Livestock exerts direct and indirect effects on plant communities, changing colonization and extinction rates of species and the surrounding environmental conditions. There is scarce knowledge on how and to what extent these effects control the floristic and functional composition of plant communities in grasslands. We performed an experiment that included several treatments simulating trampling, defoliation, faeces addition and their combinations in a Mediterranean scrub community grazing-abandoned for at least 50 years. We monitored the plots for four years, and collected data on species composition, photosynthetically active radiation (PAR) and red∶far-red ratio (R∶FR), soil moisture and compaction. We estimated community weighted means (CWM) for height, habit, life cycle, seed mass and SLA. Neither compaction nor soil moisture were modified by the treatments, while PAR and R∶FR increased in all treatments in comparison to the Control and Faeces treatments. The floristic composition of all treatments, except for Faeces, converged over time, but deviated from that of the Control. The functional traits displayed the trends expected in the presence of grazing: loss of erect species and increased cover of short species with light seeds, with rosettes and prostrate habit. However, contrary to the results in literature, SLA was lower in all the treatments than Control plots. Like the results for floristic composition, all treatments except for Faeces converged towards a similar functional composition at the end of the four year period. The results of this study show the initial evolution of a Mediterranean plant community in the presence of grazing, driven primarily by the destructive action of livestock. These actions seem to directly affect the rates of extinction/colonization, and indirectly affect the light environment but not the soil conditions. However, their effects on floristic and trait composition do not seem to differ, at least at the small spatio-temporal scale.

Delaying senescence as a response to tissue losses has been reported in some studies, but there is no information about its influence in growth compensation. We performed a first test of the relative contribution of delaying senescence after defoliation to growth compensation in Dactylis glomerata L. by means of an iterative growth analysis modified to estimate tissue losses to senescent leaves. We show that Dactylis glomerata overcompensated for relative growth rate after defoliation, mainly by slowing down senescence, and to a lesser extent by increasing the newly assimilated mass allocated to leaves.

Resource availability is an important factor affecting the capacity of compensatory growth after grazing. We performed a greenhouse experiment with Poa bulbosa, a small perennial grass of the Mediterranean and Central Asian grasslands, to test the importance of nutrient availability for compensatory growth after clipping. We also compared the results with predictions of the limited resource model (LRM). Plants were grown at low and high fertilization levels and subjected to a clipping treatment. Contrary to the LMR, we found that in Poa plants compensatory growth occurred under the high fertilization level, while it did not occur under the low level. The LMR predicts a higher tolerance for grazing in the stressful environment. Our plants showed a significant decrease in their relative growth rates (RGR) after clipping. Although the plants allocated a 32–188% greater fraction of the mass to lamina growth after clipping, this greater allocation to the leaves did not fully compensate for the initial reduction in leaf area ratio (LAR). A sensitivity analysis showed for the clipped plants under the high fertilization treatment, that changes in leaf allocation (f lam) enabled the plants to compensate for a part of the potential loss caused by defoliation. Probably, the increased biomass allocation comes largely from the bulbs. We conclude that the inconsistency of the LRM with our results originates in the lack of compensatory mechanisms in the model. To better understand how environmental conditions affect tolerance to herbivory, the effects of compensatory growth should be taken into account.

Grassland plant species have traditionally been classified into two groups, increasers and decreasers, based on changes in their relative abundance in the presence or absence of grazing. However, the ecophysiological mechanisms underlying these opposing strategies remain unclear. Seeds of ten Mediterranean species of known grazing status were germinated in different light environments to test the hypothesis that grazing selects light-dependent germination amongst grazing increasers. Two light quantity levels (Photosynthetical Active Radiation, PAR) and five light quality levels (red:far-red ratio, R:FR) were used to quantify percentages of germination of viable seeds and number of days elapsed to reach 50% of germination (T50), measured using five replicates of each light treatment and species combination. There were no differences in response to light quantity between the two grazing status groups; germination was inhibited in both groups at higher light intensities. However, increasers had lower germination percentages at low R:FR ratios, while decreasers were not affected by light quality. Germination speed (T50) was affected differently by light quality in each grazing status. There was an increase in germination speed at the lowest R:FR ratio only in increaser seeds, whereas decreaser germination speed was not affected by the treatment. PAR affected T50, regardless of grazing status; germination was faster at lower light intensities. Increaser seeds germinated faster than decreasers. These results reveal that the differential control of germination by light quality is a key factor in determining the species composition of grassland plant communities affected by herbivores in semi-arid Mediterranean grasslands.

Interacting effects of high light and drought on the performance of sun and shade phenotypes were experimentally undertaken following survival, chlorophyll fluorescence and gas exchange in 2-year-old saplings of four Mediterranean trees (Quercus ilex and Q. coccifera as water-saving species, and Pistacia lentiscus and P. terebinthus as water-spending species). Half of the saplings were grown in full sunlight and the other half in the shade (6% sunlight). Half of each combination of species-phenotype was exposed to high light during a simulated late-summer drought. Light absorptance and gas exchange were scaled up to the whole plant with the 3-D geometrical model, Y-Plant. Quercus species were more plastic and tolerated high light and water stress better than Pistacia species, surviving longer and in drier soils, and exhibiting a less pronounced photoinhibition. There was no evidence of disadvantage for shade phenotypes under high light with increasing drought. By contrast, shade phenotypes survived longer despite larger initial decreases in photochemical efficiency and higher sensitivity to drought than sun phenotypes. The enhanced control of transpiration during drought in water-saving versus water-spending species (and also in shade versus sun phenotypes in three out of the four species) allowed extended survival. Photoinhibition reduced whole crown carbon gain in high light by c. 3% and affected significantly more the shaded leaves of a given plant (reducing their carbon gain by up to 7%) than those exposed to direct sunlight. Despite this apparently minor impact, whole plant carbon gain reduction by photoinhibition negatively correlated with survival and drought tolerance. The implications for succession and forest regeneration in arid environments, particularly under a global change scenario, are discussed.

Different plant species and organs within a plant differ in their plastic response to light. These responses influence their performance and survival in relation to the light environment, which may range from full sunlight to deep shade. Plasticity, especially with regard to physiological features, is linked to a greater capacity to exploit high light and is usually low in shade-tolerant species. Among photosynthetic organs, green stems, which represent a large fraction of the total photosynthetic area of certain species, are hypothesized to be less capable of adjustment to light than leaves, because of biomechanical and hydraulic constraints. The response to light by leaves and stems of six species of leguminous, green-stemmed shrubs from dry and high-light environments was studied by growing seedlings in three light environments: deep shade, moderate shade and sun (3, 30 and 100 % of full sunlight, respectively). Survival in deep shade ranged from 2 % in Retama sphaerocarpa to 74 % in Ulex europaeus. Survival was maximal at moderate shade in all species, ranging from 80 to 98 %. The six species differed significantly in their ratio of leaf to total photosynthetic area, which influenced their light response. Survival in deep shade increased significantly with increasing ratio of leaf to total photosynthetic area, and decreased with increasing plasticity in net photosynthesis and dark respiration. Responses to light differed between stems and leaves within each species. Mean phenotypic plasticity for the variables leaf or stem specific mass, chlorophyll content, chlorophyll a/b ratio, and carotenoid to chlorophyll ratio of leaves, was inversely related to that of stems. Although mean plasticity of stems increased with the ratio of leaf to total photosynthetic area, the mean plasticity of leaves decreased. Shrubs with green stems and a low ratio of leaf to total photosynthetic area are expected to be restricted to well-lit habitats, at least during the seedling stage, owing to their inefficient light capture and the low plasticity of their stems.