Pollination limitation despite managed honeybees in South African macadamia orchards

There is growing demand for pollination services in agricultural production, which contrasts with declines of wild and managed pollinator populations. Macadamia (Macadamia integrifolia) is a mass-flowering crop that depends on pollination services and is increasingly cultivated in South Africa. We studied the crop's pollination in South African orchards considering variation in landscape context and the spatial arrangement of managed honeybees (Apis mellifera). We conducted pollination experiments and pollinator observations on macadamia trees along a distance gradient from orchard edges that bordered either near-natural or human-modified habitats. In addition, we mapped position and density of honeybee apiaries at orchard-level. Nut set of macadamia trees strongly relied on animal-mediated pollination: pollinator exclusion reduced the initial nut set (3 weeks after pollination) by 80% and the final nut set (15 weeks after pollination) by 54%. Supplemental hand-pollination of otherwise untreated flowers increased initial and final nut set by 66% and 44%, respectively, indicating substantial pollination limitation. The landscape context only weakly affected pollinator visitation to macadamia trees, with reduced visitation closer to orchard edges bordering human-modified habitats. Furthermore, we observed almost no wild pollinator species. Instead, honeybees constituted 99% of all visits, whereby honeybee visitation rates increased with a tree's connectivity to apiaries. However, neither initial nor final nut was related to visitation rates, and the final nut set was actually reduced where honeybee colony density was high, with a predicted 50% reduction in final nut set between the lowest and highest colony densities. Our study demonstrates a strong pollination limitation in South African macadamia orchards, where managed honeybees fail at delivering the increasing need for pollination services. Indeed, increasing their colony densities may further limit their pollination efficiency. A pollination management that also includes non-Apis managed pollinators and wild pollinators is possibly needed to increase nut set and provide solutions for increasing pollination service demands. In intensive macadamia orchards, this can also necessitate the need for more pollinator-friendly management practices, including habitat restoration and reduced pesticide application.