Initial success of native grasses is contingent on multiple interactions among exotic grass competition, temporal priority, rainfall and site effects.

Ecological communities are increasingly being recognized as the products of contemporary drivers and historical legacies that are both biotic and abiotic. In an attempt to unravel multiple layers of ecological contingency, we manipulated (i) competition with exotic annual grasses, (ii) the timing of this competition (temporal priority in arrival/seeding times) and (iii) watering (simulated rainfall) in a restoration-style planting of native perennial grasses. In addition, we replicated this experiment simultaneously at three sites in north-central California. Native perennial grasses had 73-99 % less cover when planted with exotic annuals than when planted alone, but this reduction was greatly ameliorated by planting the natives 2 weeks prior to the exotics. In a drought year, irrigation significantly reduced benefits of early planting so that these benefits resembled those observed in a non-drought year. There were significant differences across the three sites (site effects and interactions) in (i) overall native cover, (ii) the response of natives to competition, (iii) the strength of the temporal priority effect and (iv) the degree to which supplemental watering reduced priority effects. These results reveal the strong multi-layered contingency that underlies even relatively simple communities

Aboveground Net Primary Productivity in Grazed and Ungrazed pastures: Grazing Optimisation Hypothesis or Local Extinction of Vegetation Species

The controversy that has surrounded herbivory studies in the last few decades prompted our investigation to establish the extent to which herbivore optimisation hypothesis or compensatory growth evidence is real. We used the traditional movable cage method to collect primary productivity data on herbage, functional groups and key individual grass species in various controlled large herbivore treatments in an east African savanna. The herbivore treatments in triplicate blocks included cattle, wild herbivores with and without mega herbivores and combinations of cattle and wild herbivores also with and without mega herbivores. The findings revealed that at herbage level, most grazed treatments (four out of five) had higher productivity than the ungrazed control and three showed grazing optimisation curve at sixth polynomial degree between monthly productivity and grazing intensity (1-g/ng). At functional group level forbs productivity was higher in the ungrazed control than in any of the grazed treatments while at individual grass species level _Themeda triandra_ productivity was higher in all grazed treatments than in ungrazed control. We conclude against presence of herbivore optimisation hypothesis at herbage, functional group and species level because of lack of attributable grazing effect in grazed treatments that matches complex ecological effects in the ungrazed treatment

Herbivory and Drought Generate Short‐Term Stochasticity and Long‐Term Stability in A Savanna Understory Community

Rainfall and herbivory are fundamental drivers of grassland plant dynamics, yet few studies have examined long‐term interactions between these factors in an experimental setting. Understanding such interactions is important, as rainfall is becoming increasingly erratic and native wild herbivores are being replaced by livestock. Livestock grazing and episodic low rainfall are thought to interact, leading to greater community change than either factor alone. We examined patterns of change and stability in herbaceous community composition through four dry periods, or droughts, over 15 years of the Kenya Long‐term Exclosure Experiment (KLEE), which consists of six different combinations of cattle, native wild herbivores (e.g., zebras, gazelles), and mega‐herbivores (giraffes, elephants). We used principal response curves to analyze the trajectory of change in each herbivore treatment relative to a common initial community and asked how droughts contributed to community change in these treatments. We examined three measures of stability (resistance, variability, and turnover) that correspond to different temporal scales and found that each had a different response to grazing. Treatments that included both cattle and wild herbivores had higher resistance (less net change over 15 years) but were more variable on shorter time scales; in contrast, the more lightly grazed treatments (no herbivores or wild herbivores only) showed lower resistance due to the accumulation of consistent, linear, short‐term change. Community change was greatest during and immediately after droughts in all herbivore treatments. But, while drought contributed to directional change in the less grazed treatments, it contributed to both higher variability and resistance in the more heavily grazed treatments. Much of the community change in lightly grazed treatments (especially after droughts) was due to substantial increases in cover of the palatable grass Brachiaria lachnantha. These results illustrate how herbivory and drought can act together to cause change in grassland communities at the moderate to low end of a grazing intensity continuum. Livestock grazing at a moderate intensity in a system with a long evolutionary history of grazing contributed to long‐term stability. This runs counter to often‐held assumptions that livestock grazing leads to directional, destabilizing shifts in grassland systems

Herbivore Effects on Productivity Vary by Guild: Cattle Increase Mean Productivity While Wildlife Reduce Variability

Wild herbivores and livestock share the majority of rangelands worldwide, yet few controlled experiments have addressed their individual, additive, and interactive impacts on ecosystem function. While ungulate herbivores generally reduce standing biomass, their effects on aboveground net primary production (ANPP) can vary by spatial and temporal context, intensity of herbivory, and herbivore identity and species richness. Some evidence indicates that moderate levels of herbivory can stimulate aboveground productivity, but few studies have explicitly tested the relationships among herbivore identity, grazing intensity, and ANPP. We used a long- term exclosure experiment to examine the effects of three groups of wild and domestic ungulate herbivores (megaherbivores, mesoherbivore wildlife, and cattle) on herbaceous productivity in an African savanna. Using both field measurements (productivity cages) and satellite imagery, we measured the effects of different herbivore guilds, separately and in different combinations, on herbaceous productivity across both space and time. Results from both productivity cage measurements and satellite normalized difference vegetation index (NDVI) demonstrated a positive relationship between mean productivity and total ungulate herbivore pressure, driven in particular by the presence of cattle. In contrast, we found that variation in herbaceous productivity across space and time was driven by the presence of wild herbivores (primarily mesoherbivore wildlife), which significantly reduced heterogeneity in ANPP and NDVI across both space and time. Our results indicate that replacing wildlife with cattle (at moderate densities) could lead to similarly productive but more heterogeneous herbaceous plant communities in rangelands

Spiders (Arachnida: Araneae) from black cotton soil habitats of a highland savanna biome in Laikipia, central Kenya

Spiders were sampled at Mpala Research Centre, Laikipia, Kenya using pitfall trapping and sweep netting. Sampling was conducted from May 2001 to July 2002. A total of 10,487 individuals from 132 species belonging to 30 families were recorded. The family Salticidae had the highest number of species (24), followed by Gnaphosidae (20), Araneidae and Lycosidae (15 each), Theridiidae and Thomisidae (8 each) and Zodariidae (4). Most of the other families had fewer than 4 species. Throughout the study period, species not previously sampled emerged after rainfall peaks. Many species are apparently undescribed, highlighting the inadequate documentation of these taxa in Kenya. We suggest that the spider fauna of black cotton soil habitats is rich and useful for environmental monitoring, that further surveys using other collecting methods are needed, and that support for the conservation of this ecosystem should be continued. The study once more reveals the urgent need for taxonomic studies on Kenyan invertebrates

Elephants Mitigate the Effects of Cattle on Wildlife and Other Ecosystem Traits: Experimental Evidence

On rangelands worldwide, cattle interact with many ecosystem components, most obviously with soils, plants, and other large herbivores. Since 1995, we have been manipulating the presence of cattle, mesoherbivores, and megaherbivores (elephants and giraffes) in a series of eighteen 4-ha (10-acre) plots at the Kenya Long-term Exclosure Experiment. We have demonstrated a wide array of cattle effects on this savanna rangeland, including their reduction of grass cover, wildlife use, and soil nitrogen and phosphorus pools, but their increase of primary productivity and termite abundance. Strikingly, we demonstrate that the presence of mega-herbivores (elephants, mainly) reduces the sizes of these cattle. We provide further experimental evidence that this may be because the elephants are reducing the most desirable (N-rich) forage, causing cattle to slow their extraction of (low-N) grasses, while simultaneously reducing tree cover

Effects of mammalian herbivore declines on plant communities: observations and experiments in an African savanna

1. Herbivores influence the structure and composition of terrestrial plant communities. However, responses of plant communities to herbivory are variable and depend on environmental conditions, herbivore identity and herbivore abundance. As anthropogenic impacts continue to drive large declines in wild herbivores, understanding the context dependence of herbivore impacts on plant communities becomes increasingly important. 2. Exclosure experiments are frequently used to assess how ecosystems reorganize in the face of large wild herbivore defaunation. Yet in many landscapes, declines in large wildlife are often accompanied by other anthropogenic activities, especially land conversion to livestock production. In such cases, exclosure experiments may not reflect typical outcomes of human-driven extirpations of wild herbivores. 3. Here, we examine how plant community responses to changes in the identity and abundance of large herbivores interact with abiotic factors (rainfall and soil properties). We also explore how effects of wild herbivores on plant communities differ between large-scale herbivore exclosures and landscape sites where anthropogenic activity has caused wildlife declines, often accompanied by livestock increases. 4. Abiotic context modulated the responses of plant communities to herbivore declines with stronger effect sizes in lower-productivity environments. Also, shifts in plant community structure, composition and species richness following wildlife declines differed considerably between exclosure experiments and landscape sites in which wild herbivores had declined and were often replaced by livestock. Plant communities in low wildlife landscape sites were distinct in both composition and physical structure from both exclosure and control sites in experiments. The power of environmental (soil and rainfall) gradients in influencing plant response to herbivores was also greatly dampened or absent in the landscape sites. One likely explanation for these observed differences is the compensatory effect of livestock associated with the depression or extirpation of wildlife. 5. Synthesis. Our results emphasize the importance of abiotic environmental heterogeneity in modulating the effects of mammalian herbivory on plant communities and the importance of such covariation in understanding effects of wild herbivore declines. They also suggest caution when extrapolating results from exclosure experiments to predict the consequences of defaunation as it proceeds in the Anthropocene

Wild and domestic savanna herbivores increase smaller vertebrate diversity, but less than additively

Cattle and other livestock graze more than a quarter of the world's terrestrial area and are widely regarded to be drivers of global biodiversity declines. Studies often compare the effects of livestock presence/absence but, to our knowledge, no studies have tested for interactive effects between large wild herbivores and livestock at varying stocking rates on small-bodied wild vertebrates. We investigated the effects of cattle stocking rates (none/moderate/high) on the diversity of wildlife 0.05–1,000 kg using camera traps at a long-term exclosure experiment within a semi-arid savanna ecosystem in central Kenya. In addition, by selectively excluding wild ‘mesoherbivores’ (50–1,000 kg) and ‘megaherbivores’ (>1,000 kg; elephant and giraffe), we tested whether the presence of these two wild herbivore guilds (collectively, ‘larger wild herbivores’) mediates the effect of cattle stocking rate on habitat use and diversity of ‘smaller wildlife’ (mammals ranging between 10 and 70 cm shoulder height and birds). Our results show that cattle enhance alpha diversity of smaller wildlife (with or without larger wild herbivore presence) and of all wildlife 0.05–1,000 kg (with or without megaherbivore presence), by altering vegetation structure. However, for smaller wildlife, this effect is less pronounced in the presence of larger wild herbivores, which also shorten grass. In the absence of cattle, mesoherbivore-accessible sites showed higher alpha diversity of smaller wildlife than sites excluding mesoherbivores. Smaller wildlife habitat use was increased by high cattle stocking rates and wild mesoherbivores more in the presence of the other. Synthesis and applications. Our findings imply that grazing, whether by livestock or wildlife, can enhance local savanna wildlife diversity. The biodiversity benefits of localised increases in herbivory are likely to be due to shortened grass and associated visibility improvements (for predator avoidance/foraging). This suggests that land managers can increase local biodiversity by shortening grass, with wild or domestic herbivores (or both), at least in patches within a taller grass matrix