Large herbivores transform plant-pollinator networks in an African savanna.

Pollination by animals is a key ecosystem service1,2 and interactions between plants and their pollinators are a model system for studying ecological networks,3,4 yet plant-pollinator networks are typically studied in isolation from the broader ecosystems in which they are embedded. The plants visited by pollinators also interact with other consumer guilds that eat stems, leaves, fruits, or seeds. One such guild, large mammalian herbivores, are well-known ecosystem engineers5-7 and may have substantial impacts on plant-pollinator networks. Although moderate herbivory can sometimes promote plant diversity,8 potentially benefiting pollinators, large herbivores might alternatively reduce resource availability for pollinators by consuming flowers,9 reducing plant density,10 and promoting somatic regrowth over reproduction.11 The direction and magnitude of such effects may hinge on abiotic context-in particular, rainfall, which modulates the effects of ungulates on vegetation.12 Using a long-term, large-scale experiment replicated across a rainfall gradient in central Kenya, we show that a diverse assemblage of native large herbivores, ranging from 5-kg antelopes to 4,000-kg African elephants, limited resource availability for pollinators by reducing flower abundance and diversity; this in turn resulted in fewer pollinator visits and lower pollinator diversity. Exclusion of large herbivores increased floral-resource abundance and pollinator-assemblage diversity, rendering plant-pollinator networks larger, more functionally redundant, and less vulnerable to pollinator extinction. Our results show that species extrinsic to plant-pollinator interactions can indirectly and strongly alter network structure. Forecasting the effects of environmental change on pollination services and interaction webs more broadly will require accounting for the effects of extrinsic keystone species.

New species of Bacuma Cameron (Hymenoptera: Braconidae: Braconinae) from Kenya and West Darfur with a key to species.

Three new species of the Afrotropical braconine wasp genus Bacuma are described, and biological observations (nectar feeding) by one of them are noted. The new species are: B. kayserae Quicke & Butcher sp. nov. from Kenya, B. madiensis Quicke & Butcher sp. nov. from Uganda and B. darfurensis Quicke & Butcher sp. nov. from Sudan. A group of four large nominal species with red metasomas and finely sculptured tergites (B. granulatus, B. maculipennis, B. rufa and B. whitei) may represent a single widespread species or a pair of species separated by mesoscutum colour, or four separate but morphologically very similar species. However, given the small number of specimens available for study and the poor condition of some of these, including the types, they are not formally synonymised here. A partial key to the species of Bacuma is presented, which recognizes those species that are clearly distinct, including three new species. Interactive Lucid dichotomous and matrix keys are available on www.waspweb.org.