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Whether wild herbivores confer biotic resistance to invasion by exotic plants remains a key question in ecology. There is evidence that wild herbivores can impede invasion by exotic plants, but it is unclear whether and how this generalises across ecosystems with varying wild herbivore diversity and functional groups of plants, particularly over long-term (decadal) time frames. Using data from three long-term (13- to 26-year) exclosure experiments in central Kenya, we tested the effects of wild herbivores on the density of exotic invasive cacti, Opuntia stricta and O. ficus-indica (collectively, Opuntia), which are among the worst invasive species globally. We also examined relationships between wild herbivore richness and elephant occurrence probability with the probability of O. stricta presence at the landscape level (6150 km2). Opuntia densities were 74% to 99% lower in almost all plots accessible to wild herbivores compared to exclosure plots. Opuntia densities also increased more rapidly across time in plots excluding wild herbivores. These effects were largely driven by megaherbivores (≥1000 kg), particularly elephants. At the landscape level, modelled Opuntia stricta occurrence probability was negatively correlated with estimated species richness of wild herbivores and elephant occurrence probability. On average, O. stricta occurrence probability fell from ~0.56 to ~0.45 as wild herbivore richness increased from 6 to 10 species and fell from ~0.57 to ~0.40 as elephant occurrence probability increased from ~0.41 to ~0.84. These multi-scale results suggest that any facilitative effects of Opuntia by wild herbivores (e.g. seed/vegetative dispersal) are overridden by suppression (e.g. consumption, uprooting, trampling). Synthesis. Our experimental and observational findings that wild herbivores confer resistance to invasion by exotic cacti add to evidence that conserving and restoring native herbivore assemblages (particularly megaherbivores) can increase community resistance to plant invasions.

Mutualisms between plants and ants are common features of tropical ecosystems around the globe and can have cascading effects on interactions with the ecological communities in which they occur. In an African savanna, we assessed whether acacia ants influence nest site selection by tree-nesting birds. Birds selected nest sites in trees inhabited by ant species that vigorously defend against browsing mammals. Future research could address the extent to which hatching and fledging rates depend on the species of ant symbiont, and why ants tolerate nesting birds but not other tree associates (especially insects).

Cooperative interactions may frequently be reinforced by "partner fidelity feedback," in which high- or low-quality partners drive positive feedbacks with high or low benefits for the host, respectively. Benefits of plant-animal mutualisms for plants have been quantified almost universally in terms of growth or reproduction, but these are only two of many sinks to which a host-plant allocates its resources. By investigating how partners to host-plants impact two fundamental plant resources, carbon and water, we can better characterize plant-partner fidelity and understand how plant-partner mutualisms may be modulated by resource dynamics. In Laikipia, Kenya, four ant species compete for Acacia drepanolobium host-plants. These ants differ in multiple traits, from nectar consumption to host-plant protection. Using a 5-year ant removal experiment, we compared carbon fixation, leaf water status, and stem non-structural carbohydrate concentrations for adult ant-plants with and without ant partners. Removal treatments showed that the ants differentially mediate tree carbon and/or water resources. All three ant species known to be aggressive against herbivores were linked to benefits for host-plant resources, but only the two species that defend but do not prune the host, Crematogaster mimosae and Tetraponera penzigi, increased tree carbon fixation. Of these two species, only the nectivore C. mimosae increased tree simple sugars. Crematogaster nigriceps, which defends the tree but also castrates flowers and prunes meristems, was linked only to lower tree water stress approximated by pre-dawn leaf water potential. In contrast to those defensive ants, Crematogaster sjostedti, a poor defender that displaces other ants, was linked to lower tree carbon fixation. Comparing the effects of the four ant species across control trees suggests that differential ant occupancy drives substantial differences in carbon and water supply among host trees. Our results highlight that ant partners can positively or negatively impact carbon and/or water relations for their host-plant, and we discuss the likelihood that carbon- and water-related partner fidelity feedback loops occur across ant-plant mutualisms.

Minirhizotron imagery can be used to assess plant root health, and the amount of data for analysis motivates automation of root detection through use of neural networks. Building upon previous work, we show that we can use transfer learning from our PRMI dataset to assess root health across twelve classes of a new dataset to answer questions regarding how root health is affected by access of a tree by large herbivores, site infestation by Pheidole megacephala, and location of the tree. This dataset was collected from three paired sites at the Ol Pejeta Conservancy in Laikipia, Kenya, and consists of 20,000 images collected between September 2021 and May 2022. Each paired site represents four locations based on all four possible combinations of site infestation by Pheidole megacephala for at least 20 years and existence of herbivore-exclusion fence to keep large herbivores out. 1,332 images across all twelve classes of site and treatment combination were labeled with respective ground truths for model training. Our work uses the UNet architecture using pretrained weights on the network encoder and decoder which were obtained in 2019 in work which achieved over 99% accuracy on a dataset of peanut and switchgrass imagery. In our work, we found that training the model with our new dataset resulted in consistent performance across all classes of our new dataset, with over 99% accuracy for each class.

Invasive ants shape assemblages and interactions of native species, but their effect on fundamental ecological processes is poorly understood. In East Africa, Pheidole megacephala ants have invaded monodominant stands of the ant-tree Acacia drepanolobium, extirpating native ant defenders and rendering trees vulnerable to canopy damage by vertebrate herbivores. We used experiments and observations to quantify direct and interactive effects of invasive ants and large herbivores on A. drepanolobium photosynthesis over a 2-year period. Trees that had been invaded for ≥ 5 years exhibited 69% lower whole-tree photosynthesis during key growing seasons, resulting from interaction between invasive ants and vertebrate herbivores that caused leaf- and canopy-level photosynthesis declines. We also surveyed trees shortly before and after invasion, finding that recent invasion induced only minor changes in leaf physiology. Our results from individual trees likely scale up, highlighting the potential of invasive species to alter ecosystem-level carbon fixation and other biogeochemical cycles.

Supercolonial ants are among the largest cooperative units in nature, attaining extremely high densities. How these densities feed back into their population growth rates and how abundance and extrinsic factors interact to affect their population dynamics remain open questions. We studied how local worker abundance and extrinsic factors (rain, tree density) affect population growth rate and spread in the invasive big-headed ant, which is disrupting a keystone mutualism between acacia trees and native ants in parts of East Africa. We measured temporal changes in big-headed ant (BHA) abundance and rates of spread over 20 months along eight transects, extending from areas behind the front with high BHA abundances to areas at the invasion front with low BHA abundances. We used models that account for negative density dependence and incorporated extrinsic factors to determine what variables best explain variation in local population growth rates. Population growth rates declined with abundance, however, the strength of density dependence decreased with abundance. We suggest that weaker density dependence at higher ant abundances may be due to the beneficial effect of cooperative behavior that partially counteracts resource limitation. Rainfall and tree density had minor effects on ant population dynamics. BHA spread near 50 m/year, more than previous studies reported and comparable to rates of spread of other supercolonial ants. Although we did not detect declines in abundance in areas invaded a long time ago (> 10 years), continued monitoring of abundance at invaded sites may help to better understand the widespread collapse of many invasive ants.

Quantifying tree biomass is an important research and management goal across many disciplines. For species that exhibit predictable relationships between structural metrics (e.g. diameter, height, crown breadth) and total weight, allometric calculations produce accurate estimates of above-ground biomass. However, such methods may be insufficient where inter-individual variation is large relative to individual biomass and is itself of interest (for example, variation due to herbivory). In an East African savanna bushland, we analysed photographs of small (2 = 0.86 versus R2 = 0.68). When tested on trees in herbivore-exclusion plots versus unfenced (open) plots, our predictive equation based on photos confirmed higher above-ground biomass in the exclusion plots than in unfenced (open) plots (P < 0.001), in contrast to no significant difference based on the allometric equation (P = 0.43). As such, our new technique based on photographs offers an accurate and cost-effective complement to existing methods for tree biomass estimation at small scales with potential application across a wide variety of settings.

Across the globe, biological invasions have disrupted mutualisms, producing reverberating consequences for ecosystems. Although invasive species frequently trigger mutualism disruptions, few studies have quantified the demographic mechanisms by which mutualism breakdown may generate population effects. In a Kenyan savanna, the invasive big-headed ant (Pheidole megacephala) has disrupted a foundational mutualism between the monodominant whistling-thorn tree (Acacia drepanolobium) and native ants (Crematogaster spp.) that deter browsing by large mammalian herbivores. We conducted experiments to quantify the demographic consequences of this mutualism disruption in the presence and absence of large mammalian herbivores. Invasion by P. megacephala exacerbated population declines of A. drepanolobium, primarily through decreased survival and reproduction of adult trees. However, these fitness reductions were small compared to those resulting from the presence of large mammalian herbivores, which negatively impacted growth and survival. Contrary to expectation, the expulsion of metabolically costly Crematogaster mutualists by P. megacephala did not result in higher population growth rates for trees protected from large mammalian herbivores. Our results suggest that invasive P. megacephala may impose a direct metabolic cost to trees exceeding that of native mutualists while providing no protection from browsing by large mammalian herbivores. Across landscapes, we expect that invasion by P. megacephala will reduce A. drepanolobium populations, but that the magnitude and demographic pathways of this effect will hinge on the presence and abundance of browsers.

Diverse communities of large mammalian herbivores (LMH), once widespread, are now rare. LMH exert strong direct and indirect effects on community structure and ecosystem functions, and measuring these effects is important for testing ecological theory and for understanding past, current, and future environmental change. This in turn requires long-term experimental manipulations, owing to the slow and often nonlinear responses of populations and assemblages to LMH removal. Moreover, the effects of particular species or body-size classes within diverse LMH guilds are difficult to pinpoint, and the magnitude and even direction of these effects often depends on environmental context. Since 2008, we have maintained the Ungulate Herbivory Under Rainfall Uncertainty (UHURU) experiment, a series of size-selective LMH exclosures replicated across a rainfall/productivity gradient in a semiarid Kenyan savanna. The goals of the UHURU experiment are to measure the effects of removing successively smaller size classes of LMH (mimicking the process of size-biased extirpation) and to establish how these effects are shaped by spatial and temporal variation in rainfall. The UHURU experiment comprises three LMH-exclusion treatments and an unfenced control, applied to nine randomized blocks of contiguous 1-ha plots (n = 36). The fenced treatments are MEGA (exclusion of megaherbivores, elephant and giraffe), MESO (exclusion of herbivores ≥40 kg), and TOTAL (exclusion of herbivores ≥5 kg). Each block is replicated three times at three sites across the 20-km rainfall gradient, which has fluctuated over the course of the experiment. The first 5 years of data were published previously (Ecological Archives E095-064) and have been used in numerous studies. Since that publication, we have (1) continued to collect data following the original protocols, (2) improved the taxonomic resolution and accuracy of plant and small-mammal identifications, and (3) begun collecting several new data sets. Here, we present updated and extended raw data from the first 12 years of the UHURU experiment (2008-2019). Data include daily rainfall data throughout the experiment; annual surveys of understory plant communities; annual censuses of woody-plant communities; annual measurements of individually tagged woody plants; monthly monitoring of flowering and fruiting phenology; every-other-month small-mammal mark-recapture data; and quarterly large-mammal dung surveys. There are no copyright restrictions; notification of when and how data are used is appreciated and users of UHURU data should cite this data paper when using the data.

Environmental circumstances shaping soil microbial communities have been studied extensively, but due to disparate study designs it has been difficult to resolve whether a globally consistent set of predictors exists, or context-dependency prevails. Here, we used a network of 18 grassland sites (11 sampled across regional plant productivity gradients) to examine i) if the same abiotic or biotic factors predict both large- and regional-scale patterns in bacterial and fungal community composition, and ii) if microbial community composition differs consistently with regional plant productivity (low vs high) across different sites. We found that there is high congruence between predictors of microbial community composition across spatial scales; bacteria were predominantly associated with soil properties and fungi with plant community composition. Moreover, there was a microbial community signal that clearly distinguished high and low productivity soils that was shared across worldwide distributed grasslands suggesting that microbial assemblages vary predictably depending on grassland productivity.

Cascading effects of high trophic levels onto lower trophic levels have been documented in many ecosystems. Some studies also show evidence of extended trophic cascades, in which guilds dependent on lower trophic levels, but uninvolved in the trophic cascade themselves, are affected by the trophic cascade due to their dependence on lower trophic levels. Top-down effects of large mammals on plants could lead to a variety of extended trophic cascades on the many guilds dependent on plants, such as pollinators. In this study, floral-visitor and floral abundances and assemblages were quantified within a series of 1-ha manipulations of large-mammalian herbivore density in an African savanna. Top-down effects of large mammals on the composition of flowers available for floral visitors are first shown, using regressions of herbivore activity on metrics of floral and floral-visitor assemblages. An extended trophic cascade is also shown: the floral assemblage further altered the assemblage of floral visitors, according to a variety of approaches, including a structural equation modelling approach (model with an extended trophic cascade was supported over a model without, AICc weight = 0.984). Our study provides support for extended trophic cascades affecting floral visitors, suggesting that trophic cascades can have impacts throughout entire communities.

Biological invasions are both a pressing environmental challenge and an opportunity to investigate fundamental ecological processes, such as the role of top predators in regulating biodiversity and food-web structure. In whole-ecosystem manipulations of small Caribbean islands on which brown anole lizards (Anolis sagrei) were the native top predator, we experimentally staged invasions by competitors (green anoles, Anolis smaragdinus) and/or new top predators (curly-tailed lizards, Leiocephalus carinatus). We show that curly-tailed lizards destabilized the coexistence of competing prey species, contrary to the classic idea of keystone predation. Fear-driven avoidance of predators collapsed the spatial and dietary niche structure that otherwise stabilized coexistence, which intensified interspecific competition within predator-free refuges and contributed to the extinction of green-anole populations on two islands. Moreover, whereas adding either green anoles or curly-tailed lizards lengthened food chains on the islands, adding both species reversed this effect—in part because the apex predators were trophic omnivores. Our results underscore the importance of top-down control in ecological communities, but show that its outcomes depend on prey behaviour, spatial structure, and omnivory. Diversity-enhancing effects of top predators cannot be assumed, and non-consumptive effects of predation risk may be a widespread constraint on species coexistence. Whole-ecosystem manipulations of Caribbean islands occupied by brown anoles, involving the addition of competitors (green anoles) and/or top predators (curly-tailed lizards), demonstrate that predator introductions can alter the ecological niches and destabilize the coexistence of competing prey species.

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 (

The extinction of 80% of megaherbivore (>1,000 kg) species towards the end of the Pleistocene altered vegetation structure, fire dynamics and nutrient cycling world-wide. Ecologists have proposed (re)introducing megaherbivores or their ecological analogues to restore lost ecosystem functions and reinforce extant but declining megaherbivore populations. However, the effects of megaherbivores on smaller herbivores are poorly understood. We used long-term exclusion experiments and multispecies hierarchical models fitted to dung counts to test (a) the effect of megaherbivores (elephant and giraffe) on the occurrence (dung presence) and use intensity (dung pile density) of mesoherbivores (2–1,000 kg), and (b) the extent to which the responses of each mesoherbivore species was predictable based on their traits (diet and shoulder height) and phylogenetic relatedness. Megaherbivores increased the predicted occurrence and use intensity of zebras but reduced the occurrence and use intensity of several other mesoherbivore species. The negative effect of megaherbivores on mesoherbivore occurrence was stronger for shorter species, regardless of diet or relatedness. Megaherbivores substantially reduced the expected total use intensity (i.e. cumulative dung density of all species) of mesoherbivores, but only minimally reduced the expected species richness (i.e. cumulative predicted occurrence probabilities of all species) of mesoherbivores (by

Summary Pollination by animals is a key ecosystem service 1 , 2 and interactions between plants and their pollinators are a model system for studying ecological networks, 3 , 4 yet plant-pollinator networks are typically studied in isolation from the broader ecosystems in which they are embedded. The plants visited by pollinators also interact with other consumer guilds that eat stems, leaves, fruits, or seeds. One such guild, large mammalian herbivores, are well-known ecosystem engineers 5 , 6 , 7 and may have substantial impacts on plant-pollinator networks. Although moderate herbivory can sometimes promote plant diversity, 8 potentially benefiting pollinators, large herbivores might alternatively reduce resource availability for pollinators by consuming flowers, 9 reducing plant density, 10 and promoting somatic regrowth over reproduction. 11 The direction and magnitude of such effects may hinge on abiotic context—in particular, rainfall, which modulates the effects of ungulates on vegetation. 12 Using a long-term, large-scale experiment replicated across a rainfall gradient in central Kenya, we show that a diverse assemblage of native large herbivores, ranging from 5-kg antelopes to 4,000-kg African elephants, limited resource availability for pollinators by reducing flower abundance and diversity; this in turn resulted in fewer pollinator visits and lower pollinator diversity. Exclusion of large herbivores increased floral-resource abundance and pollinator-assemblage diversity, rendering plant-pollinator networks larger, more functionally redundant, and less vulnerable to pollinator extinction. Our results show that species extrinsic to plant-pollinator interactions can indirectly and strongly alter network structure. Forecasting the effects of environmental change on pollination services and interaction webs more broadly will require accounting for the effects of extrinsic keystone species.

Biological invasions can lead to the reassembly of communities and understanding and predicting the impacts of exotic species on community structure and functioning are a key challenge in ecology. We investigated the impact of a predatory species of invasive ant, Pheidole megacephala, on the structure and function of a foundational mutualism between Acacia drepanolobium and its associated acacia-ant community in an East African savanna. Invasion by P. megacephala was associated with the extirpation of three extrafloral nectar-dependent Crematogaster acacia ant species and strong increases in the abundance of a competitively subordinate and locally rare acacia ant species, Tetraponera penzigi, which does not depend on host plant nectar. Using a combination of long-term monitoring of invasion dynamics, observations and experiments, we demonstrate that P. megacephala directly and indirectly facilitates T. penzigi by reducing the abundance of T. penzigi's competitors (Crematogaster spp.), imposing recruitment limitation on these competitors, and generating a landscape of low-reward host plants that favor colonization and establishment by the strongly dispersing T. penzigi. Seasonal variation in use of host plants by P. megacephala may further increase the persistence of T. penzigi colonies in invaded habitat. The persistence of the T. penzigi-A. drepanolobium symbiosis in invaded areas afforded host plants some protection against herbivory by elephants (Loxodonta africana), a key browser that reduces tree cover. However, elephant damage on T. penzigi-occupied trees was higher in invaded than in uninvaded areas, likely owing to reduced T. penzigi colony size in invaded habitats. Our results reveal the mechanisms underlying the disruption of this mutualism and suggest that P. megacephala invasion may drive long-term declines in tree cover, despite the partial persistence of the ant-acacia symbiosis in invaded areas.

Summary Large mammalian herbivores (LMH) strongly shape the composition and architecture of plant communities. A growing literature shows that negative direct effects of LMH on vegetation frequently propagate to suppress the abundance of smaller consumers. Indirect effects of LMH on the behaviour of these consumers, however, have received comparatively little attention despite their potential ecological significance. We sought to understand (i) how LMH indirectly shape small-mammal habitat use by altering the density and distribution of understory plants; (ii) how these effects vary with climatic context (here, seasonality in rainfall); and (iii) the extent to which behavioural responses of small mammals are contingent upon small-mammal density. We tested the effects of a diverse LMH community on small-mammal habitat use using four years of spatially explicit small-mammal-trapping and vegetation data from the UHURU Experiment, a replicated set of LMH exclosures in semi-arid Kenyan savanna. Small-mammal habitat use was positively associated with tree density and negatively associated with bare (unvegetated) patches in all plots and seasons. In the presence of LMH, and especially during the dry season, small mammals consistently selected tree cover and avoided bare patches. In contrast, when LMH were excluded, small mammals were weakly associated with tree cover and did not avoid bare patches as strongly. These behavioural responses of small mammals were largely unaffected by changes in small-mammal density associated with LMH exclusion. Our results show that LMH indirectly affect small-mammal behaviour, and that these effects are influenced by climate and can arise via density-independent mechanisms. This raises the possibility that anthropogenic LMH declines might interact with changing patterns of rainfall to alter small-mammal distribution and behaviour, independent of numerical responses by small mammals to these perturbations. For example, increased rainfall in East Africa (as predicted in many recent climate-model simulations) may relax constraints on small-mammal distribution where LMH are rare or absent, whereas increased aridity and/or drought frequency may tighten them. This article is protected by copyright. All rights reserved.

Supercolonial ants are among the largest cooperative units in nature, attaining extremely high densities. How these densities feed back into their population growth rates and how abundance and extrinsic factors interact to affect their population dynamics remain open questions. We studied how local worker abundance and extrinsic factors (rain, tree density) affect population growth rate and spread in the invasive big-headed ant, which is disrupting a keystone mutualism between acacia trees and native ants in parts of East Africa. We measured temporal changes in big-headed ant (BHA) abundance and rates of spread over 20 months along eight transects, extending from areas behind the front with high BHA abundances to areas at the invasion front with low BHA abundances. We used models that account for negative density dependence and incorporated extrinsic factors to determine what variables best explain variation in local population growth rates. Population growth rates declined with abundance, however, the strength of density dependence decreased with abundance. We suggest that weaker density dependence at higher ant abundances may be due to the beneficial effect of cooperative behavior that partially counteracts resource limitation. Rainfall and tree density had minor effects on ant population dynamics. BHA spread near 50 m/year, more than previous studies reported and comparable to rates of spread of other supercolonial ants. Although we did not detect declines in abundance in areas invaded a long time ago ( 10 years), continued monitoring of abundance at invaded sites may help to better understand the widespread collapse of many invasive ants.

Founder populations often show rapid divergence from source populations after colonizing new environments. Epigenetic modifications can mediate phenotypic responses to environmental change and may be an important mechanism promoting rapid differentiation in founder populations. Whereas many long-term studies have explored the extent to which divergence between source and founder populations is genetically heritable versus plastic, the role of epigenetic processes during colonization remains unclear. To investigate epigenetic modifications in founding populations, we experimentally colonized eight small Caribbean islands with brown anole lizards (Anolis sagrei) from a common source population. We then quantitatively measured genome-wide DNA methylation in liver tissue using reduced representation bisulfite sequencing of individuals transplanted onto islands with high- versus low-habitat quality. We found that lizard sex and habitat quality explained a significant proportion of epigenetic variation. Differentially methylated cytosines mapped to genes that encode proteins with functions likely to be relevant to habitat change (e.g., signal transduction, immune response, circadian rhythm). This study provides experimental evidence of a relationship between epigenetic responses and the earliest stages of colonization of novel environments in nature and suggests that habitat quality influences the nature of these epigenetic modifications.

The role of arbuscular mycorrhizal fungi (AMF) in enhancing plant tolerance to drought is well known. However, the degree to which AMF-plant symbioses are locally adapted has been suggested but is less well understood, especially at small spatial scales. Here, we examined the effects of two arbuscular mycorrhizal fungal communities on drought tolerance of Themeda triandra, a native African perennial bunchgrass. In our study area, mound building activities of Odontotermes sp. termites produce heterogeneous habitat, particularly with respect to water availability, and do so over small spatial scales (

Positive indirect effects of consumers on their resources can stabilize food webs by preventing overexploitation, but the coupling of trophic and non-trophic interactions remains poorly integrated into our understanding of community dynamics. Elephants engineer African savanna ecosystems by toppling trees and breaking branches, and although their negative effects on trees are well documented, their effects on small-statured plants remain poorly understood. Using data on 117 understory plant taxa collected over 7 yr within 36 1-ha experimental plots in a semi-arid Kenyan savanna, we measured the strength and direction of elephant impacts on understory vegetation. We found that elephants had neutral effects on most (83-89%) species, with a similar frequency of positive and negative responses among the remainder. Overall, estimated understory biomass was 5-14% greater in the presence of elephants across a range of rainfall levels. Whereas direct consumption likely accounts for the negative effects, positive effects are presumably indirect. We hypothesized that elephants create associational refuges for understory plants by damaging tree canopies in ways that physically inhibit feeding by other large herbivores. As predicted, understory biomass and species richness beneath elephant-damaged trees were 55% and 21% greater, respectively, than under undamaged trees. Experimentally simulated elephant damage increased understory biomass by 37% and species richness by 49% after 1 yr. Conversely, experimentally removing elephant damaged branches decreased understory biomass by 39% and richness by 30% relative to sham-manipulated trees. Camera-trap surveys revealed that elephant damage reduced the frequency of herbivory by 71%, whereas we detected no significant effect of damage on temperature, light, or soil moisture. We conclude that elephants locally facilitate understory plants by creating refuges from herbivory, which countervails the direct negative effects of consumption and enhances larger-scale biomass and diversity by promoting the persistence of rare and palatable species. Our results offer a counterpoint to concerns about the deleterious impacts of elephant "overpopulation" that should be considered in debates over wildlife management in African protected areas: understory species comprise the bulk of savanna plant biodiversity, and their responses to elephants are buffered by the interplay of opposing consumptive and non-consumptive interactions.

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.

Herbivores alter plant biodiversity (species richness) in many of the world’s ecosystems, but the magnitude and the direction of herbivore effects on biodiversity vary widely within and among ecosystems. One current theory predicts that herbivores enhance plant biodiversity at high productivity but have the opposite effect at low productivity. Yet, empirical support for the importance of site productivity as a mediator of these herbivore impacts is equivocal. Here, we synthesize data from 252 large-herbivore exclusion studies, spanning a 20-fold range in site productivity, to test an alternative hypothesis—that herbivore-induced changes in the competitive environment determine the response of plant biodiversity to herbivory irrespective of productivity. Under this hypothesis, when herbivores reduce the abundance (biomass, cover) of dominant species (for example, because the dominant plant is palatable), additional resources become available to support new species, thereby increasing biodiversity. By contrast, if herbivores promote high dominance by increasing the abundance of herbivory-resistant, unpalatable species, then resource availability for other species decreases reducing biodiversity. We show that herbivore-induced change in dominance, independent of site productivity or precipitation (a proxy for productivity), is the best predictor of herbivore effects on biodiversity in grassland and savannah sites. Given that most herbaceous ecosystems are dominated by one or a few species, altering the competitive environment via herbivores or by other means may be an effective strategy for conserving biodiversity in grasslands and savannahs globally. Fil: Koerner, Sally E.. University of North Carolina at Greensboro. Department of Biology; Estados Unidos Fil: Smith, Melinda D.. State University of Colorado - Fort Collins; Estados Unidos Fil: Burkepile, Deron E.. University of California; Estados Unidos Fil: Hanan, Niall P.. New Mexico State University; Estados Unidos Fil: Avolio, Meghan L.. University Johns Hopkins; Estados Unidos Fil: Collins, Scott L.. University of New Mexico; Estados Unidos Fil: Knapp, Alan K.. State University of Colorado - Fort Collins; Estados Unidos Fil: Lemoine, Nathan P.. State University of Colorado - Fort Collins; Estados Unidos Fil: Forrestel, Elisabeth J.. University of California at Davis; Estados Unidos Fil: Eby, Stephanie. Northeastern University; Estados Unidos Fil: Thompson, Dave I.. University of the Witwatersrand; Sudáfrica. Kruger National Park. South African Environmental Observation Network. Ndlovu Node. Scientific Services; Sudáfrica Fil: Aguado-Santacruz, Gerardo A.. Tecnológico Nacional de México; México Fil: Anderson, John P.. New Mexico State University; Estados Unidos Fil: Anderson, T. Michael. University Wake Forest; Estados Unidos Fil: Angassa, Ayana. Botswana University of Agriculture and Natural Resources; Sudáfrica. Hawassa University; Etiopía Fil: Bagchi, Sumanta. Indian Institute of Science; India Fil: Bakker, Elisabeth S.. Netherlands Institute of Ecology; Países Bajos Fil: Bastin, Gary. Alice Springs; Australia Fil: Baur, Lauren E.. University of New Mexico; Estados Unidos Fil: Beard, Karen H.. State University of Utah; Estados Unidos Fil: Beever, Erik A.. United States Geological Survey; Estados Unidos. State University of Montana; Estados Unidos Fil: Bohlen, Patrick J.. University of Central Florida; Estados Unidos Fil: Boughton, Elizabeth H.. MacArthur Agroecology Research Center; Estados Unidos Fil: Canestro, Don. University of California; Estados Unidos Fil: Cesa, Ariela. Instituto Nacional de Tecnología Agropecuaria. Centro Regional Buenos Aires; Argentina Fil: Chaneton, Enrique Jose. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura. Universidad de Buenos Aires. Facultad de Agronomía. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura; Argentina Fil: Cheng, Jimin. Northwest A&F University; China Fil: D?Antonio, Carla M.. University of California; Estados Unidos Fil: Deleglise, Claire. Université Grenoble Alpes; Francia Fil: Dembélé, Fadiala. Institut de Formation et de Recherche Appliquee; Sudáfrica Fil: Dorrough, Josh. Ecosystem Mangement Science; Austria Fil: Eldridge, David J.. University of New South Wales; Australia Fil: Fernandez Going, Barbara. University of Texas at Austin; Estados Unidos Fil: Fernández Lugo, Silvia. Universidad de La Laguna; España Fil: Fraser, Lauchlan H.. Thompson Rivers University; Canadá Fil: Freedman, Bill. Dalhousie University Halifax; Canadá Fil: García Salgado, Gonzalo. Dalhousie University Halifax; Canadá Fil: Goheen, Jacob R.. University of Wyoming; Estados Unidos Fil: Guo, Liang. Northwest A&F University; China Fil: Husheer, Sean. New Zealand Forest Surveys; Nueva Zelanda Fil: Karembé, Moussa. Université des Sciences; Malí Fil: Knops, Johannes M. H.. Universidad de Nebraska - Lincoln; Estados Unidos Fil: Kraaij, Tineke. Nelson Mandela University; Sudáfrica Fil: Kulmatiski, Andrew. State University of Utah; Estados Unidos Fil: Kytöviita, Minna Maarit. Universidad de Jyvaskyla; Finlandia Fil: Lezama, Felipe. Universidad de la República; Uruguay Fil: Loucougaray, Gregory. Université Grenoble Alpes; Francia Fil: Loydi, Alejandro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Bahía Blanca. Centro de Recursos Naturales Renovables de la Zona Semiárida. Universidad Nacional del Sur. Centro de Recursos Naturales Renovables de la Zona Semiárida; Argentina Fil: Milchunas, Dan G.. State University of Colorado - Fort Collins; Estados Unidos Fil: Milton, Suzanne J.. South African Environmental Observation Network; Sudáfrica Fil: Morgan, John W.. La Trobe University; Australia Fil: Moxham, Claire. Arthur Rylah Institute for Environmental Research; Australia Fil: Nehring, Kyle C.. State University of Utah; Estados Unidos Fil: Olff, Han. University of Groningen; Países Bajos Fil: Palmer, Todd M.. University of Florida; Estados Unidos Fil: Rebollo, Salvador. Universidad de Alcalá; España Fil: Riginos, Corinna. The Nature Conservancy; Estados Unidos Fil: Risch, Anita C.. Swiss Federal Institute for Forest, Snow and Landscape Research; Suiza Fil: Rueda, Marta. Consejo Superior de Investigaciones Científicas. Estación Biológica de Doñana; España Fil: Sankaran, Mahesh. National Centre For Biological Sciences; India. University of Leeds; Reino Unido Fil: Sasaki, Takehiro. Yokohama National University; Japón Fil: Schoenecker, Kathryn A.. Fort Collins Science Center and Colorado State University; Estados Unidos Fil: Schultz, Nick L.. Federation University; Australia Fil: Schütz, Martin. Swiss Federal Institute for Forest, Snow and Landscape Research; Suiza Fil: Schwabe, Angelika. Universitat Technische Darmstadt; Alemania Fil: Siebert, Frances. North West University; Sudáfrica Fil: Smit, Christian. University of Groningen; Países Bajos Fil: Stahlheber, Karen A.. University of Wisconsin Green Bay; Estados Unidos Fil: Storm, Christian. Universitat Technische Darmstadt; Alemania Fil: Strong, Dustin J.. United States Department of Agriculture. Agriculture Research Service; Estados Unidos Fil: Su, Jishuai. Northwest A&F University; China Fil: Tiruvaimozhi, Yadugiri V.. National Centre for Biological Sciences; India Fil: Tyler, Claudia. University of California; Estados Unidos Fil: Val, James. Office of Environment and Heritage; Australia Fil: Vandegehuchte, Martijn L.. University of Ghent; Bélgica. Swiss Federal Institute for Forest, Snow and Landscape Research; Suiza Fil: Veblen, Kari E.. State University of Utah; Estados Unidos Fil: Vermeire, Lance T.. United States Department of Agriculture. Agriculture Research Service; Estados Unidos Fil: Ward, David. Kent State University; Estados Unidos Fil: Wu, Jianshuang. Lhasa National Ecological Research Station; China Fil: Young, Truman P.. University of California at Davis; Estados Unidos. Mpala Research Centre; Kenia Fil: Yu, Qiang. Chinese Academy Of Agricultural Sciences; China. Institute of Agricultural Resources and Regional Planning; China Fil: Zelikova, Tamara Jane. University Of Wyoming; . University of Wyoming; Estados Unidos

Biological invasions are both a pressing environmental challenge and an opportunity to investigate fundamental ecological processes, such as the role of top predators in regulating biodiversity and food-web structure. In whole-ecosystem manipulations of small Caribbean islands on which brown anole lizards (Anolis sagrei) were the native top predator, we experimentally staged invasions by competitors (green anoles, Anolis smaragdinus) and/or new top predators (curly-tailed lizards, Leiocephalus carinatus). We show that curly-tailed lizards destabilized the coexistence of competing prey species, contrary to the classic idea of keystone predation. Fear-driven avoidance of predators collapsed the spatial and dietary niche structure that otherwise stabilized coexistence, which intensified interspecific competition within predator-free refuges and contributed to the extinction of green-anole populations on two islands. Moreover, whereas adding either green anoles or curly-tailed lizards lengthened food chains on the islands, adding both species reversed this effect-in part because the apex predators were trophic omnivores. Our results underscore the importance of top-down control in ecological communities, but show that its outcomes depend on prey behaviour, spatial structure, and omnivory. Diversity-enhancing effects of top predators cannot be assumed, and non-consumptive effects of predation risk may be a widespread constraint on species coexistence.

International audience; There is tremendous diversity of interactions between plants and other species. These relationships range from antagonism to mutualism. Interactions of plants with members of their ecological community can lead to a profound metabolic reconfiguration of the plants’ physiology. This reconfiguration can favour beneficial organisms and deter antagonists like pathogens or herbivores. Determining the cellular and molecular dialogue between plants, microbes, and insects, and its ecological and evolutionary implications is important for understanding the options for each partner to adopt an adaptive response to its biotic environment. Moving forward, understanding how such ecological interactions are shaped by environmental change and how we potentially mitigate deleterious effects will be increasingly important. The development of integrative multidisciplinary approaches may provide new solutions to the major ecological and societal issues ahead of us. The rapid evolution of technology provides valuable tools and opens up novel ways to test hypotheses that were previously unanswerable, but requires that scientists master these tools, understand potential ethical problems flowing from their implementation, and train new generations of biologists with diverse technical skills. Here, we provide brief perspectives and discuss future promise and challenges for research on insect–plant interactions building on the 16th International Symposium on Insect–Plant interactions (SIP) meeting that was held in Tours, France (2–6 July 2017). Talks, posters, and discussions are distilled into key research areas in insect–plant interactions, highlighting the current state of the field and major challenges, and future directions for both applied and basic research.

Interspecific interactions are important structuring forces in ecological communities. Interactions can be disturbed when species are lost from a community. When interactions result in fitness gains for at least one participating organism, that organism may experience reduced fitness as a result of interaction disturbance. However, many species exhibit traits that enable individuals to persist and reproduce in spite of such disruptions, resulting in resilience to interaction disturbance. Such traits can result in interaction generalization, phenotypic and behavioral plasticity, and adaptive capacity. We discuss examples of these traits and use case studies to illustrate how restoration practitioners can use a trait-based approach to examine species of concern, identify traits that are associated with interspecific interactions and are relevant to resilience, and target such traits in restoration. Restoration activities that bolster interaction resilience could include, for example, reintroducing or supporting specific functional groups or managing abiotic conditions to reduce interaction dependence by at-risk species (e.g. providing structural complexity offering shelter and cover). Resilience may also be an important consideration in species selection for restoration. Establishment of resilient species, able to persist after interaction disturbance, may be essential to restoring to a functioning ecological community. Once such species are present, they could help support more specialized species that lack resilience traits, such as many species of concern. Understanding the conditions under which processes linked to resilience may enable species to persist and communities to reform following interaction disturbance is a key application of community ecology to ecological restoration.

Increasingly, the restoration of large carnivores is proposed as a means through which to restore community structure and ecosystem function via trophic cascades. After a decades-long absence, African wild dogs (Lycaon pictus) recolonized the Laikipia Plateau in central Kenya, which we hypothesized would trigger a trophic cascade via suppression of their primary prey (dik-dik, Madoqua guentheri ) and the subsequent relaxation of browsing pressure on trees. We tested the trophic-cascade hypothesis using (1) a 14-year time series of wild dog abundance; (2) surveys of dik-dik population densities conducted before and after wild dog recovery; and (3) two separate, replicated, herbivore-exclusion experiments initiated before and after wild dog recovery. The dik-dik population declined by 33% following wild dog recovery, which is best explained by wild dog predation. Dik-dik browsing suppressed tree abundance, but the strength of suppression did not differ between before and after wild dog recovery. Despite strong, top-down limitation between adjacent trophic levels (carnivore- herbivore and herbivore-plant), a trophic cascade did not occur, possibly because of a time lag in indirect effects, variation in rainfall, and foraging by herbivores other than dik-dik. Our ability to reject the trophic-cascade hypothesis required two important approaches: (1) temporally replicated herbivore exclusions, separately established before and after wild dog recovery; and (2) evaluating multiple drivers of variation in the abundance of dik-dik and trees. While the restoration of large carnivores is often a conservation priority, our results suggest that indirect effects are mediated by ecological context, and that trophic cascades are not a foregone conclusion of such recoveries.

The search for predictions of species diversity across environmental gradients haschallenged ecologists for decades. The humped-back model (HBM) suggests that plantdiversity peaks at intermediate productivity; at low productivity few species can toleratethe environmental stresses, and at high productivity a few highly competitive speciesdominate. Over time the HBM has become increasingly controversial, and recent studiesclaim to have refuted it. Here, by using data from coordinated surveys conductedthroughout grasslands worldwide and comprising a wide range of site productivities, weprovide evidence in support of the HBM pattern at both global and regional extents. Therelationships described here provide a foundation for further research into the local,landscape, and historical factors that maintain biodiversity. Fil: Fraser, Lauchlan H.. Thompson Rivers University; Estados Unidos Fil: Pither, Jason. University of British Columbia; Canadá Fil: Jentsch, Anke. University Of Bayreuth; Alemania Fil: Sternberg, Marcelo. Tel Aviv University; Israel Fil: Zobel, Martín. University Of Tartu.; Estonia Fil: Cabido, Marcelo Ruben. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina Fil: Díaz, Sandra Myrna. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina Fil: Enrico, Lucas. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina Fil: Nabinger, Carlos. Universidade Federal do Rio Grande do Sul; Brasil Fil: Naseri, Kamal. Ferdowsi University of Mashhad; Irán Fil: Overbeck, Gerhard E.. Universidade Federal do Rio Grande do Sul; Brasil Fil: Palmer, Todd M.. University of Florida; Estados Unidos Fil: Parsons, Sheena. University of Kansas; Estados Unidos Fil: Pesek, Mary. University of Kansas; Estados Unidos Fil: Pillar, Valério D.. Universidade Federal do Rio Grande do Sul; Brasil Fil: Pringle, Robert M.. Princeton University; Estados Unidos Fil: Roccaforte, Kathy. University of Kansas; Estados Unidos Fil: Schmidt, Amanda. Thompson Rivers University; Canadá Fil: Shang, Zhanhuan. Lanzhou University; China Fil: Stahlmann, Reinhold. University Of Bayreuth; Alemania Fil: Stotz, Gisela C.. University of Alberta; Canadá Fil: Sugiyama, Shu-ichi. Hirosaki University; Japón Fil: Szentes, Szilárd. Szent István University; Hungría Fil: Thompson, Don. Lethbridge Research Centre; Canadá Fil: Tungalag, Radnaakhand. National University of Mongolia; Mongolia Fil: Undrakhbold, Sainbileg. National University of Mongolia; Mongolia Fil: van Rooyen, Margaretha. University of Pretoria; Sudáfrica Fil: Wellstein, Camilla. Free University of Bozen Bolzano; Italia Fil: Wilson, J. Bastow. University Of Otago; Canadá. Crown Research Institutes. Landcare Research; Nueva Zelanda Fil: Zupo, Talita. Universidad Estadual Paulista; Brasil

Invasive species can indirectly affect ecosystem processes via the disruption of mutualisms. The mutualism between the whistling thorn acacia (Acacia drepanolobium) and four species of symbiotic ants is an ecologically important one; ants strongly defend trees against elephants, which can otherwise have dramatic impacts on tree cover. In Laikipia, Kenya, the invasive big-headed ant (Pheidole megacephala) has established itself at numerous locations within the last 10-15 years. In invaded areas on five properties, we found that three species of symbiotic Crematogaster ants were virtually extirpated, whereas Tetraponera penzigi co-occurred with P. megacephala. T. penzigi appears to persist because of its nonaggressive behavior; in a whole-tree translocation experiment, Crematogaster defended host trees against P. megacephala, but were extirpated from trees within hours. In contrast, T. penzigi retreated into domatia and withstood invading ants for .30 days. In the field, the loss of defensive Crematogaster ants in invaded areas led to a five- to sevenfold increase in the number of trees catastrophically damaged by elephants compared to un-invaded areas. In savannas, tree cover drives many ecosystem processes and provides essential forage for many large mammal species; thus, the invasion of big-headed ants may strongly alter the dynamics and diversity of East Africa's whistling thorn savannas by disrupting this system's keystone acacia-ant mutualism.

Many species of large wildlife have declined drastically worldwide. These reductions often lead to profound shifts in the ecology of entire communities and ecosystems. However, the effects of these large-wildlife declines on other taxa likely hinge upon both underlying abiotic properties of these systems and on the types of secondary anthropogenic changes associated with wildlife loss, making impacts difficult to predict. To better understand how these important contextual factors determine the consequences of large-wildlife declines on other animals in a community, we examined the effects of three common forms of large-wildlife loss (removal without replacement [using fences], removal followed by replacement with domestic stock, and removal accompanied by crop agricultural use) on small-mammal abundance, diversity, and community composition, in landscapes that varied in several abiotic attributes (rainfall, soil fertility, land-use intensity) in central Kenya. We found that small-mammal communities were indeed heavily impacted by all forms of large-wildlife decline, showing, on average: (1) higher densities, (2) lower species richness per site, and (3) different species assemblages in sites from which large wildlife were removed. However, the nature and magnitude of these effects were strongly context dependent. Rainfall, type of land-use change, and the interaction of these two factors were key predictors of both the magnitude and type of responses of small mammals. The strongest effects, particularly abundance responses, tended to be observed in low-rainfall areas. Whereas isolated wildlife removal primarily led to increased small-mammal abundance, wildlife removal associated with secondary uses (agriculture, domestic stock) had much more variable effects on abundance and stronger impacts on diversity and composition. Collectively, these results (1) highlight the importance of context in determining the impacts of large-wildlife decline on small-mammal communities, (2) emphasize the challenges in extrapolating results from controlled experimental studies to predict the effects of wildlife declines that are accompanied by secondary land-uses, and (3) suggest that, because of the context-dependent nature of the responses to large-wildlife decline, large-wildlife status alone cannot be reliably used to predict small-mammal community changes.

Intraspecific variation in plant defense phenotype is common and has wide-ranging ecological consequences. Yet prevailing theories of plant defense allocation, which primarily account for interspecific differences in defense phenotype, often fail to predict intraspecific patterns. Furthermore, although individual variation in defense phenotype is often attributed to ecological interactions, few general mechanisms have been proposed to explain the ubiquity of variable defense phenotype within species. Here, we show experimentally that associational refuges and induced resistance interact to create predictable intraspecific variation in defense phenotype in African savanna plants. Physically defended species from four families (Acanthaceae, Asparagaceae, Cactaceae, and Solanaceae) growing in close association with spinescent Acacia trees had 39-78% fewer spines and thorns than did isolated conspecifics. For a subset of these species, we used a series of manipulative experiments to show that this variability is maintained primarily by a reduction in induced responses among individuals that seldom experience mammalian herbivory, whether due to association with Acacia trees or to experimental herbivore exclusion. Unassociated plants incurred 4- to 16-fold more browsing damage than did associated individuals and increased spine density by 16-38% within one month following simulated browsing. In contrast, experimental clipping induced no net change in spine density among plants growing beneath Acacia canopies or inside long-term herbivore exclosures. Associated and unassociated individuals produced similar numbers of flowers and seeds, but seedling recruitment and survival were vastly greater in refuge habitats, suggesting a net fitness benefit of association. We conclude that plant-plant associations consistently decrease defense investment in this system by reducing both the frequency of herbivory and the intensity of induced responses, and that inducible responses enable plants to capitalize on such associations in heterogeneous environments. Given the prevalence of associational and induced defenses in plant communities worldwide, our results suggest a potentially general mechanism by which biotic interactions might predictably shape intraspecific variation in plant defense phenotype.

Many species of large wildlife have declined drastically worldwide. These reductions often lead to profound shifts in the ecology of entire communities and ecosystems. However, the effects of these large-wildlife declines on other taxa likely hinge upon both underlying abiotic properties of these systems and on the types of secondary anthropogenic changes associated with wildlife loss, making impacts difficult to predict. To better understand how these important contextual factors determine the consequences of large-wildlife declines on other animals in a community, we examined the effects of three common forms of large- wildlife loss (removal without replacement (using fences), removal followed by replacement with domestic stock, and removal accompanied by crop agricultural use) on small-mammal abundance, diversity, and community composition, in landscapes that varied in several abiotic attributes (rainfall, soil fertility, land-use intensity) in central Kenya. We found that small- mammal communities were indeed heavily impacted by all forms of large-wildlife decline, showing, on average: (1) higher densities, (2) lower species richness per site, and (3) different species assemblages in sites from which large wildlife were removed. However, the nature and magnitude of these effects were strongly context dependent. Rainfall, type of land-use change, and the interaction of these two factors were key predictors of both the magnitude and type of responses of small mammals. The strongest effects, particularly abundance responses, tended to be observed in low-rainfall areas. Whereas isolated wildlife removal primarily led to increased small-mammal abundance, wildlife removal associated with secondary uses (agriculture, domestic stock) had much more variable effects on abundance and stronger impacts on diversity and composition. Collectively, these results (1) highlight the importance of context in determining the impacts of large-wildlife decline on small-mammal communities, (2) emphasize the challenges in extrapolating results from controlled experimental studies to predict the effects of wildlife declines that are accompanied by secondary land-uses, and (3) suggest that, because of the context-dependent nature of the responses to large-wildlife decline, large-wildlife status alone cannot be reliably used to predict small-mammal community changes.

Predicting how species' abundances and ranges will shift in response to climate change requires a mechanistic understanding of how multiple factors interact to limit population growth. Both abiotic stress and species interactions can limit populations and potentially set range boundaries, but we have a poor understanding of when and where each is most critical. A commonly cited hypothesis, first proposed by Darwin, posits that abiotic factors (e.g., temperature, precipitation) are stronger determinants of range boundaries in apparently abiotically stressful areas ("stress" indicates abiotic factors that reduce population growth), including desert, polar, or high-elevation environments, whereas species interactions (e.g., herbivory, competition) play a stronger role in apparently less stressful environments. We tested a core tenet of this hypothesis-that population growth rate is more strongly affected by species interactions in less stressful areas-using experimental manipulations of species interactions affecting a common herbaceous plant, Hibiscus meyeri (Malvaceae), across an aridity gradient in a semiarid African savanna. Population growth was more strongly affected by four distinct species interactions (competition with herbaceous and shrubby neighbors, herbivory, and pollination) in less stressful mesic areas than in more stressful arid sites. However, contrary to common assumptions, this effect did not arise because of greater density or diversity of interacting species in less stressful areas, but rather because aridity reduced sensitivity of population growth to these interactions. Our work supports classic predictions about the relative strength of factors regulating population growth across stress gradients, but suggests that this pattern results from a previously unappreciated mechanism that may apply to many species worldwide.

Both large-wildlife loss and climatic changes can independently influence the prevalence and distribution of zoonotic disease. Given growing evidence that wildlife loss often has stronger community-level effects in low-productivity areas, we hypothesized that these perturbations would have interactive effects on disease risk. We experimentally tested this hypothesis by measuring tick abundance and the prevalence of tick-borne pathogens ( Coxiella burnetii and Rickettsia spp . ) within long-term, size-selective, large-herbivore exclosures replicated across a precipitation gradient in East Africa. Total wildlife exclusion increased total tick abundance by 130% (mesic sites) to 225% (dry, low-productivity sites), demonstrating a significant interaction of defaunation and aridity on tick abundance. When differing degrees of exclusion were tested for a subset of months, total tick abundance increased from 170% (only mega-herbivores excluded) to 360% (all large wildlife excluded). Wildlife exclusion differentially affected the abundance of the three dominant tick species, and this effect varied strongly over time, likely due to differences among species in their host associations, seasonality, and other ecological characteristics. Pathogen prevalence did not differ across wildlife exclusion treatments, rainfall levels, or tick species, suggesting that exposure risk will respond to defaunation and climate change in proportion to total tick abundance. These findings demonstrate interacting effects of defaunation and aridity that increase disease risk, and they highlight the need to incorporate ecological context when predicting effects of wildlife loss on zoonotic disease dynamics.

While foundation species can stabilize ecosystems at landscape scales, their ability to persist is often underlain by keystone interactions occurring at smaller scales. Acacia drepanolobium is a foundation tree, comprising >95% of woody cover in East African black-cotton savanna ecosystems. Its dominance is underlain by a keystone mutualistic interaction with several symbiotic ant species in which it provides housing (swollen thorns) and carbohydrate-rich nectar from extra-floral nectaries (EFN). In return, it gains protection from catastrophic damage from mega-herbivores. Crematogaster mimosae is the ecologically dominant symbiotic ant in this system, also providing the highest protection services. In addition to tending EFN, C. mimosae tend scale insects for carbohydrate-rich honeydew. We investigated the role of scale insects in this specialized ant-plant interaction. Specifically, does this putatively redundant third partner strengthen the ant-plant mutualism by making the ant a better protector of the tree? Or does it weaken the mutualism by being costly to the tree while providing no additional benefit to the ant-plant mutualism? We coupled observational surveys with two scale-manipulation experiments and found evidence that this third partner strengthens the ant-plant mutualism. Trees with scale insects experimentally removed experienced a 2.5X increase in elephant damage compared to trees with scale insects present over 10 months. Reduced protection was driven by scale removal causing a decrease in ant colony size and per capita baseline activity and defensive behavior. We also found that ants increased scale-tending and the density of scale insects on trees when EFN were experimentally reduced. Thus, in this system, scale insects and EFN are likely complementary, rather than redundant, resources with scale insects benefitting ants when EFN production is low (such as during annual dry periods in this semi-arid ecosystem). This study reveals that a third-partner strengthens an ant-plant mutualism that serves to stabilize a whole ecosystem.

Ant-plant protection symbioses, in which plants provide food and/or shelter for ants in exchange for protection from herbivory, are model systems for understanding the ecology of mutualism. While interactions between ants, host plants, and herbivores have been intensively studied, we know little about how plant-plant interactions influence the dynamics of these mutualisms—despite strong evidence that plants compete for resources, that hosting ants can be costly, and that host-plant provisioning to ants can therefore be constrained by resource availability. We used field experiments in a semi-arid Kenyan savanna to examine interactions between the ant-plant Acacia drepanolobium, neighboring grasses, and two species of symbiotic acacia-ants with divergent behaviors: Crematogaster mimosae, an aggressive symbiont that imposes high costs to host trees via consumption of extrafloral nectar, and Tetraponera penzigi, a less-protective symbiont that imposes lower costs because it does not consume nectar. We hypothesized that by competing with acacias for resources, neighboring grasses (a) reduce hosts’ ability to support costly C. mimosae, while having little or no effect on the ability of hosts to support low-cost T. penzigi, and (b) reduce sapling growth rates irrespective of ant occupant. We factorially manipulated the presence/absence of grasses and the identity of ant occupants on saplings and evaluated effects on colony survivorship and sapling growth rates over 40 weeks. Contrary to prediction, the high-cost/high-reward nectar-dependent mutualist C. mimosae had higher colony-survival rates on saplings with grass neighbors present. Grasses appear to have indirectly facilitated the survival of C. mimosae by reducing water stress on host plants; soils under saplings shaded by grasses had higher moisture content, and these saplings produced more active nectaries than grass-removal saplings. Consistent with prediction, survival of low-cost/low-reward T. penzigi did not differ significantly between grass-removal treatments. Saplings occupied by low-cost/low-reward T. penzigi grew 100% more on average than saplings occupied by high-cost/high-reward C. mimosae, demonstrating that mutualist-partner identity strongly and differentially influences demographic rates of young plants. In contrast, contrary to prediction, grass neighbors had no significant net impact on sapling growth rates. Our results suggest that neighboring plants can exert strong and counterintuitive effects on ant-plant protection symbioses, highlighting the need to integrate plant-plant interactions into our understanding of these mutualisms. This article is protected by copyright. All rights reserved.

Sterilisation of parasites prevents host reproduction, thereby diverting host resources to their own benefit. Previous theory predicts that parasites should evolve maximum virulence, yet hosts are often incompletely sterilised. Whereas prior attempts to resolve this paradox have sought evolu- tionary explanations, we present theory and experiments showing that incomplete sterilisation can arise from ecologically driven fluctuations in parasite load. The African ant-plant Acacia drepa- nolobium reproduced more when occupied by small colonies of the sterilising symbiont Crematog- aster nigriceps. In nature, small colonies result from interference competition between ant colonies; these territorial conflicts thus provide intermittent windows of opportunity for host reproduction. Our mean-field model shows that numerical insufficiency of parasites can produce partial sterilisation of host populations, creating the appearance of reduced virulence even if ants have evolved to sterilise completely. This general framework helps explain both the apparent ubiq- uity of partial sterilisation parasitism and the ability of these symbiotic associations to persist.

Many tropical plants are defended by ants, and the costs and benefits of these mutualisms can vary across gradients of herbivory, soil fertility, latitude, and other environmental factors. Yet despite an abundant literature documenting thermal constraints on ant activity and behavior, we know little about whether temperature variation can influence the benefits conferred by ants to plants. We evaluated the effects of dawn-to-dusk fluctuations in temperature on patrolling and aggressive behavior in four arboreal ant mutualists of Acacia drepanolobium trees in central Kenya. We found that ant aggressive behavior significantly increased with branch surface temperature, primarily in the two most aggressive ant species: Crematogaster mimosae and C. nigriceps workers attacked a simulated herbivore at higher rates as surface temperature rose. In a browsing experiment, we found that goats browsed more frequently and for longer durations on C. mimosae-defended trees during cooler times of day, while goat browsing on plants from which ants had been removed was not affected by temperature. Our study demonstrates temperature-dependence in the efficacy of ant defense against herbivory and suggests that these ant-plants may be more vulnerable to herbivory during cooler hours of the day, when many native browsers are most active.

Determining how competing species coexist is essential to understanding patterns of biodiversity. Indirect facilitation, in which a competitively dominant species exerts a positive effect on one competitor by more strongly suppressing a third, shared competitor, is a potentially potent yet understudied mechanism for competitive coexistence. Here we provide evidence for indirect facilitation in a guild of four African Acacia ant species that compete for nesting space on the host plant Acacia drepanolobium, showing that a competitively dominant acacia ant species indirectly creates establishment opportunities for the most subordinate species that may help to maintain diversity. Using long-term observational data and field experiments, we demonstrate that the competitively dominant ant species outcompetes two competitively intermediate species, while tolerating colonies of the subordinate competitor; this creates opportunities for local colonization and establishment of colonies of the subordinate species within the dominant species' territories. Host plants occupied by this subordinate species are then more likely to be colonized by the intermediate species, which in turn are more likely to be displaced by the dominant species. This process has the potential to generate a cyclical succession of ant species on host trees, contributing to stable coexistence within this highly competitive community.