The coppicing of a savanna tree species (Terminalia sericea) in relation to resource manipulation and disturbance
The growth and recovery of trees from disturbances such as fire and browsing is driven by the intensity of the disturbances and the availability of resources. In savannas, resprouting has become recognized as a key functional trait in plant ecology over the past decade. Although this may indeed be the case, there is still limited information about the physiology and growth strategies of resprouting trees. Available information about the influence of disturbance comes from ecosystems that are in many ways different from tropical and sub-tropical savannas. Therefore it is important to know and understand post-disturbance tree responses and limitations so as to establish sustainable use and management practices. This thesis reports the findings of a study, conducted in the Mpumalanga province of South Africa, aimed at achieving a better understanding of the influences of resource availability (water and nutrients) and disturbances (herbivory and repeated cutting) on the coppicing of a widely distributed savanna tree species that is both ecologically and economically important; Terminalia sericea. To investigate the effects of resource availability, cut trees were exposed to different levels of water and nutrient (nitrogen and phosphorus) supplementation over a period of two years in a factorial experimental design. A number of coppice regrowth variables (e.g. shoot production, resprout shoot diameter and shoot length) were measured monthly, while the phenological responses (e.g. timing of leaf discolouration and fruit presence) were monitored every two weeks. The effects of disturbances were investigated in two separate experiments, in which cut trees were exposed to a five month browsing period and different cutting frequencies, respectively. Coppice regrowth variables were monitored for 12 months in trees exposed to browsing and for five months in trees exposed to different cutting frequencies. The effect of multiple cutting cycles on total non-structural carbohydrates and leaf chemistry (carbon, nitrogen and phosphorus) was measured. There was evidence of self-thinning of coppice shoots within cut trees in all experiments in the second year of growth, with supplemented trees also recording lower shoot numbers. With shoot production higher in trees that received no resource addition, water and nutrients interacted synergistically, doubling shoot diameter and shoot length for supplemented trees after 12-months of addition. The majority of the phenophases monitored peaked in the wet growing season. Supplemented trees changed colour from the typical green to senescent yellow later in the growing season than unsupplemented trees. Fruit presence occurred in the second year after cutting for supplemented trees. Shoot length and shoot diameter for unbrowsed trees were twice those for browsed trees, with leaf nitrogen and phosphorus content significantly higher for browsed trees compared to unbrowsed trees. Trees subjected to multiple cutting cycles recorded half the TNC levels, and half the resprout shoot diameter and shoot length of trees cut only once. Results from this study demonstrate that self-thinning (i.e., negative change in shoot number) is not primarily under resource control. By contrast, the findings suggest that shoot growth characteristics, the timing and duration of phenological stages in coppicing trees are resource-limited in savannas. Browsing induces an initial compensatory response through higher shoot production in browsed trees and should be kept minimal because, in the long run, browsed cut trees would take longer to recover lost biomass compared to unbrowsed cut trees. Repeated cutting significantly depleted non-structural carbohydrate reserves in stems, implying that repeatedly cut trees rely heavily on non-structural carbohydrate reserves for regrowth. The significantly lower shoot diameter and shoot length in repeatedly cut trees imply that the coppicing ability of a tree reduces as the cutting frequencies increase. Such information can be vital in establishing the competitive growth ability of T. sericea in a multiple-species ecosystem affected by changes in resource availability as well as natural disturbances. With reference to multiple cut trees, a form of cutting or harvesting strategy should be in place that allows for sustainable regeneration of the study species. A threshold in terms of number of cutting cycles a tree can tolerate based on either maximum or minimum levels of reserve carbohydrates should also be established, as this has a direct effect on the coppice growth and survival.