Lignin signatures show the effects of changes in heather and bracken cover on the composition of organic matter in a moorland soil profile

Bracken (Pteridium aquilinum) is aggressively displacing heather (Calluna vulgaris) on many moorlands in Britain. We investigated the use of lignin derivatives to identify the distribution of soil organic matter (SOM) derived from bracken in moorland soil profiles formed under heather. Phenylpropanoids extracted from recently senesced litters, roots and SOM, using alkaline CuO oxidation, showed distinct signatures for bracken and heather, with vanillyl moieties dominating bracken litter extracts and vanillyl and syringyl dominating heather litter extracts. Ratios of vanillyl and syringyl concentrations characterised the SOM derived from heather and bracken better than the concentrations of the individual moieties. The analysis showed up to a depth of 5 cm under pure bracken cover, and at the interface between heather and bracken, the SOM was largely derived from bracken litter but below that depth SOM was apparently derived from heather. The use of these methods to identify the plant origin of SOM not only enables understanding the effects of changing vegetation cover on organic matter dynamics in moorland soils but could also facilitate management techniques in moorland/heathland restoration which involve the removal of comparatively nutrient-rich SOM derived from bracken. Received: 30 November 1997 / Accepted: 11 April 1998     

Effects of grazing and topography on long-term vegetation changes in a Mediterranean ecosystem in Israel

The dynamics of Mediterranean vegetation over 28 years was studied in the Northern Galilee Mountains, Israel, in order to identify and quantify the major factors affecting it at the landscape scale. Image analysis of historical and current aerial photographs was used to produce high resolution digital vegetation maps (pixel size = 30 cm) for an area of 4 km² in the Galilee Mountains, northern Israel. GIS tools were used to produce corresponding maps of grazing regime, topographic indices and other relevant environmental factors. The effects of those factors were quantified using a multiple regression analyses. Major changes in the vegetation occurred during the period studied (1964–1992); tree cover increased from 2% in 1964 to 41% in 1992, while herbaceous vegetation cover decreased from 56% in 1964 to 24% in 1992. Grazing, topography and initial vegetation cover were found to significantly affect present vegetation patterns. Both cattle grazing and goat grazing reduced the rate of increase in tree cover, yet even intensive grazing did not halt the process. Grazing affected also the woody-herbaceous vegetation dynamics, reducing the expansion of woody vegetation. Slope, aspect, and the interaction term between these two factors, significantly affected vegetation pattern. Altogether, 56% and 72% of the variability in herbaceous and tree cover, respectively, was explained by the regression models. This study indicates that spatially explicit Mediterranean vegetation dynamics can be predicted with fair accuracy using few biologically important environmental variables.     

Seed bank and vegetation composition of forest stands of varying age in central Belgium: consequences for regeneration of ancient forest vegetation

Vegetation composition differs significantly between ancientand recent forest, due to slow colonization capacity of typical forest speciesand the higher abundance of early successional species in recent forest.However, little is known about differences in persistent seed bank compositionbetween ancient and recent forest and about the interaction between seed bankand vegetation in relation with forest age. We surveyed the seed bank and theunderstorey vegetationcomposition in transects from ancient to recent forest. Seed bank and fieldlayer vegetation characteristics and similarity between seed bank andvegetationwere analysed in relation to recent forest age and distance to the ancientforest. A total of 39 species and 14,911 seedlings germinated, whichcorresponds with a seed density of 12,426 seeds/m².Total seed density is significantly higher in the youngest recent forest parcel(55 years). Also the seed bank composition in the youngest forest parceldifferssignificantly from the other parcels. After a longer period of reforestation,the seed bank approaches that of the ancient forest, suggesting seed bankdepletion, although the seed bank is permanently replenished to some extent byseed bank forming species from local disturbances. Seed bank composition doesnot change significantly with distance to the ancient forest. Similaritybetween seed bank and vegetation composition, nomatter the forest age, is very low, but decreases with increasing forest age.The most frequent species in the vegetation are absent in the seed bank andvice versa. The contribution of forest species is highin the vegetation and they almost not occur in the seed bank, while species offorest edges and clearings, and species of disturbed environments are morefrequent in the seed bank. The seed bank is mainly composed of earlysuccessional species of former forest stages or species which temporary occurinsmall-scale disturbances. The seed bank may enhance the negative effects ofearlysuccessional, mainly competitive species to the forest species richness in therecent forest. In this respect, forest management should minimise forestdisturbances, to prevent germination of competitive species form buriedseeds.     

Effects of simultaneous changes in light, nutrients, and herbivory levels, on the structure and function of a subtropical turtlegrass meadow

The interactive effects of light, nutrients, and simulated herbivory on the structure and functioning of a subtropical turtlegrass bed were analyzed monthly from May to October 2001 in Perdido Bay, FL. For each of the three factors, two levels were evaluated in a factorial design with four replicates per treatment. The variables included: light, at ambient and 40% reduction; nutrients, at ambient and 2× ambient concentrations; and herbivory, with no herbivory and simulated effects of a density of 15 sea urchins/m². In practice, light levels turned out to be 40% of surface PAR for ambient conditions, and 16% for shaded plots. Biomass removed as herbivory represented, on average, slightly less than 20% of the above-ground biomass. Separate three-way ANOVAs found no significant three-way interactions for any of the response variables, and few two-way interactions. There were no significant nutrient effects on turtlegrass above-ground biomass, although nutrient additions produced significant decreases in epibiont biomass, and net above-ground primary production (NAPP); significant increases in below-ground biomass during the peak of the growing season. Shoot density and average number of leaves per shoot increased significantly, while the C/N ratio of the oldest leaf in the enriched plots decreased significantly. Light reduction significantly negatively affected all response variables, except below-ground biomass, shoot density and leaf length. Herbivory had isolated and inconsistent significant effects on below-ground biomass, shoot density, average number of leaves per shoot, and leaf length and width. Overall, our results indicate that nutrients are not limiting in Perdido Bay, and that nutrient additions had mostly detrimental effects. Light appeared to be the most important variable limiting seagrasses growth and abundance, and as with terrestrial plants, seagrasses seemed to respond more to light and nutrients than to herbivory. However, it is essential that additional tests of the single and interactive effects of the three key factors of light, nutrients and herbivory be done to evaluate the generality of our work, since our study is the first of its kind in seagrass meadows.     

Nitrogen and phosphorus limitation in a coastal barrier salt marsh: the implications for vegetation succession

1 A factorial fertilizer experiment was conducted in a 15-year-old coastal barrier salt marsh with a low soil nitrogen content, and in an older 100-year-old marsh with a higher nitrogen content. Plots were fertilized at high and low marsh elevations in both marshes. Nitrogen and phosphorus were applied at low and high concentrations both separately and in combination in each of 3 successive years.
2 Nitrogen limited above-ground plant growth in both young and old salt marshes in all years. Phosphorus limitation of plant growth was apparent in the first year in the young marsh and in the last year in both marshes. In young marshes with low soil organic matter, phosphorus limitation may occur. In addition, phosphorus limitation occurs at both successional stages when a marsh is saturated with nitrogen.
3 Plant species that are typical of nitrogen-rich habitats and late successional stages significantly increased in biomass after fertilization. Limonium vulgare , a low stature species of early and intermediate successional stages, decreased in biomass, whereas the taller Elymus pycnanthus and Artemisia maritima increased. After 3 years of fertilization, plant species composition in a young marsh was similar to the species composition in an unfertilized older marsh. Fertilization of a 100-year-old marsh, however, still resulted in a change in plant species composition, suggesting that succession was still occurring and that, overall, plants in marshes of different age are similar in their response to fertilization.     

Studies on decomposition ofCeratophyllum demersum litter under laboratory and field conditions: losses of dry mass and nutrients,qualitative changes in organic compounds and consequences for ambient water and sediments

A study was made of decomposition ofCeratophyllum demersum litter over a 17-day period under controlled conditions of temperature and oxygen (5, 10 and 18 °C; aerobic and anaerobic) and over a 169-day period in the field (Lake Vechten, The Netherlands). Litter, water and sediment were sampled on the 0, 2, 4, 7 and 17th day under controlled conditions and on the 0, 17, 49, 127 and 169th day in the field. The litter was analyzed quantitatively for dry mass, ash, carbon, nitrogen, phosphorus and qualitatively of organic composition by pyrolysis mass spectrometry. The water was analyzed for the elemental concentrations of organic carbon (total and dissolved), nitrogen (total, ammonia and particulate) and phosphorus (total and orthophosphate) and for the concentrations of photosynthetic pigments and bacteria. The sediment was analyzed for the elemental concentrations of nitrogen, carbon and phosphorus, and for bacterial numbers.The pattern of litter mass loss fitted an exponential model fairly well. Mass decreased faster under controlled aerobic than under anaerobic conditions and the decrease was stimulated by increasing temperature, relatively more in the range of 5 to 10 °C (by 20%) than in the range of 10 of 18 °C (by 2%). The residual mass ranged from 73 to 43% of initial under controlled aerobic conditions and from 84 to 65% under anaerobic conditions after 17 days. It decreased far less in the field, to 38% of initial mass in the field after 169 days.The litter initially lost a carbohydrate fraction by leaching in all treatments. The protein content decreased initially as well but increased subsequently at increasing temperature stimulated under anaerobic conditions. The changes in organic composition were correlated with those in nitrogen but not with those in carbon and phosphorus contents. The organic composition of litter incubated in the field differed from that of litter incubated in the laboratory. The field residues contained less proteinaceous material than the laboratory residues.The changes in carbon, nitrogen and phosphorus concentrations in the litter showed different patterns. The carbon concentration generally increased, the nitrogen concentration initially dropped and increased subsequently, and the phosphorus concentration initially dropped and remained relatively constant subsequently. Chemical immobilization of the decomposition process may have occurred in the laboratory, but was unlikely in the field.Carbon, nitrogen and phosphorus left the litter initially largely in particulate form and were recovered in the water. The ratio dissolved: total nutrient concentration was lower under controlled aerobic than under anaerobic conditions. Increasing temperature stimulated bacterial use of dissolved organic carbon and nitrogen. A rapid nutrient flow occurred from macrophyte litter, via water to sediment.The phytoplankton biomass in the water was greatly stimulated by substances freed from the decomposing litter. Diatoms increased generally relatively more than green algae, predominating alternatively with green algae under aerobic conditions and continuously under anaerobic conditions. Bacterial numbers in the water initially increased, partly due to transgression of bacteria from the sediment-water interface to the water and partly due to an actual increase in community biomass. The bacteria returned largely to the sediment-water interface, stimulated by increasing temperature, as most of the substrate readily usable by them had left the litter in the litter-bag and was associated with the upper sediment layers.It is feasible that the annual die-off of theC. demersum population of Lake Vechten barely affects nutrient cycling in the lake, because the contribution to the nutrient pools of the lake when fully mixed is only small. However, small particles originating from decomposingC. demersum litter may influence the lake considerably by decreasing water transparency and serving as a food source for filter-feeders and detritivorous macrofauna.     

Effects of weed control on species composition of aquatic plants and bank plants and macrofauna in ditches

For several years, the effect of ditch management measures on the species composition of aquatic and bank flora and macrofauna were studied in a polder with a peat soil. The mowing basket increased the presence of floating plants whereas the ditch-scoop increased filamentous algae and submerged plant species. Monospecific stands of aquatic plants had fewer macrofaunal species than stands with many plant species. Direct and indirect effects of management measures on the macrofauna composition were shown.     

Selection harvest in temperate deciduous forests: impact on herb layer richness and composition

Herb layer richness and composition of four forest types and three different management treatments were investigated in 16 deciduous forest stands of northern Germany. Specifically, we compared the species richness and composition occurring in mature forest stands that were single-tree and group selection harvested to those in unmanaged reference stands. Mean species richness of all herb layer species increased significantly with increasing harvest severity. When analyzing plant groups separately, it became obvious that this overall pattern was not consistent. While a negative relationship was detected between vernal herb richness and harvest severity, group selection harvest significantly increased species richness of summer herbaceous forest species and generalists. Woody species richness was not related to harvest severity. Community composition of the spring aspect was not significantly affected by selection harvest, whereas herb layer species composition in the summer aspect differed significantly among the three harvest treatments. A dominance of highly competitive shrub and generalist species was confined to some parts of the most intensively harvested stands. Overall, our results indicate that the herb layer community was not severely adversely affected by selection harvest at the intensities used in the studied stands. It is suggested that selection harvest systems may be feasible tools with which to conserve local forest vascular plant diversity and at the same time to meet the demand for timber products. However, information about forest history and the implementation of the selection harvest system are basic requirements when interpreting the results of studies on understorey response to selection harvest.     

Relative importance of nutrients and mortality factors on prokaryotic community composition in two lakes of different trophic status: microcosm experiments

The effect of nutrient resources (N and P enrichment) and of different grazing communities on the prokaryotic community composition (PCC) was investigated in two freshwater ecosystems: Sep reservoir (oligomesotrophic) and lake Aydat (eutrophic). An experimental approach using microcosms was chosen, that allowed control of both predation levels, by size fractionation of predators, and resources, by nutrient amendments. Changes in PCC were monitored by fluorescent in situ hybridization (FISH) and terminal-restriction fragment length polymorphism (T-RFLP). The main mortality agents were (i) heterotrophic nanoflagellates and virus-like particles in Aydat and (ii) cladocerans in Sep. All the nutritional elements assayed (N-NO3, P-PO4 and N-NH4) together with prokaryotic production (PP) always accounted for a significant part of the variations in PCC. Overall, prokaryotic diversity was mainly explained by resources in Sep, by a comparable contribution of resources and mortality factors in lake Aydat and, to a lesser extent, by the combined action of both.     

A fence-line contrast reveals effects of heavy grazing on plant diversity and community composition in Namaqualand,South Africa

Changes in plant species richness and community composition were investigated across a fence separating heavily grazed communal and lightly grazed commercial farming systems in Namaqualand, South Africa. No significant differences in plant species richness between communal and commercial farming systems were detected either locally within individual plots or overall across all plots. Within-plot, richness of species tolerant of grazing, such as annuals and geophytes, has increased, while the richness of large palatable shrub species has decreased on the communal rangeland. In terms of plant cover, species' responses to grazing were strongly associated with growth form. Annuals and geophytes formed the majority of grazing increasers, while large, presumably palatable, shrubs and leaf succulents were characteristic grazing decreasers. An investigation into population processes of five shrub species revealed that heavy grazing on the communal rangeland has resulted in: reduced size of palatable shrub species; reduced flower production and seedling recruitment of palatable species; increased density and recruitment of the unpalatable shrub, Galenia africana. Reductions in shrub volume, reproductive output and seedling recruitment were most marked in the palatable shrub Osteospermum sinuatum and were in the order of 90%. The results are further discussed in terms of their relevance to rangeland dynamics and the current land use practices of the region.