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Research Highlight: Lundgren, E. J., Ramp, D., Middleton, O. S., Wooster, E. I. F., Kusch, E., Balisi, M., Ripple, W. J., Hasselerharm, C. D., Sanchez, J. N., Mills, M. & Wallach, A. D. (2022) A novel trophic cascade between cougars and feral donkeys shapes desert wetlands. Journal of Animal Ecology, (91, 2010-2022). https://doi.org/10.1111/1365-2656.13766. Despite being absent from most regions of the contemporary world, megafauna species dominated the dynamics of ecological communities until the late Pleistocene. Trophic rewilding is a promising approach to restoring megafauna interactions, their functional roles and the consequent trophic cascades. Unintentional rewilding with large non-native herbivores, such as equids, offers outstanding opportunities for ecologists to understand the outcomes of using replacement species to restore the ecological functions of extinct native megafauna. In this context, it is relevant to understand the extent to which extant native predators can impose top-down control on non-native megaherbivores and how trophic cascades arising from novel predator-prey interactions influences biodiversity and ecosystem functions. In Death Valley National Park (United States), Lundgren et al. depict a natural experiment showing compelling evidence of native cougars overcoming ecological naïveté-that is, the mismatches between predator and prey species that do not share a common evolutionary history-and are now successfully preying on younger individuals of feral donkeys. These non-native donkeys, whose growing wild populations threaten the native biota, became cougars' most frequent dietary item in that region. In areas with cougars, donkeys changed their spatiotemporal foraging patterns, becoming diurnal and less active. On the other hand, donkeys remain more active and forage throughout the day and night in areas without cougars. The cougar-donkey interaction triggered a behaviourally mediated trophic cascade emerging from a 'landscape of fear', that is, from the perception of spatial heterogeneity in predation risk by donkeys. Areas with cougars have less trampled ground, fewer donkey trails, and much more canopy cover and vegetation around water. Donkeys concentrate their activity mostly in topologically plain terrains lacking proper sites for the ambush behaviour of cougars and with more intense human presence, likely acting as a shield against the predators. Lundgren et al. present a promising model system for studying the effects of fearful grazers on community structure in the context of novel ecological interactions being established in the Anthropocene. Whether the emerging cascade is transient or persistent, the relative roles of consumptive and non-consumptive effects as underlying mechanisms, and their consequences for food web structure, functioning and stability, are questions of general interest. Addressing them can help us to elucidate the costs and benefits of using non-native megaherbivores in the functional restoration of permanently invaded ecosystems.

Pesquisa em Destaque: Lundgren, E. J., Ramp, D., Middleton, O. S., Wooster, E. I. F., Kusch, E., Balisi, M., Ripple, W. J., Hasselerharm, C. D., Sanchez, J. N., Mills, M. & Wallach, A. D. (2022) A novel trophic cascade between cougars and feral donkeys shapes desert wetlands. Journal of Animal Ecology, 00:00-00. https://doi.org/10.1111/1365-2656.13766. Apesar de estarem ausentes da maioria das regiões do mundo contemporâneo, as espécies da megafauna dominaram a dinâmica das comunidades ecológicas até o Pleistoceno tardio. A refaunação trófica é uma abordagem promissora para a restauração das interações da megafauna, de seus papéis funcionais e das cascatas tróficas delas decorrentes. A refaunação não-intencional com grandes herbívoros não-nativos, tais como equídeos, oferece oportunidades extraordinárias para os ecólogos entenderem as consequências do uso de espécies substitutas para restaurar as funções ecológicas da extinta megafauna nativa. Nesse contexto, é relevante entendermos o quanto os predadores nativos atuais podem impor controle de cima para baixo sobre mega-herbívoros não-nativos, e como cascatas tróficas oriundas de novas interações predador-presa influenciam a biodiversidade e as funções ecossistêmicas. No Parque Nacional do Vale da Morte (Estados Unidos), Lundgren et al. reportam um experimento natural mostrando evidências contundentes de que onças pardas superaram a ingenuidade ecológica - i.e., os desajustes entre predadores e presas que não compartilham uma história evolutiva comum - e agora estão predando os indivíduos mais jovens de burros ferais com sucesso. Esses burros exóticos, cujas populações crescentes ameaçam a biota nativa, se tornaram o item mais frequente da dieta das onças pardas nessa região. Em áreas com onças pardas, os burros mudaram seus padrões espaço-temporais de forrageio, tornando-se diurnos e menos ativos. Por outro lado, os burros permanecem mais ativos e forrageiam durante o dia e à noite em áreas sem onças pardas. A interação entre onças pardas e burros desencadeou uma cascata trófica comportamentalmente mediada que emerge de uma "paisagem de medo", isto é, da percepção dos burros sobre a heterogeneidade espacial do risco de predação. Áreas com onças pardas tiveram menos solo pisoteado, menos trilhas de burros, e muito mais cobertura de dossel e vegetação no entorno da água. Os burros concentram a maior parte de suas atividades em terrenos topologicamente planos, os quais não apresentam lugares apropriados para o comportamento de emboscada das onças pardas e têm presença humana mais intensa, provavelmente agindo como escudos contra predadores. Lundgren et al. apresentam um sistema modelo promissor para o estudo de efeitos de pastadores temorosos sobre a estrutura comunitária no contexto de novas interações ecológicas em estabelecimento no Antropoceno. Se a cascata trófica emergente é transiente ou persistente, os papéis relativos dos efeitos diretos e indiretos como mecanismos subjacentes, e suas consequências para a estrutura, funcionamento e estabilidade da teia trófica são questões de interesse geral. Respondê-las pode nos ajudar a elucidar os custos e benefícios de usar mega-herbívoros não-nativos na restauração funcional de ecossistemas permanentemente invadidos.

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Clustering of species with similar niches or traits occurs in communities, but the mechanisms behind this pattern are still unclear. In the emergent neutrality model, species with similar niches and competitive ability self-organise into clusters. In the hidden-niche model, unaccounted-for niche differences stabilise coexistence within clusters. Finally, clustering may occur through alliances of species that facilitate each other. We tested these hypotheses using population-growth models that consider interspecific interactions parameterised for 35 species using field data. We simulated the expected community dynamics under different species-interaction scenarios. Interspecific competition was weaker within rather than between clusters, suggesting that differences in unmeasured niche axes stabilise coexistence within clusters. Direct facilitation did not drive clustering. In contrast, indirect facilitation seemingly promoted species alliances in clusters whose members suppressed common competitors in other clusters. Such alliances have been overlooked in the literature on clustering, but may arise easily when within cluster competition is weak.

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Predators can influence prey through direct consumption as well as through non-consumptive effects (NCEs). NCEs usually occur mediated by behavioral changes in the prey upon detection of predator cues. Such changes may involve reduction of feeding with a variety of physiological consequences. We evaluated NCEs from an intertidal predatory snail (Acanthina monodon) on a dominant habitat-forming mussel species (Perumytilus purpuratus) from the southeastern Pacific coast. We tested whether A. monodon exerts negative NCEs on clearance rate, oxygen consumption rate, biodeposit production, and between-valve gap size in P. purpuratus. We found that waterborne predator cues triggered a decrease in these variables except biodeposit production. However, the organic content of the biodeposits increased in the presence of predator cues. The snail's physical contact with the mussels strengthened the negative NCEs on between-valve gap size. Since P. purpuratus is a dominant filter-feeder and foundation species in rocky intertidal habitats, predator NCEs on this species might indirectly influence ecosystem-level processes and community structure.

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Host plant selection by herbivores is driven by a complex array of cues, including leaf traits and previous leaf damage. Herbivore-associated cues to host selection at the plant and leaf scale aid understanding of mechanisms responsible for host preference that might translate into increased performance, as well as processes structuring herbivore populations mediated by interactions. We investigated how changes induced by a galling insect in the tropical fern Cyathea phalerata act as repellent or attractant cues for sawfly feeding and the effects of leaf size on herbivory levels. We recorded gall abundance, damage by chewers, leaf size, plant nutritional quality, phenolic concentration and leaf anatomical traits between galled and non-galled leaf samples. Galled samples contained less N, higher levels of phenolics and higher C/N ratio. However, leaf-chewing damage did not differ between galled and non-galled leaves. The gall structure was avoided by chewers, as it had high concentrations of phenolics, lignification and suberization. Larger leaves sustained higher gall abundance, but leaf size did not have a significant effect on chewer damage. A co-occurrence index calculated for both guilds indicated that galls and chewers exhibited a distribution that did not differ from random, reinforcing that the two guilds on C. phalerata do not show patterns of repulsion such as those maintained by interspecific competition. Sawflies dismissing chemical cues indicate that the increase in phenolics caused by galling insects does not generate increased protection of the galled pinnules. Our results highlight ferns as key resources for herbivores and as a potential plant group to study new research avenues on plant-insect interactions.

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The biotic resistance hypothesis asserts that native species may hinder the invasion of exotic species, which can occur either directly or indirectly by influencing interactions between exotic and local species. Aphid-tending ants may play a key role in the indirect biotic resistance to plant invasion. Ants may protect aphids, thus increasing their negative effect on exotic plants, but may also deter chewing herbivores, thus benefiting exotic plants. We studied native aphid-tending ants (Dorymyrmex tener, Camponotus distinguendus, and Dorymyrmex richteri) on exotic nodding thistles (Carduus thoermeri), which are attacked by thistle aphids (Brachycaudus cardui) and thistle-head weevils (Rhinocyllus conicus). We evaluated the impact of ants, aphids, and weevils on thistle seed set. We compared ant species aggressiveness towards aphid predators and weevils and performed ant-exclusion experiments to determine the effects of ants on aphid predators and weevils. We analysed whether ant species affected thistle seed set through their effects on aphids and/or weevils. The ant D. tener showed the most aggressive behaviour towards aphid predators and weevils. Further, D. tener successfully removed aphid predators from thistles but did not affect weevils. Excluding D. tener from thistles increased seed set. Analyses supported a negative indirect pathway between the aggressive D. tener and thistle seed set through aphid populations, while the other ant species showed no indirect effects on thistle reproduction. Therefore, aggressive aphid-tending ants may enhance biotic resistance by increasing aphid infestation on exotic invasive plants. This study highlights the importance of indirect biotic resistance in modulating the success of invasive species.

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Introduction: The enemy hypothesis postulates that gall traits protect galling insects against natural enemies. Galls show a huge range of sizes, colors and ornaments, which vary even intraspecifically. However, galling insects are targets of various organisms that attack them directly or indirectly. In this context, to consider only one gall trait to investigate gall structure acting against only one guild of enemies can conceal an understanding of the community-level interactions. Objective: Herein, we take these ideas into consideration to investigate the conspicuous galls induced by Palaeomystella oligophaga Becker and Adamski 2008 (Lepidoptera) on Macairea radula (Bonpl.) (Melastomataceae) as a model system. We characterize this system through categorization of the different enemy guilds present in the community. We identified them to the lowest taxonomic level possible and determined the kind of interaction responsible for galling insects' deaths. Considering the enemy hypothesis and the selection of secondary characteristics, we also aimed to determine which of the multiple gall traits influence the survival success of galling insects in a multi-enemy context. Methods: We inspected galls and characterized the enemy guilds affecting the galling insect and the mortality rates produced by each one of them. Next, we tested whether the distinct gall traits measured (parenchyma thickness, color, projections) promote galling insect survival with respect to each enemy. Results: The mortality induced by indirect enemies (organisms that interact with gall tissues and can interact secondarily with galling insect) was 47.3 %, being higher than that caused by parasitoids and predators (31.5 %). Despite the gall's structural complexity, live galling insects showed the smallest occurrence (21.2 %). Parenchyma thickness was negatively related to Calliephialtes parasitoids, Gelechiidae cecidophages and predation signals. Conclusions: We demonstrated that the attacks to gall tissues by the cecidophages represented the highest threat to P. oligophaga survival, being higher than the mortality caused by direct enemies. That is, the gall traits were not as efficient as supposed to protect the galling insect from the attack of natural enemies. Nevertheless, we also demonstrated that parenchyma thickness can be negatively related to some organisms, especially direct enemies. Other traits hypothesized as defensive (e.g. projections, coloration) may simply play no role.

Introducción: La hipótesis del enemigo postula que las características de la agalla protegen al agallero contra los enemigos naturales. Las agallas muestran una gran variedad de tamaños, colores y adornos, que varían incluso de forma intraespecífica. Sin embargo, los agalleros son objetivos de varios organismos que los atacan directa o indirectamente. En este contexto, considerar solo una característica para investigar la estructura de la agalla actúando contra un solo gremio de enemigos puede ocultar una comprensión de las interacciones a nivel comunitario. Objetivos: Para investigar las ideas presentadas usamos las agallas conspicuas inducidas por Palaeomystella oligophaga Becker y Adamski 2008 (Lepidoptera) en Macairea radula (Bonpl.) (Meslastomataceae) como sistema modelo. Describimos este sistema a través de la categorización de los diferentes gremios enemigos presentes en la comunidad. Los identificamos al nivel taxonómico más bajo posible y determinamos el tipo de interacción responsable de la muerte de los agalleros. Teniendo en cuenta la hipótesis del enemigo y la selección de características secundarias, también buscamos determinar cuáles de las múltiples caracteristicas de la agalla influyen en el éxito de supervivencia de los agalleros en un contexto de enemigos múltiples. Métodos: Inspeccionamos las agallas y caracterizamos los gremios enemigos que afectan al agallero y las tasas de mortalidad producidas por cada uno de ellos. Luego, probamos si las distintas caracteristicas de las agallas medidas (grosor del parénquima, color, proyecciones) promueven la supervivencia de los agalleros con respecto a cada enemigo. Resultados: La mortalidad indirecta inducida por los cecidofagos fue del 47.3 %, superior a la causada por los parasitoides y los depredadores (31.5 %). Apesar de la complejidad estructural de la agalla, los agalleros vivos mostraron la menor presencia (21.2 %). El grosor del parénquima se relacionó negativamente con los parasitoides de Calliephialtes, los cecidófagos de Gelechiidae y la depredación. Conclusiones: Demostramos que los ataques a los tejidos biliares por cecidófagos representaron la mayor amenaza para la supervivencia de P. oligophaga. Esto es interesante porque, de acuerdo con la hipótesis del enemigo, la estructura de las agallas debería proporcionar protección para los insectos agalleros en lugar de atraer a los insectos que se alimentan de la agalla misma. Sin embargo, también demostramos que el grosor del parénquima puede estar relacionado negativamente con algunos organismos, especialmente los enemigos directos. Otras caracteristicas hipotéticas como defensivas (por ejemplo, proyecciones, coloración) pueden simplemente no desempeñar ningún papel.

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Theory suggests that non-trophic interactions can be a major mechanism behind community stability and persistence, but community-level empirical data are scarce, particularly for effects on species interactions mediated through changes in the physical environment. Here, we explored how ecosystem engineering effects can feed back to the engineer, not only modulating the engineer's population density (node modulation) but also affecting its interactions with other species (link modulation). Gall induction can be viewed as ecosystem engineering since galls serve as habitat for other species. In a community-level field experiment, we generated treatments with reduced or elevated ecosystem engineering by removing or adding post-emergence galls to different plots of their host plant in the Brazilian Cerrado. We tested the effect of post-emergence galls on the galler, as well as on the galler-parasitoid and galler-aphid interactions. The manipulation of post-emergence galls had little effect on the galler-abundance and survivorship were not affected, and gall volume changed only slightly-but modified interactions involving the galler, parasitoid wasps and inquiline aphids. Aphid inquilines negatively affected density-dependent parasitism rates (interaction modification) likely by killing parasitised galling larvae. Post-emergence galls interfered with aphid inquilinism-likely by the provision of alternative habitat for aphids-and thus interfered with the negative effect of aphids on parasitism (modification of an interaction modification). This work is one of the few studies to demonstrate experimentally the role played by environment-mediated interaction modification at a community level in the field. Moreover, by manipulating a species' ecosystem engineering effect (post-emergence galls) instead of the species itself, we demonstrate the novel result that populations can be regulated by non-trophic effects initiated by their own activities that alter their interaction with other species. This reveals that indirect interactions mediated via the environment offer new pathways of feedback loops for population regulation. Our results indicate that interaction modification has the potential to be a key regulatory mechanism underlying interaction variation in nature, and play a major role in community structure, dynamics and stability.

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Over the past few years, our knowledge of how ecological interactions shape the structure and dynamics of natural communities has rapidly advanced. Plant chemical traits play key roles in these processes because they mediate a diverse range of direct and indirect interactions in a community-wide context. Many chemically mediated interactions have been extensively studied in industrial cropping systems, and thus have focused on simplified, pairwise and linear interactions that rarely incorporate a community perspective. A contrasting approach considers the agroecosystem as a functioning whole, in which food production occurs. It offers an opportunity to better understand how plant chemical traits mediate complex interactions which can enhance or hinder ecosystem functions. In this paper, we argue that studying chemically mediated interactions in agroecosystems is essential to comprehend how agroecosystem services emerge and how they can be guaranteed through ecosystem management. First, we discuss how plant chemical traits affect and are affected by ecological interactions. We then explore research questions and future directions on how studying chemical mediation in complex agroecosystems can help us understand the emergence and management of ecosystem services, specifically biological control and pollination.

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Predators that forage at boundaries between ecosystems can affect prey from adjacent ecosystems, thereby triggering consumptive and non-consumptive cascading effects, which may affect diversity and food web structure across ecosystems. In the present study, we manipulated the access of insectivorous birds, lizards, and anurans to tank bromeliads in scrub vegetation in southern Brazil. We measured cascading effects on the community structure of aquatic invertebrates inhabiting bromeliad leaves and on the ecosystem processes of decomposition rate and bromeliad growth. The exclusion of terrestrial vertebrate predators increased the biomass of Odonate and Tabanid apex predators, which shifted the body size structure of the assemblage and generated inverted biomass pyramids that were top-heavy. Within bromeliads with larger aquatic predators, the species composition and abundance of other aquatic invertebrates also changed, resulting in higher abundance of mesopredators and scrapers, and lower abundance of shredders. Under those conditions, the detritus decomposition rate decreased, and bromeliads produced more leaves, perhaps because of the higher deposition of nitrogenous waste by mesopredators. Our results highlight that the effects of terrestrial vertebrate predators can propagate across aquatic ecosystems, altering species composition, body size structure, food web organization, and ecosystem function.

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Predators control prey populations and influence communities and the functioning of ecosystems through a combination of consumptive and nonconsumptive effects. These effects can be locally confined to one ecosystem but can also be extended to neighboring ecosystems. In this study, we investigated the nonconsumptive effects of terrestrial avian predators on the communities of aquatic invertebrates inhabiting bromeliads and on the functioning of these natural ecosystems. Bromeliads with stuffed birds placed nearby showed a decrease in aquatic damselfly larvae abundance and biomass, and we can infer that these changes were caused by antipredator responses. These larvae, which are top predators in bromeliad ecosystems, changed the composition of the entire aquatic invertebrate community. While total species richness, mesopredator richness, and shredder abundance increased in the presence of birds, scraper biomass decreased, possibly as a consequence of the increase in mesopredator richness. High scraper biomass in the absence of birds may have accelerated detrital decomposition, making more nutrients available for bromeliads, which grew more. These results show that nonconsumptive effects triggered by terrestrial predators can cascade down to lower trophic levels and dramatically affect the functioning of aquatic ecosystems, which can in turn alter nutrient provision to terrestrial ecosystems.

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Studies on the robustness of ecological communities suggest that the loss or reduction in abundance of individual species can lead to secondary and cascading extinctions. However, most such studies have been simulation-based analyses of the effect of primary extinction on food web structure. In a field experiment we tested the direct and indirect effects of reducing the abundance of a common species, focusing on the diverse and self-contained assemblage of arthropods associated with an abundant Brazilian shrub, Baccharis dracunculifolia D.C. (Asteraceae). Over a 5-month period we experimentally reduced the abundance of Baccharopelma dracunculifoliae (Sternorrhyncha: Psyllidae), the commonest galling species associated with B. dracunculifolia, in 15 replicate plots paired with 15 control plots. We investigated direct effects of the manipulation on parasitoids attacking B. dracunculifoliae, as well as indirect effects (mediated via a third species or through the environment) on 10 other galler species and 50 associated parasitoid species. The experimental manipulation significantly increased parasitism on B. dracunculifoliae in the treatment plots, but did not significantly alter either the species richness or abundance of other galler species. Compared to control plots, food webs in manipulated plots had significantly lower values of weighted connectance, interaction evenness and robustness (measured as simulated tolerance to secondary extinction), even when B. dracunculifoliae was excluded from calculations. Parasitoid species were almost entirely specialized to individual galler species, so the observed effects of the manipulation on food web structure could not have propagated via the documented trophic links. Instead, they must have spread either through trophic links not included in the webs (e.g. shared predators) or non-trophically (e.g. through changes in habitat availability). Our results highlight that the inclusion of both trophic and non-trophic direct and indirect interactions is essential to understand the structure and dynamics of even apparently discrete ecological communities.

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Ecosystem engineering is increasingly recognized as a relevant ecological driver of diversity and community composition. Although engineering impacts on the biota can vary from negative to positive, and from trivial to enormous, patterns and causes of variation in the magnitude of engineering effects across ecosystems and engineer types remain largely unknown. To elucidate the above patterns, we conducted a meta-analysis of 122 studies which explored effects of animal ecosystem engineers on species richness of other organisms in the community. The analysis revealed that the overall effect of ecosystem engineers on diversity is positive and corresponds to a 25% increase in species richness, indicating that ecosystem engineering is a facilitative process globally. Engineering effects were stronger in the tropics than at higher latitudes, likely because new or modified habitats provided by engineers in the tropics may help minimize competition and predation pressures on resident species. Within aquatic environments, engineering impacts were stronger in marine ecosystems (rocky shores) than in streams. In terrestrial ecosystems, engineers displayed stronger positive effects in arid environments (e.g. deserts). Ecosystem engineers that create new habitats or microhabitats had stronger effects than those that modify habitats or cause bioturbation. Invertebrate engineers and those with lower engineering persistence (<1 year) affected species richness more than vertebrate engineers which persisted for >1 year. Invertebrate species richness was particularly responsive to engineering impacts. This study is the first attempt to build an integrative framework of engineering effects on species diversity; it highlights the importance of considering latitude, habitat, engineering functional group, taxon and persistence of their effects in future theoretical and empirical studies.

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