonal variation in groundwater composition was studied in an old dune slack complex on the Wadden Sea island of Schiermonnikoog that recently had lost practically all rare basiphilous plant species. In order to assess its restoration perspectives the groundwater acid neutralization capacity (ANC _aq ) of various sites was discussed in relation to the decalcification depth, amount of organic matter and amount of nitrogen in the topsoil layer. The decalcification depth reflected the (former) hydrological regime quite well; shallow in the exfiltration zone and deep in the infiltration part of the slack. The pattern of HCO₃ ^–, SO₄ ^2– and pH reflected the functioning of a flow-through lake, even when the slack was not flooded. Relics of low productivity (mesotrophic) vegetation (Samolo-Littorelletum) with rare species were related to low amounts of organic matter and nitrogen in the topsoil layer. At this site the variation in HCO₃ ^– concentration was also relatively low during the year. It was hypothesized that a regular supply of acid buffering components from calcareous soil layers buffers the pH at a high level and slows down the accumulation of organic matter thus creating opportunities for the rare basiphilous plant species. A possible biological control mechanism for these processes was discussed. A reconstruction of the former, present and future eco-hydrological situation was made of the dune slack.     

Physiological responses to Cd and Zn in two Cd/Zn hyperaccumulating Thlaspi species

In a model hyperaccumulation study a Cd/Zn hyperaccumulator Thlaspi caerulescens accession Ganges and a recently reported Cd/Zn hyperaccumulator Thlaspi praecox grown in increasing Cd and Zn concentrations in the substrate and in field collected polluted soil were compared. Plant biomass, concentrations of Cd and Zn, total chlorophylls and anthocyanins, antioxidative stress parameters and activities of selected antioxidative enzymes were compared. Increasing Cd, but not Zn in the substrate resulted in the increase of biomass of roots and shoots of T. praecox and T. caerulescens. The two species hyperaccumulated Cd in the shoots to a similar extent, whereas T. caerulescens accumulated more Zn in the shoots than T. praecox. Cadmium amendment decreased total chlorophyll concentration and glutathione reductase activity, and increased non-protein thiols concentration only in T. praecox, suggesting that it is less tolerant to Cd than T. caerulescens. In the field-contaminated soil, T. caerulescens accumulated higher Cd concentrations; but as T. praecox produced higher biomass, both species have similar ability to extract Cd.     

The ectomycorrhizal flora of primary and secondary stands of Pinus sylvestris in relation to soil conditions and ectomycorrhizal succession

Abstract. Ectomycorrhizal species composition and sporocarp abundance in two 15–20-yr old primary stands of Pinus sylvestris in the central part of the Netherlands was compared with those in two 16- and 27-yr old secondary stands of P. sylvestris in the Northeast of the country. The trees of the primary stands were spontaneously seeded in a drifting sand area. Only thin litter and humus layers were present. The trees of the 16-yr old secondary stand were planted on podzolic sandy soil and those of the 27-yr old stand on non-podzolic sandy soil. In both secondary stands thick litter and humus layers had developed. The litter and humus partly originated from former stands at the same sites and partly from the present stands. In the secondary stands the thick litter and humus layers and herb vegetation were removed (‘sod-cutting’) in order to simulate the thin litter and humus layers in the primary stands. Control treatments were present. Surveys in 1991, 1992 and 1993 showed that sod-cutting enhanced both abundance and diversity of ectomycorrhizal fungi. However, species richness and diversity were higher in the primary stands than in the secondary ones, also even in the sod-cut plots. High species richness and diversity were associated with low concentrations of nitrogen and relatively high pH in the litter and humus layers, and in the mineral soil.     

Genetic diversity and ecological differentiation in the endangered fen orchid (<Emphasis Type="Italic">Liparis loeselii</Emphasis>)

Liparis loeselii is a rare and endangered orchid occurring in Europe and north-east America. Genetic diversity and structure of this species in north-west France and the United Kingdom were investigated using amplified fragment length polymorphisms (AFLPs). Although clonality and autogamy are common in L. loeselii, we found moderate to important variability within populations. We observed a significant genetic differentiation between populations occurring in dune slacks and fens. This may be correlated with leaf shape as dune slack individuals are sometimes treated as the distinct variety L. loeselii var. ovata. Genetic differentiation between populations was generally low suggesting that gene flow can occur over long distances and possibly across the English Channel. These results show that populations from dune slacks and fens should be managed separately and that geographically distant populations may be equivalent.     

Restoration of Wet Dune Slacks on the Dutch Wadden Sea Islands: Recolonization After Large-Scale Sod Cutting

The effects of sod cutting were studied in a dune area on the Dutch Wadden Sea Island of Texel. Sod cutting was carried out in a range of different dune slacks in order to restore dune slack vegetation with many endangered Red List species. Sod cutting removed approximately 96% of the soil seed bank. Species abundant in the seed bank, notably Juncacea, also had a high frequency in the vegetation that established during the first year after the restoration measures. Many other species not registered in the seed bank or in the former vegetation also appeared. Species richness in the monitored plots exceeded that of uncut reference plots after a few years. Colonization rates were higher than extinction rates in most plots, indicating that a stable state has not been reached after 5 years. Differences in species richness between slacks appeared to be related to the occurrence of source areas nearby and availability of dispersal agents, such as flooding and animals.     

The relationship between vegetation dynamics and water table in tropical dune slacks

Abstract. Inundation of tropical dune slacks is an irregular phenomenon produced by consecutive years with high precipitation and also by intense tropical storms. Our hypothesis was that the distribution, structure and composition of vegetation in tropical dune slacks have changed over time as a result of various factors, principally their topographical position with respect to the water table, frequency and duration of inundation, and historical fluctuations in climate. We tested this hypothesis in two different slacks that represent the slack community at the study site. Inundation caused changes in species composition and distribution, as well as in vegetation structure. In one slack, inundation was considered as disturbance because its intensity increased when areas remained inundated for longer periods of time, causing death of vegetation cover and favouring invasion of more aggressive, secondary species (Pluchea odorata, Ambrosia artemisiifolia, Panicum maximum). When inundation lasted less than three months, rhizomatous herbs remained (Cyperus articulatus, Lippia nodiflora, Hydrocotyle bonariensis) inhibiting the succession process. The results suggest that the different degrees of inundation and the irregular frequency and intensity of inundation in the slacks under study, have favoured different successional tendencies.     

Vegetation zonation in dune slacks on the Leba Bar,Polish Baltic Sea coast

Vegetation zones in dune slacks on the Leba Bar on the Polish Baltic Sea coast are described. Plant colonization starts on moist sand in eastern ends of slacks, in the wake of receding mobile dunes. Communities on deflation ribs, dominated by psammophytic grasses, sedges and lichens, are structurally simple. Communities in troughs, dominated by species of fresh habitats, become structurally more complex in a western direction and they culminate with the Pinus silvestris forest. The main lines of vegetation variation correlate with groundwa-ter depth, micro topography, soil profile build-up and substrate acidification.     

Does the seed bank contribute to the restoration of species‐rich vegetation in wet dune slacks?

Questions: What is the contribution of the seed bank to restoration of species‐rich vegetation in oligotrophic wet dune slacks? Does the restoration management affect the seed bank? Location: Calcareous coastal dune slacks at the west coast of The Netherlands. Methods: Species composition of the seed bank and the above‐ground vegetation was sampled in dune slacks that had a variable extent of groundwater level rise in combination with either topsoil removal or mowing. Results: The seed bank had a high potential for restoration of species‐rich vegetation: 60 species were found in the seed bank of which 14 were characteristic of oligotrophic, wet dune vegetation. While topsoil removal almost completely removed the seed bank, groundwater level rise did not permanently submerge the seed bank of species of oligotrophic, wet conditions. Changes in abundance in the established vegetation were unrelated to species abundance in the seed bank. Of all new species establishments in the vegetation relevés, 76% occurred where the species was not found in the seed bank. The chance that presence of a species in the seed bank led to establishment the following year was only 11%. Conclusion: The seed bank was not the dominant source for newly establishing species following the large disturbance that was induced by restoration management. Changes in species abundance after the restoration impact were not related to species abundance in the seed bank, but to ongoing succession and current year dispersal. To attain a high number of new establishments, restoration projects should preferably be planned in the proximity of refuge populations, rather than relying on the seed bank alone.     

土壤水分及其垂直分布对内蒙古草原冰草和冷蒿竞争的影响

内蒙古退化冷蒿草原围封恢复演替3-4年,植物群落更替出现半灌木冷蒿（Artemisia frigida）向禾草冰草（Agropyron cristatum）群落突然转变的现象,这对内蒙古草原植被恢复具有积极的作用,但目前关于冰草-冷蒿演替和竞争机制的研究鲜见报道。水分作为草原植被生长的主要限制因子,是影响群落演替的重要因素。气候干旱和过度放牧导致草原土壤水分垂直分布发生明显变化。由于冰草和冷蒿的根系分布深浅不同,土壤水分垂直变化可能会影响冰草-冷蒿的竞争。为探讨土壤水分及其垂直分布对冰草和冷蒿竞争的影响,本盆栽试验设置冰草、冷蒿单种以及混种处理,并进行不同土壤深度（0-30cm,30-60cm）的水分处理（上干下湿、上湿下干、上干下干）,结果表明：（1）与冷蒿相比,冰草受土壤水分变化影响显著,相对于上干下干处理,冰草在湿润（上干下湿或者上湿下干）处理的地上、地下生物量均显著增加,而冷蒿没有显著变化;（2）上干下湿和上湿下干处理间,冰草、冷蒿的地上、地下总生物量和根长差异均不显著;（3）不同水分处理,冰草和冷蒿的根系均可分布在30-60cm土壤中,且土壤30-60cm层冰草单种的根生物量和根长显著高于冷蒿单种;（4）相对竞争强度和竞争攻击力系数表明：湿润处理冰草的竞争力大于冷蒿;上干下干处理冷蒿的竞争力大于冰草。土壤水分变化引起冰草、冷蒿的明显竞争,上湿下干与上干下湿处理间冰草与冷蒿竞争力差异不显著。由于冰草和冷蒿均为多年生物种,冰草-冷蒿的竞争实验仍需在今后的研究中反复地验证。     

Changes in biomass and spatial distribution of <Emphasis Type="Italic">Elodea nuttallii</Emphasis> (Planch.) St. John,an invasive submerged plant,in oligomesotrophic Lake Kizaki from 1999 to 2002

Distribution of pure Elodea nuttallii vegetation was surveyed from 1999 to 2002, immediately after the most recent expansion of the species in Lake Kizaki, Japan. During 2001 and 2002, areas of E. nuttallii vegetation rapidly diminished and the summer plant height decreased wherever the vegetation remained. The organic matter content, total phosphorus, and extracted P of the sediment from the vegetation bed were measured. A linear relationship was observed between the extracted P in the sediment and the biomass. The extracted P significantly decreased in the shallow littoral vegetation bed, where the biomass clearly diminished. A fertilization experiment using the shallow littoral sediment collected in the vegetation bed was conducted in 2001. In this experiment, apical shoots of E. nuttallii were planted in pots with fertilized sediment (nitrogen, phosphorus, and potassium additions). The growth of E. nuttallii shoots was significantly enhanced by enrichment with phosphorus alone. The ecological implication of sediment phosphorus limitation is discussed in relation to the cause of decline in the E. nuttallii population in Lake Kizaki.     

Effects of trait plasticity on aboveground biomass production depend on species identity in experimental grasslands

Experimental studies with grassland species found a positive relationship between species richness and community biomass production, however the response of individual species was highly variable. The mechanisms behind these patterns are poorly understood. Here we studied aboveground biomass production and plasticity of growth characteristics of four legumes with similar morphology (Lotus corniculatus, Medicago lupulina, Onobrychis viciifolia, Trifolium hybridum) in experimental grasslands varying in species richness (1, 2, 4, 8, 16 and 60) and composition. We identified O. viciifolia and T. hybridum as species that reached higher biomass production in mixtures than expected from monoculture yields, while L. corniculatus and M. lupulina mostly had lower yields than expected. Variation of morphological traits across the species-richness gradient was lowest in the highly competitive O. viciifolia, but increased in the smaller species. The tall-growing O. viciifolia achieved higher biomass production by both, a higher number of plant individuals and an increase in mean mass per individual. Mean shoot number per individual remained constant, but individuals produced heavier shoots. The medium-sized T. hybridum also increased the number of plant individuals, but mean mass per individual did not respond to community species richness. The average mass per shoot was increased in mixtures, but the species developed less shoots per individual. Shoot length and stem weight ratio of T. hybridum increased with community species richness. Morphological changes in the less successful L. corniculatus and M. lupulina with a smaller growth stature were similarly directed as those of T. hybridum. The observed morphological changes are known as typical shade-avoidance mechanisms in dense vegetation. Our study shows that stress responses to changes in resource availability may be a mechanism to enforce higher aboveground biomass production of individual species in mixtures, but it depends on species identity whether trait plasticity is large enough to exceed stress-induced growth limitations.