Consistent with studies affirming the influence of vegetation connectivity on predatory arthropod movement and predation range, our results illustrate how vegetation connectivity facilitates A. sericeasur foraging mobility and pest removal. In coffee systems, higher degrees of vegetation connectivity are associated with shade trees, as well as more heterogeneous habitat complexity and variability in plant structure. In other studies, ants generally increase predation services in shaded systems as compared to monocultures and, in coffee plants, more effectively remove CBB in shaded coffee systems as compared to sun monoculture systems. Interestingly, most studies find the opposite effect of structural complexity on parasitoid behavior, with higher degrees of plant structural complexity leading to decreased parasitoid foraging efficiency. This negative relationship between parasitism and habitat complexity transfers to coffee systems, where the parasitic phorid flies exert a greater inhibiting effect on Azteca ants in simple, low-shade farms than in complex, high-shade farms. Together with the aforementioned study, our combined results illustrate how habitat complexity at the landscape scale and vegetation connectivity at the plot scale dually facilitate A. sericeasur-mediated pest removal: by facilitating ant mobility and by reducing the efficiency of the parasitoid that interferes with their pest removal ability. In order for A. sericeasur to provide ant-mediated pest removal services, coffee agroforests must include enough shade trees to provide sufficient habitats for ant nests. Planting coffee plants close enough to shade trees to allow for direct connectivity and leaving some vegetation connections between coffee plants and shade trees rather than chopping them or relying on herbicides can facilitate ant-provided ecosystem services by providing foraging paths through naturally occurring structural connectivity.

By enhancing the A. sericeasur effectiveness in controlling CBB populations, vegetation connectivity can potentially reduce chemical pesticide use. Our results offer management insight into one piece of a complex ecological puzzle. Because A. sericeasur tend C. viridis, square black flower bucket they could indirectly reduce coffee plant growth by contributing to high-scale densities and an associated damaging sooty mold. However, high densities of C. viridis also beneficially attract Lecanicillium lecanii, which attacks coffee leaf rust, a devastating coffee fungal disease. Moreover, the CBB is regarded as a far more damaging coffee pest than C. viridis. Furthermore, facilitating the mobility of A. sericeasur as a single ant species is not necessarily the most effective pest management approach, as higher ant diversity can improve pest control through the cooperation of complementary predatory species. Enhanced A. sericeasur activity on coffee plants could alter the behavior of other ant species, which could have positive or negative effects on overall pest control services due to spatial complementarity or potential negative interactions between predators. However, studies find that increasing connectivity generally increases species richness, and so, vegetation connections that increase A. sericeasur mobility likely facilitate the mobility of other predatory ants in coffee systems, even by providing alternative paths to avoid aggressive altercations with A. sericeasur. Although A. sericeasur occupies only 3–5% of the shade trees at our research site, other ants known to contribute to CBB regulation would likely also use vegetation pathways, facilitating additional pest control. Future research should examine how vegetation connectivity impacts the abundance and diversity of other ant species on coffee plants and the associated spatial complementarity between specific predators of the CBB. Future studies could also investigate how phorid attacks on Azteca vary on different foraging pathways to better understand the mechanisms behind their preference for vegetation pathways.

Connectivity affects arboreal ant distribution, behavior, and interactions with other organisms in agroecosystems, profoundly impacting ant community diversity and ant provided ecosystem services. Our results demonstrate how vegetation connectivity increases A. sericeasur activity, recruitment to resources, and CBB removal, and that naturally occurring vegetation connectivity, in the form of branches and natural substrates, accounts for this enhancement. As climate change increases coffee’s susceptibility to CBB damage, agroecological and economically feasible forms of pest control are increasingly necessary for coffee-producing communities. Farm management conducive to forest conservation, habitat and structural complexity, and the associated higher degrees of vegetation connectivity will facilitate ant-provided pest control services in coffee agroecosystems.Wild birds provide many ecosystem services that are economically, ecologically, and culturally important to humans . On a global scale, insectivorous birds consume an estimated 400–500 million tons of insects annually and have the capacity to decrease arthropod populations and increase crop yields of both temperate and tropical farms . While these beneficial effects are not always observed , attention has focused on promoting avian diversity and abundance on farms to leverage these benefits . The fact that birds consume agricultural pests does not ensure that they can control them, in the sense of substantially reducing densities of rapidly-growing pests. Here, we evaluate the capacity of birds to suppress agricultural pests, specifically the coffee berry borer, aninvasive pest found in almost every coffee-producing region worldwide. The coffee berry borer is one of the most economically significant pests of coffee worldwide , causing an estimated annual global loss of US $500 million . These small beetles damage coffee crops when a female bores into a coffee cherry and excavates chambers for larvae to grow, consuming the coffee bean. Control of CBB can be accomplished by spraying fungal bioinsecticide Beauvaria bassinia, increasing harvest frequency or continually removing, by hand, over-ripe and fallen cherries, which serve as reservoirs for infestations .

The last, and most laborious, control method appears to be the most economically effective In addition to human-mediated control, natural predators such as ants, parasitoid wasps, and nematodes are being explored as potential bio-control agents . Birds have also been identified as a significant biological control agent of CBB . Field experiments in Central America have shown that CBB infestation dramatically decreases when birds are present . For example, Karp et al. reported that bird predation suppresses CBB infestation by 50% and saves farmers US $75– 310/ha per year; another estimate values bird predation at US $584/ha . Suppression is done by both resident foliage-gleaning insectivores, such as rufous-capped warblers , and Neotropical migrants like the yellow warbler . Similar to other agriculture systems, avian abundance is higher on farms with heterogenous landscapes in close proximity to native habitat , suggesting low-intensity shade coffee farms are better not only for supporting biodiversity, but also in providing pestmediating ecosystem services . Several lines of evidence support the notion that birds depredate CBB in coffee plantations, and that their effects are biologically significant. Firstly, we know that a variety of bird species consume CBB from assays of avian fecal and regurgitant samples , though the detection rate is quite low . Low detection rates might be due to low consumption rates; detectability of DNA in feces depends on number of CBB eaten, and time since feeding, as well as fecal mass . Secondly, bird and bat exclosure experiments are associated with greater CBB infestation within enclosures . At the same time, it is not clear how birds can effectively suppress CBB at most sites, and throughout the season. Exclosure experiments that report avian suppression appear to be at sites with relatively low CBB infestations , whereas coffee-producing regions with more recent introduction of CBB have infestations of up to 500,000 CBB in a season . We also do not know whether suppression is effective throughout the reproductive cycle of the CBB, or just when abundances are relatively low. Finally, CBB field traps often capture large numbers of CBB, even in the presence of birds . Consequently, while there is clear evidence that birds consume CBB, the degree to which CBB populations can be suppressed is less clear, particularly because of the species’ population growth potential . Here, square black flower bucket wholesale we use a CBB population growth model to assess the capacity of birds at naturally occurring densities to reduce CBB populations, as a function of a starting infestation size. We created an age-based population growth model for CBB using data from a life-stage transition matrix published by Mariño et al. . We converted their matrix into a female-only, daily time-step, deterministic Leslie matrix; we could not estimate population growth directly from the original matrix because it did not use a common time step . We incorporated a skewed adult sex ratio to mimic real populations , and added a life-stage for dispersing females, the stage at which CBB are vulnerable to predation by birds. Since the entire CBB lifecycle occurs within the coffee cherry, CBB are vulnerable to predation by birds for a short time window when adult females disperse between plants and burrow into a new cherry .

Birds do not eat coffee cherries, with the exception of the Jacu , which is found in southeastern South America. Consequently, we assumed that only adult CBB females are vulnerable to bird predation. With our Leslie matrix, we projected population growth for a closed population during a single CBB breeding season. We projected growth at three levels of initial starting populations of CBB , calculated from published estimates of CBB densities from alcohol lure traps in coffee farms from Colombia, Hawaii and Costa Rica. We then determined the degree to which dispersing female survival rate would have to be decreased to result in a 50% depression in the adult population size at the end of the coffee season at all three infestation levels. Finally, we assessed the plausibility of this degree of CBB suppression by birds as a function of avian energy requirements, reported avian densities on coffee farms, prey composition of avian diets, estimated caloric value of CBB, and the starting population size of CBB females.Coffee phenology is directly related to rainfall patterns that differ among coffee producing regions, leading to distinct seasons, and timing of harvest. Our model assumes environmental conditions of Costa Rica, and thus describe the coffee phenology of this region. In regions of Costa Rica with marked seasonality, coffee flowering is triggered during the dry to wet season transition by the onset of acute precipitation . Areas with relatively consistent rain patterns have more continuous flowering events and a longer harvest season In the Central Valley of Costa Rica, flowering typically begins in March, with three flowering events spread over a month . Flowers are short-lived, lasting only a few days before fruit begin to develop. Maturation of coffee cherries is slow, with immature green cherries taking up to 240 days to develop into red, ripe fruit that is ready for harvest in mid-October through January . After harvest, coffee plants are left to recuperate until flowering is initiated again the following year by the next onset of rain.Following the coffee flowering period and initiation of cherry growth, adult female CBB emerge and disperse via flight in search of new cherries to colonize . Timing of emergence appears to be driven primarily by relative humidity and temperature, with dispersal peaks occurring around the end of the coffee harvest, from December through March . Females begin ovipositing in chambers carved out of the coffee endosperm roughly 120–150 days after coffee flowering, when the dry content of the seed is 20% or higher . It is this dispersal period, and subsequent drilling into the coffee cherry, when CBB are vulnerable to predation by birds, as the remainder of the CBB life cycle occurs within the coffee cherry. There are five main CBB developmental stages: egg, larva, pupa, juvenile, and adult. Females can oviposit daily for up to 40 days, averaging 1–2 eggs per day . After a week, eggs hatch and larva take 17 days to develop into pupa. Following pupation , juveniles emerge and reach sexual maturity after about 4 days . The length of the CBB life cycle can be slowed and accelerated depending on average temperature ; the developmental times used here are based on 25 C rearing conditions . Offspring sex ratio is skewed toward females, ranging from 1:5 to 1:494 . Since males are flightless, mating occurs between siblings within the natal cherry. Fertilized females then disperse to colonize other cherries, though multigenerational oviposition within the natal cherry is possible. The prolonged maturation of the coffee crop allows continual reproduction, with 2–8 CBB generations feasible in a single season if environmental conditions and food availability be favorable . With the removal of cherries during harvest, adult CBB will enter diapause in coffee cherries that remain on the plant or fall to the ground .