Kevin P. Kirkman

Grassland productivity limited by multiple nutrients

Terrestrial ecosystem productivity is widely accepted to be nutrient limited1. Although nitrogen (N) is deemed a key determinant of aboveground net primary production (ANPP)2,3, the prevalence of co-limitation by N and phosphorus (P) is increasingly recognized4–8. However, the extent to which terrestrial productivity is co-limited by nutrients other than N and P has remained unclear. Here, we report results from a standardized factorial nutrient addition experiment, in which we added N, P and potassium (K) combined with a selection of micronutrients (K+μ), alone or in concert, to 42 grassland sites spanning five continents, and monitored ANPP. Nutrient availability limited productivity at 31 of the 42 grassland sites. And pairwise combinations of N, P, and K+μ co-limited ANPP at 29 of the sites. Nitrogen limitation peaked in cool, high latitude sites. Our findings highlight the importance of less studied nutrients, such as K and micronutrients, for grassland productivity, and point to significant variations in the type and degree of nutrient limitation. We suggest that multiple-nutrient constraints must be considered when assessing the ecosystem-scale consequences of nutrient enrichment.

Abundance of introduced species at home predicts abundance away in herbaceous communities

Many ecosystems worldwide are dominated by introduced plant species, leading to loss of biodiversity and ecosystem function. A common but rarely tested assumption is that these plants are more abundant in introduced vs. native communities, because ecological or evolutionary-based shifts in populations underlie invasion success. Here, data for 26 herbaceous species at 39 sites, within eight countries, revealed that species abundances were similar at native (home) and introduced (away) sites - grass species were generally abundant home and away, while forbs were low in abundance, but more abundant at home. Sites with six or more of these species had similar community abundance hierarchies, suggesting that suites of introduced species are assembling similarly on different continents. Overall, we found that substantial changes to populations are not necessarily a pre-condition for invasion success and that increases in species abundance are unusual. Instead, abundance at home predicts abundance away, a potentially useful additional criterion for biosecurity programmes.

Species abundance at home predicts abundance away in grasslands

Many ecosystems worldwide are dominated by introduced plant species, leading to loss of biodiversity and ecosystem function. A common but rarely tested assumption is that these plants are more abundant in introduced vs. native communities, because ecological or evolutionary-based shifts in populations underlie invasion success. Here, data for 26 herbaceous species at 39 sites, within eight countries, revealed that species abundances were similar at native (home) and introduced (away) sites - grass species were generally abundant home and away, while forbs were low in abundance, but more abundant at home. Sites with six or more of these species had similar community abundance hierarchies, suggesting that suites of introduced species are assembling similarly on different continents. Overall, we found that substantial changes to populations are not necessarily a pre-condition for invasion success and that increases in species abundance are unusual. Instead, abundance at home predicts abundance away, a potentially useful additional criterion for biosecurity programmes.

Controls of aboveground net primary production in mesic savanna grasslands: an inter-hemispheric comparison.

Patterns and controls of annual aboveground net primary productivity (ANPP) are fundamental metrics of ecosystem functioning. It is generally assumed, but rarely tested, that determinants of ANPP in one region within a biome will operate similarly throughout that biome, as long as physiognomy and climate are broadly consistent. We tested this assumption by quantifying ANPP responses to fire, grazing history, and nitrogen (N) addition in North American (NA) and South African (SA) savanna grasslands. We found that total ANPP responded in generally consistent ways to fire, grazing history, and N addition on both continents. Annual fire in both NA and SA consistently stimulated total ANPP (28–100%) relative to unburned treatments at sites with deep soils, and had no effect on ANPP in sites with shallow soils. Fire did not affect total ANPP in sites with a recent history of grazing, regardless of whether a single or a diverse suite of large herbivores was present. N addition interacted strongly and consistently with fire regime in both NA and SA. In annually burned sites that were not grazed, total ANPP was stimulated by N addition (29–39%), but there was no effect of N fertilization in the absence of fire. In contrast, responses in forb ANPP to fire and grazing were somewhat divergent across this biome. Annual fire in NA reduced forb ANPP, whereas grazing increased forb ANPP, but neither response was evident in SA. Thus, despite a consistent response in total ANPP, divergent responses in forb ANPP suggest that other aspects of community structure and ecosystem functioning differ in important ways between these mesic savanna grasslands.

Which grazing management practices are most appropriate for maintaining biodiversity in South African grassland

The relative benefit of different grazing management practices for maintaining grassland biodiversity integrity was assessed. Practices considered were grazing system, stocking rate, animal type, grazing season, fire–grazing interaction, plus burning regime. Sparse empirical data indicated that stocking rate, cattle-to-sheep ratio, and increased fire intervals can affect plant diversity. Certain species behave as increaser or decreaser species in response to stocking rate or cattle-to-sheep ratio. Plant diversity appears robust in the face of variation in fire regime except for preclusion of fire. Multicriteria analysis was used to assess the relative impact of grazing systems (continuous, conventional, or high-intensity grazing by sheep, cattle, or sheep plus cattle) on grassland biodiversity integrity. Selected indicators covered landscape composition, structure and functioning. High-intensity systems and continuous sheep grazing have the most, and continuous grazing with beef the least, negative impact, depending on stocking rate. The relation between veld condition and plant diversity needs to be tested further before veld condition is used as a management tool for maintaining biodiversity. Proposed research priorities include inventory of management systems used, rapid expansion of the empirical data base, an improved understanding of patterns of plant diversity across the biome, and evaluation of the pyro–grazing diversity paradigm.

Plant community response to loss of large herbivores differs between North American and South African savanna grasslands

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.

Landscape-scale feeding patterns of African elephant inferred from carbon isotope analysis of feces

The African elephant (Loxodonta africana) is a large-bodied, generalist herbivore that eats both browse and grass. The proportions of browse and grass consumed are largely expected to reflect the relative availability of these resources. We investigated variations in browse (C3 biomass) and grass (C4) intake of the African elephant across seasons and habitats by stable carbon isotope analysis of elephant feces collected from Kruger National Park, South Africa. The results reflect a shift in diet from higher C4 grass intake during wet season months to more C3 browse-dominated diets in the dry season. Seasonal trends were correlated with changes in rainfall and with nitrogen (%N) content of available grasses, supporting predictions that grass is favored when its availability and nutritional value increase. However, switches to dry season browsing were significantly smaller in woodland and grassland habitats where tree communities are dominated by mopane (Colophospermum mopane), suggesting that grasses were favored here even in the dry season. Regional differences in diet did not reflect differences in grass biomass, tree density, or canopy cover. There was a consistent relationship between %C4 intake and tree species diversity, implying that extensive browsing is avoided in habitats characterized by low tree species diversity and strong dominance patterns, i.e., mopane-dominated habitats. Although mopane is known to be a preferred species, maintaining dietary diversity appears to be a constraint to elephants, which they can overcome by supplementing their diets with less abundant resources (dry season grass). Such variations in feeding behavior likely influence the degree of impact on plant communities and can therefore provide key information for managing elephants over large, spatially diverse, areas.

Stable isotope series from elephant ivory reveal lifetime histories of a true dietary generalist

Longitudinal studies have revealed how variation in resource use within consumer populations can impact their dynamics and functional significance in communities. Here, we investigate multi-decadal diet variations within individuals of a keystone megaherbivore species, the African elephant (Loxodonta africana), using serial stable isotope analysis of tusks from the Kruger National Park, South Africa. These records, representing the longest continuous diet histories documented for any extant species, reveal extensive seasonal and annual variations in isotopic—and hence dietary—niches of individuals, but little variation between them. Lack of niche distinction across individuals contrasts several recent studies, which found relatively high levels of individual niche specialization in various taxa. Our result is consistent with theory that individual mammal herbivores are nutritionally constrained to maintain broad diet niches. Individual diet specialization would also be a costly strategy for large-bodied taxa foraging over wide areas in spatio-temporally heterogeneous environments. High levels of within-individual diet variability occurred within and across seasons, and persisted despite an overall increase in inferred C4 grass consumption through the twentieth century. We suggest that switching between C3 browsing and C4 grazing over extended time scales facilitates elephant survival through environmental change, and could even allow recovery of overused resources.

Inducible defences in Acacia sieberiana in response to giraffe browsing

Acacia trees comprise a principal component of the diet of many African browsers. The resultant browsing pressure has led to the evolution of both physical and chemical responses in Acacia trees. In an observational study, we investigated the physical and chemical defenses in Acacia sieberiana var. woodii in response to different intensities of giraffe herbivory. Trees with high browsing intensity had significantly longer spines, smaller leaves and higher total cyanide (prussic acid) concentrations than trees with low browsing intensity. No significant differences were present in the density of spines and leaves between high and low browsing intensity sites. Similarly, no significant differences were found in the concentrations of condensed tannins and total leaf nitrogen. However, comparisons with other Acacia species revealed that the sampled A. sieberiana trees had substantially higher condensed tannin concentrations than many other species. It therefore appears that condensed tannins may form a high constitutive, or fixed, defence in A. sieberiana in this population, while prussic acid and spine length (with a decrease in leaf size) form important inducible defences.

Smoke Solutions and Temperature Influence the Germination and Seedling Growth of South African Mesic Grassland Species

Abstract Fire, natural or of anthropologic origin, is a recurrent phenomenon in South African mesic grassland. The species composition of these grasslands is sensitive to fire, particularly fire frequency. However, the mechanisms involved in influencing species composition are not fully understood. Currently there is a general suggestion that plant-derived smoke and smoke-isolated biologically active butenolide compound provide an important germination cue for a range of Poaceae species. Studies also show that these smoke solutions play a role in vegetative growth of many plants. We examined if this fire-response syndrome is related to the effect of plant-derived smoke-water (1∶500 v/v) and smoke-derived butenolide compound (10−8 M) on seed germination and seedling growth of six major constituent species of the grassland. In addition, the interaction of the smoke solutions with temperature was examined by incubating seeds at a range of temperatures. Treating seeds with smoke-water and butenolide, the germination rate and final germination percentage were greater in three of the six species. Themeda triandra Forssk. and Tristachya leucothrix Trin. ex Nees showed the greatest response, with final germination increased from 43% to 67% and 35% to 63%, respectively. With increasing temperature (> 30°C), Aristida junciformis Trin. & Rupr., Hyparrhenia hirta (L.) Staph, and Panicum maximum Jacq. responded positively to the test solutions. In nearly all the species tested, smoke-water–treated seeds produced significantly longer shoots or roots. However, the degree of response varied from species to species and across different temperatures. Findings from this study suggest that plant-derived smoke and its interaction with temperature may significantly influence the germination and seedling growth of the South African mesic grassland species, which can further alter the grassland composition.