Seed predators limit plant recruitment in Neotropical savannas

Despite the well‐documented impacts of consumers on seed abundance the link between seed predation and plant population dynamics remains poorly understood because experimental studies linking patterns of predation with seedling establishment are rare. We used experimental manipulations with six woody plant species to elucidate the effects of seed predator type, habitat, and plant species identity on rates of seed predation and seedling recruitment in the Neotropical savannas known as the Cerrado. We found that seed predation rates are consistently high across a diversity of local habitat types, with important inter‐habitat variation in seed predation for three of the six species used in our experiments. We also found that seed predation has a clear demographic signal – experimentally excluding predators resulted in higher rates of seedling establishment over the course of two seasons. Because the intensity of seed predation varied between species and habitats, it may play a role in structuring local patterns of plant abundance and community composition. Finally, our results lend support to the recent hypothesis that herbivores have major and underappreciated impacts in Neotropical savannas, and that top–down factors can influence the demography of plants in this extensive and biodiversity‐rich biome in previously unexplored ways.     

Linking traits of foraging animals to spatial patterns of plants: social and solitary ants generate opposing patterns of surviving seeds

Foraging traits of seed predators are expected to impact the spatial structure of plant populations, community dynamics and diversity. Yet, many of the key mechanisms governing distance- or density-dependent seed predation are poorly understood. We designed an extensive set of field experiments to test how seed predation by two harvester ant species interact with seed dispersal in shaping the spatial patterns of surviving seeds. We show that the Janzen–Connell establishment pattern can be generated by central-place foragers even if their focal point is located away from the seed source. Furthermore, we found that differences in the social behaviour of seed predators influence their sensitivity to seed density gradients and yield opposing spatial patterns of surviving seeds. Our results support the predictions of a recent theoretical framework that unifies apparently opposing plant establishment patterns, and suggest that differences in foraging traits among seed predators can drive divergent pathways of plant community dynamics.     

Frugivore loss limits recruitment of large-seeded trees

Although global declines in frugivores may disrupt seed dispersal mutualisms and inhibit plant recruitment, quantifying the likely reduction in plant regeneration has been difficult and rarely attempted. We use a manipulative factorial experiment to quantify dependence of recruitment on dispersal (i.e. fruit pulp removal and movement of seed away from parental area) in two large-seeded New Zealand tree species. Complete dispersal failure would cause a 66 to 81 per cent reduction in recruitment to the 2-year-old seedling stage, and synergistic interactions with introduced mammalian seed and seedling predators increase the reduction to 92 to 94 per cent. Dispersal failure reduced regeneration through effects on seed predation, germination and (especially) seedling survival, including distance- and density-dependent (Janzen-Connell) effects. Dispersal of both species is currently largely dependent on a single frugivore, and many fruits today remain uneaten. Present-day levels of frugivore loss and mammal seed and seedling predators result in 57 to 84 per cent fewer seedlings after 2 years. Our study demonstrates the importance of seed dispersal for local plant population persistence, and validates concerns about the community consequences of frugivore declines.     

Enemies mediate distance- and density-dependent mortality of tree seeds and seedlings: a meta-analysis of fungicide,insecticide and exclosure studies

Conspecific negative distance- and density-dependence is often assumed to be one of the most important mechanisms controlling forest community assembly and species diversity globally. Plant pathogens, and insect and mammalian herbivores, are the most common natural enemy types that have been implicated in this phenomenon, but their general effects at different plant life stages are still unclear. Here, we conduct a meta-analysis of studies that involved robust manipulative experiments, using fungicides, insecticides and exclosures, to assess the contributions of different natural enemy types to distance- and density-dependent effects at seed and seedling stages. We found that distance- and density-dependent mortality caused by natural enemies was most likely at the seedling stage and was greater at higher mean annual temperatures. Conspecific negative distance- and density-dependence at the seedling stage is significantly weakened when fungicides were applied. By contrast, negative conspecific distance- and density-dependence is not a general pattern at the seed stage. High seed mass reduced distance- and density-dependent mortality at the seed stage. Seed studies excluding only large mammals found significant negative conspecific distance-dependent mortality, but exclusion of all mammals resulted in a non-significant effect of conspecifics. Our study suggests that plant pathogens are a major cause of distance- and density-dependent mortality at the seedling stage, while the impacts of herbivores on seedlings have been understudied. At the seed stage, large and small mammals, respectively, weaken and enhance negative conspecific distance-dependent mortality. Future research should identify specific agents of mortality, investigate the interactions among different enemy types and assess how global change may affect natural enemies and thus influence the strength of conspecific distance- and density-dependence.     

Biotic resistance to plant invasion in grassland: Does seed predation increase with resident plant diversity?

Seed predation impacts heavily on plant populations and community composition in grasslands. In particular, generalist seed predators may contribute to biotic resistance, i.e. the ability of resident species in a community to reduce the success of non-indigenous plant invaders. However, little is known of predators’ preferences for seeds of indigenous or non-indigenous plant species or how seed predation varies across communities. We hypothesize that seed predation does not differ between indigenous and non-indigenous plant species and that seed predation is positively related to plant species diversity in the resident community. The seed removal of 36 indigenous and non-indigenous grassland species in seven extensively or intensively managed hay meadows across Switzerland covering a species-richness gradient of 18–50 plant species per unit area (c. 2 m²) was studied. In mid-summer 2011, c. 24,000 seeds were exposed to predators in Petri dishes filled with sterilized soil, and the proportions of seeds removed were determined after three days’ exposure. These proportions varied among species (9.2–62.5%) and hay meadows (17.8–48.6%). Seed removal was not related to seed size. Moreover, it did not differ between indigenous and non-indigenous species, suggesting that mainly generalist seed predators were active. However, seed predation was positively related to plant species richness across a gradient in the range of 18–38 species per unit area, representing common hay meadows in Switzerland. Our results suggest that generalist post-dispersal seed predation contributes to biotic resistance and may act as a filter to plant invasion by reducing the propagule pressure of non-local plant species.     

Dispersal rates affect species composition in metacommunities of Sarracenia purpurea inquilines

Dispersal among local communities can have a variety of effects on species composition and diversity at local and regional scales. Local conditions (e.g., resource and predator densities) can have independent effects, as well as interact with dispersal, to alter these patterns. Based on metacommunity models, we predicted that local diversity would show a unimodal relationship with dispersal frequency. We manipulated dispersal frequencies, resource levels, and the presence of predators (mosquito larvae) among communities found in the water-filled leaves of the pitcher plant Sarracenia purpurea. Diversity and abundance of species of the middle trophic level, protozoa and rotifers, were measured. Increased dispersal frequencies significantly increased regional species richness and protozoan abundance while decreasing the variance among local communities. Dispersal frequency interacted with predation at the local community scale to produce patterns of diversity consistent with the model. When predators were absent, we found a unimodal relationship between dispersal frequency and diversity, and when predators were present, there was a flat relationship. Intermediate dispersal frequencies maintained some species in the inquiline communities by offsetting extinction rates. Local community composition and the degree of connectivity between communities are both important for understanding species diversity patterns at local and regional scales.     

Rodent seed predation: effects on seed survival, recruitment, abundance, and dispersion of bird-dispersed tropical trees

Tropical tree species vary widely in their pattern of spatial dispersion. We focus on how seed predation may modify seed deposition patterns and affect the abundance and dispersion of adult trees in a tropical forest in India. Using plots across a range of seed densities, we examined whether seed predation levels by terrestrial rodents varied across six large-seeded, bird-dispersed tree species. Since inter-specific variation in density-dependent seed mortality may have downstream effects on recruitment and adult tree stages, we determined recruitment patterns close to and away from parent trees, along with adult tree abundance and dispersion patterns. Four species (Canarium resiniferum, Dysoxylum binectariferum, Horsfieldia kingii, and Prunus ceylanica) showed high predation levels (78.5-98.7%) and increased mortality with increasing seed density, while two species, Chisocheton cumingianus and Polyalthia simiarum, showed significantly lower seed predation levels and weak density-dependent mortality. The latter two species also had the highest recruitment near parent trees, with most abundant and aggregated adults. The four species that had high seed mortality had low recruitment under parent trees, were rare, and had more spaced adult tree dispersion. Biotic dispersal may be vital for species that suffer density-dependent mortality factors under parent trees. In tropical forests where large vertebrate seed dispersers but not seed predators are hunted, differences in seed vulnerability to rodent seed predation and density-dependent mortality can affect forest structure and composition.     

Predators alter the scaling of diversity in prey metacommunities

Although predator effects on the number of locally coexisting species are well understood, there are few formal predictions of how these local predator effects influence patterns of prey diversity at larger spatial scales. Building on the theory of island biogeography, we develop a simple model that describes how predators can alter the scaling of diversity in prey metacommunities and compares the effects of generalist and specialist predators on regional prey diversity. Generalist predators, which consume prey randomly with respect to species identity, are predicted to reduce α‐diversity and increase β‐diversity thereby maintaining regional diversity (γ‐diversity). Alternatively, specialist predators, which filter out prey species intolerant of predators, are predicted to reduce bothα‐diversity andβ‐diversity by causing the same prey species to be extirpated in each locality, resulting in regional prey species extinctions and lower γ‐diversity. These distinct effects of generalist and specialist predators on prey diversity at different spatial scales are uniquely shaped by the extent of predation within those metacommunities. Overall, our model results make general predictions for how different types of predators can differentially affect prey diversity across spatial scales, allowing a more complete understanding of the possible implications of predator eradications or introductions for biodiversity.     

Top predator control of plant biodiversity and productivity in an old-field ecosystem

Abstract Predators can have strong indirect effects on plants by altering the way herbivores impact plants. Yet, many current evaluations of plant species diversity and ecosystem function ignore the effects of predators and focus directly on the plant trophic level. This report presents results of a 3‐year field experiment in a temperate old‐field ecosystem that excluded either predators, or predators and herbivores and evaluated the consequence of those manipulations on plant species diversity (richness and evenness) and plant productivity. Sustained predator and predator and herbivore exclusion resulted in lower plant species evenness and higher plant biomass production than control field plots representing the intact natural ecosystem. Predators had this diversity‐enhancing effect on plants by causing herbivores to suppress the abundance of a competitively dominant plant species that offered herbivores a refuge from predation risk.     

Post-dispersal seed predation: consequences for plant demography and evolution

Post-dispersal seed predation is only one of many factors underlying plant demography and evolution. Nevertheless, the generalist feeding habits of many post-dispersal seed predators and the limited ability of plants either to compensate for or to respond to post-dispersal seed losses directly suggest that post-dispersal seed predation may have a considerable impact on plant populations. Seed predators probably have little direct influence on the demography of plants that regenerate exclusively by vegetative means or are buffered by a large active seed bank, but such species are only a minority in most plant communities.In general, ants are significant post-dispersal seed predators in arid and semi-arid ecosystems while they act mainly as seed dispersers rather than as predators in temperate ecosystems. Although studies have probably underestimated the importance of invertebrates and birds as seed predators, rodents appear to have greater potential to influence seed dynamics, and are particularly important in temperate ecosystems. For example, production of mast seed crops is more effective at satiating specialist invertebrate seed predators than generalist vertebrates, and recruitment may be limited by post-dispersal seed predation even during mast years.Both spatial variation in post-dispersal seed predation and differences in predation between species are important elements which facilitate the coexistence of different plant species. Where microsites are limiting, selective post-dispersal seed predators can influence pre-emptive competition for these microsites. Seed size determines the extent of density-dependent predation and the exploitation of buried seed. This suggests that post-dispersal seed predators may also play a role in the evolution of seed characteristics. However, conclusions regarding the ecological and evolutionary impact of post-dispersal seed predators will remain speculative without a more substantial empirical base.     

Regeneration of Salix arbuscula and Salix lapponum within a Large Mammal Exclosure: The Impacts of Microsite and Herbivory

Over‐grazing or browsing by large herbivores may result in the loss of individual plant species or entire plant communities. Restoration schemes often involve exclusion of large mammals, but the resulting changes in vegetation may alter other important ecological processes such as regeneration, via changes in microsite availability for seed germination or increases in populations of seedling predators. Working within a large fenced area from which large mammals were excluded, we experimentally tested the effects of microsite, small herbivores, and their interactions on post‐dispersal seed and early seedling mortality of one nationally scarce (Salix arbuscula) and one nationally rare (S. lapponum) species of montane willow. Seeds were sown in three different microsites: natural vegetation, mown vegetation (mimicking grazed sward), and bare ground. Small exclosures and slug pellets were used to examine the effects of small mammal and slug predation, respectively. Survival of seedlings was monitored during the summer following planting. The presence of bare ground, rather than the absence of herbivores, was of over‐riding importance for early seedling survival and establishment. Protecting seedlings from small mammals made no difference to the levels of survival; however, protecting seedlings from slugs (Arion spp.) resulted in approximately 45% of seedlings surviving until the end of the summer compared to only 30% when seedlings were available to slugs. Although excluding large herbivores may increase seed production of existing individuals, the impacts of changes to plant communities on processes such as regeneration need to be considered if restoration projects are to be fully successful.     

Seed predation of two alien Opuntia species invading Mediterranean communities

Invasibility depends on the interaction of the introduced species with the abiotic and biotic factors of the recipient community. In particular, the biotic resistance posed by native herbivores has been claimed to be of great importance in controlling plant invasion. We investigated fruit and seed predation of two exotic Opuntia species within and between Mediterranean communities in order to determine how patterns of predation matched patterns of invasion. Predators were small mammals, presumably mice, which could consume more than 50% of the seeds produced. Predators could be equally effective in consuming fruit and single seeds. O. maxima fruits were slightly preferred to O. stricta fruits, but predators did not distinguish between seeds. Seed predation was more intense in invaded than in non-invaded communities. However, there was a high spatial variation in seed predation that did not always match patterns of invasion, suggesting that seed predation alone is not a good predictor of community invasibility to Opuntia. According to these results invasibility to Opuntia is limited in some (but not all) communities by native mice. Seed losses by predation were high for both species. However, we estimated that more than 75% of seeds dispersed by birds to non-invaded areas are not predated.     

Seed Dispersers,Seed Predators,and Browsers Act Synergistically as Biotic Filters in a Mosaic Landscape

In this study, we analize the functional influence of animals on the plants they interact with in a mediterranean mountain. We hypothesise that seed dispersers, seed predators, and browsers can act as biotic filters for plant communities. We analyse the combined effects of mutualistic (seed dispersal) and antagonistic (seed predation, herbivory) animal interactions in a mosaic landscape of Mediterranean mountains, basing our results on observational and experimental field. Most of the dispersed seeds came from tree species, whereas the population of saplings was composed predominantly of zoochorous shrub species. Seed predators preferentially consumed seeds from tree species, whereas seeds from the dominant fleshy-fruited shrubs had a higher probability of escaping these predators. The same pattern was repeated among the different landscape units by browsers, since they browsed selectively and far more intensely on tree-species saplings than on the surrounding shrubs. In synthesis, our work identifies the major biotic processes that appear to be favoring a community dominated by shrubs versus trees because seed dispersers, predators, and herbivores together favored shrub dispersal and establishment versus trees.     

Spatial concordance at a regional scale in the regeneration process of a circum-Mediterranean relict (Buxus balearica): connecting seed dispersal to seedling establishment

Both microhabitat and temporal scales have significant effects on the regeneration process of plant species. However, the consistency of those patterns at large scales is largely unknown, despite of being essential from both an ecological and a conservation viewpoint. In this study, we examined the spatio‐temporal variation in the process of regeneration through seeds of the relict Mediterranean shrub Buxus balearica. By means of observations and experiments, we quantified the losses at different recruitment phases (seed rain, seed predation, seedling emergence and establishment) both at a small spatial scale (distance and microhabitat effects) and at a temporal scale (four years), and compared them at a regional scale (10 different localities). We additionally studied transition probabilities between phases in two populations and measured the size structure of 18 localities encompassing all the distributional range of the species. The results show that seed shadow follows a leptokurtic pattern which is consistently maintained until the seedling phase (i.e. there is spatial concordance). Despite post‐dispersal factors are spatially uncoupled, they do not remove the gradient established in the seed shadow. Although the importance of different seed predators varied between regions, post‐dispersal seed predation (ca 80%), together with seedling emergence (usually lower than 10%, some years even nil), and seedling mortality during the first summer (ca 70%) were – consistently in all populations – the most critical bottlenecks in the regeneration of the species. Moreover, an absence of seed bank prevents the buffering of dramatic losses during such phases. Watering of sown seeds increased seedling emergence and survival, and number of leaves per seedling, and survival was consistently higher under vegetation coverage. These results suggest that hydric stress notably contributes to limit the regeneration of this once widespread species evolved under a subtropical climate, which nowadays appears in old‐structured populations throughout its distributional area.     

Post-dispersal seed predation of three grassland species in a plant diversity experiment

Aims Post-dispersal seed predation is an important ecosystem process because it can influence the seed's fate after the initial dispersal from the mother plant and subsequently transform communities. Even at small scales, post-dispersal seed predation can vary greatly depending on seed identity, granivorous taxa or microhabitat structure. However, little is known about the role of plant species richness and functional group richness in post-dispersal seed predation. The overall aim of this study was to test whether increasing plant species richness or plant functional group richness affects the rate and variability of post-dispersal seed predation. We additionally investigated the influence of vegetation structure and seed species identity on the rate and variability of post-dispersal seed predation and whether the influence of different granivorous taxa changed with increasing plant species richness.Methods We conducted seed removal experiments along a long-term experimental plant diversity gradient, comprising plots with monocultures to 60 species mixtures of common grassland species in Jena, Germany, in August 2011. We studied seeds of Onobrychis viciifolia, Pastinaca sativa and Trifolium pratense in exclusion experiments (seed cafeterias), an experimental setup that allowed access either for arthropods or slugs or for all granivorous taxa. Traditionally, seeds removed from seed cafeterias were classified as consumed but we used traceable fluorescent-coloured seeds to obtain more accurate predation rates by subtracting recovered seeds from overall removed seeds. The effect of multiple vegetation variables on mean and variability of seed predation rates was analysed using generalized mixed-effect models and linear regressions, respectively.Important findings Rates of recovered seeds were low but contributed to significant differences between seed predation rates and removal rates of seeds in some treatments. Seed predation rates were not directly correlated with increasing plant species richness or plant functional group richness but were influenced byseed species identity and granivorous taxa. Vegetation variables such as vegetation height and cover were significantly associated with seed predation rates. Depending on the seed species and/or the granivorous taxa, different vegetation variables correlated with seed predation rates. Our results indicate that effects of plant functional group richness and multiple vegetation variables on the magnitude of post-dispersal seed predation varied with seed identity and seed predator taxa. A direct effect of plant species and plant functional group richness could be shown on the variability of post-dispersal seed predation for some seed species and their respective predators. Thus, the changes in magnitude of post-dispersal seed predation with increasing plant species richness could potentially impact the fitness of some plant species and thereby influence plant community structure.     

Rodent seed predation as a biotic filter influencing exotic plant abundance and distribution

Biotic resistance is commonly invoked to explain why many exotic plants fail to thrive in introduced ranges, but the role of seed predation as an invasion filter is understudied. Abiotic conditions may also influence plant populations and can interact with consumers to determine plant distributions, but how these factors jointly influence invasions is poorly understood. In central Argentina’s Caldenal savannas, we experimentally examined how seed predation and water availability influenced recruitment/establishment of nine exotic plant invaders over 2 years. We then explored how seed predation patterns related to invasion patterns. Excluding rodent seed predators dramatically increased seedling recruitment for eight of nine exotic species (by 100–300 % in most cases) and increased young/adult plant abundance for four species in one or both years. Adding water to ameliorate drought tended to increase seedling numbers for most species, but these trends were not significant. Vegetation surveys revealed that exotic plant richness was 50 % lower in matrix habitat compared with disturbed roadsides and that cover of the two most aggressive invaders, which were both strongly suppressed by seed predation, was 75–80 % lower in matrix than roadside habitats. Seed offerings indicated seed removal by rodents was 11 times greater in intact matrix habitat compared with roadsides. Rodent seed predation represents a significant source of biotic resistance to plant invasions. Ubiquitous disturbances such as road construction can disrupt this filter. The widely recognized role that disturbance plays in facilitating invasions, which is largely attributed solely to reduced plant competition, may also arise from disruption of top–down controls.     

Potential for ants and vertebrate predators to shape seed-dispersal dynamics of the invasive thistles <Emphasis Type="Italic">Cirsium arvense</Emphasis> and <Emphasis Type="Italic">Carduus nutans</Emphasis> in their introduced range (North America)

The population dynamics of invasive plants are influenced by positive and negative associations formed with members of the fauna present in the introduced range. For example, mutualistic associations formed with pollinators or seed dispersers may facilitate invasion, but reduced fitness from attack by native herbivores can also suppress it. Since population expansion depends on effective seed dispersal, interactions with seed dispersers and predators in a plant species introduced range may be of particular importance. We explored the relative contributions of potential seed dispersers (ants) and vertebrate predators (rodents and birds) to seed removal of two diplochorous (i.e., wind- and ant-dispersed), invasive thistles, Cirsium arvense and Carduus nutans, in Colorado, USA. We also conducted behavior trials to explore the potential of different ant species to disperse seeds, and we quantified which potential ant dispersers were prevalent at our study locations. Both ants and vertebrate predators removed significant amounts of C. arvense and C. nutans seed, with the relative proportion of seed removed by each guild varying by location. The behavior trials revealed clear seed preferences among three ant species as well as differences in the foragers’ abilities to move seeds. In addition, two ant species that acted as potential dispersal agents were dominant at the study locations. Since local conditions in part determined whether dispersers or predators removed more seed, it is possible that some thistle populations benefit from a net dispersal effect, while others suffer proportionally more predation. Additionally, because the effectiveness of potential ant dispersers is taxon-specific, changes in ant community composition could affect the seed-dispersal dynamics of these thistles. Until now, most studies describing dispersal dynamics in C. arvense and C. nutans have focused on primary dispersal by wind or pre-dispersal seed predation by insects. Our findings suggest that animal-mediated dispersal and post-dispersal seed predation deserve further consideration.     

Patch size,shape and edge distance influence seed predation on a palm species in the Atlantic forest

Seed predation is an important ecological process that affects the abundance, diversity and distribution of plant species, and it is known to be influenced by defaunation and forest fragmentation. Most studies on seed predation in human‐modified landscapes do not take into account the different spatial scales in which this process operates. In this study, we evaluated how variables at three distinct spatial scales affected the seed predation of a palm that provides a keystone resource to the frugivore community, the queen palm Syagrus romanzoffiana. Thirteen landscapes that vary in forest cover, number of fragments and patch sizes were sampled in the Brazilian Atlantic forest. We also evaluated the contribution of the three main groups of seed predators: squirrels, terrestrial rodents and invertebrates. Our results indicate that seed predation is more affected by fragment and local variables than by landscape influences. In addition, the size of the fragment, its shape and the distance from the nearest forest edge were the main predictors of the proportion of predated seeds. Moreover, the two main seed predators (squirrels and invertebrates) responded to the same fragment and local variables. Because most of the Atlantic forest consists of small fragments, we expect that the seed predation of this keystone palm should be high in most of its distribution, with potential consequences for the frugivore community.     

Distance-dependence in two Amazonian palms: effects of spatial and temporal variation in seed predator communities

Animals aid population growth and fitness in tropical forest communities through dispersal and negatively impact populations through seed predation. The interaction between dispersal and seed predation can produce distance- or density-dependence; powerful mechanisms for maintaining species diversity incorporated in the Janzen–Connell model. Large mammals, the highest biomass seed predators of intact Amazonian communities and at risk due to human disturbance, are potentially central to these interactions. This study tests the Janzen–Connell model and investigates the impact of mammalian seed predators on seedling recruitment and maintenance of tree diversity. Patterns of both vertebrate and invertebrate seed predation and seedling recruitment were studied in the two most abundant canopy tree species in western Amazonia (Arecaceae: Astrocaryum murumuru and Iriartea deltoidea). We specifically examined effects of both spatial and temporal variation of the highest biomass seed predator in southwest Amazonian forests, the white-lipped peccary (Tayassu pecari), on recruitment through disturbed and undisturbed sites and through a fortuitous 12 year natural extinction and recolonization event of T. pecari. Distance-dependent seedling recruitment was found in Astrocaryum and Iriartea at both sites. However, the median distance of seedlings was ~1.5× farther from reproductive adults in both palms at the undisturbed site. The number of Iriartea seeds escaping predation increased 6,000% in both space and time due to the decline of T. pecari abundance. The results demonstrate that Janzen–Connell effects are stronger in intact ecosystems and tie these mechanistically to changes in seed predator abundance. This study shows that anthropogenic changes in mammal communities decrease the magnitude of Janzen–Connell effects in Amazonian forests and may result in decreases in tree diversity.     

Parasitism on seed predators overcomes the detrimental effects of defoliation on plant fitness in a tritrophic system

It is widely recognised that the interaction between plants and herbivores cannot be completely understood if the natural enemies of the latter are not included. Most studies looking at the effects of herbivores and their enemies on plant fitness only consider one herbivore species or guild; however, plants in nature usually face the attack of more than one herbivore guild simultaneously and these herbivores may have a non-additive effect on the attraction with bodyguards and plant fitness. In this study, we asked whether folivory affects the activity of parasitoids on seed predators and whether this effect cascades down to plant fitness. We assessed these questions in a tritrophic system: the plant Ruellia nudiflora, its pre-dispersal seed predators and the parasitoids of the latter. Plants were submitted to either 50 % artificial defoliation or no defoliation (control). The number of seeds, fruit production and parasitoid incidence was assessed periodically in both sets of plants. Parasitoids indirectly and positively affected seed number, while defoliation had a direct negative effect on the number of seeds and an indirect negative effect on parasitoid incidence. However, the combined effect of defoliation and seed predation increased the indirect positive effect of the parasitoids on seed production, which overcame the negative effects of defoliation.