Positive plant interactions in the Iberian Southeast: Mechanisms, environmental gradients, and ecosystem function

Semi-arid ecosystems are privileged sites to address the effects of plant interactions on community structure and dynamics because environmental conditions are demanding and may change quickly, altering in significant ways the balance between positive and negative effects among neighboring plants. Plant interaction processes have been well documented in the semi-arid region of Southeast Spain over the last 15 years. In this article we review the growing body of research on plant–plant interactions available from this area, highlighting its importance in increasing our knowledge on this field of study. This review has been organized in five sections, i) facilitation mechanisms; ii) the nurse effect; iii) the balance of interactions and environmental, ontogenic, temporal, and spatial gradients; iv) the effects of facilitation on biodiversity; and, v) facilitation and ecosystem functioning. Mechanisms of facilitation in these systems are relatively well known, but not completely explored. In these environments competition, mainly for water, is intense between neighbors and switches in intensity from belowground to aboveground as productivity increases. By contrast, facilitation may decrease quickly with increasing productivity, although the balance between facilitation and competition is not fully understood, and is further complicated because shifts can also be driven by factors such as life history or physiology of interacting species. Positive interactions are critical for maintaining biodiversity in some ecosystems in SE Spain, but their role as a driver of ecosystem functioning is less clear. Research on plant–plant interactions in this region has been highly influential and has contributed to our overall understanding of plant community dynamics. Despite the important progress achieved during the last 15 years, there is still substantial scope for exploring the effects of plant interactions at the ecosystem level, and their role as modulators of disturbances such as the current global environmental change.     

Stress resistance strategy in an arid land shrub: interactions between developmental instability and fractal dimension

This paper investigates allocation of energy to mechanisms that generate and preserve architectural forms (i.e. developmental stability, complexity of branching patterns) and productivity (growth and reproduction) in response to environmental disturbances (i.e. grazing and resource availability). The statistical error in translational symmetry was used to detect random intra-individual variability during development. This can be thought of as a measure of developmental instability caused by stress. Additionally, we use changes in fractal complexity and shoot distribution of branch structures as an alternate indicator of stress. These methods were applied to Anthyllis cytisoides L., a semi-arid environment shrub, to ascertain the effect of grazing and slope exposure on developmental traits in a 2×2 factorial design. The results show that A. cytisoides maintains developmental stability at the expense of productivity. Anthyllis cytisoides was developmentally more stable when grazed and when on south-facing, as opposed to north-facing slopes. On the contrary, shoot length, leaf area, fractal dimension and reproductive-to-vegetative allocation ratio were larger in north- than in south-facing slopes. As a consequence, under extreme xeric conditions, shrub mortality increased in north-facing slopes, especially when not grazed. The removal of transpiring area and the reduction of plant competition favoured developmental stability and survival in grazed plants. Differences between grazed and ungrazed plants were most evident in more mesic (north-facing) areas.     

Climatic change and rainfall patterns: Effects on semi-arid plant communities of the Iberian Southeast

The structure and functioning of semi-arid ecosystems are strongly influenced by precipitation patterns. Water availability in such environments is highly pulsed, and discrete rainfall events interspersed with drought periods are important components of the annual water supply. Plant communities do not only respond to rainfall quantity, but also to variations in time, so that relatively small changes in rainfall frequency (i.e., pulsed inputs) may have strong effects on communities. Within the Mediterranean basin, climate change models forecast a decrease in mean annual precipitation and more extreme events (i.e., less rainy days and longer drought periods between events), along with seasonal changes. However, little is known on the consequences of these future precipitation changes on plant communities, especially in semi-arid environments. Here, we summarize the few experiments that have manipulated rainfall patterns in arid and semi-arid areas worldwide, and introduce the first results of a pioneer, long-term rainfall exclusion in the semi-arid southeast region of the Iberian Peninsula. The experiment is not only manipulating the amount of rainfall, but also its frequency and seasonal distribution in a grassland-shrubland in the Tabernas desert (Almería, Spain). This work monitored the effect of precipitation changes on different ecosystem processes for five years, at the species and community level, concluding that this plant community (as other communities studied in the same area) exhibited great resilience to changes in rainfall availability, likely caused by plant adaptation to large intra- and inter-annual precipitation variability.     

Abiotic conditions, neighbour interactions, and the distribution of Stipa tenacissima in a semiarid mountain range

Abiotic conditions and interactions among plants are important factors determining plant geographical distributions. With the aim of analyzing the distribution of a perennial grass species in a semiarid mountain range, we experimentally tested above- and belowground effects of shrubs on the grass along a gradient of aridity. We measured growth, physiology and reproductive output of Stipa tenacissima tussocks surrounded by intact or manipulated neighbour shrubs. Performance of Stipa improved under mesic conditions, but the net effect of shrubs on Stipa did not change along the gradient. Belowground, we did not find significant effects, suggesting belowground resource partitioning between tussocks and shrubs. Aboveground net effects of shrubs on Stipa were generally positive and independent from abiotic conditions. The improved performance of Stipa at the mesic end of gradient suggests that its absolute physiological optimum lies out of its realized range. Overall, the arid end of the geographical distribution of Stipa in this mountain range may be determined by environmental severity. The coincidence of the upper limit of Stipa with the lower tree limit suggests that the more mesic end of Stipa distribution may depend on other factors; most likely competition with trees, which, unlike shrubs, could be excluding Stipa tenacissima from forests.     

Colonization processes in semi-arid Mediterranean old-fields

Semi-arid Mediterranean ecosystems undergo a slow process of secondary succession after disturbance which is dependent on rainfall. We tested the hypothesis that colonization of abandoned lands in semi-arid SE Spain was slowed down by seed dispersal and species-specific responses to climatic variability. In field and glasshouse experiments we analysed germination of the two main colonizer species, their appearance in the field as seedlings, their relationships with climate and the resulting population structure. Seed germination of the two dominant legume species, Anthyllis cytisoides and Retama sphaerocarpa, was enhanced by scarification, but triggered in the field by temperature and rainfall, respectively. There was no evidence of allelopathic effects affecting germination in the field. Population structure reflected high inter-annual variability in shrub establishment, which was related to rainfall in spite of self-thinning and other processes that modeled current shrub numbers. Colonization of abandoned lands in semi-arid, Mediterranean environments in SE Spain seems to be primarily limited by seed dispersal, but also affected by climatic variability, which produced temporally separated recruitment events.