Decline of acid-sensitive plant species in heathland can be attributed to ammonium toxicity in combination with low pH

The effects of increasing ammonium concentrations in combination with different pH levels were studied on five heathland plant species to determine whether their occurrence and decline could be attributed to ammonium toxicity and/or pH levels. Plants were grown in growth media amended with four different ammonium concentrations (10, 100, 500 and 1000 micromol l(-1)) and two pH levels resembling acidified (pH 3.5 or 4) and weakly buffered (pH 5 or 5.5) situations. Survival of Antennaria dioica and Succisa pratensis was reduced by low pH in combination with high ammonium concentrations. Biomass decreased with increased ammonium concentrations and decreasing pH levels. Internal pH of the plants decreased with increasing ammonium concentrations. Survival of Calluna vulgaris, Deschampsia flexuosa and Gentiana pneumonanthe was not affected by ammonium. Moreover, biomass increased with increasing ammonium concentrations. Biomass production of G. pneumonanthe reduced at low pH levels. A decline of acid-sensitive species in heathlands was attributed to ammonium toxicity effects in combination with a low pH.     

Resistance of three co-occurring resprouter Erica species to highly frequent disturbance

The resistance to experimental, highly frequent disturbance has been analysed in three congeneric, strong-resprouter species (Erica australis, E. scoparia and E. arborea) that co-occur in heath-dominated communities of the northern side of the Strait of Gibraltar, southern Spain. To do so, mature individuals of the three species from a long undisturbed location were clipped at the ground level every sixth month during two years. The relationship between the resprouted biomass dry weight (as indicative of the resprouting vigour) and the upper surface area of the lignotuber along the experiment was established separately for each species at each clipping event by means of linear regressions analysis. The resprouting vigour of the three species was compared by means of independent one-way ANOVAs within each clipping event. Resprouting vigour decreased after recurrent clippings in the three species. Nevertheless, significant differences between species in this loss of resprouting vigour were detected, being E. scoparia the most resistant to the experimental, highly frequent clipping. It is concluded that experimental levels of recurrent disturbance may help to find out differences in resilience within similar (taxonomically, morfologically and/or ecologically), strong-resprouter plant species. Considering the history of forestry management in the nothern side of the Strait of Gibraltar, differences in this regard between the three Erica species may contribute to explain their somewhat segregated ecological distribution in this region.     

Vertical structure of Erica umbellata,a representative species of European Ibero-Atlantic dry heaths

The canopy structure of Erica umbellata was studied in order to (a) quantify biomass allocation among several organ types, (b) analyse the possible changes in vertical structure related to season and plant size, and (c) evaluate the effectiveness of non-destructive measures to estimate biomass for a species that is declining in some areas, due to fire and other human disturbances. The study was conducted in NW Spain, sampling E. umbellata plants belonging to three size groups. Destructive (biomass) and non-destructive (frequency, height, diameter) measures were used to characterise the vertical distribution and abundance of photosynthetic, woody, reproductive and dead organs. Allometric equations were calculated to estimate total mass using non-destructive measurements. As E. umbellata increases in size, those organs with a higher renewal rate (leaves, new stems, flowers) increase in the upper strata. Seasonal differences are recorded for the reproductive organs and new stems. A sharp decrease in the green/total phytomass ratio (from 0.51 to 0.17) is observed as plant size increases. Organ biomass can be accurately predicted from total weight and it is also possible to estimate the total weight from non-destructive measures, which provides an easier way of recording data in the field.     

The steppes of Tierra del Fuego: floristic and growthform patterns controlled by soil fertility and moisture

Northern Tierra del Fuego is the insular extreme of the Magellanic steppe, a unique example of a cold-temperate oceanic grassland in South America. We surveyed this poorly known region with a landscape-based procedure to classify plant communities and investigate environmental gradients. Sampling sites were selected with the aid of aerial photos, and floristic composition, soil, topography and landform type were recorded in each site. Classification was achieved with Two-Way Indicator Species Analysis and gradients were analyzed with Detrended Canonical Correspondence Analysis. Two broad groups of communities were separated by the presence of typical acidophilous dwarf shrubs: acidophilous/mesotrophic and neutrophilous vegetation. Along the main floristic gradient, ericoid dwarf shrubs were replaced by forbs and grasses as soil acidity decreased. The acidic habitats were on Pleistocene glacial and alluvial deposits, and had coarse texture, poorly decomposed organic matter, and low base status. The eutrophic habitats were on Tertiary rocks of medium to fine texture and had well-decomposed organic matter and high base status. A second axis of variation was associated to latitudinal position and slope aspect, revealing a precipitation/evaporation gradient. Scrubs dominated by midheight shrubs, with species from the southern forests, were at the humid extreme (higher latitudes/south aspects); and short grass steppes, with taxa commonly found further North in the continent, at the xeric end (lower latitudes/north aspects). Although fertility and moisture seem to control the overall floristic gradient, the extreme vegetation types of the fertility gradient, both having a very simple structure, could be attributed to grazing disturbance. Floristics and ecology of the Fuegian steppe resemble to a certain degree some boreal oceanic systems.     

Relating species abundance distributions to species-area curves in two Mediterranean-type shrublands

Abstract. Based on both theoretical and empirical studies there is evidence that different species abundance distributions underlie different species‐area relationships. Here I show that Australian and Californian shrubland communities (at the scale from 1 to 1000 m²) exhibit different species‐area relationships and different species abundance patterns. The species‐area relationship in Australian heathlands best fits an exponential model and species abundance (based on both density and cover) follows a narrow log normal distribution. In contrast, the species‐area relationship in Californian shrublands is best fit with the power model and, although species abundance appears to fit a log normal distribution, the distribution is much broader than in Australian heathlands. I hypothesize that the primary driver of these differences is the abundance of small‐stature annual species in California and the lack of annuals in Australian heathlands. Species‐area is best fit by an exponential model in Australian heathlands because the bulk of the species are common and thus the species‐area curves initially rise rapidly between 1 and 100 m². Annuals in Californian shrublands generate very broad species abundance distributions with many uncommon or rare species. The power function is a better model in these communities because richness increases slowly from 1 to 100 m² but more rapidly between 100 and 1000 m² due to the abundance of rare or uncommon species that are more likely to be encountered at coarser spatial scales. The implications of this study are that both the exponential and power function models are legitimate representations of species‐area relationships in different plant communities. Also, structural differences in community organization, arising from different species abundance distributions, may lead to different species‐area curves, and this may be tied to patterns of life form distribution.     

Combination of herbivore removal and nitrogen deposition increases upland carbon storage

Ecosystem carbon (C) accrual and storage can be enhanced by removing large herbivores as well as by the fertilizing effect of atmospheric nitrogen (N) deposition. These drivers are unlikely to operate independently, yet their combined effect on aboveground and belowground C storage remains largely unexplored. We sampled inside and outside 19 upland grazing exclosures, established for up to 80 years, across an N deposition gradient (5–24 kg N ha^?1 yr^?1) and found that herbivore removal increased aboveground plant C stocks, particularly in moss, shrubs and litter. Soil C storage increased with atmospheric N deposition, and this was moderated by the presence or absence of herbivores. In exclosures receiving above 11 kg N ha^?1 year^?1, herbivore removal resulted in increased soil C stocks. This effect was typically greater for exclosures dominated by dwarf shrubs (Calluna vulgaris) than by grasses (Molinia caerulea). The same pattern was observed for ecosystem C storage. We used our data to predict C storage for a scenario of removing all large herbivores from UK heathlands. Predictions were made considering herbivore removal only (ignoring N deposition) and the combined effects of herbivore removal and current N deposition rates. Predictions including N deposition resulted in a smaller increase in UK heathland C storage than predictions using herbivore removal only. This finding was driven by the fact that the majority of UK heathlands receive low N deposition rates at which herbivore removal has little effect on C storage. Our findings demonstrate the crucial link between herbivory by large mammals and atmospheric N deposition, and this interaction needs to be considered in models of biogeochemical cycling.     

Ericoid mycorrhizal association: ability to adapt to a broad range of habitats

Ericoid mycorrhizal fungi are symbiotically associated with the roots of members of the Ericaceae which include genera such as Calluna, Epacris, Erica, Rhododendron and Vaccinium. These ericoid mycorrhizal associations have adapted to a broad range of habitats, from mor humus soils of the northern hemisphere to sandy soils occurring in the southern hemisphere. They also play an important part in enabling plants like Calluna vulgaris (L.) Hull in the northern hemisphere to colonize mine spoils which are inhospitable environments of toxic waste for growth of most plants. The mechanisms of utilizing complex forms of nitrogen and phosphorus and providing protection against toxic metals are described. These mechanisms carried out by ericoid mycorrhizal associations enable host plants to establish in diverse habitats.