Density-dependent outcomes in a digestive mutualism between carnivorous Roridula plants and their associated hemipterans.

Recent studies have shown that mutualisms often have variable outcomes in space and time. In particular, the outcomes may be dependent on the density of the partners with unimodal or saturating outcomes providing stability to the mutualism. We examine density-dependent outcomes of an obligate, species-specific mutualism between a South African carnivorous plant (Roridula dentata) and a hemipteran (Pameridea) that facilitates prey digestion, but also sucks plant sap. Plants occur in sandy, leached, nitrogen-poor soils and have no digestive enzymes to digest prey. Instead they rely on obligately dependent hemipterans to supply nitrogen by digesting prey for them and defecating on their leaves. We documented the densities of Pameridea on Roridula in the field. In the greenhouse, we manipulated the hemipteran densities on Roridula and measured the mean relative growth rates of plants with differing hemipteran densities. Plants exhibited a unimodal response to the density of their mutualist partners. Those with no hemipterans had negative growth rates, suggesting that hemipterans are important in facilitating nitrogen absorption. Plants with intermediate hemipteran densities had positive growth rates but growth rates were negative under very high hemipteran densities. Our research provides support for variable and unimodal outcomes in mutualism. Unimodal outcomes may be particularly important in obligate mutualisms and this is one of the few studied outside of pollinating seed parasite mutualisms. In this system, extrinsic factors such as other predators may affect the mutualism by altering the numbers of hemipterans.