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1. Given the role of fire in shaping ecosystems, especially grasslands and savannas, it is important to understand its broader impact on these systems. Post-fire stimulation of plant nutrients is thought to benefit grazing mammals and explain their preference for burned areas. However, fire also reduces vegetation height and increases visibility, thereby potentially reducing predation risk. Consequently, fire may be more beneficial to smaller herbivores, with higher nutritional needs and greater risks of predation. 2. We tested the impacts of burning on different sized herbivores' habitat preference in Serengeti National Park, as mediated by burning's effects on vegetation height, live : dead biomass ratio and leaf nutrients. 3. Burning caused a less than 4 month increase in leaf nitrogen (N), and leaf non-N nutrients [copper (Cu), potassium (K), and magnesium (Mg)] and a decrease in vegetation height and live : dead biomass. During this period, total herbivore counts were higher on burned areas. Generally, smaller herbivores preferred burned areas more strongly than larger herbivores. 4. Unfortunately, it was not possible to determine the vegetation characteristics that explained burned area preference for each of the herbivore species observed. However, total herbivore abundance and impala (Aepyceros melampus) preference for burned areas was due to the increases in non-N nutrients caused by burning. 5. These findings suggest that burned area attractiveness to herbivores is mainly driven by changes to forage quality and not potential decreases in predation risk caused by reductions in vegetation height.

Mechanistic explanations of herbivore spatial distribution have focused largely on either resource-related (bottom-up) or predation-related (top-down) factors. We studied direct and indirect influences on the spatial distributions of Serengeti herbivore hotspots, defined as temporally stable areas inhabited by mixed herds of resident grazers. Remote sensing and variation in landscape features were first used to create a map of the spatial distribution of hotspots, which was tested for accuracy against an independent data set of herbivore observations. Subsequently, we applied structural equation modeling to data on soil fertility and plant quality and quantity across a range of sites. We found that hotspots in Serengeti occur in areas that are relatively flat and located away from rivers, sites where ungulates are less susceptible to predation. Further, hotspots tend to occur in areas where hydrology and rainfall create conditions of relatively low-standing plant biomass, which, coupled with grazing, increases forage quality while decreasing predation risk. Low-standing biomass and higher leaf concentrations of N, Na, and Mg were strong direct predictors of hotspot occurrence. Soil fertility had indirect effects on hotspot occurrence by promoting leaf Na and Mg. The results indicate that landscape features contribute in direct and indirect ways to influence the spatial distribution of hotspots and that the best models incorporated both resource- and predation-related factors. Our study highlights the collective and simultaneous role of bottom-up and top-down factors in determining ungulate spatial distributions.

Herbivory and fire shape plant community structure in grass-dominated ecosystems, but these disturbance regimes are being altered around the world. To assess the consequences of such alterations, we excluded large herbivores for seven years from mesic savanna grasslands sites burned at different frequencies in North America (Konza Prairie Biological Station, Kansas, USA) and South Africa (Kruger National Park). We hypothesized that the removal of a single grass-feeding herbivore from Konza would decrease plant community richness and shift community composition due to increased dominance by grasses. Similarly, we expected grass dominance to increase at Kruger when removing large herbivores, but because large herbivores are more diverse, targeting both grasses and forbs, at this study site, the changes due to herbivore removal would be muted. After seven years of large-herbivore exclusion, richness strongly decreased and community composition changed at Konza, whereas little change was evident at Kruger. We found that this divergence in response was largely due to differences in the traits and numbers of dominant grasses between the study sites rather than the predicted differences in herbivore assemblages. Thus, the diversity of large herbivores lost may be less important in determining plant community dynamics than the functional traits of the grasses that dominate mesic, disturbance-maintained savanna grasslands.

In areas with diverse herbivore communities such as African savannas, the frequency of disturbance by fire may alter the top–down role of different herbivore species on plant community dynamics. In a seven year experiment in the Kruger National Park, South Africa, we examined the habitat use of nine common herbivore species across annually burned, triennially burned and unburned areas. We also used two types of exclosures (plus open access controls) to examine the impacts of different herbivores on plant community dynamics across fire disturbance regimes. Full exclosures excluded all herbivores > 0.5 kg (e.g. elephant, zebra, impala) while partial exclosures allowed access only to animals with shoulder heights ≤ 0.85 m (e.g. impala, steenbok). Annual burns attracted a diverse suite of herbivores, and exclusion of larger herbivores (e.g. elephant, zebra, wildebeest) increased plant abundance. When smaller species, mainly impala, were also excluded there were declines in plant diversity, likely mediated by a decline in open space available for colonization of uncommon plant species. Unburned areas attracted the least diverse suite of herbivores, dominated by impala. Here, herbivore exclusion, especially of impala, led to strong declines in plant richness and diversity. With no fire disturbance, herbivore exclusion led to competitive exclusion via increases in plant dominance and light limitation. In contrast, on triennial burns, herbivore exclusion had no effect on plant richness or diversity, potentially due to relatively little open space for colonization across exclosure treatments but also little competitive exclusion due to the intermediate fire disturbance. Further, the diverse suite of grazers and browsers on triennial burns may have had a compensating effect of on the diversity of grasses and forbs. Ultimately, our work shows that differential disturbance regimes can result in differential consumer pressure across a landscape and result in heterogeneous patterns in top–down control of community dynamics.

1. The indicator theory of sexual selection suggests that mating displays honestly signal aspects of fitness. While rarely studied, kinematic (locomotor) performance is an excellent candidate for an honest indicator, as mating displays of many animals include rapid or extended locomotion that may be physiologically correlated with performance traits that impact survival. 2. We investigate the indicator value of display locomotion of wild-caught male guppies, Poecilia reticulata, by examining relationships between mating display kinematic traits, anti-predator kinematic traits, and survival during a subsequent staged encounter with a natural predator, the pike cichlid Crenicichla alta. 3. We first compared guppy display kinematics with subsequent survival, and found that display body angle and angular speed positively predicted survival. We next compared anti-predator kinematic and tactical traits with survival, to identify traits that might link mating displays to survival. We measured anti-predator traits in two tests, first in response to a standardized stimulus (fast start test), and second in response to the live predator (encounter test). Guppy fast start speed and encounter speed, time in refuges, and approach distance (response distance) all positively predicted survival, while encounter swim duration negatively predicted survival. These data provided our final hypothesis, that these particular anti-predator traits would be correlated with mating display kinematics. However, we detected only one of eight predicted correlations, a negative relationship between display body angle and encounter swim duration that may reflect an energy trade-off. 4. We conclude that courtship locomotor performance can be an honest survival indicator in guppies, and that the mechanism linking courtship to survival merits further study. These results suggest that courtship locomotion may contribute to viability impacts on the evolution of animal mate choice, and support others in suggesting that these traits may reward greater attention in sexual and natural selection studies.

Aim: Theory suggests that as ecological systems approach regime shifts, they become increasingly slow in recovering from perturbations. This phenomenon, known as critical slowing down [CSD], leads to spatial and temporal signatures in ecological state variables, thus potentially offering early indicators of regime shifts. Indicators using temporal dynamics have been empirically validated in laboratory microcosms and other well-mixed systems, but tests of spatial indicators of regime shifts at large spatial scales in the field are rare due to the relative absence of high-resolution data and difficulties in experimental manipulations. Here, we test theoretical predictions of CSD-based spatial indicators using large-scale field data from the Serengeti-Mara grassland-woodland system. Location: Serengeti-Mara ecosystem, Tanzania and Kenya. Time period: Year 2000 Major taxa studied: Vegetation Method: We used a space-for-time substitution method to empirically test the validity of CSD-based spatial indicators, i.e., we computed indicators along a spatial [in lieu of temporal] gradient of ecological states. First we used a model of vegetation dynamics to determine if our space-for-time substitution method was appropriate. Then we tested for CSD-based spatial indicators using high-resolution spatial vegetation [30 m] and rainfall [2.5 km] data from the Serengeti-Mara ecosystem. Results: Our model predicts that CSD-based indicators increase along a spatial gradient of alternative vegetation states. Empirical analyses suggest that grasslands and woodlands occur as alternative stable states in the Serengeti-Mara ecosystem with rainfall as one of the potential drivers of transitions between these states. We found that four indices of CSD showed the theoretically expected increasing trends along spatial gradients of grasslands to woodlands: spatial variance, spatial skewness, spatial correlation at lag-1 and spatial spectra at low frequencies. Main conclusions: Our results suggest that CSD-based spatial indicators can offer early warning signals of critical transitions in large-scale ecosystems.

We sought to understand the role that water availability (expressed as an aridity index) plays in determining regional and global patterns of richness and evenness, and in turn how these water availability–diversity relationships may result in different richness–evenness relationships at regional and global scales. We examined relationships between water availability, richness and evenness for eight grassy biomes spanning broad water availability gradients on five continents. Our study found that relationships between richness and water availability switched from positive for drier (South Africa, Tibet and USA) vs. negative for wetter (India) biomes, though were not significant for the remaining biomes. In contrast, only the India biome showed a significant relationship between water availability and evenness, which was negative. Globally, the richness–water availability relationship was hump-shaped, however, not significant for evenness. At the regional scale, a positive richness–evenness relationship was found for grassy biomes in India and Inner Mongolia, China. In contrast, this relationship was weakly concave-up globally. These results suggest that different, independent factors are determining patterns of species richness and evenness in grassy biomes, resulting in differing richness–evenness relationships at regional and global scales. As a consequence, richness and evenness may respond very differently across spatial gradients to anthropogenic changes, such as climate change.

1. Productivity and herbivory often interact to shape plant community composition and species richness with levels of production mediating the impact of herbivory. However, differences in herbivore traits such as size, feeding guild and dietary requirements may result in different impacts of diverse herbivore guilds across productivity gradients. 2. We used size-selective herbivore exclosures to separate the effects of herbivory by larger herbivores, such as elephant, Burchell's zebra and blue wildebeest from those of medium/smaller herbivores, such as impala and warthog, on herbaceous plant communities. These exclosures were established along a 10-fold productivity gradient, ranging from 90 to 950 g m⁻² of standing plant biomass in the Kruger National Park, South Africa. 3. Exclusion of all herbivores generally increased plant species richness at low productivity, but decreased richness at high productivity. Exclusion of medium/smaller herbivores (e.g. impala, warthog) showed stronger effects on plant richness, particularly loss of forbs, at higher productivity rather than at lower productivity. In contrast, exclusion of larger herbivores had stronger effects on plant richness, typically with increasing forb richness, at low rather than high productivity. 4. The change in species richness appeared linked to changes in light availability following herbivore exclusion. Strong increases in shading led to declines in species richness while more moderate increases in shading led to increases in species richness, possibly due to amelioration of heat and water stress by modest increases in shading. 5. Increasing plant dominance, which likely alters multiple mechanisms of plant interactions, was correlated with declines in plant richness following herbivore exclusion. The impact of increasing dominance on plant richness operated independent of productivity, with the exclusion of impala appearing particularly important in driving this relationship. 6. Synthesis. We show that the impact of herbivore losses on plant diversity will be strongly situation dependent and will vary with the herbivores lost (e.g. larger vs. smaller, grazers vs. browsers), plant functional type (e.g. grasses vs. forbs) and environmental context (e.g. productivity). Although larger herbivores are often emphasized for their strong impacts on community dynamics and ecosystem processes, we show that smaller, abundant herbivores can exert strong top-down control on plant communities.

Fire and grazing, key determinants of structure and function of savanna grasslands worldwide, have been extensively altered by humans. We used existing long-term manipulations of fire and grazing in North American and South African mesic savanna grasslands, as well as new experiments, to determine whether the impacts of fire and grazing by large herbivores differed between these systems. We found that despite a body of literature suggesting that these savanna grasslands respond uniquely to fire and grazing, their ecosystem responses (aboveground productivity) were generally similar. In contrast, plant-community responses to fire and herbivores diverged strongly between systems. The differences in plant-community responses, as well as convergence in ecosystem function, were underpinned by a common mechanism: the response of grass dominance to changing fire and grazing regimes. As a result, divergent responses of plant communities to altered fire and grazing regimes did not preclude convergence in ecosystem function.

Fire and herbivory are important determinants of nutrient availability in savanna ecosystems. Fire and herbivory effects on the nutritive quality of savanna vegetation can occur directly, independent of changes in the plant community, or indirectly, via effects on the plant community. Indirect effects can be further subdivided into those occurring because of changes in plant species composition or plant abundance (i.e., quality versus quantity). We studied relationships between fire, herbivory, rainfall, soil fertility, and leaf nitrogen (N), phosphorus (P), and sodium (Na) at 30 sites inside and outside of Serengeti National Park. Using structural equation modeling, we asked whether fire and herbivory influences were largely direct or indirect and how their signs and strengths differed within the context of natural savanna processes. Herbivory was associated with enhanced leaf N and P through changes in plant biomass and community composition. Fire was associated with reduced leaf nutrient concentrations through changes in plant community composition. Additionally, fire had direct positive effects on Na and nonlinear direct effects on P that partially mitigated the indirect negative effects. Key mechanisms by which fire reduced plant nutritive quality were through reductions of Na‐rich grasses and increased abundance ofThemeda triandra, which had below‐average leaf nutrients.