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A growing body of literature recognizes that pairwise species interactions are not necessarily an appropriate metaphorical molecule of community ecology. Two examples are intransitive competition and nonlinear higher‐order effects. While these two processes have been discussed extensively, the explicit analysis of how the two of them behave when simultaneously part of the same dynamic system has not yet been explored theoretically. A concrete situation exists on coffee farms in Puerto Rico in which three ant species form an intransitive competitive triplet, and that triplet is strongly influenced, nonlinearly, by a fly parasitoid that modifies the competitive ability of one of the species. Using this arrangement as a template, we explore the dynamical consequences with a simple ordinary differential equation (ODE) model. Results are complicated and include alternative periodic and chaotic attractors. The qualitative structures of those complications, however, may be approximately retrieved from the basic natural history of the system.

The coffee leaf rust disease (CLR), caused by the fungusHemileia vastatrix, is an economically significant phytopathogen of cultivatedCoffeaspecies. Since coffee plants with CLR drop their infected leaves, rain splash from infected leaf litter could be one wayH. vastatrixspores are dispersed, but this mechanism has not been demonstrated. Here we experimentally verify that splash from leaf litter can lead toH. vastatrixdispersal and infection. In a semi-controlled experiment, we compared the infection rates of pairs of potted uninfected, susceptibleC. arabicaplants with leaf litter that was either infected with CLR (experimental treatment) or without CLR (control). Plants with CLR litter became infected sooner than control plants by a median difference of 2.5 weeks. On average, CLR litter treatment plants had 18% of their leaves infected, while control plants had 9% of leaves infected, though later patterns could reflect leaf turnover and reinfection. Future research should investigate the dynamics of leaf litter splash for CLR in the field. Possible management measures to limit the effect of splash from leaf litter could include planting cover crops or restoring natural groundcover, techniques known to provide additional benefits as understood in general practices of agroecology.

Most species are embedded in multi-interaction networks. Consequently, theories focusing on simple pair-wise interactions cannot predict ecological and/or evolutionary outcomes. This study explores how cascading higher-order interactions (HOIs) would affect the population dynamics of a focal species. Employing a system that involves a myrmecophylic beetle, a parasitic wasp that attacks the beetle, an ant, and a parasitic fly that attacks the ant, the study explores how none, one, and two HOIs affect the parasitism and the sex ratio of the beetle. We conducted mesocosm experiments to examine these HOIs on beetle survival and sex ratio and found that the 1st degree HOI does not change the beetle’s survival rate or sex ratio. However, the 2nd degree HOI significantly reduces the beetle’s survival rate and changes its sex ratio from even to strongly female-biased. We applied Bayes’ theorem to analyze the per capita survival probability of female vs. male beetles and suggested that the unexpected results might arise from complex eco-evolutionary dynamics involved with the 1st and 2nd degree HOIs. Field data suggested the HOIs significantly regulate the sex ratio of the beetle. As the same structure of HOIs appears in other systems, we believe the complexity associated with the 2nd degree HOI would be more common than known and deserve more scientific attention.

Natural pest control is an alternative to pesticide use in agriculture, and may help to curb insect declines and promote crop production. Nonconsumptive interactions in natural pest control that historically have received far less attention than consumptive interactions, may have distinct impacts on pest damage suppression and may also mediate positive multipredator interactions. Additionally, when nonconsumptive effects are driven by natural enemy aggression, variation in alternative resources for enemies may impact the strength of pest control. Here we study control of the coffee berry borer (CBB),Hypothenemus hampei, by a keystone arboreal ant species,Azteca sericeasur, which exhibits a nonconsumptive effect on CBB by throwing them off coffee plants. We conducted two experiments to investigate: (1) if the strength of this behavior is driven by spatial or temporal variability in scale insect density (an alternative resource thatAztecatends for honeydew), (2) if this behavior mediates positive interactions betweenAztecaand other ground‐foraging ants, and (3) the effect this behavior has on the overall suppression of CBB damage in multipredator scenarios. Our behavioral experiment showed that nearly all interactions betweenAztecaand CBB are nonconsumptive and that this behavior occurs more frequently in the dry season and with higher densities of scale insects on coffee branches. Our multipredator experiment revealed that borers thrown off coffee plants byAztecacan survive and potentially damage other nearby plants but may be suppressed by ground‐foraging ants. Although we found no non‐additive effects betweenAztecaand ground‐foraging ants on overall CBB damage, together, both species resulted in the lowest level of plant damage with the subsequent reduction in “spillover” damage caused by thrown CBB, indicating spatial complementarity between predators. These results present a unique case of natural pest control, in which damage suppression is driven almost exclusively by nonconsumptive natural enemy aggression, as opposed to consumption or prey behavioral changes. Furthermore, our results demonstrate the variability that may occur in nonconsumptive pest control interactions when natural enemy aggressive behavior is impacted by alternative resources, and also show how these nonconsumptive effects can mediate positive interactions between natural enemies to enhance overall crop damage reduction.