Vegetative and chemical responses to simulated leaf browsing during the growth season, and their subsequent effect on herbivory, were studied on Combretum apiculatum Sonder (Combretaceae) in Botswana. Treatments (50% and 100% leaf and shoot apex removal) were performed just before the shoot growth curve levelled out, and responses recorded 3 months later, just before leaf fall. Compared to controls, defoliation treatments, removing apical dominance, reduced growth in tree height and increased shoot mortality, although the production of lateral shoots increased. At the end of the trial, there was no difference in total length of annual shoots between treatment groups. Significant refoliation occurred only after 100% defoliation. Refoliated leaves were smaller and the 100% defoliated trees had a lower final leaf biomass. Total leaf biomass production was, however, equal for all treatment groups. Refoliated leaves contained higher levels of N, lower levels of aciddetergent fibre (ADF) and total phenolics, and showed a trend towards lower levels of condensed tannins, compared to leaves on control trees. Such chemical changes may be due to either carbon stress or to younger physiological age of new leaves. In spite of the observed potential increase in food quality, we found no evidence of increased levels of insect or ungulate herbivory on refoliated leaves, which, at least for insect herbivory, may be explained by the reduction in temporal availability of leaves. We conclude that the single severe defoliation was not detrimental to C. apiculatum in the short-term, although the resource loss and induced compensatory growth may produce negative effects during subsequent growth seasons.

Increased shoot growth has recurrently been recorded following real or simulated shoot browsing, although responses may vary between different woody species. We investigated morphological responses to simulated dry-season browsing on Colophospermum mopane, Acacia tortilis, Grewia flava, Peltophorum africanum and Ziziphus mucronata, which differ in growth rate, occurrence of spines and use by mammalian herbivores. Treatment groups included a control, four clipping intensities and cutting at ground level. Responses were monitored during the following growth season on three annual shoots, located in the bottom, middle and top height interval, on each sapling. All five species showed compensatory growth responses, although no obvious relation was found between the strength of responses and growth rate or natural browsing pressure. The general pattern was an increased shoot length, shoot diameter, number of lateral shoots and shoot biomass with increased clipping intensity. These responses are probably an effect of the reduction of competing growing points and disrupted apical dominance. Growth responses were stronger in shoots with higher location on the sapling, which may reflect the pressure on saplings to attain a certain height to avoid browsing, reproduce and increase resistance to fire. We found no evidence of the number of spines being an induced defence in either of the two spinescent species studied.

No abstract

J. 2004. Defensive traits of savanna trees Á/ the role of shoot exposure to browsers. Á/ Oikos 107: 161 Á/171.To investigate patterns of plant defence, a study was conducted on 13 tree species in Botswana. We tested the hypotheses that (1) shoots on small, young trees are more defended than shoots at the same height on tall, mature individuals and (2) shoots within browsing height are more defended than shoots above the reach of browsers, on mature trees. Plant traits studied were total phenolics, tannin activity, nitrogen (N), acid-detergent fibre (ADF), and degree of spinescence (in three species). We found some support for the first hypothesis. Overall young trees had a significantly higher concentration of total phenolics than mature trees and on species level these differences were significant for three of the 13 species. However, we found no differences in tannin activity, spinescence or ADF concentration between young and mature trees, and four species had higher levels of N in young trees, suggesting higher overall palatability. Contrary to our second hypothesis, shoots within browsing height were less chemically defended (total phenolics and tannin activity) than shoots above browsing height in six of 13 species. Two species had less ADF in lower shoots, and no significant differences were found in N or spinescence for any species. This study included a broad range of tree species at two sites and the results indicate that general theories of plant defence, originating from boreal and northern temperate forests, cannot be simply applied to savanna ecosystems.

Studies on the role of mammalian herbivore saliva in plant^animal interactions have mostly focused on graminoid species and bovine saliva. A trial was performed in Botswana with clipping treatments to simulate browsing of shoots and the application of goat saliva on the woody species Combretum apiculatum Sonder (Combretaceae). Treatments were performed during early growing season while shoot growth was rapid, and responses of trees were recorded later in the same season. Clipped shoots with saliva had signi¢cantly enhanced shoot growth (tripled in length) and leaf production (2.7 times more leaves) compared to clipped shoots without saliva. However, unclipped shoots still grew more than clipped shoots, with or without saliva treatment.Re¨sumeL es e¨tudes du roª le de la salive des mammife© res herbivores dans les interactions plantes-animal se sont principalement axe¨es sur les gramine¨es et sur la salive des espe© ces bovines. On a fait un essai au Botswana, qui consistait a© couper de jeunes tiges pour en stimuler la repousse et a© appliquer de la salive de che© vre sur une espe© ce ligneuse, Combretum apiculatum Sonder (Combretaceae). Ces traitements ont e¨te¨applique¨s au de¨but de la pe¨riode de repousse, quand la croissance est rapide, et ona enregistrel a re¨ponse des arbres plus tard dans la meª me saison. Les tiges coupe¨es enduites de salive connaissaient une croissance signi¢cativement plus rapide (une longueur triple) et une augmentation de la production de feuilles (2.7 fois plus de feuilles), compare¨aux tiges coupe¨es sans salive. Ceci dit, les tiges non coupe¨es poussaient encore plus vite que les tiges coupe¨es, avec ou sans salive.