Two centuries of vegetation succession in an inland sand dune area,central Netherlands

Questions: (a) What are the rates and directions of vegetation succession in an inland sand‐dune system? (b) What are the differences in successional trajectories in different relief types? and (c) Is it possible to preserve the last areas of still active dunes and under what circumstances? Location: The study sites were located in the northern part of the Veluwe Region, central Netherlands; longitude 5°44′ E, latitude 52°20′ N, altitude 9 to 24 m a.s.l. Methods: Vegetation and relief mapping was conducted in three permanent plots, 200 m × 200 m in size, in 1988 and 2003. Phytosociological relevés (2400) were recorded in each 10 m × 10 m subplots. Age of woody species was determined by wood coring. Geographic Information System, ordination analyses, and TWINSPAN were used for data exploration and elaboration. Results: A total of 70 vascular plants and 19 bryophytes were recorded over successional stages spanning approximately190 years. The following dominant species formed the sequence of successional stages, but not all participated in all relief types: Ammophila arenaria, Festuca arenaria, Corynephorus canescens, Festuca ovina and Agrostis capillaris, and pine forest dominated in its herb layer at first by Deschampsia flexuosa and later by either Empetrum nigrum, Vaccinium myrtillus or Vaccinium vitis‐idaea. Conclusions: The successional trajectory is basically unidirectional for more than 100 years; no clear multiple successional pathways were observed, as is frequent in coastal dunes. Successional divergence was observed after approximately 130 years in the composition of the herb layer in the closed pine forest. The obvious vegetation heterogeneity in the still active sand‐blown area is related to differences in timing of vegetation establishment on particular relief types, thus the succession exhibits a terrain‐dependent asynchronous character. We conclude that the last patches of still‐active sand dunes can be preserved only by repeated strong artificial disturbances.     

Transition from heathland to grassland initiated by the heather beetle

This paper discusses the transition from heather-dominated vegetation to one dominated by grass. The dominating species involved are Calluna vulgaris and Deschampsia flexuosa. The study covered three years, 1980–82, and was done at the Hoorneboeg heathland in the Netherlands. Several factors account for this transition: damage caused by the chrysomelid heather beetle Lochmaea suturalis Thomson, soil type, and interaction between beetle damage and Deschampsia when already present underneath or in the immediate vicinity of a Calluna canopy. There was a time-lag between beetle damage and grass establishment or expansion. In fact, this transition concerns a type of vegetation succession and can be treated as such. This succession therefore involves several causative factors external to the system Calluna-Deschampsia. The successional process is conceived as a stochastic one defined by transition probabilities which depend on a time component, a space component and a logistic response component. All these components are discussed in this paper, but the discussion of yet another component, the crowding of Deschampsia itself, will be treated separately.     

Phylogenetic relationships of Deschampsia antarctica (Poaceae): Insights from nuclear ribosomal ITS

Deschampsia antarctica E. Desv. is the only monocot in the Antarctic floristic zone. We evaluated the phylogenetic relationships of Deschampsia antarctica to other grasses using parsimony as the optimality criterion. Five different sets of gap, transversion and transitions costs were explored to analyze the effect of parameter choice on the phylogenetic results. Both internal transcribed spacers (ITS1 and ITS2) and the 5.8S subunit of nuclear ribosomal DNA were included in the analysis. A total of 43 species were analyzed including seven species of Deschampsia. Deschampsia antarctica forms a well supported group with five species of Deschampsia. Deschampsia does not appear monophyletic as D. flexuosa (L.) Trin. is not included in this clade. The clade to which D. antarctica belongs is sister to some Aveneae in all analyses. This study is the first contribution that evaluates the phylogenetic position of D. antarctica in relation to other species of Deschampsia.     

Carbon and nitrogen in soil and vegetation at sites differing in successional age

We studied vegetation and soil development during primary succession in an inland drift sand area in the Netherlands. We compared five sites at which primary succession had started at different moments in the past, respectively 0, 10, 43 and 121 years ago, and a site at which succession had not yet started. In the three younger sites the vegetation was herbaceous, whereas in the two older sites a pine forest had formed. Forest formation was accompanied by the development of an FH-layer in the soil, an increase in the amount of soil organic matter, and an increase in nitrogen mineralisation rate from 1.9 to 18 g N m^–2 yr^–1. Soil moisture content also increased, whereas pH showed a steady decrease with site age. The vegetation changed from a herbaceous vegetation dominated by mosses and lichens and the grass species Corynephorus canescens and Festuca ovina towards a pine forest with an understorey vegetation dominated by Deschampsia flexuosa and, at the oldest site, with dwarf shrubs Empetrum nigrum and Vaccinium myrtillus. At the same time the total amounts of carbon and nitrogen of the ecosystem increased, with a relatively stronger increase of the carbon pool. The establishment of trees during succession greatly affects the dynamics of the ecosystem, especially its carbon dynamics.     

Reciprocal interactions between plants and soil in an upland grassland

Through the production of litter, plants with different life history strategies are predicted to both affect and be affected by the properties of soil. Competitive species are expected to increase the fertility of, and have a positive growth feedback with, soil, whereas stress-tolerant species should decrease fertility but show no growth feedback. We maintained monocultures of competitive (Lolium perenne and Agrostis capillaris) and stress-tolerant (Festuca ovina and Nardus stricta) grasses on an unproductive grassland for six years. The Nardus soil developed significantly greater inorganic nitrogen than the Agrostis and Festuca soil, and significantly greater soil moisture content than the Festuca soil. However, there were no differences in organic matter content, phosphate or bulk density between the soil types. In a greenhouse assay, each species was grown in soil cores from the different monocultures as well as natural turf. There were significant differences in growth between plant species and soil types. As expected, L. perenne produced the greatest amount of biomass. However, plants grown on Nardus soil were twice as large and had a 21% lower root allocation than plants grown on any of the other soil types. Lolium perenne, A. capillaris and F. ovina had significant negative growth feedbacks with their own soil (−0.460, −0.821 and −0.792, respectively) and N. stricta had a significant positive feedback (0.560). This study highlights the difficulties of predicting how plant traits will affect soil properties.     

Mycorrhizae transfer carbon from a native grass to an invasive weed: evidence from stable isotopes and physiology

Invasive exotic weeds pose one of the earth's most pressing environmental problems. Although many invaders completely eliminate native plant species from some communities, ecologists know little about the mechanisms by which these exotics competitively exclude other species. Mycorrhizal fungi radically alter competitive interactions between plants within natural communities, and a recent study has shown that arbuscular mycorrhizal (AM) fungi provide a substantial competitive advantage to spotted knapweed, Centaurea maculosa, a noxious perennial plant that has spread throughout much of the native prairie in the northwestern U.S. Here we present evidence that this advantage is potentially due to mycorrhizally mediated transfer of carbon from a native bunchgrass, Festuca idahoensis, to Centaurea. Centaurea maculosa, Festuca idahoensis (Idaho fescue, C₃), and Bouteloua gracilis (blue gramma, C₄) were grown in the greenhouse either alone or with Centaurea in an incomplete factorial design with and without AM fungi. Centaurea biomass was 87–168% greater in all treatments when mycorrhizae were present in the soil (P < 0.0001). However, Centaurea biomass was significantly higher in the treatment with both mycorrhizae and Festuca present together than in any other treatment combination (P < 0.0001). This high biomass was attained even though Centaurea photosynthetic rates were 14% lower when grown with Festuca and mycorrhizae together than when grown with Festuca without mycorrhizae. Neither biomass nor photosynthetic rates of Centaurea were affected by competition with the C₄ grass Bouteloua either with or without mycorrhizae. The stable isotope signature of Centaurea leaves grown with Festuca and mycorrhizae was more similar to that of Festuca, than when Centaurea was grown alone with mycorrhizae (P = 0.06), or with Festuca but without mycorrhizae (P = 0.09). This suggests that carbon was transferred from Festuca to the invasive weed. We estimated that carbon transferred from Festuca by mycorrhizae contributed up to 15% of the aboveground carbon in Centaurea plants. Our results indicate that carbon parasitism via AM soil fungi may be an important mechanism by which invasive plants out compete their neighbors, but that this interaction is highly species-specific.     

Glycine uptake in heath plants and soil microbes responds to elevated temperature, CO2 and drought

Temperate terrestrial ecosystems are currently exposed to climatic and air quality changes with increased atmospheric CO₂, increased temperature and prolonged droughts. The responses of natural ecosystems to these changes are focus for research, due to the potential feedbacks to the climate. We here present results from a field experiment in which the effects of these three climate change factors are investigated solely and in all combinations at a temperate heath dominated by heather (Calluna vulgaris) and wavy hair-grass (Deschampsia flexuosa).Climate induced increases in plant production may increase plant root exudation of dissolved organic compounds such as amino acids, and the release of amino acids during decomposition of organic matter. Such free amino acids in soil serve as substrates for soil microorganisms and are also acquired as nutrients directly by plants. We investigated the magnitude of the response to the potential climate change treatments on uptake of organic nitrogen in an in situ pulse labelling experiment with ¹⁵N¹³C₂-labelled glycine (amino acid) injected into the soil.In situ root nitrogen acquisition by grasses responded significantly to the climate change treatments, with larger ¹⁵N uptake in response to warming and elevated CO₂ but not additively when the treatments were combined. Also, a larger grass leaf biomass in the combined T and CO₂ treatment than in individual treatments suggest that responses to combined climate change factors cannot be predicted from the responses to single factors treatments.The soil microbes were superior to plants in the short-term competition for the added glycine, as indicated by an 18 times larger ¹⁵N recovery in the microbial biomass compared to the plant biomass. The soil microbes acquired glycine largely as an intact compound (87%), with no effects of the multi factorial climate change treatment through one year.     

Litter effects of two co-occurring alpine species on plant growth, microbial activity and immobilization of nitrogen

We measured the litter chemistry of two co‐dominant alpine species, Acomastylis rossii, a forb characterized by a low growth rate and N uptake capacity, and Deschampsia caespitosa, a grass characterized by a high growth rate and N uptake capacity, and examined the effect litter of these two species had on the growth of Deschampsia phytometers in a greenhouse. We also examined the influence of litter from the two species on microbial respiration and immobilization of N, in two separate laboratory microcosm experiments and in the field. We hypothesized that Acomastylis litter would reduce plant growth more than Deschampsia litter, corresponding with either 1) suppression of microbial activity and thus a decrease in N mineralization, or 2) by stimulation of microbial biomass and increasing microbial immobilization of N. Relative to Deschampsia litter, Acomastylis litter had higher total water soluble organic carbon (DOC), and higher total phenolic concentration. Deschampsia litter had 30 times higher carbohydrate (primarily glucose and fructose) concentrations than Acomastylis litter. Soil respiration, microbial biomass N, and consumption of DOC and N were higher with the Acomastylis litter treatment than the Deschampsia litter treatment in experimental microcosms, and both respiration and microbial biomass N were higher in field soils under canopies dominated by Acomastylis relative to those dominated by Deschampsia. These results indicate that phenolics in Acomastylis are a C source for soil microorganisms, rather than an inhibitor of microbial activity. Deschampsia phytometers grew significantly less, had higher root: shoot biomass ratios, and acquired less nitrogen in the Acomastylis litter treatment relative to the control and Deschampsia litter treatments. The results of these experiments indicate that Acomastylis litter influences soil N cycling by increasing microbial activity and N immobilization, which may influence N supply to neighboring plants. This mechanism has the potential to influence competitive interactions between Acomastylis and its neighbors.     

Contrasted Responses of Two Understorey Species to Direct and Indirect Effects of a Canopy Gap

Positive associations between adult trees and understorey species have been explained either by direct or indirect facilitation. We tested both models by comparing the performance of two understorey species with contrasted stress-tolerance abilities Galium odoratum and Deschampsia flexuosa. Individuals of both species were transplanted in the four combinations of two treatments (gap and removal of an herbaceous competitor, Molinia caerulea). Our experiment demonstrated that direct facilitation of adult trees may explain the restricted occurrence of the shade-demanding Galium within closed forest communities. In contrast, the shade-tolerant Deschampsia was subjected to additional competition within the forest, likely because adult trees had a higher negative effect on light availability and a similar negative effect on nitrogen availability within the forest than did Molinia in the gaps.     

Root turnover as determinant of the cycling of C,N, and P in a dry heathland ecosystem

Root production and turnover were studied using sequential core sampling and observations in permanent minirhizotrons in the field in three dry heathland stands dominated by the evergreen dwarfshrub Calluna vulgaris and the grasses Deschampsia flexuosa and Molinia caerulea, respectively. Root biomass production, estimated by core sampling, amounted to 160 (Calluna), 180 (Deschampsia) and 1380 (Molinia) g m^-2 yr^-1, respectively. Root biomass turnover rate in Calluna (0.64 yr^-1) was lower compared with the grasses (Deschampsia: 0.96 yr^-1; Molinia 1.68yr^-1)). Root length turnover rate was 0.75–0.77 yr^-1 (Deschampsia) and 1.17–1.49 yr^-1 (Molinia), respectively. No resorption of N and P from senescing roots was observed in either species. Input of organic N into the soil due to root turnover, estimated using the core sampling data, amounted to 1.8 g N m^-2 yr^-1(^Calluna), 1.7 g N m^-2 yr^-1 (Deschampsia) and 19.7 g N m^-2 yr^-1 (Molinia), respectively. The organic P input was 0.05, 0.07 and 0.55 g P M^-2 yr^-1, respectively. Using the minirhizotron turnover estimates these values were20–22% (Deschampsia) and 11–30% (Molinia) lower.When the biomass turnover data were used, it appeared that in the Molinia stand root turnover contributed 67% to total litter production, 87% to total litter nitrogen loss and 84% to total litter phosphorus loss. For Calluna and Deschampsia these percentages were about three and two times lower, respectively.This study shows that (1) Root turnover is a key factor in ecosystem C, N, and P cycling; and that (2) The relative importance of root turnover differs between species.     

Nitrification in Dutch heathland soils

A survey was conducted over a range of 17 Dutch heathland locations, subdivided into 41 sites dominated by either dwarf-shrubs (Calluna vulgaris or Erica tetralix) or grass species (Deschampsia flexuosa or Molinia caerulea). Among the habitats of the dominant plant species relatively little differences in general soil properties were observed. The P status of Deschampsia sites was relatively high as well as the NO₃ ^–-N concentrations in the 0–10 cm layer (FH included) at the grass-dominated sites. At sites with a dead or degenerating dwarf-shrub vegetation, NH₄ ⁺-N concentrations reached very high levels.Net production of nitrate was observed during incubation of intact 0–10 cm soil cores (FH-layer included) in the laboratory for all sites, even though in some instances, particularly at Calluna and Erica sites, no nitrate was initially measured. Generally, a higher nitrification rate was found for the grass-dominated sites, and for Deschampsia in particular. The net production of nitrate was highly significantly correlated with net N mineralization, being a reasonable predictor of nitrification in a simple regression model (R²=0.47; P<0.001). Net nitrification was also significantly correlated with the NO₃ ^–-N initially present at the start of the growing season (R=0.65; P<0.001) and with the labile organic P content of the soil (R=0.65; P<0.001). By including initial NO₃ ^–-N and labile organic P, together with net N mineralization and pH, in a multiple regression model, net nitrate production could be predicted with a much higher precision (R²=0.75; P<0.001). Although apparent nitrification was not significantly correlated with pH, the latter contributed significantly to the multiple regression equation for the prediction of the former.The influence of the labile organic P pool may act via its positive correlation with microbial biomass, thus more or less reflecting the potential mineralization/nitrifying capacity of a particular site.     

Nutrient pools and fluxes of the ground vegetation in coniferous forests due to fertilizing,liming and amelioration

The effects of fertilization and amelioration treatments on some nutrient pools and fluxes of ground vegetation in mature pine and spruce stands on acid soils in South Germany are described. In N-limited pine forests with moderate canopy density and with Deschampsia flexuosa an additional N-accumulation in biomass of 20–40 kg ha^-1 occurred 3 years after pure N-fertilization. The N, P, K-cycling through ground vegetation was stimulated more than 10 years by a combined N + CaCO₃ + P treatment leading toa shift in dominance from cryptogams and Ericaceae towards Deschampsia flexuosa and ruderal species like Epilobium angustifolium. The effect of a lupine treatment (combined with initial soil preparation, liming and P supply) was far stronger than the effect of the other experimental procedures. But the fertilizer and amelioration effects on the herb layer of pine forests tended to decline after two decades for different reasons.The shade-tolerant ground vegetation in a nitrogen-saturated spruce forest was not able to prevent heavy additional nitrate losses from upper mineral soil after dolomitic liming. But the Ca, Mg and K fluxes through ground vegetation were strongly elevated in the third year after treatment.     

Effects of prescribed burning on plant available nutrients in dry heathland ecosystems

Heathland management is an important tool with which to modify ecosystem impacts caused by atmospheric nutrient deposition. Since changes in nutrient availability as a result of management measures affect the outcomes of heathland succession and species competition, studies on this issue are important from both a nature conservation and management point of view. This study reports the effects of prescribed burning on nutrient availability in dry heathland soils and the nutrient content of the two competing heathland species Calluna vulgaris and Deschampsia flexuosa, with particular reference to N and P. We hypothesise that winter prescribed burning leads to additional N availability, which enhances the importance of P in the context of nutrient limitation in heathland ecosystems. In the nature reserve “Lueneburg Heath” (NW Germany) we examined the availability of nutrients in the humus horizons and in the leachate as well as the relevant C:element ratios in Calluna and Deschampsia before and after a burning experiment. Our results show that prescribed burning resulted in drastically increased NH₄⁺ availability in the O-horizon. We observed only short-term effects (for NO₃⁻, PO₄³⁻, Mg) and insignificant effects on the availability of other nutrients (K, Ca). As a consequence of an increased nutrient availability in the humus horizons and a limited nutrient uptake by plants after burning, leaching increased significantly for N, Ca, K, and Mg after burning treatment. No significant changes were found in the foliar C:N ratios for either species after prescribed burning, although Deschampsia showed an increased deficiency for all the other nutrients, particularly for P, as expressed by increased foliar C:P and N:P ratios. By contrast, the nutrient content of Calluna did not change significantly, suggesting that prescribed burning favours the competitive capacity of Calluna as against Deschampsia. We assume that water shortage as a result of changes in the microclimate was mainly responsible for the deterioration of the nutrient content of Deschampsia. This gives Calluna a competitive advantage, enabling it to out-compete Deschampsia on burned heathlands, with respect to the key factor P-limitation.     

Habitat characteristics and the effect of various nutrient solutions on growth and mineral nutrition ofArnica montana L. grown on natural soil

Arnica montana, one of the character species of the replacement plant communityViolion caninae on sandy acid humic podzol, is declining in the Netherlands since 1950. Locally, it is even extinct.This process of decline may be attributed to (i) autonomic succession; (ii) increased rate of acidification of the soil and/or (iii) change in competitive relations amongArnica and more productive plant species, such as the successive dominantDeschampsia flexuosa. This paper examines the last two hypothesized factors, supposedly being regulated by atmospheric input of N-input, by (a) describing the habitat characteristics of a seemingly still healthy individual-rich population ofArnica and (b) growingArnica andDeschampsia in mixed humic podzol soil (mainly A₁ horizon) fertilized with variously-composed nutrient solutions in order to assess the nutrient supplying capacity of the podzol soil and species-specific nutritional demands related to their respective growth capabilities.The results suggest that an increased rate of soil acidification yielding extra supply of ionic Al and Mn is presumably of less importance. The implications of growth rate differences amongArnica andDeschampsia as related to their nutritional demands are likely far more important.Arnica grows more slowly thanDeschampsia in fertilized humus podzol. The latter species has a much more efficient use of nutrients. Furthermore, both species differ in K-nutrition if NH₄ ⁺ is the dominant N-source, a situation normally occurring in acid podzolic soils.Deschampsia possesses the capability to adapt its metabolic nutrition (avoiding nutritional imbalances) to a wide range of nutrient supplies thereby maintaining a rather constant level of growth.Arnica, on the contrary, lacks this capability. These results are discussed in the framework of competitive relations among co-occurring plant species in the plant communityViolion caninae. It is concluded that maintenance ofArnica and likely other character species of that alliance — all characterized by a low growth rate — will only be achieved when the plant community is properly managed by trampling, mowing or light levels of grazing (low stocking rate). Of prime importance is to maintain a low and relatively open vegetation structure at a relatively low level of nutrient supply.     

Competition between two grass species with and without grazing over a productivity gradient

Soil nutrient-level and herbivory are predicted to have opposing effects on the allocation pattern of the competitive dominant plant species. Lower stem and higher leaf allocation are favoured when plants are grazed, whereas a higher stem allocation is favoured at high nutrient levels. Grazing by hares and geese can prevent invasion of the tall Elymus athericus, into short vegetation of Festuca rubra, at unproductive stages of salt-marsh succession but not at more productive stages. We hypothesise that the negative effect of herbivory on Elymus decreases due to increasing soil nitrogen levels and shifts the competitive balance towards this species. We tested how simulated grazing and nitrogen availability affected the competitive balance between adult plants of both grass species in a greenhouse experiment. Elymus had a higher above-ground biomass production, invested relatively more in stem and root tissue and had a larger shoot length than Festuca. The above-ground relative yield of Elymus in mixtures of both species increased with increasing nitrogen levels. This indicates that Elymus was the superior competitor at high soil fertility. Although clipping removed relatively more biomass from Elymus than from Festuca and exceeded the observed biomass removal in field conditions, it did not change the competitive balance between both species. Decreasing effects of herbivory due to increasing nitrogen levels are not a likely explanation for the invasion of Elymus in productive marshes. The results suggest that once Elymus has established it can easily invade vegetation dominated by Festuca irrespective of grazing by herbivores such as hares and geese. Herbivory by small herbivores may mainly retard the invasion of this plant by influencing establishment itself.     

果园间作模式下杏树与苜蓿的根系分布特征及其土壤理化性质研究

该研究采用田间小区试验,设计杏树(Prunus armeniaca)下清耕(CK)和杏树间作紫花苜蓿(T)2个处理,实地采集测定各样地不同土层紫花苜蓿的根系生物量以及杏树的侧根系生物量,并测定土壤pH、电导率及其土壤有机质和速效氮含量,分析果园间作模式下紫花苜蓿对果树侧根系垂直分布特征及其土壤理化性质的影响,为果园间作苜蓿模式的推广提供理论依据。结果表明:(1)CK与T处理下的杏树侧根生物量在土壤中的垂直分布都主要集中在20~60cm土层,其生物量分别为750.8g和737.6g,分别占总侧根生物量的64.4%和64.5%;紫花苜蓿根系生物量分布呈倒金字塔型,且主要分布在0~40cm土层(166.3g),其中0~20cm土层的根系生物量最高(97.4g),占根系总生物量的35.8%。(2)与CK处理相比,T处理可有效增加果园表层土壤的有机质含量、速效态氮含量、硝态氮含量和铵态氮含量,其中,在0~20cm土层分别显著增加17.1%、40.8%、28.5%和40.8%,在20~40cm土层分别显著增加36.1%、23.1%、60.2%和23.8%,并显著降低了表层土壤电导率,但对土壤pH无显著影响。研究认为,杏园间作牧草紫花苜蓿虽然杏树与苜蓿根系会发生较小资源的竞争,但有利于改善林下土壤的理化性质和养分状况,能够有效促进果树的生长发育。     

Plant nutrient mobilization in temperate heathland responds to elevated CO2, temperature and drought

Temperate terrestrial ecosystems are currently exposed to increased atmospheric CO₂ and progressive climatic changes with increased temperature and periodical drought. We here present results from a field experiment, where the effects of these three main climate change related factors are investigated solely and in all combinations at a temperate heathland. Significant responses were found in the top soils below the two dominant species (Calluna vulgaris and Deschampsia flexuosa). During winter incubation, microbial immobilization of N and ammonification rate decreased in response to warming in Deschampsia soil, and microbial immobilization of N and P decreased in warmed Calluna soil. Warming tended to increase microbial N and P in Calluna but not in Deschampsia soil in fall, and more microbial C was accumulated under drought in Calluna soil. The effects of warming were often counteracted or erased when combined with CO₂ and drought. Below Deschampsia, the net nitrification rate decreased in response to drought and, while phosphorus availability and microbial P immobilization decreased, but nitrification increased in response to elevated CO₂. Furthermore, leaf litter decomposition of both species decreased in response to drought. These complex changes in availability and release of nutrients from soil organic matter turnover and mineralization in response to elevated CO₂ and climate change may influence the future plant carbon sequestration and species composition at temperate heathlands.     

Intergeneric relationships ofLolium,Festuca, andVulpia (Poaceae) and their phylogeny

Morphological and seed protein analyses of 26 species of the generaLolium, Festuca andVulpia confirmed their close systematic affinities. Six inflorescence characters readily differentiatedFestuca fromLolium. Protein similarities betweenFestuca of sect.Bovinae and cross-pollinated species ofLolium, coupled with cytogenetic and crossability data, substantiate that they should be united into one genus.Vulpia had phenetic similarities with sect.Scariosae, Montanae andOvinae ofFestuca. Lolium, Festuca, andVulpia are most likely derived from a common ancestral form which was close toFestuca pratensis andLolium perenne.     

Experimental manipulation of succession in heathland ecosystems

An experiment was carried out in two heathland ecosystems, one dominated by Calluna vulgaris and the other by Molinia caerulea, to analyse the effects of soil organic matter accumulation and nutrient mineralization on plant species dynamics during succession. The experiment included one treatment that received nutrient solution and two treatments where the rate of soil organic matter accumulation was reduced by removing litter or accelerated by adding litter. In a fourth treatment the C. vulgaris litter produced in the C. vulgaris-dominated plots was replaced by litter of M. caerulea and vice versa. Treatments were applied over 8 years. Addition of nutrient solution caused C. vulgaris to decline, and grass species to increase sharply, compared to the control plots. Addition of litter enhanced both N mineralization and the biomass of M. caerulea and Deschampsia flexuosa but reduced the biomass of C. vulgaris. The effects of replacing C. vulgaris litter by M. caerulea litter, or vice versa, on N mineralization and species dynamics could not be attributed to differences between the decomposability of the different litter materials that were transferred. The results confirm the hypothesis that increased litter inputs accelerate the rate of species replacement during succession.     

Root herbivores,pathogenic fungi,and competition between Centaurea maculosa and Festuca idahoensis

We used a common garden experiment to evaluate the isolated and combined effects of a biocontrol agent, the insect (Agapeta zoegana, Lepidoptera), and a native North American fungal pathogen (Sclerotinia sclerotiorum) on competition between the noxious weed Centaurea maculosa and the native Festuca idahoensis. In 0.5-m² plots with 24 plants per plot, competition between Centaurea and Festuca was highly asymmetrical, with Centaurea strongly reducing the final biomass and reproduction of Festuca, and Festuca having no effect, or possibly a positive effect, on Centaurea. The direct effects of the biological control agents differed entirely. All Centaurea individuals died in plots receiving Sclerotinia, but Agapeta did not significantly reduce the growth of Centaurea, and apparently stimulated a compensatory reproductive response in the weed. Individual Centaurea plants that had been damaged by Agapeta produced more flowerheads, and the number of Centaurea plants with Agapeta root damage in a plot was positively correlated with total Centaurea biomass. These differences in the direct effects of the consumers were reflected in their indirect effects. In plots where Sclerotinia killed Centaurea (strong direct effects) Festuca growth and reproduction was equal to that in Festuca plots without Centaurea and the reproductive output of Festuca increased substantially (strong indirect effects). However, in the absence of Sclerotinia, the application of Agapeta did not significantly decrease Centaurea biomass (weak direct effects) and actually stimulated small, but significant decreases in Festuca reproduction and trends towards lower Festuca biomass (weak and opposite indirect effects –Agapeta does not eat Festuca). If the direct effects of biocontrol agents on Centaurea are weak, as suggested by our results, natives are unlikely to be released from the competitive effects of Centaurea, and natives may suffer from Centaurea's compensatory response to herbivory. This revised version was published online in August 2006 with corrections to the Cover Date.