Scale-dependent bi-trophic interactions in a semi-arid savanna: how herbivores eliminate benefits of nutrient patchiness to plants.

The scale of resource heterogeneity may influence how resources are locally partitioned between co-existing large and small organisms such as trees and grasses in savannas. Scale-related plant responses may, in turn, influence herbivore use of the vegetation. To examine these scale-dependent bi-trophic interactions, we varied fertilizer [(nitrogen (N)/phosphorus (P)/potassium (K)] applications to patches to create different scales of nutrient patchiness (patch size 2 × 2 m, 10 × 10 m, or whole-plot 50 × 50 m) in a large field experiment in intact African savanna. Within-patch fertilizer concentration and the total fertilizer load per plot were independently varied. We found that fertilization increased the leaf N and P concentrations of trees and grasses, resulting in elevated utilization by browsers and grazers. Herbivory off-take was particularly considerable at higher nutrient concentrations. Scale-dependent effects were weak. The net effect of fertilization and herbivory was that plants in fertilized areas tended to grow less and develop smaller rather than larger standing biomass compared to plants growing in areas that remained unfertilized. When all of these effects were considered together at the community (plot) level, herbivory completely eliminated the positive effects of fertilization on the plant community. While this was true for all scales of fertilization, grasses tended to profit more from coarse-grained fertilization and trees from fine-grained fertilization. We conclude that in herbivore-dominated communities, such as the African savanna, nutrient patchiness results in the herbivore community profiting rather more than the plant community, irrespective of the scale of patchiness. At the community level, the allometric scaling theory's prediction of plant-and probably also animal-production does not hold or may even be reversed as a result of complex bi-trophic interactions.

Soil nutrient status determines how elephant utilize trees and shape environments.

1. Elucidation of the mechanism determining the spatial scale of patch selection by herbivores has been complicated by the way in which resource availability at a specific scale is measured and by vigilance behaviour of the herbivores themselves. To reduce these complications, we studied patch selection by an animal with negligible predation risk, the African elephant. 2. We introduce the concept of nutrient load as the product of patch size, number of patches and local patch nutrient concentration. Nutrient load provides a novel spatially explicit expression of the total available nutrients a herbivore can select from. 3. We hypothesized that elephant would select nutrient-rich patches, based on the nutrient load per 2500 m(2) down to the individual plant scale, and that this selection will depend on the nitrogen and phosphorous contents of plants. 4. We predicted that elephant would cause more adverse impact to trees of lower value to them in order to reach plant parts with higher nutrient concentrations such as bark and root. However, elephant should maintain nutrient-rich trees by inducing coppicing of trees through re-utilization of leaves. 5. Elephant patch selection was measured in a homogenous tree species stand by manipulating the spatial distribution of soil nutrients in a large field experiment using NPK fertilizer. 6. Elephant were able to select nutrient-rich patches and utilized Colophospermum mopane trees inside these patches more than outside, at scales ranging from 2500 down to 100 m(2) . 7. Although both nitrogen and phosphorus contents of leaves from C. mopane trees were higher in fertilized and selected patches, patch choice correlated most strongly with nitrogen content. As predicted, stripping of leaves occurred more in nutrient-rich patches, while adverse impact such as uprooting of trees occurred more in nutrient-poor areas. 8. Our results emphasize the necessity of including scale-dependent selectivity in foraging studies and how elephant foraging behaviour can be used as indicators of change in the availability of nutrients.

Large herbivores may alter vegetation structure of semi-arid savannas through soil nutrient mediation.

In savannas, the tree-grass balance is governed by water, nutrients, fire and herbivory, and their interactions. We studied the hypothesis that herbivores indirectly affect vegetation structure by changing the availability of soil nutrients, which, in turn, alters the competition between trees and grasses. Nine abandoned livestock holding-pen areas (kraals), enriched by dung and urine, were contrasted with nearby control sites in a semi-arid savanna. About 40 years after abandonment, kraal sites still showed high soil concentrations of inorganic N, extractable P, K, Ca and Mg compared to controls. Kraals also had a high plant production potential and offered high quality forage. The intense grazing and high herbivore dung and urine deposition rates in kraals fit the accelerated nutrient cycling model described for fertile systems elsewhere. Data of a concurrent experiment also showed that bush-cleared patches resulted in an increase in impala dung deposition, probably because impala preferred open sites to avoid predation. Kraal sites had very low tree densities compared to control sites, thus the high impala dung deposition rates here may be in part driven by the open structure of kraal sites, which may explain the persistence of nutrients in kraals. Experiments indicated that tree seedlings were increasingly constrained when competing with grasses under fertile conditions, which might explain the low tree recruitment observed in kraals. In conclusion, large herbivores may indirectly keep existing nutrient hotspots such as abandoned kraals structurally open by maintaining a high local soil fertility, which, in turn, constrains woody recruitment in a negative feedback loop. The maintenance of nutrient hotspots such as abandoned kraals by herbivores contributes to the structural heterogeneity of nutrient-poor savanna vegetation.

Extensive browsing by a conventional grazer? Stable carbon isotope analysis reveals extraordinary dietary flexibility among Sanga cattle of North Central Namibia.

Intraspecies dietary flexibility, such as variable consumption of graze vs. browse in herbivores, has received scant attention on a spatial scale despite growing evidence of substantial variability within and among populations, especially in bovids. Here, we report on extraordinary differences in cattle diet among two communal pasture areas across seasons in northern Namibia: King Nehale (KN, open grassland) and Okongo (OK, dense woodland). Percentage C3 browse and C4 grass consumption was determined from δ(13)C values of dung samples, using a Bayesian stable-isotope mixing model (SIAR - stable isotope analysis in R). During the wet and early dry season, KN cattle consumed 11 and 19% browse, respectively, and the OK cattle consumed 84% browse. At the end of the dry season, the browse intake of KN cattle increased to 33% while that of OK cattle decreased to 55%. Vegetation structure influenced the graze/browse consumption strongly in both areas. A better understanding of this extraordinary dietary flexibility is imperative as anthropogenically driven habitat change is projected to lead to the extinction of perceived grazing specialists.

Determinants of persistence and tolerance of carnivores on Namibian ranches: implications for conservation on Southern African private lands.

Changing land use patterns in southern Africa have potential to dramatically alter the prospects for carnivore conservation. Understanding these influences is essential for conservation planning. We interviewed 250 ranchers in Namibia to assess human tolerance towards and the distribution of large carnivores. Cheetahs (Acinonyx jubatus), leopards (Panthera pardus) and brown hyaenas (Hyaena brunnea) were widely distributed on Namibian farmlands, spotted hyaenas (Crocuta crocuta) had a narrower distribution, and wild dogs (Lycaon pictus) and lions (Panthera leo) are largely limited to areas near source populations. Farmers were most tolerant of leopards and least tolerant of lions, wild dogs and spotted hyaenas. Several factors relating to land use correlated consistently with carnivore-presence and landowner tolerance. Carnivores were more commonly present and/or tolerated where; wildlife diversity and biomass were higher; income from wildlife was higher; income from livestock was lower; livestock biomass was lower; in conservancies; game fencing was absent; and financial losses from livestock depredation were lower. Efforts to create conditions whereby the costs associated with carnivores are lowest, and which confer financial value to them are likely to be the most effective means of promoting carnivore conservation. Such conditions are achieved where land owners pool land to create conservancies where livestock are replaced with wildlife (or where livestock husbandry is improved) and where wildlife generates a significant proportion of ranch income. Additional measures, such as promoting improved livestock husbandry and educational outreach efforts may also help achieve coexistence with carnivores. Our findings provide insights into conditions more conducive to the persistence of and tolerance towards large carnivores might be increased on private (and even communal) lands in Namibia, elsewhere in southern and East Africa and other parts of the world where carnivore conservation is being attempted on private lands.