Herbivory Regimes Research Articles

Native lagomorphs facilitate noxious weeds in a semi-arid rangeland

Rangeland management has traditionally focused on the grazing effects of livestock, or on wild ungulates because they are potential competitors with livestock. However, herbivory by smaller herbivores, such as lagomorphs, could play a much greater role in shaping the plant community than is commonly thought. Being selective feeders with high mass-specific metabolic rates, we hypothesized that lagomorphs impose an herbivory regime on the rangeland that differs from that of ungulates and drives the plant community toward a different composition. We used an 8-year exclosure experiment with three plot types (complete exclosure, partial exclosure, open rangeland), each 5.95 m2, to separate the effects of grazing by lagomorphs (jackrabbits and cottontails) from the effects of the large grazers (mainly cattle and bison) on vegetation structure and plant community composition. We replicated the experiment across 20 sites in a semiarid rangeland in the western US. The plant community in the complete exclosure plots (lagomorphs and ungulates excluded) developed a fivefold greater standing crop of grass than the open rangeland but supported the lowest biomass, density, and aerial cover of forbs. Partial exclosure plots (ungulates excluded, lagomorphs included) developed a community with higher forb phytomass (33.6% increase relative to open rangeland) and lower forb species richness (16.4% decrease relative to open rangeland). This effect was mainly caused by the encroachment of noxious weeds, including Salsola tragus (prickly Russian thistle/tumbleweed), which covered up to 37% of the ground in plots that lagomorphs had exclusive access to. We infer that selective grazing by lagomorphs facilitates noxious weeds that are elsewhere controlled to some extent by (a) the coarse grazing and trampling effects of large ungulates and (b) competition with native herbaceous species. For semiarid rangelands that have been degraded by livestock grazing, our complete and partial exclosures show that a reduction in stocking rate might not return the rangeland to its former state due to a hysteresis effect resulting from the facilitation of noxious weeds by lagomorphs. Rangeland management should be informed by an understanding of the entire food web, which includes the direct and indirect grazing effects of small herbivores such as lagomorphs.

Herbivory modifies plant symbiont number and impact on host plant performance in the field.

Species interactions are a unifying theme in ecology and evolution. Both fields are currently moving beyond their historical focus on isolated pairwise relationships to understand how ecological communities affect focal interactions. Additional species can modify both the number of interactions and the fitness consequences of each interaction (i.e., selection). Although only selection affects the evolution of the focal interaction, the two are often conflated, limiting our understanding of the evolution of multispecies interactions. We manipulated aboveground herbivory on the legume Medicago lupulina in the field and quantified its effect on number of symbionts and the per-symbiont impact on plant performance in two belowground symbioses: mutualistic rhizobia bacteria (Ensifer meliloti) and parasitic root-knot nematodes (Meloidogyne hapla). We found that herbivores modified the number of rhizobia nodules, as well as the benefit per nodule. However, each effect was specific to a distinct herbivory regime: natural herbivory affected nodule number, whereas leafhoppers (Cicadellidae) weakened the per nodule benefit. We did not detect any effect of herbivory on nematode gall number or the cost of infection. Our data demonstrate that distinguishing between symbiont number from the fitness consequences of symbiosis is crucial to accurately infer how pairwise interactions will evolve in a community.

Experimental insect suppression causes loss of induced, but not constitutive, resistance in Solanumcarolinense.

Spatiotemporal variation in herbivory is a major driver of intraspecific variation in plant defense. Comparatively little is known, however, about how changes in herbivory regime affect the balance of constitutive and induced resistance, which are often considered alternative defensive strategies. Here, we investigated how nearly a decade of insect herbivore suppression affected constitutive and induced resistance in horsenettle (Solanum carolinense), a widespread herbaceous perennial. We allowed replicated horsenettle populations to respond to the presence or absence of herbivores by applying insecticide to all plants in half of 16 field plots. Horsenettle density rapidly increased in response to insecticide treatment, and this effect persisted for at least 4 years after the cessation of herbivore suppression. We subsequently grew half-sibling families from seeds collected during and shortly after insecticide treatment in a common garden and found strong effects of insect suppression on induced resistance. Feeding trials in field mesocosms with false Colorado potato beetles (Leptinotarsa juncta), a common specialist herbivore, revealed that multiyear herbivore suppression drove rapid attenuation of induced resistance: offspring of plants from insect-suppression plots exhibited a near-complete loss of induced resistance to beetles, whereas those from control plots incurred ~70% less damage after experimental induction. Plants from insect-suppression plots also had ~40% greater constitutive resistance compared with those from control plots, although this difference was not statistically significant. We nonetheless detected a strong trade-off between constitutive and induced resistance across families. In contrast, the constitutive expression of trypsin inhibitors (TI), an important chemical defense trait in horsenettle, was reduced by 20% in the offspring of plants from insect-suppression plots relative to those from control plots. However, TIs were induced to an equal extent whether or not insect herbivores had been historically suppressed. Although several defense and performance traits (prickle density, TI concentration, resistance against false Colorado potato beetles and flea beetles, biomass, and seed mass) varied markedly across families, no traits exhibited significant pairwise correlations. Overall, our results indicate that, whereas the divergent responses of multiple defense traits to insect suppression led to comparatively small changes in overall constitutive resistance, they significantly reduced induced resistance against false Colorado potato beetle.

Herbivore‐induced branching increases sapling survival in temperate forest canopy gaps

Abstract Tree architectures reflect the main abiotic and biotic selection pressures determining tree growth and survival. Studies have shown that trees growing in herbivore‐dominated ecosystems, such as savannas, develop denser, more divaricate ‘cage’‐like architectures in response to chronic browsing pressure (also known as ‘brown world’ architectures). In contrast, trees growing under resource‐limited conditions develop traits that allow them to better compete for abiotic resources (referred to as ‘green world’ architectures); for example, light limitation can induce trees to form more pole‐like structures that allow them to grow faster in height. In temperate forests, abiotic drivers have traditionally been assumed to be the main factors affecting plant architecture. However, gap dynamics typically characterize old‐growth temperate forests, where disturbances such as storms create gaps with temporary high‐light availability, but also attracting intense mammalian browsing. Under such conditions, one would expect tree saplings to exhibit high architectural plasticity enabling them to develop a ‘cage’ in response to high herbivore pressure, but when herbivore pressure declines switch to a more pole‐like structure that allows them to grow quickly in height and compete for light. We tested the ability of five dominant European tree species to induce a ‘cage’ architecture (hereafter referred to as ‘cage plasticity’) under contrasting light and herbivory regimes in a full‐factorial experiment in Białowieża Forest. We planted 720 young tree saplings in six replicated blocks in which we manipulated ungulate browsing (exclosure vs. control) and light levels (gap vs. closed forest). Two out of the five species (hornbeam Carpinus betulus and lime Tilia cordata) were able to develop a cage when exposed to ungulate herbivores but only under high‐light conditions. These two species had the highest survival rates in the herbivory treatment over a 10‐year period. Synthesis. Cage plasticity in temperate forest tree species may reflect an overlooked adaptation to mammal herbivory originating in a past when large mammals exerted strong selection pressures in European ecosystems. Our findings challenge the view of closed‐canopy forest communities being solely driven by abiotic drivers and call for a re‐evaluation of the importance of large herbivores in shaping the functional traits of temperate tree species.

Fire and herbivory drive fungal and bacterial communities through distinct above- and belowground mechanisms

Fire and herbivory are important natural disturbances in grassy biomes. Both drivers are likely to influence belowground microbial communities but no studies have unravelled the long-term impact of both fire and herbivory on bacterial and fungal communities. We hypothesized that soil bacterial communities change through disturbance-induced shifts in soil properties (e.g. pH, nutrients) while soil fungal communities change through vegetation modification (biomass and species composition). To test these ideas, we characterised soil physico-chemical properties (pH, acidity, C, N, P and exchangeable cations content, texture, bulk density, moisture), plant species richness and biomass, microbial biomass and bacterial and fungal community composition and diversity (using 16S and ITS rRNA amplicon sequencing, respectively) in six long-term (18 to 70 years) ecological research sites in South African savanna and grassland ecosystems. We found that fire and herbivory regimes profoundly modified soil physico-chemical properties, plant species richness and standing biomass. In all sites, an increase in woody biomass (ranging from 12 to 50%) was observed when natural disturbances were excluded. The intensity and direction of changes in soil properties were highly dependent on the topo-pedo-climatic context. Overall, fire and herbivory shaped bacterial and fungal communities through distinct driving forces: edaphic properties (including Mg, pH, Ca) for bacteria, and vegetation (herbaceous biomass and woody cover) for fungi. Fire and herbivory explained on average 7.5 and 9.8% of the fungal community variability, respectively, compared to 6.0 and 5.6% for bacteria. The relatively small changes in microbial communities due to natural disturbance is in stark contrast to dramatic vegetation and edaphic changes and suggests that soil microbial communities, having evolved with disturbance, are resistant to change. This represents both a buffer to short-term anthropogenic-induced changes and a restoration challenge in the face of long-term changes.

Complex causes and consequences of rangeland greening in South America – multiple interacting natural and anthropogenic drivers and simultaneous ecosystem degradation and recovery trends

Land-surface greening has been reported globally over the past decades. While often seen to represent ecosystem recovery, the impacts on biodiversity and society can also be negative. Greening has been widely reported from rangelands, where drivers and processes are complex due to its high environmental heterogeneity and societal dynamics. Here, we assess the complexity behind greening and assess its links to various drivers in an iconic, heterogeneous rangeland area, the Iberá Wetlands and surroundings, in Argentina. Time-series satellite imagery over the past 19 years showed overall net greening, but also substantial local browning both in protected and unprotected areas, linking to land use, temporal changes in surface water, fire, and weather. We found substantial woody expansion mainly in the unprotected land, with 37% contributed by tree plantations and the remaining 63% by spontaneous woody expansion, along with widespread transitions from terrestrial land to seasonal surface water. Fire occurrences tended to reduce greening with unprotected areas experiencing widespread and frequent fire. However, protected areas had more browning in unburnt areas than burned areas. Temporal variation in annual precipitation and temperature tended to nonlinearly influence fire occurrences with an interplay of human fire management, further shaping the vegetation greening, pointing to high complexity behind the observed rangeland greening involving interactions among local drivers. Our findings highlight that the observed overall greening is an outcome of multiple trends with clear negative impacts on biodiversity and the local livestock-oriented culture (notably expanding tree plantations) and spontaneous vegetation dynamics, partly involving spontaneous woody expansion. The latter has positive potential for biodiversity and ecosystem services in terms of woodland recovery, but can become negative in such a natural savanna region if expansions develop on a too broad scale, highlighting the importance of ensuring recovery of natural fire and herbivory regimes in protected areas along with sustainable rangeland management elsewhere.

Changes in Planktivory and Herbivory Regimes in a Shallow South American Lake (Lake Blanca Chica, Argentina) Over the Last 250 Years

Shallow lakes are vulnerable ecosystems impacted by human activities and climate change. The Cladocera occupy a central role in food webs and are an excellent paleoecological indicator of food web structure and trophic status. We conducted a paleolimnological study in Lake Blanca Chica (Argentina) to detect changes on the planktivory and herbivory regimes over the last 250 years. Generalized additive models were fitted to the time series of fish predation indicators (ephippial abundance and size, mucrone size, fish scales, and the planktivory index) and pheophorbide a concentration. The cladoceran assemblage changed from littoral-benthic to pelagic species dominance and zooplankton switched from large-bodied (Daphnia) to small-bodied grazers (Bosmina) ca. 1900 due to increased predation. The shift in planktivory regime (ca. 1920–1930), indicated by fish scales and the planktivory index, as well as herbivory (ca. 1920–1950), was triggered by eutrophication. Changes in planktivory affected the size structure of Bosmina, reducing its body size. This study describes the baseline for the lake as well as the profound changes in the composition and size structure of the zooplankton community due to increased predation and the shift in the planktivory regime. These findings will provide a reference status for future management strategies of this ecosystem.

Linking spatial patterns of terrestrial herbivore community structure to trophic interactions.

Large herbivores influence ecosystem functioning via their effects on vegetation at different spatial scales. It is often overlooked that the spatial distribution of large herbivores results from their responses to interacting top-down and bottom-up ecological gradients that create landscape-scale variation in the structure of the entire community. We studied the complexity of these cascading interactions using high-resolution camera trapping and remote sensing data in the best-preserved European lowland forest, Białowieża Forest, Poland. We showed that the variation in spatial distribution of an entire community of large herbivores is explained by species-specific responses to both environmental bottom-up and biotic top-down factors in combination with human-induced (cascading) effects. We decomposed the spatial variation in herbivore community structure and identified functionally distinct landscape-scale herbivory regimes ('herbiscapes'), which are predicted to occur in a variety of ecosystems and could be an important mechanism creating spatial variation in herbivory maintaining vegetation heterogeneity.

Hyper-arid tall shrub species have differing long-term responses to browsing management

Hyper-arid rangeland vegetation is typically dominated by large woody species which are often overlooked in herbivory studies. Long-term responses of tall shrub populations to herbivory change are poorly understood in the Arabian Peninsula. Population and size of 1559 individuals from four shrub species were assessed over an 11-year period under two herbivory regimes, one in which domestic livestock (camels) were replaced by semi-wild ungulates (Oryx and gazelles) before, and the other during, the study period. Each shrub species exhibited a different response to the change in herbivory. Populations of Calotropis procera decreased dramatically. Populations of both Calligonum polygonoides and Lycium shawii increased through sexual reproduction, but the spatial distribution of recruits indicated different modes of seed dispersal. Average lifespans were estimated at 22 and 20 years respectively. The persistence strategy of Leptadenia pyrotechnica was similar to tree species of this habitat in that vegetative regrowth was prioritized over recruitment, and average lifespan was estimated at 95 years. Shrub responses to changes in ungulate management are therefore species-specific. The response of individual plant size was faster than the response of population size, which was limited by slow sexual recruitment (L. pyrotechnica) or localized seed dispersal (C. polygonoides).

Ungulate Herbivory Is Correlated with High Aspen Suckering Density but Reductions in Aspen Growth Rates and Recruitment

Aspen is a foundation species that provides vital habitat for hundreds of plant and animal species. However, changing ungulate herbivory regimes may be altering recruitment success and resilience of aspen forests. The objective of this observational study was to quantify the impacts of ungulate herbivory on aspen recruitment potential. We sampled 36 aspen dominant stands on the Fishlake National Forest in Utah for browse of apical meristems, suckering density (< 180 cm in height), and recruitment density (≥ 180 cm). Our analysis indicated that for each 10% of apical meristems browsed, annual vertical and radial growth rates and recruitment density were reduced by 1.2 cm, 0.02 mm, and 17 ha−1, respectively. In contrast, aspen suckering density increased by 1 230 suckers ha−1, with each 10% browse of apical meristems. This suggests that ungulates contribute to aspen suckers persisting in the understory for longer periods and may alter recruitment success of aspen stands. Aspen that were under high ungulate browse pressure were associated with lower growth rates and persisted for twice as long in the understory. For example, a 5-yr-old sucker was predicted to be 115 cm without browse of apical meristems and 60 cm with 100% browse of apical meristems. Further, aspens of the same height with 100% browse of apical meristems were nearly 4 yr older on average than aspens with no evidence of browse of apical meristems. Approximately 34% of aspen suckers had persisted in the understory < 100 cm for ≥ 6 yr, and 7% of suckers had persisted in the understory for ≥ 10 yr. Our results suggest that high rates of meristem removal are correlated with increased aspen suckering yet reduced aspen growth and recruitment.

Heritable plant phenotypes track light and herbivory levels at fine spatial scales.

Organismal phenotypes often co-vary with environmental variables across broad geographic ranges. Less is known about the extent to which phenotypes match local conditions when multiple biotic and abiotic stressors vary at fine spatial scales. Bittercress (Brassicaceae: Cardamine cordifolia), a perennial forb, grows across a microgeographic mosaic of two contrasting herbivory regimes: high herbivory in meadows (sun habitats) and low herbivory in deeply shaded forest understories (shade habitats). We tested for local phenotypic differentiation in plant size, leaf morphology, and anti-herbivore defense (realized resistance and defensive chemicals, i.e., glucosinolates) across this habitat mosaic through reciprocal transplant-common garden experiments with clonally propagated rhizomes. We found habitat-specific divergence in morphological and defensive phenotypes that manifested as contrasting responses to growth in shade common gardens: weak petiole elongation and attenuated defenses in populations from shade habitats, and strong petiole elongation and elevated defenses in populations from sun habitats. These divergent phenotypes are generally consistent with reciprocal local adaptation: plants from shade habitats that naturally experience low herbivory show reduced investment in defense and an attenuated shade avoidance response, owing to its ineffectiveness within forest understories. By contrast, plants from sun habitats with high herbivory show shade-induced elongation, but no evidence of attenuated defenses canonically associated with elongation in shade-intolerant plant species. Finally, we observed differences in flowering phenology between habitat types that could potentially contribute to inter-habitat divergence by reducing gene flow. This study illuminates how clonally heritable plant phenotypes track a fine-grained mosaic of herbivore pressure and light availability in a native plant.

Modulation of different kelp life stages by herbivory: compensatory growth versus population decimation

Partitioning the effects of herbivory on different life stages of primary producers is key to understanding the population-wide consequences of herbivory. We assessed the performance of microscopic (MiS <1 mm) juveniles, macroscopic (MaS) juveniles and adult kelp (Laminaria ochroleuca) under contrasting herbivory regimes through a herbivore exclusion field experiment. The abundance of MiS and the survival of MaS decreased by 67 and 63%, respectively, when herbivorous fishes and sea urchins were present. Blade growth (linear and area) of adult kelp displayed contrasting patterns under herbivore pressure: a 60% increase and a 46% decrease, respectively. These results indicate that while herbivory severely reduces juvenile survival, it may also induce compensatory growth (measured as linear growth) in adult kelp. In summary, we here demonstrate how herbivory affects all sporophyte life stages of the kelp L. ochroleuca. This is likely to have important implications for situations where historical patterns of herbivore presence and herbivory are changing, such as is increasingly the case in many temperate regions due to warming around the world.

Herbivory impacts of elk, deer and cattle on aspen forest recruitment along gradients of stand composition, topography and climate

The abundance and composition of large ungulate communities are changing dramatically across terrestrial ecosystems due to human activities. This has resulted in novel herbivory regimes that create strong top-down effects on biological communities, and can reduce ecosystem resilience. Biotic and abiotic conditions that influence the population structure and behavior of ungulate herbivores are highly variable across space and time, making it difficult to predict ungulate impacts across complex landscapes. Where multiple ungulate species co-exist, it can be challenging to differentiate species impacts on plant communities. We examined the effects of cattle, American elk, and mule deer on aspen regeneration and recruitment along gradients of forest composition and physiographic conditions in 3 National Forests of Utah, USA. Aspen regeneration and recruitment increased with winter precipitation, elevation and higher abundance of overstory aspen. Binned fecal count densities of elk, mule deer, and cattle showed strong inverse correlations with aspen height (elk: r2=0.83; deer: r2=0.92; cattle: r2=0.35) and aspen recruitment (elk: r2=0.51; deer: r2=0.59). Aspen stem density was negatively correlated with elk (r2=0.35) and cattle fecal densities (r2=0.71). We found that 60% of the sites we surveyed had low ungulate activity and good aspen regeneration, 32% were intermediate use sites that warrant additional monitoring, and 8% of sites showed high ungulate use that exhibited aspen regeneration failure and lack of recruitment where targeted management is recommended. We conclude that ungulate impacts in aspen forests should be considered on a site by site basis and should primarily be evaluated based on the total abundance of the ungulate community with precipitation, elevation and stand composition as secondary factors.

Induced response to herbivory in stinging hair traits of Japanese nettle (Urtica thunbergiana) seedlings in two subpopulations with different browsing pressures by sika deer

Evolutionary theories predict that natural selection favors inducible defense when the risk of predation is unpredictable. In this context, the magnitude of the induced defense in populations experiencing intermittent herbivory is predicted to be larger than that in populations experiencing constant herbivory when there is genetic differentiation between populations. To test this prediction, we conducted a clipping experiment to investigate induced response to shoot damage by the stinging hair traits of Japanese nettle (Urtica thunbergiana) seedlings. For this purpose, we studied two nettle subpopulations, one under constant browsing and another under intermittent browsing by sika deer in Nara Park, central Japan. The clipping experiment demonstrates that both subpopulations exhibited induced defenses in response to the clipping of the shoot apex as the number and length of stinging hairs increased after clipping. The subpopulation experiencing intermittent browsing exhibited smaller trait values and larger induced defenses, indicated by the number of stinging hairs on the upper leaf surface and the length of stinging hairs on both leaf surfaces compared with the subpopulation experiencing constant browsing. These results are consistent with the prediction and suggest that genetic differentiation of the induced defense between subpopulations is caused by adaptation to the herbivory regime. We discuss other plausible factors affecting the magnitude of the induced defense of the nettle subpopulations.

Large herbivores promote habitat specialization and beta diversity of African savanna trees.

Edaphic variation in plant community composition is widespread, yet its underlying mechanisms are rarely understood and often assumed to be physiological. In East African savannas, Acacia tree species segregate sharply across soils of differing parent material: the ant-defended whistling thorn, A.drepanolobium (ACDR), is monodominant on cracking clay vertisols that are nutrient rich but physically stressful, whereas poorly defended species such as A.brevispica (ACBR) dominate on nutrient-poor but otherwise less-stressful sandy loams. Using a series of field experiments, we show that large-mammal herbivory interacts with soil properties to maintain this pattern. In the absence of large herbivores, transplanted saplings of both species established on both soil types. Browsers strongly suppressed survival and growth of ACDR saplings on sandy soil, where resource limitation constrained defensive investment. On clay soil, ACBR saplings established regardless of herbivory regime, but elephants prevented recruitment to maturity, apparently because trees could not tolerate the combination of biotic and abiotic stressors. Hence, each tree species was filtered out of one habitat by browsing in conjunction with different edaphic factors and at different ontogenetic stages. Browser abundance was greater on sandy soil, where trees were less defended, consistent with predicted feedbacks between plant community assembly and herbivore distributions. By exploring two inversely related axes of soil "quality" (abiotic stress and nutrient content), our study extends the range of mechanisms by which herbivores are known to promote edaphic specialization, illustrates how the high cost of a protection mutualism can constrain the realized niche of host trees, and shows that large-scale properties of savanna ecosystems are shaped by species interactions in cryptic ways that mimic simple abiotic determinism. These results suggest that ongoing declines in large-herbivore populations may relax spatial heterogeneity in plant assemblages and reduce the beta diversity of communities.

Palaeoecological signatures of vegetation change induced by herbivory regime shifts on subantarctic Enderby Island

The stratigraphic relationships of palaeoecological proxies and use of changepoint analyses to determine the cause and effect relationships between past events has allowed a better understanding of the relative contributions of humans and environmental drivers to Late Quaternary extinctions and of their effects on terrestrial ecosystems. Few studies, however, have validated these approaches at localities where past interactions between vegetation communities and large herbivores are well-documented. Here, we use a peat core from subantarctic Enderby Island to present the first study tracing the spores of dung fungi alongside pollen at a site where the history of mammalian herbivore introductions (and subsequent eradication), and their effects on the vegetation, are precisely known. We find a strong connection between spore influx rates of the dung-fungus Sporormiella and PCA axis 1 of the pollen assemblages, suggesting that past vegetation change caused by herbivore introductions and eradications at the core site can be readily deduced from the palaeoecological record. The response of the vegetation community to the removal of herbivores was so rapid, however, that a difference in timing between changepoints relating to specific pollen taxa, the overall pollen community, and the decline of Sporormiella spores, could not be resolved in our record, despite a sampling resolution of <5 years. We suggest that further case-studies, spanning different vegetation and herbivore communities, are required to provide increased confidence in inferences drawn about cause-and-effect relationships using proxy changepoint offsets in palaeoecological records.

A continent-wide assessment of the form and intensity of large mammal herbivory in Africa.

Megafaunal extinctions and a lack of suitable remote sensing technology impede our understanding of both the ecological legacy and current impacts of large mammal herbivores in the Earth system. To address this, we reconstructed the form and intensity of herbivory pressure across sub-Saharan Africa ~1000 years ago. Specifically, we modeled and mapped species-level biomass for 92 large mammal herbivores using census data, species distributions, and environmental covariates. Trait-based classifications of these species into herbivore functional types, and analyses of their biomass surfaces, reveal four ecologically distinct continental-scale herbivory regimes, characterized by internally similar forms and intensities of herbivory pressure. Associations between herbivory regimes, fire prevalence, soil nutrient status, and rainfall provide important insights into African ecology and pave the way for integrating herbivores into global-scale studies.

How herbivores affect ecosystems

Ecology Abiotic forces, such as fire and water, have powerful effects on ecosystem structure and function. Animals that eat plants also have strong effects in natural systems, but their impacts are harder to assess. Hempson et al. measured how herbivores affect vegetation across Africa (see the Perspective by Gill). Four distinct herbivory regimes emerge from the analysis, characterized by forest antelopes, arid-region gazelles, high-diversity savannah fauna, and bulk feeders (such as elephants), which have had equivalent impact to those of fire and water on shaping ecosystems. Science , this issue p. [1056][1]; see also p. [1036][2] [1]: /lookup/doi/10.1126/science.aac7978 [2]: /lookup/doi/10.1126/science.aad6760

Spinescent East African savannah acacias also have thick bark, suggesting they evolved under both an intense fire and herbivory regime

Spinescent East African savannah acacias also have thick bark, suggesting they evolved under both an intense fire and herbivory regime

Non-native megaherbivores: the case for novel function to manage plant invasions on islands.

There is a heated debate about whether all non-native species are 'guilty until proven innocent', or whether some should be accepted or even welcomed. Further fanning the flames, I here present a case where introductions of carefully vetted, non-native species could provide a net conservation benefit. On many islands, native megaherbivores (flightless birds, tortoises) recently went extinct. Here, rewilding with carefully selected non-native species as ecological replacements is increasingly considered a solution, reinstating a herbivory regime that largely benefits the native flora. Based on these efforts, I suggest that restoration practitioners working on islands without a history of native megaherbivores that are threatened by invasive plants should consider introducing a non-native island megaherbivore, and that large and giant tortoises are ideal candidates. Such tortoises would be equally useful on islands where eradication of invasive mammals has led to increased problems with invasive plants, or on islands that never had introduced mammalian herbivores, but where invasive plants are a problem. My proposal may seem radical, but the reversibility of using giant tortoises means that nothing is lost from trying, and that indeed much is to be gained. As an easily regulated adaptive management tool, it represents an innovative, hypothesis-driven 'innocent until proven guilty' approach.