Impact of feral water buffalo and fire on growth and survival of mature savanna trees: An experimental field study in Kakadu National Park, northern Australia

The impact of feral Asian water buffalo (Bubalus bubalis) and season of fire on growth and survival of mature trees was monitored over 8 years in the eucalypt savannas of Kakadu National Park. Permanently marked plots were paired on either side of a 25-km-long buffalo-proof fence at three locations on an elevational gradient, from ridge-top to the edge of a floodplain; buffalo were removed from one side of the fence. All 750 trees ? 1.4 m height were permanently marked; survival and diameter of each tree was measured annually; 26 species were grouped into four eco-taxonomic groups. The buffalo experiment was maintained for 7 years; trees were monitored an additional year. Fires were excluded from all sites the first 3 years, allowed to occur opportunistically for 4 years and excluded for the final year. Fires were of two main types: low-intensity early dry season and high-intensity late dry season. Growth rates of trees were size-specific and positively related to diameters as exponential functions; trees grew slowest on the two ends of the gradient. Eucalypt mortality rates were 1.5 and 3 times lower than those of pantropics and of arborescent monocots, respectively, but the relative advantage was lost with fires or buffalo grazing. Without buffalo grazing, ground level biomass was 5-8 t ha -1 compared with 2-3 t ha -1, within 3 years. In buffalo-absent plots, trees grew significantly slower on the dry ridge and slope, and had higher mortality across the entire gradient, compared with trees in buffalo-present plots. At the floodplain margin, mortality of small palms was higher in buffalo-present sites, most likely due to associated heavy infestations of weeds. Low-intensity fires produced tree growth and mortality values similar to no-fire, in general, but, like buffalo, provided a 'fertilization' effect for Eucalyptus miniata and Eucalyptus tetrodonta, increasing growth in all size classes. High-intensity fires reduced growth and increased mortality of all functional groups, especially the smallest and largest (>35 cm d.b.h.) trees. When buffalo and fires were excluded in the final year, there were no differences in growth or mortality between paired sites across the environmental gradient. After 8 years, the total numbers of trees in buffalo-absent plots were only 80% of the number in buffalo-present plots, due to relatively greater recruitment of new trees in buffalo-present plots; fire-sensitive pantropics were particularly disadvantaged. Since the removal of buffalo is disadvantageous, at least over the first years, to savanna tree growth and survival due to a rebound effect of the ground-level vegetation and subsequent changes in fire-vegetation interactions, process-orientated management aimed at reducing fuel loads and competitive pressure may be required in order to return the system to a previous state. The 'footprint' of 30 years of heavy grazing by buffalo has implications for the interpretation of previous studies on fire-vegetation dynamics and for current research on vegetation change in these savannas.