skip to main content


Title: Herbivory and Drought Reduce the Temporal Stability of Herbaceous Cover by Increasing Synchrony in a Semi-arid Savanna
Ecological stability in plant communities is shaped by bottom-up processes like environmental resource fluctuations and top-down controls such as herbivory, each of which have demonstrated direct effects but may also act indirectly by altering plant community dynamics. These indirect effects, called biotic stability mechanisms, have been studied across environmental gradients, but few studies have assessed the importance of top-down controls on biotic stability mechanisms in conjunction with bottom-up processes. Here we use a long-term herbivore exclusion experiment in central Kenya to explore the joint effects of drought and herbivory (bottom-up and top-down limitation, respectively) on three biotic stability mechanisms: (1) species asynchrony, in which a decline in one species is compensated for by a rise in another, (2) stable dominant species driving overall stability, and (3) the portfolio effect, in which a community property is distributed among multiple species. We calculated the temporal stability of herbaceous cover and biotic stability mechanisms over a 22-year time series and with a moving window to examine changes through time. Both drought and herbivory additively reduced asynchronous dynamics, leading to lower stability during droughts and under high herbivore pressure. This effect is likely attributed to a reduction in palatable dominant species under higher herbivory, which creates space for subordinate species to fluctuate synchronously in response to rainfall variability. Dominant species population stability promoted community stability, an effect that did not vary with precipitation but depended on herbivory. The portfolio effect was not important for stability in this system. Our results demonstrate that this system is naturally dynamic, and a future of increasing drought may reduce its stability. However, these effects will in turn be amplified or buffered depending on changes in herbivore communities and their direct and indirect impacts on plant community dynamics.  more » « less
Award ID(s):
1931224
NSF-PAR ID:
10335422
Author(s) / Creator(s):
; ; ; ; ; ; ; ; ;
Date Published:
Journal Name:
Frontiers in Ecology and Evolution
Volume:
10
ISSN:
2296-701X
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract

    Arid ecosystems are strongly limited by water availability, and precipitation plays a major role in the dynamics of all species in arid regions, as well as the ecosystem processes that occur there. However, understanding how biotic interactions mediate long‐term responses of dryland ecosystems to rainfall remains very fragmented. We report on a unique large‐scale field experiment spanning 25 yr and three trophic levels (plants, small mammal herbivores, predators) in a dryland ecosystem in the northern Chilean Mediterranean Region where we assessed how biotic interactions influence the long‐term plant community responses to precipitation. As the most persistent ecological changes in dryland systems may result from changes in the structure, cover, and composition of the perennial vegetation, we emphasized the interplay between bottom‐up and top‐down controls of perennial plants in our analyses. Rainfall was the primary factor affecting the dynamics of, and interactions among, plants and small mammals. Ephemeral plant cover dynamics closely tracked short‐term annual rainfall, but seemed unaffected by top‐down controls (herbivory). In contrast, the response of the perennial plant cover to precipitation was mediated by (1) a complex interplay between subtle top‐down (herbivory) controls that become more apparent in the long‐term, (2) competition with ephemeral plants during wet years, and (3) an indirect effect of predators on subdominant shrubs and perennial herbs. This long‐term field experiment highlights how climate‐induced responses of arid perennial vegetation are influenced by interactions across trophic levels and temporal scales. In the face of global change, understanding how multi‐trophic controls mediate dryland vegetation responses to climate is essential to properly managing the conservation of biodiversity in arid systems.

     
    more » « less
  2. Abstract

    Animals exert both direct and indirect controls over elemental cycles, linking primary producer‐based (green) and decomposer‐based (brown) food webs through top‐down trophic interactions and bottom‐up element regeneration. Where animals are aggregated at high biomass, they create hotspots of elemental cycling. The relative importance of animal control on elemental cycling depends on animal biomass, species functional traits (i.e. feeding mode and stoichiometry), and their overlap.

    We evaluated how animal community complexity affects the mechanisms regulating energy flow to the brown food web. We conducted a mesocosm experiment where we varied the biomass and overlap of animals with different life history and stoichiometric traits (stream fish and mussels) and measured how this influenced the quantity and fraction of labile carbon available to microbes. We used linear models and structural equation modelling to evaluate direct (excretion) and indirect (herbivory, nutrient availability, and nutrient stoichiometry) effects of animals on bioavailable dissolved organic carbon (BDOC) concentration.

    In experimental stream mesocosms, we found support for both direct (DOC excretion) and indirect (grazing) animal influences on BDOC concentration. Although we found that snail, fish, and mussel biomass increased nutrient concentrations, neither nutrient concentration nor stoichiometry had a significant effect on BDOC concentration. This has been due to the high background nutrient concentration context of our stream mesocosm water. Snails, probably due to their high biomass and small body size, exerted a significant positive direct control on BDOC concentration. Fish and mussels exerted a significant negative indirect control on BDOC via their effects (grazing and bioturbation) on algal biomass.

    Our results imply that primary consumers with different feeding strategies provide a key mechanism regulating the flow of DOC into the brown food web through direct (excretion) and indirect (grazing) controls on primary producers. This highlights that animals can provide important controls on the production of bioavailable organic energy supporting microbes in aquatic ecosystems, but the importance of these controls depends on the nutrient context and the distribution of primary producer and animal biomasses.

     
    more » « less
  3. 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
  4. Abstract

    Top‐down effects of predators and bottom‐up effects of resources are important drivers of community structure and function in a wide array of ecosystems. Fertilization experiments impose variation in resource availability that can mediate the strength of predator impacts, but the prevalence of such interactions across natural productivity gradients is less clear. We studied the joint impacts of top‐down and bottom‐up factors in a tropical mangrove forest system, leveraging fine‐grained patchiness in resource availability and primary productivity on coastal cays of Belize. We excluded birds from canopies of red mangrove (Rhizophoraceae:Rhizophora mangle) for 13 months in zones of phosphorus‐limited, stunted dwarf mangroves, and in adjacent zones of vigorous mangroves that receive detrital subsidies. Birds decreased total arthropod densities by 62%, herbivore densities more than fivefold, and reduced rates of leaf and bud herbivory by 45% and 52%, respectively. Despite similar arthropod densities across both zones of productivity, leaf and bud damage were 2.0 and 4.3 times greater in productive stands. Detrital subsidies strongly impacted a suite of plant traits in productive stands, potentially making leaves more nutritious and vulnerable to damage. Despite consistently strong impacts on herbivory, we did not detect top‐down forcing that impacted mangrove growth, which was similar with and without birds. Our results indicated that both top‐down and bottom‐up forces drive arthropod community dynamics, but attenuation at the plant‐herbivore interface weakens top‐down control by avian insectivores.

     
    more » « less
  5. Abstract

    Pulsed fluxes of organisms across ecosystem boundaries can exert top‐down and bottom‐up effects in recipient food webs, through both direct effects on the subsidized trophic levels and indirect effects on other components of the system. While previous theoretical and empirical studies demonstrate the influence of allochthonous subsidies on bottom‐up and top‐down processes, understanding how these forces act in conjunction is still limited, particularly when an allochthonous resource can simultaneously subsidize multiple trophic levels. Using the Lake Mývatn region in Iceland as an example system of allochthony and its potential effects on multiple trophic levels, we analyzed a mathematical model to evaluate how pulsed subsidies of aquatic insects affect the dynamics of a soil–plant–arthropod food web. We found that the relative balance of top‐down and bottom‐up effects on a given food web compartment was determined by trophic position, subsidy magnitude, and top predators’ ability to exploit the subsidy. For intermediate trophic levels (e.g., detritivores and herbivores), we found that the subsidy could either alleviate or intensify top‐down pressure from the predator. For some parameter combinations, alleviation and intensification occurred sequentially during and after the resource pulse. The total effect of the subsidy on detritivores and herbivores, including top‐down and bottom‐up processes, was determined by the rate at which predator consumption saturated with increasing size of the allochthonous subsidy, with greater saturation leading to increased bottom‐up effects. Our findings illustrate how resource pulses to multiple trophic levels can influence food web dynamics by changing the relative strength of bottom‐up and top‐down effects, with bottom‐up predominating top‐down effects in most scenarios in this subarctic system.

     
    more » « less