The ratio of germanium to silicon in plant phytoliths: quantification of biological discrimination under controlled experimental conditions

Slight differences in the chemical behavior of germanium (Ge) and silicon (Si) during soil weathering enable Ge/Si ratios to be used as a tracer of Si pathways. Mineral weathering and biogenic silicon cycling are the primary modifiers of Ge/Si ratios, but knowledge of the biogenic cycling component is based on relatively few studies. We conducted two sets of greenhouse experiments in order to better quantify the range and variability in Ge discrimination by plants. Graminoid species commonly found in North American grassland systems, Agropyron smithii, Schizachyrium scoparium, and Andropogon gerardii were grown under controlled hydroponic environmental conditions. Silicon leaf contents were positively correlated with solution Si and ambient temperature but not with nutrient solution pH, electrical conductivity, or species. The Ge/Si ratio incorporated into phytoliths shows a distribution coefficient [(Ge/Si)_phytolith/(Ge/Si)_solution] of about 0.2 and is remarkably invariant between species, photosynthetic pathway, and solution temperature. Ge seems to be discriminated against during the uptake and translocation of Si to the opal deposition sites by about a factor of five. In the second experiment, a wider range of graminoid species (Agropyron smithii, Bouteloua gracilis, Buchloe dactyloides, Oryzopsis hymenoides, Schizachyrium scoparium and Andropogon gerardii) were grown in two different soil mediums. Plant phytoliths showed a distribution factor of about 0.4 for field grown grasses, and 0.6 for potting soil grown grasses with no clear trends among the species. Evidence of the direction and degree of biological Ge discrimination during plant uptake provides a geochemical finger print for plants and improves the utility of Ge/Si ratios in studies of terrestrial weathering and links between Si cycles in terrestrial and marine systems.     

The effects of reindeer grazing on the composition and species richness of vegetation in forest–tundra ecotone

We conducted an 8-year exclosure experiment (1999–2006) in a forest–tundra ecotonal area in northwestern Finnish Lapland to study the effects of reindeer grazing on vegetation in habitats of variable productivity and microhabitat structure. The experimental sites included tundra heath, frost heath and riparian habitats, and the two latter habitats were characterized by hummock-hollow ground forms. The total cover of vegetation, cover of willow (Salix spp.), dwarf birch (Betula nana), dwarf shrubs, forbs and grasses (Poaceae spp.) increased in exclosures in all habitats. The increase in the total cover of vegetation and in the covers of willow and dwarf birch tended to be greatest in the least productive tundra heath. Opposing to the increase in the dominant vascular plant groups, the cover and species number of bryophytes decreased in exclosures. We conclude that the effects of reindeer grazing on vegetation composition depend on environmental heterogeneity and the responses vary among plant groups. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Temporal trends in species composition and plant traits in natural grasslands of Uruguay

Abstract. We report the successional trends of the major life‐forms (graminoids and forbs) in natural grasslands of Uruguay over a 9‐yr period after the removal of domestic herbivores. For the whole community, species richness and diversity decreased over the successional period. In graminoids we observed clear temporal trajectories in floristic composition; the rate of floristic change decreased with time and was associated with a shift in plant traits. The exclusion of large herbivores promoted erect and tall grasses with narrow leaves and greater seed length, vegetative growth constrained to the cool season and increased frequency of annual species. Forbs did not show a clear temporal trend in species composition, but there was, nevertheless, a plot‐specific species turnover of this functional group that was reflected in their attributes. Species spreading by means of rhizomes, with vegetative growth restricted to the warm season. Species with larger seeds increased under grazing exclusion, as did annual and nitrogen‐fixing forbs. The floristic changes induced by cattle exclusion occurred early in the succession. This early high rate of change has practical implications for management and conservation programs of the natural grasslands of Uruguay. Additionally, the shift in plant traits may be helpful in devising simple indicators of grazing impact.     

Effects of fertilisation and irrigation on ‘foliar afterlife’ in alpine tundra

Question: How do increases in soil nutrient and water availability alter the nutrient fluxes through the resorption and litter decomposition pathways and how do they affect litter nutrient pools in a low‐productive alpine tundra ecosystem? Location: An alpine lichen‐rich tundra on Mt. Malaya Khati‐para in the NW Caucasus, Russia (43°27’ N, 41°42’ E; altitude 2800 m a.s.l.). Methods: We conducted a 4‐year fertilisation (N, P, N+P, lime) and irrigation experiment, and analysed the responses of nutrient resorption from senescing leaves, leaf litter quality and decomposability of six pre‐dominant vascular plant species, total plant community litter production and litter (nutrient) accumulation. Results: Vascular plant litter [N] and [P] increased 1.5 and 10 fold in response to N and P additions, due to increased concentrations of the nutrients in fresh leaves and unchanged or reduced resorption efficiency. Litter decomposability was not affected by nutrient amendments. Fertilisation enhanced litter production (180%; N+P treatment) and litter accumulation (80%; N+P), owing to tremendously increased production and low decomposability of graminoids. Together with increased litter [N] and [P] this led to great increases in total litter nutrient pools. Conclusions: Due to increased production of graminoids, nutrients added to the alpine tundra soil were mostly immobilised in recalcitrant, nutrient‐rich litter. This suggests that changing species composition in low productive ecosystems may act as an internal buffer mechanism, which under increased soil nutrient availability prevents the community from rapidly acquiring features typical of a high productive ecosystem such as high decomposability and high nutrient availability.     

Vegetation variation within and among palustrine wetlands along an altitudinal gradient in KwaZulu-Natal,South Africa

The variation in graminoid species composition and diversity and the distribution of photosynthetic pathways among 66 wetlands in KwaZulu-Natal, South Africa, and within six of these wetlands was described and related to measured physical parameters, using multivariate and univariate techniques. Altitude, which ranged from 550 m to 2120 m, accounted for most variation among wetlands, with an almost complete turnover of species along this gradient. Landform setting was less important in explaining overall species composition, but relationships of individual species were revealed (e.g. Eleocharis dregeana showed an affinity for depressions). Within a wetland there was an almost complete turnover of species along a gradient of wetness, as described using soil morphological criteria. Most species were consistently associated with the same wetness zones across different wetland sites, e.g., Phragmites australis with the wettest zone, Pycreus macranthus with the intermediate zone, and Eragrostis plana with the least wet zone. The occurrence and abundance of different photosynthetic pathway types depended on altitude and degree of wetness. At high altitudes, C₃ sedges, notably Carex acutiformis, dominated the wettest zone and C₃ and C₄ grasses and sedges dominated the intermediate and least wet zones. At mid altitudes, C₃ and C₄ sedges and C₃ grasses dominated the wettest zone, C₃ and C₄ grasses and sedges dominated the intermediate zone and C₄ grasses dominated the least wet zone. Low altitude sites showed a similar distribution of photosynthetic pathways as mid-altitude sites, but C₃ species were less abundant. Species richness was positively associated with the log of wetland size and, at the level of an individual wetland, species richness and evenness were found to be consistently greater in the intermediate and least wet zone compared with the wettest zone. The management implications of the results are discussed in the light of continuing anthropogenic loss of wetlands in the study area and global climate change.     

Endozoochorous dispersal, seedling emergence and fruiting success in disturbed and undisturbed successional stages of sheep-grazed inland sand ecosystems

This study examines the seedling emergence and fruiting success of plant species endozoochorously dispersed by sheep in a field experiment in a Koelerion glaucae area. We designed a factorial experiment (faeces/soil disturbance/successional stage/year) that mimics sheep-mediated microsites and analysed all occurring vascular plant species from September 2002–December 2004. The faeces samples were collected in the same ecosystem and additionally analysed in a common-garden experiment (seedling emergence potential). According to the latter, 28 vascular plant taxa were identified (on average 124 seedlings per 100 g air-dry sheep faeces). In the field, 15 species emerged directly out of faeces, only five of which were able to set seeds (the threatened species Medicago minima, Phleum arenarium, Silene conica, Vicia lathyroides and the non-threatened Vulpia myuros). Graminoid competitors were not able to set seedlings in the field but did so in the common-garden experiment (mainly Carex hirta). The total numbers of seedlings emerging out of faeces (5% of the potential) and reaching fruit ripeness (0.4%) were very low.In the stage of seedling emergence, ten of 42 species were significantly influenced by faeces, 11 species by disturbance and two species by an interaction of the two treatment factors (Bromus tectorum, Saxifraga tridactylites). In the fruiting stage, two species profited by faeces (Bromus tectorum, Vulpia myuros) and two species by disturbance (Erophila verna, Corynephorus canescens). Overall, the density and diversity of fruiting individuals were significantly influenced by both treatment factors. The faeces factor showed a strong dependence on year (increasing effect on both dependent variables in the first year, decreasing effect in the second year). The disturbance factor is dependent particularly on successional stage (increasing effect especially in the later successional stage).Our study revealed the paradox that threatened species with low nutrient demands are the most successful direct colonisers of sheep-faeces deposits. For them to reach the fruiting stage is rare but is of importance if new habitats are to be made accessible.     

More efficient estimation of plant biomass

Question: The optimal use of the point intercept method (PIM) for efficient estimation of plant biomass has not been addressed although PIM is a commonly used method in vegetation analysis. In this study we compare results achieved using PIM at a range of efforts, we assess a method for calculating these results that are new with PIM and we provide a formula for planning the optimal use of PIM. Location: Northern Norway. Methods: We collected intercept data at a range of efforts, i.e. from one to 100 pins per 0.25 m² plots, on three plant growth forms in a mountain meadow. After collection of intercept data we clipped and weighed the plant biomass. The relationship between intercept frequency and weighed biomass (b) was estimated using both a weighted linear regression model (WLR) and an ordinary linear regression model (OLR). The accuracy of the estimate of biomass achieved by PIM at different efforts was assessed by running computer simulations at different pin densities. Results: The relationship between intercept frequency and weighed biomass (b) was far better estimated using WLR compared to the normally used OLR. Efforts above 10 pins per 0.25 m² plot had a negligible effect on the accuracy of the estimate of biomass achieved by PIM whereas the number of plots had a strong effect. Moreover, for a given level of accuracy, the required number of plots varied depending on plant growth form. We achieved similar results to that of the computer simulations when applying our WLR based formula. Conclusion: This study shows that PIM can be applied more efficiently than was done in previous studies for the purpose of plant biomass estimation, where several plots should be analysed but at considerably less effort per plot. Moreover, WLR rather than OLR should be applied when estimating biomass from intercept frequency. The formula we have deduced is a useful tool for planning plant biomass analysis with PIM.