skip to main content
US FlagAn official website of the United States government
dot gov icon
Official websites use .gov
A .gov website belongs to an official government organization in the United States.
https lock icon
Secure .gov websites use HTTPS
A lock ( lock ) or https:// means you've safely connected to the .gov website. Share sensitive information only on official, secure websites.


Title: The past, present, and future of herbivore impacts on savanna vegetation
Abstract Herbivory is a key process structuring vegetation in savannas, especially in Africa where large mammal herbivore communities remain intact. Exclusion experiments consistently show that herbivores impact savanna vegetation, but effect size variation has resisted explanation, limiting our understanding of the past, present and future roles of herbivory in savanna ecosystems.Synthesis of vegetation responses to herbivore exclusion shows that herbivory decreased grass abundance by 57.0% and tree abundance by 30.6% across African savannas.The magnitude of herbivore exclusion effects scaled with herbivore abundance: more grazing herbivores resulted in larger grass responses and more browsing herbivores in larger tree responses. However, existing experiments are concentrated in semi‐arid savannas (400–800‐mm rainfall) and soils data are mostly lacking, which makes disentangling environmental constraints a challenge and priority for future research.Observed herbivore impacts were ~2.1× larger than existing estimates modelled based on consumption. Wildlife metabolic rates may be higher than are usually used for estimating consumption, which offers one clear avenue for reconciling estimated herbivore consumption with observed herbivore impacts. Plant‐soil feedbacks, plant community composition, and the phenological or demographic timing of herbivory may also influence vegetation productivity, thereby magnifying herbivore impacts.Because herbivore abundance so closely predicts vegetation impact, changes in herbivore abundance through time are likely predictive of the past and future of their impacts. Grazer diversity in Africa has declined from its peak 1 million years ago and wild grazer abundance has declined historically, suggesting that grazing likely had larger impacts in the past than it does today.Current wildlife impacts are dominated by small‐bodied mixed feeders, which will likely continue into the future, but the magnitude of top‐down control may also depend on changing climate, fire and atmospheric CO2.Synthesis. Herbivore biomass determines the magnitude of their impacts on savanna vegetation, with effect sizes based on direct observation that outstrip existing modelled estimates across African savannas. Findings suggest substantial ecosystem impacts of herbivory and allow us to generate evidence‐based hypotheses of the past and future impacts of herbivores on savanna vegetation.  more » « less
Award ID(s):
1802453 1826666
PAR ID:
10446376
Author(s) / Creator(s):
 ;  ;  ;  ;  
Publisher / Repository:
Wiley-Blackwell
Date Published:
Journal Name:
Journal of Ecology
Volume:
109
Issue:
8
ISSN:
0022-0477
Page Range / eLocation ID:
p. 2804-2822
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract 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. 
    more » « less
  2. Abstract Many savannas are experiencing increased cover of trees and shrubs, resulting in reduced herbaceous productivity, shifts in savanna functional structure and potential reductions in ecotourism. Clearing woody plants has been suggested as an effective management strategy to mitigate these effects and restore these systems to an open state with higher rates of grass production and herbivory. This study investigated the effectiveness of repeated shrub clearing as a tool to mitigate bush encroachment in a semi‐arid savanna in southern Africa.We present data from a 7‐year experiment in the Mthimkhulu Game Reserve bordering Kruger National Park, South Africa.Colophospermum mopanestems and resprouting shoots were basally cut 2–3 times per year (2015–2022) in three pairs of treatment and control plots of 60 × 60 m. We monitored changes in soil moisture, grass biomass and herbivore activity via dung counts. We assessedC. mopanephysiological responses to repeated cutting using non‐structural carbohydrates and stable water isotopes to infer changes to energy storage and functional rooting depth, respectively.The cleared treatment had higher soil moisture and grass biomass than the control treatment. Dung counts showed impala and buffalo visited the cleared treatment more frequently than the control treatment.Repeated cutting had limited effects onC. mopanesurvival in the first 2–3 years after initial clearing, but 80% of individuals were dead after 7 years. Repeatedly cutC. mopanehad lower belowground starch concentrations and used water from shallower soil depths thanC. mopanein control plots.Synthesis and applications. Repeated cutting increased soil moisture availability and grass biomass, and attracted charismatic grazing herbivores. While more costly than once‐off clearing methods, this practice created more employment opportunities for a neighbouring rural community. Transforming portions of the ecosystem to a grass‐dominated state may increase ecotourism potential through improved game viewing in open systems. 
    more » « less
  3. Abstract Despite wide recognition of the importance of anthropogenically driven changes in large herbivore communities—including both declines in wildlife and increases in livestock—there remain large gaps in our knowledge about the impacts of these changes on plant communities, particularly when combined with concurrent changes in climate. Considering these prominent forms of global change in tandem enables us to better understand controls on savanna vegetation structure and diversity under real‐world conditions.We conducted a field experiment using complete and semi‐permeable herbivore exclosures to explore the difference in plant communities among sites with wild herbivores only, with cattle in addition to wild herbivores, and with no large herbivores. To understand variation in effects across climatic contexts, the experiment was replicated at three locations along a topoclimatic gradient in California. Critically, this is the first such experiment to compare cattle and wildlife impacts along an environmental gradient within a single controlled experiment.Vegetation structure responded strongly to herbivore treatment regardless of climate. Relative to the isolated effects of wildlife, exclusion of all large herbivores generally increased structural components related to cover and above‐ground biomass while the addition of cattle led to reductions in vegetation cover, litter, shading and standing biomass. Furthermore, wildlife had a consistent neutral or positive effect on plant diversity, while the effect of livestock addition was context dependent. Cattle had a neutral to strongly negative effect at low aridity, but a positive effect at high aridity. These results suggest that (a) herbivore effects can override climate effects on vegetation structure, (b) cattle addition can drive different effects on diversity and (c) herbivore effects on diversity are modulated by climate.Synthesis. Our results illustrate very distinctive shifts in plant communities between two realistic forms of change in ungulate herbivore assemblages—livestock addition and large herbivore losses—particularly for plant diversity responses, and that these responses vary across climatic contexts. This finding has important implications for the management and protection of plant biodiversity given that over a quarter of the Earth's land area is managed for livestock and climate regimes are changing globally. 
    more » « less
  4. Abstract Large grazers modify vegetated ecosystems and are increasingly viewed as keystone species in trophic rewilding schemes. Yet, as their ecosystem influences are context‐dependent, a crucial challenge is identifying where grazers sustain, versus undermine, important ecosystem properties and their resilience.Previous work in diverse European saltmarshes found that, despite changing plant and invertebrate community structure, grazers do not suppress below‐ground properties, including soil organic carbon (SOC). We hypothesised that, in contrast, eastern US saltmarshes would be sensitive to large grazers as extensive areas are dominated by a single grass,Spartina alterniflora. We predicted that grazers would reduce above‐ and below‐groundSpartinabiomass, suppress invertebrate densities, shift soil texture and ultimately reduce SOC concentration.We tested our hypotheses using a replicated 51‐month large grazer (horse) exclusion experiment in Georgia, coupled with observations of 14 long‐term grazed sites, spanning ~1000 km of the eastern US coast.Grazer exclusion quickly led to increasedSpartinaheight, cover and flowering, and increased snail density. Changes in vegetation structure were reflected in modified soil texture (reduced sand, increased clay) and elevated root biomass, yet we found no response of SOC. Large grazer exclusion also reduced drought‐associated vegetation die‐off.We also observed vegetation shifts in sites along the eastern US seaboard where grazing has occurred for hundreds of years. Unlike in the exclusion experiment, long‐term grazing was associated with reduced SOC. A structural equation model implicated grazing by revealing reduced stem height as a key driver of reduced soil organic carbon.Synthesis: These results illustrate the context dependency of large grazer impacts on ecosystem properties in coastal wetlands. In contrast to well‐studied European marshes, eastern US marshes are dominated and structured by a single foundational grass species resulting in vegetation and soil properties being more sensitive to grazing. Coastal systems characterised by a single foundation species might be inherently vulnerable to large grazers and lack resilience in the face of other disturbances, underlining that frameworks to explain and predict large grazer impacts must account for geographic variation in ecosystem structure. 
    more » « less
  5. African savannas are the last stronghold of diverse large-mammal communities, and a major focus of savanna ecology is to understand how these animals affect the relative abundance of trees and grasses. However, savannas support diverse plant life-forms, and human-induced changes in large-herbivore assemblages—declining wildlife populations and their displacement by livestock—may cause unexpected shifts in plant community composition. We investigated how herbivory affects the prevalence of lianas (woody vines) and their impact on trees in an East African savanna. Although scarce (<2% of tree canopy area) and defended by toxic latex, the dominant liana,Cynanchum viminale(Apocynaceae), was eaten by 15 wild large-herbivore species and was consumed in bulk by native browsers during experimental cafeteria trials. In contrast, domesticated ungulates rarely ate lianas. When we experimentally excluded all large herbivores for periods of 8 to 17 y (simulating extirpation), liana abundance increased dramatically, with up to 75% of trees infested. Piecewise exclusion of different-sized herbivores revealed functional complementarity among size classes in suppressing lianas. Liana infestation reduced tree growth and reproduction, but herbivores quickly cleared lianas from trees after the removal of 18-y-old exclosure fences (simulating rewilding). A simple model of liana contagion showed that, without herbivores, the long-term equilibrium could be either endemic (liana–tree coexistence) or an all-liana alternative stable state. We conclude that ongoing declines of wild large-herbivore populations will disrupt the structure and functioning of many African savannas in ways that have received little attention and that may not be mitigated by replacing wildlife with livestock. 
    more » « less