Elephants limit aboveground carbon gains in African savannas

Understanding the drivers of vegetation carbon dynamics is essential for climate change mitigation and effective policy formulation. However, most efforts focus on abiotic drivers of plant biomass change, with little consideration for functional roles performed by animals, particularly at landscape scales. We combined repeat airborne Light Detection and Ranging with measurements of elephant densities, abiotic factors, and exclusion experiments to determine the relative importance of drivers of change in aboveground woody vegetation carbon stocks in Kruger National Park, South Africa. Despite a growing elephant population, aboveground carbon density (ACD) increased across most of the landscape over the 6‐year study period, but at fine scales, bull elephant density was the most important factor determining carbon stock change, with ACD losses recorded only where bull densities exceeded 0.5 bulls/km². Effects of bull elephants were, however, spatially restricted and landscape dependent, being especially pronounced along rivers, at mid‐elevations, and on steeper slopes. In contrast, elephant herds and abiotic drivers had a comparatively small influence on the direction or magnitude of carbon stock change. Our findings demonstrate that animals can have a substantive influence on regional‐scale carbon dynamics and warrant consideration in carbon cycling models and policy formulation aimed at carbon management and climate change mitigation.     

Increased tree densities in South African savannas: >50 years of data suggests CO2 as a driver

For the past century, woody plants have increased in grasslands and savannas worldwide. Woody encroachment may significantly alter ecosystem functioning including fire regimes, herbivore carrying capacity, biodiversity and carbon storage capacity. Traditionally, increases in woody cover and density have been ascribed to changes in the disturbance regime (fire and herbivores) or rainfall. Increased atmospheric CO₂ concentrations may also contribute, by increasing growth rates of trees relative to grasses. This hypothesis is still heavily debated because usually potential CO₂ effects are confounded by changes in land use (disturbance regime). Here we analyse changes in woody density in fire experiments at three sites in South African savannas where the disturbance regime (fire and herbivores) was kept constant for 30 and 50 years. If global drivers had significant effects on woody plants, we would expect significant increases in tree densities and biomass over time under the constant disturbance regime. Woody density remained constant in a semiarid savanna but tripled in a mesic savanna between the 1970s and 1990s. At the third site, a semiarid savanna near the southern limits of the biome, tree density doubled from the mid 1990s to 2010. Interpretation of the causes is confounded by population recovery after clearing, but aerial photograph analysis on adjacent non‐cleared areas showed an accompanying 48% increase in woody cover. Increased CO₂ concentrations are consistent with increased woody density while other global drivers (rainfall) remained constant over the duration of the experiments. The absence of a response in one semiarid savanna could be explained by a smaller carbon sink capacity of the dominant species, which would therefore benefit less from increased CO₂. Understanding how savannas and grasslands respond to increased CO₂ and identifying the causes of woody encroachment are essential for the successful management of these systems.     

Relationship between group dynamics and spatial distribution of African elephants in a semi‐arid environment

Group dynamics related to distribution of African elephants in the semi‐arid environment of Tsavo East National Park, Kenya was studied between 2007 and 2010. I examined the seasonal distribution of lone bulls, bull groups, family units and mixed groups and group size. Lone bulls were widely distributed in the dry season and localized in the wet season, whereas bull groups were localized in both seasons. On average bull group size did not differ with seasons or areas, but larger groups were observed in preferred areas in the dry season. Family units were widely distributed in the dry season and localized in the wet season, whereas mixed groups were localized in both seasons. Although family units and mixed groups tended to be large in all areas in the wet season, large groups were also observed in the dry season. Large family units were widely distributed, whereas large mixed groups were localized. The relationship between group dynamics and distribution was attributed to the combination of social, ecological and poaching factors. This study provided insight into distribution patterns that can be applied to the security of the Tsavo elephant population.     

No effects of fire,large herbivores and their interaction on regrowth of harvested trees in two West African savannahs

Theory and empirical evidence for the impacts of fire and herbivory in savannahs is well established – they are top‐down disturbances that maintain savannahs in disequilibrium states away from potential tree cover. In African savannahs, the demand for fuelwood is extremely high, so tree harvest likely also has an impact, both directly and indirectly, on tree cover, density and biomass. Many savannah trees resprout vigorously from the base after harvest. However, harvested trees regenerate as saplings susceptible to fire and browsing, so harvest may have important demographic consequences. Here, we report the effects of tree harvest, and its interaction with fire and herbivory, on savannah dynamics by analysing woody regrowth following a harvest in arid Sahelian and mesic Guinean savannahs in Mali, West Africa. Tree harvest resulted in an overall reduction in wood production per tree compared to growth in nonharvested trees. Regrowth, either biomass or height, did not differ among fire and herbivory treatments. Our results suggest that the resprouting abilities that savannah trees have evolved to cope with frequent fire are essential for surviving tree harvest and subsequent disturbance. In these savannahs, regrowth is rapid enough in the first growing season to escape the impact of dry season fires.     

Responses of woody vegetation to exclusion of large herbivores in semi‐arid savannas

The Nkuhlu large‐scale long‐term exclusion experiment in Kruger National Park was designed to study the long‐term effects of large herbivores on vegetation. One treatment excludes elephants, another excludes all herbivores larger than hares and another one comprises an open, control area. Vegetation monitoring was implemented in 2002 when a baseline survey was conducted prior to exclusion. Monitoring was repeated 5 years after exclusion. Data from the surveys were analysed to establish how structure and composition of woody vegetation had changed 5 years after herbivore exclusion. The analysis showed that neither plant assemblage nor mean vegetation height had changed significantly since exclusion. However, both species richness and density of woody plants increased 5 years after exclusion of all large herbivores, but not after the exclusion of elephants alone. One already common species, Dichrostachys cinerea, became more common after excluding all large herbivores compared with either no exclusion or elephant exclusion, possibly leading to competitive suppression of other species. Species other than D. cinerea tended to either increase or decrease in density, but the changes were insufficient to induce significant shifts in the overall assemblage of woody plants. The results indicate that after 5 years of exclusion, the combined assemblage of large herbivores, and not elephants alone, could induce changes in species richness and abundances of woody plants, but the effect was so far insufficient to induce measureable shifts in the assemblages of woody plants. It is possible that assemblages will change with time and increasing elephant numbers may amplify future changes.     

Flood‐deposited wood creates regeneration niches for riparian vegetation on a semi‐arid South African river

Question : The formation of large woody debris (LWD) piles during floods has significant impacts on riparian succession through pioneering plants often establishing in association with wood. We assess the importance of LWD for seed regeneration of riparian plants after a century‐scale flood disturbance in a semi‐arid environment. Location : The Sabie River within Kruger National Park in the semi‐arid northeast of South Africa. Methods : Our approach was to quantify the riparian soil seed bank, to record the frequency of establishment of riparian plants in woody debris piles, and to conduct experimental out‐plantings of common riparian trees in plots with and without LWD. Results : We found the abundance and diversity of seedlings were higher in soils taken from wood piles than from open reference areas, and most seedlings were herbaceous species. Surveys indicated that numbers of seedlings recorded within woody debris were significantly greater than in open reference areas or within established vegetation. Seedling establishment in various cover‐types also varied for different riparian tree species. Experimental out‐planting of seedlings of two riparian tree species (Philenoptera violacea and Combretum erythrophyllum) revealed that, after 433 days, planted seedlings survived only in woody debris piles. Conclusion : LWD formed after a large flood creates heterogeneous patches that may influence post‐disturbance regeneration of riparian vegetation by providing a variety of environmental niches for seedlings establishment. We suspect that higher seedling survival in LWD is due to increased moisture (particularly in the dry season) and nutrients, and protection from seasonal flooding and herbivory.     

Fire season affects size and architecture of Colophospermum mopane in southern African savannas

Wildfires may be started naturally by lightning or artificially by humans. In the savanna regions of southern Africa, lightning fires tend to occur at the start of the wet season, during October and November, while anthropogenic fires are usually started during the dry season, between July and August. A long-term field manipulation experiment initiated in the Kruger National Park in 1952 was used to explore whether this seasonal divergence affects tree abundance, spatial pattern, size and architecture. After 44 years of prescribed burning treatments that simulated the seasonal incidence of lightning and anthropogenic fires, mean densities of the locally-dominant shrub, Colophospermum mopane, were 638 and 500 trees ha^–1 respectively. Trees in burnt plots had aggregated distributions while trees in unburnt plots had random distributions. Significant differences (p < 0.001) were recorded in a range of morphological parameters including tree height, canopy diameter, mean stem circumference and number of stems. The incidence of resprouting also differed significantly between treatments, with burnt trees containing a high proportion of coppiced stems. The differences in tree size and architecture between the mid-dry season and early-wet season burning plots suggest that anthropogenic fires applied during July and August cannot substitute for a natural lightning fire regime. Anthropogenic fire yields a landscape that is shorter, more scrubby and populated by numerous coppiced shrubs than the landscape generated by natural lightning fire conditions.     

Calibrating a camera trap–based biased mark–recapture sampling design to survey the leopard population in the N'wanetsi concession,Kruger National Park,South Africa

Estimating large carnivore abundance can be challenging. A biased leopard (Panthera pardus) population survey was conducted in the N'wanetsi concession in the Kruger National Park (KNP), South Africa, using motion‐sensitive camera traps from April to August 2008. Survey effort included 88 trapping occasions and 586 trap days. The survey yielded 24 leopard photographs, comprising fourteen adults of eleven males and three females. The capture rate was determined to be 24.4 trap days per leopard. Estimates of population abundance stabilized at approximately 500 trap days. Precision of population estimates began to stabilize after 378 trap days. We estimated that there were nineteen leopards in an area of 150 km². Leopard density was estimated at 12.7 leopards per 100 km². We explore the possibility of employing the methods used in this study to survey the leopard population in the KNP and surrounding areas.     

Indirect effects of domestic and wild herbivores on butterflies in an African savanna

Indirect interactions driven by livestock and wild herbivores are increasingly recognized as important aspects of community dynamics in savannas and rangelands. Large ungulate herbivores can both directly and indirectly impact the reproductive structures of plants, which in turn can affect the pollinators of those plants. We examined how wild herbivores and cattle each indirectly affect the abundance of a common pollinator butterfly taxon, Colotis spp., at a set of long‐term, large herbivore exclosure plots in a semiarid savanna in central Kenya. We also examined effects of herbivore exclusion on the main food plant of Colotis spp., which was also the most common flowering species in our plots: the shrub Cadaba farinosa. The study was conducted in four types of experimental plots: cattle‐only, wildlife‐only, cattle and wildlife (all large herbivores), and no large herbivores. Across all plots, Colotis spp. abundances were positively correlated with both Cadaba flower numbers (adult food resources) and total Cadaba canopy area (larval food resources). Structural equation modeling (SEM) revealed that floral resources drove the abundance of Colotis butterflies. Excluding browsing wildlife increased the abundances of both Cadaba flowers and Colotis butterflies. However, flower numbers and Colotis spp. abundances were greater in plots with cattle herbivory than in plots that excluded all large herbivores. Our results suggest that wild browsing herbivores can suppress pollinator species whereas well‐managed cattle use may benefit important pollinators and the plants that depend on them. This study documents a novel set of ecological interactions that demonstrate how both conservation and livelihood goals can be met in a working landscape with abundant wildlife and livestock.     

Landscape‐scale effects of fire on bird assemblages: does pyrodiversity beget biodiversity?

Aim A common strategy for conserving biodiversity in fire‐prone environments is to maintain a diversity of post‐fire age classes at the landscape scale, under the assumption that ‘pyrodiversity begets biodiversity’. Another strategy is to maintain extensive areas of a particular seral state regarded as vital for the persistence of threatened species, under the assumption that this will also cater for the habitat needs of other species. We investigated the likely effects of these strategies on bird assemblages in tree mallee vegetation, characterized by multi‐stemmed Eucalyptus species, where both strategies are currently employed. Location The semi‐arid Murray Mallee region of south‐eastern Australia. Methods We systematically surveyed birds in 26 landscapes (each 4‐km diameter), selected to represent gradients in the diversity of fire age classes and the proportion of older vegetation (> 35 years since fire). Additional variables were measured to represent underlying vegetation‐ or fire‐mediated properties of the landscape, as well as its biogeographic context. We used an information‐theoretic approach to investigate the relationships between these predictor variables and the species richness of birds (total species, threatened species and rare species). Results Species richness of birds was not strongly associated with fire‐mediated heterogeneity. Species richness was associated with increasing amounts of older vegetation in landscapes, but not with the proportion of recently burned vegetation in landscapes. Main conclusions The preference of many mallee birds for older vegetation highlights the risk of a blanket application of the ‘pyrodiversity begets biodiversity’ paradigm. If application of this paradigm involved converting large areas from long unburned to recently burned vegetation to increase fire‐mediated heterogeneity in tree mallee landscapes, our findings suggest that this could threaten birds. This research highlights the value of adopting a landscape‐scale perspective when evaluating the utility of fire‐management strategies intended to benefit biodiversity.     

Is there a temporal niche separation in the leaf phenology of savanna trees and grasses?

Aim It has been proposed that, in tropical savannas, trees deploy their leaves earlier in the growing season and grasses deploy their leaves later. This hypothesis implies a mechanism that facilitates the coexistence of trees and grasses in savannas. If true, this hypothesis would also allow algorithms to use differences in the phenological timing of grass and tree leaves to partition the relative contribution of grasses and trees to net primary production. In this study we examine whether a temporal niche separation between grasses and trees exists in savanna. Location A semi‐arid, subtropical savanna, Kruger National Park, South Africa. Methods We use a multi‐spectral camera to track through an entire growing season the normalized difference vegetation index (NDVI) of individual canopies of grasses and trees at eight sites arranged along a precipitation and temperature gradient. Results Among trees, we identified two distinct phenological syndromes: an early flushing syndrome and a late‐flushing syndrome. Leaf flush in the tree strategies appears to pre‐empt rainfall, whereas grass leaf flush follows the rain. The growing season of trees is 20 (late‐flushing trees) to 27 (early flushing trees) days longer than that of the grasses. Main conclusions We show that grasses and trees have different leaf deployment strategies. Trees deployed leaves at lower temperatures than grasses and retained them for longer at the end of the growing season. The timing of the increase in NDVI is, however, similar between grasses and late‐flushing trees and this complicates the separation of grass and tree signals from multi‐spectral satellite imagery.     

Collective decision‐making promotes fitness loss in a fusion‐fission society

While collective decision‐making is recognised as a significant contributor to fitness in social species, the opposite outcome is also logically possible. We show that collective movement decisions guided by individual bison sharing faulty information about habitat quality promoted the use of ecological traps. The frequent, but short‐lived, associations of bison with different spatial knowledge led to a population‐wide shift from avoidance to selection of agricultural patches over 9 years in and around Prince Albert National Park, Canada. Bison were more likely to travel to an agricultural patch for the first time by following conspecifics already familiar with agricultural patches. Annual adult mortality increased by 12% due to hunting of bison on agricultural lands. Maladaptive social behaviour accordingly was a major force that contributed to a ~50% population decline in less than a decade. In human‐altered landscapes, social learning by group‐living species can lead to fitness losses, particularly in fusion‐fission societies.     

The effects of water‐level manipulation on the benthic invertebrates of a managed reservoir

1. Reservoir creation and management can enhance many ecological services provided by freshwater ecosystems, but may alter the natural conditions to which aquatic biota have adapted. Benthic macroinvertebrates often reflect environmental conditions, and this community may be particularly susceptible to water‐level changes that alter sediment exposure, temperature regime, wave‐induced sediment redistribution and basal productivity. 2. Using a before–after control–impact experimental design, we assessed changes in macroinvertebrate community structure corresponding with changes in water‐level management in two lentic systems in the Voyageurs National Park, Minnesota, U.S.A. Littoral zone (depths 1–5 m) benthic macroinvertebrate assemblages were sampled in Rainy Lake (control system) and Namakan Reservoir (impact system) in 1984–85, and again in 2004–05 following a change in water‐level management that began in January 2000. The new regime reduced the magnitude of winter drawdown in Namakan Reservoir from 2.5 to 1.5 m, and allowed the reservoir to fill to capacity in late May, a month earlier than under the prior regime. Rainy Lake water levels were not altered substantially. 3. We found changes in macroinvertebrate community structure in Namakan Reservoir relative to Rainy Lake at 1–2 m depths but not at 3–5 m depths. These shallower depths would have been most directly affected by changes in sediment exposure and ice formation. 4. In 2004–05, Namakan Reservoir benthos showed lower overall abundance, more large‐bodied taxa and an increase in non‐insect invertebrates relative to 1984–85, without corresponding changes in Rainy Lake. 5. Changes in the benthic community in Namakan may reflect cooler water in spring and early summer as well as lower resource availability (both autochthonous production and allochthonous inputs) under the new regime.     

Effect of Tree‐Fall Gaps on Fruit‐Feeding Nymphalid Butterfly Assemblages in a Peruvian Rain Forest

One of the main natural disturbances that affects the structure of rain forests is treefalls, frequently resulting in gaps. Tree‐fall gaps can bring drastic changes in environmental conditions compared with the undisturbed understory. We investigated the effect of tree‐fall gaps on fruit‐feeding butterfly (Nymphalidae) species assemblages in an undisturbed lowland rain forest in southeastern Peru. We used fruit‐baited traps suspended 2 m above ground in 15 tree‐fall gaps ranging in area from 100 to 1000 m² and in adjacent undisturbed understory. Our data support the hypothesis that tree‐fall gap and understory habitats are utilized by different butterfly species assemblages. There were morphological differences between gap and understory species, where the understory species had a larger wing area to thoracic volume. Vegetation structure and composition were important factors affecting the butterfly assemblages. Most of the butterfly species showed an avoidance of vines and a strong association with the presence of trees and shrubs in gaps. There were also differences among gap assemblages that increased with gap size. Some of the species that were associated with gaps have been considered as canopy species. Other gap species in the present study, however, are known to feed on fruits and/or use host plants mainly, or only, occurring in gaps, implicating that the gap assemblage is a mix of canopy species and those unique to gaps. This indicates that, in an undisturbed Amazon forest, tree‐fall gaps may contribute to maintain species diversity by creating a mosaic of specific habitats and resources that favors different butterfly assemblages.