Modeling elephant-mediated cascading effects of water point closure.

Wildlife management to reduce the impact of wildlife on their habitat can be done in several ways, among which removing animals (by either culling or translocation) is most often used. There are, however, alternative ways to control wildlife densities, such as opening or closing water points. The effects of these alternatives are poorly studied. In this paper, we focus on manipulating large herbivores through the closure of water points (WPs). Removal of artificial WPs has been suggested in order to change the distribution of African elephants, which occur in high densities in national parks in Southern Africa and are thought to have a destructive effect on the vegetation. Here, we modeled the long-term effects of different scenarios of WP closure on the spatial distribution of elephants, and consequential effects on the vegetation and other herbivores in Kruger National Park, South Africa. Using a dynamic ecosystem model, SAVANNA, scenarios were evaluated that varied in availability of artificial WPs; levels of natural water; and elephant densities. Our modeling results showed that elephants can indirectly negatively affect the distributions of meso-mixed feeders, meso-browsers, and some meso-grazers under wet conditions. The closure of artificial WPs hardly had any effect during these natural wet conditions. Under dry conditions, the spatial distribution of both elephant bulls and cows changed when the availability of artificial water was severely reduced in the model. These changes in spatial distribution triggered changes in the spatial availability of woody biomass over the simulation period of 80 years, and this led to changes in the rest of the herbivore community, resulting in increased densities of all herbivores, except for giraffe and steenbok, in areas close to rivers. The spatial distributions of elephant bulls and cows showed to be less affected by the closure of WPs than most of the other herbivore species. Our study contributes to ecologically informed decisions in wildlife management. The results from this modeling exercise imply that long-term effects of this intervention strategy should always be investigated at an ecosystem scale.

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.

Trophic rewilding revives biotic resistance to shrub invasion.

Trophic rewilding seeks to rehabilitate degraded ecosystems by repopulating them with large animals, thereby re-establishing strong top-down interactions. Yet there are very few tests of whether such initiatives can restore ecosystem structure and functions, and on what timescales. Here we show that war-induced collapse of large-mammal populations in Mozambique's Gorongosa National Park exacerbated woody encroachment by the invasive shrub Mimosa pigra-considered one of the world's 100 worst invasive species-and that one decade of concerted trophic rewilding restored this invasion to pre-war baseline levels. Mimosa occurrence increased between 1972 and 2015, a period encompassing the near extirpation of large herbivores during the Mozambican Civil War. From 2015 to 2019, mimosa abundance declined as ungulate biomass recovered. DNA metabarcoding revealed that ruminant herbivores fed heavily on mimosa, and experimental exclosures confirmed the causal role of mammalian herbivory in containing shrub encroachment. Our results provide mechanistic evidence that trophic rewilding has rapidly revived a key ecosystem function (biotic resistance to a notorious woody invader), underscoring the potential for restoring ecological health in degraded protected areas.

War-induced collapse and asymmetric recovery of large-mammal populations in Gorongosa National Park, Mozambique.

How do large-mammal communities reassemble after being pushed to the brink of extinction? Few data are available to answer this question, as it is rarely possible to document both the decline and recovery of wildlife populations. Here we present the first in-depth quantitative account of war-induced collapse and postwar recovery in a diverse assemblage of large herbivores. In Mozambique's Gorongosa National Park, we assembled data from 15 aerial wildlife counts conducted before (1968-1972) and after (1994-2018) the Mozambican Civil War (1977-1992). Pre-war total biomass density exceeded 9,000 kg km-2, but populations declined by >90% during the war. Since 1994, total biomass has substantially recovered, but species composition has shifted dramatically. Formerly dominant large herbivores-including elephant (Loxodonta africana), hippo (Hippopotamus amphibius), buffalo (Syncerus caffer), zebra (Equus quagga), and wildebeest (Connochaetes taurinus)-are now outnumbered by waterbuck (Kobus ellipsiprymnus) and other small to mid-sized antelopes. Waterbuck abundance has increased by an order of magnitude, with >55,000 individuals accounting for >74% of large-herbivore biomass in 2018. By contrast, elephant, hippo, and buffalo, which totaled 89% of pre-war biomass, now comprise just 23%. These trends mostly reflect natural population growth following the resumption of protection under the Gorongosa Restoration Project; reintroductions (465 animals of 7 species) accounted for a comparatively small fraction of the total numerical increase. Waterbuck are growing logistically, apparently as-yet unchecked by interspecific competition or predation (apex-carnivore abundance has been low throughout the post-war interval), suggesting a community still in flux. Most other herbivore populations have increased post-war, albeit at differing rates. Armed conflict remains a poorly understood driver of ecological change; our results demonstrate the potential for rapid post-war recovery of large-herbivore biomass, given sound protected-area management, but also suggest that restoration of community structure takes longer and may require active intervention.

Ecology of grazing lawns in Africa.

Grazing lawns are a distinct grassland community type, characterised by short-stature and with their persistence and spread promoted by grazing. In Africa, they reveal a long co-evolutionary history of grasses and large mammal grazers. The attractiveness to grazers of a low-biomass sward lies in the relatively high quality of forage, largely due to the low proportion of stem material in the sward; this encourages repeat grazing that concomitantly suppresses tall-grass growth forms that would otherwise outcompete lawn species for light. Regular grazing that prevents shading and maintains sward quality is thus the cornerstone of grazing lawn dynamics. The strong interplay between abiotic conditions and disturbance factors, which are central to grazing lawn existence, can also cause these systems to be highly dynamic. Here we identify differences in growth form among grazing lawn grass species, and assess how compositional differences among lawn types, as well as environmental variables, influence their maintenance requirements (i.e. grazing frequency) and vulnerability to degradation. We also make a clear distinction between the processes of lawn establishment and lawn maintenance. Rainfall, soil nutrient status, grazer community composition and fire regime have strong and interactive influences on both processes. However, factors that concentrate grazing pressure (e.g. nutrient hotspots and sodic sites) have more bearing on where lawns establish. Similarly, we discuss the relevance of enhanced rates of nitrogen cycling and of sodium levels to lawn maintenance. Grazer community composition and density has considerable significance to grazing lawn dynamics; not all grazers are adapted to foraging on short-grass swards, and differences in body size and relative mouth dimensions determine which species are able to convert tall-grass swards into grazing lawns under different conditions. Hence, we evaluate the roles of different grazers in lawn dynamics, as well as the benefits that grazer populations derive from having access to grazing lawns. The effects of grazing lawns can extend well beyond their borders, due to their influence on grazer densities, behaviour and movements as well as fire spread, intensity and frequency. Variation in the area and proportion of a landscape that is grazing lawn can thus have a profound impact on system dynamics. We provide a conceptual model that summarises grazing lawn dynamics, and identify a rainfall range where we predict grazing lawns to be most prevalent. We also examine the biodiversity associated with grazing lawn systems, and consider their functional contribution to the conservation of this biodiversity. Finally, we assess the utility of grazing lawns as a resource in a rangeland context.