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

Abstract

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.