Seedling responses to moderate and severe herbivory: a field-clipping experiment with a keystone Mediterranean palm.

Plant-ungulate interactions are critical in shaping the structure of Mediterranean plant communities. Nevertheless, there is a dearth of knowledge on how plant intrinsic and extrinsic factors mediate the sign and strength of plant-ungulate interactions. This is most relevant when addressing natural or assisted restoration of plant communities in human-disturbed areas. We conducted field-clipping experiments simulating how different intensities of ungulate herbivory may affect the natural regeneration and establishment of the Mediterranean dwarf palm (Chamaerops humilis), a keystone species in Mediterranean ecosystems. We quantified seedling survival and size in two human-disturbed sites (SW Spain) where wild and domestic ungulates exert high herbivory pressure on vegetation. Severe clipping and seedling aging reduced rates of seedling survival. In contrast, moderate clipping did not affect seedling survival, suggesting a certain degree of C. humilis tolerance to herbivory. Severe clipping reduced seedling height strongly but not seedling diameter, and these effects seem to have decreased seedling survival. Nurse shrubs increased seedling size, which likely improved seedling survival. We also found seedling compensatory growth which varied between study sites. Field-clipping experiments can help disentangle effects of plant extrinsic and intrinsic factors on the sign and strength of plant-ungulate interactions and their ecological consequences on the dynamics of human-disturbed ecosystems. We call attention to the importance of appropriately managing scenarios of severe herbivory and summer droughts, particularly frequent in Mediterranean ecosystems, as synergic effects of such key drivers can negatively affect the structure and dynamics of plant communities and endanger their conservation.

Nutrients cause grassland biomass to outpace herbivory.

Human activities are transforming grassland biomass via changing climate, elemental nutrients, and herbivory. Theory predicts that food-limited herbivores will consume any additional biomass stimulated by nutrient inputs ('consumer-controlled'). Alternatively, nutrient supply is predicted to increase biomass where herbivores alter community composition or are limited by factors other than food ('resource-controlled'). Using an experiment replicated in 58 grasslands spanning six continents, we show that nutrient addition and vertebrate herbivore exclusion each caused sustained increases in aboveground live biomass over a decade, but consumer control was weak. However, at sites with high vertebrate grazing intensity or domestic livestock, herbivores consumed the additional fertilization-induced biomass, supporting the consumer-controlled prediction. Herbivores most effectively reduced the additional live biomass at sites with low precipitation or high ambient soil nitrogen. Overall, these experimental results suggest that grassland biomass will outstrip wild herbivore control as human activities increase elemental nutrient supply, with widespread consequences for grazing and fire risk.

Using n-alkanes to estimate diet composition of herbivores: a novel mathematical approach.

N-alkanes are long-chain saturated hydrocarbons occurring in plant cuticles that can be used as chemical markers for estimating the diet composition of herbivores. An important constraint of using n-alkanes to estimate diet composition with currently employed mathematical procedures is that the number of markers must be equal or larger than the number of diet components. This is a considerable limitation when dealing with free-ranging herbivores feeding on complex plant communities. We present a novel approach for the estimation of diet composition using n-alkanes which applies equally to cases where the number of markers is lower, equal or greater than the number of plant species in the diet. The model uses linear programming to estimate the minimum and maximum proportions of each plant in the diet, and avoids the need for grouping species in order to reduce the number of estimated dietary components. We illustrate the model with two data sets of n-alkane content of plants and faeces obtained from a sheep grazing experiment conducted in Australia and a red deer study in Portugal. Our results are consistent with previous studies on those data sets and provide additional information on the proportions of individual species in the diet. Results show that sheep included in the diet high proportions of white clover (from 0.25 to 0.37), and relatively high proportions of grasses (e.g. brome from 0.14 to 0.26) but tended to avoid Lotus spp. (always less than 0.04 of the diet). For red deer we found high proportions of legumes (e.g. Trifolium angustifolium and Vicia sativa reaching maximum proportions of 0.42 and 0.30 of the diet, respectively) with grasses being less important and Cistus ladanifer, a browse, also having relevance (from 0.21 to 0.42 of the diet).