Ant–plant–homopteran mutualism: how the third partner affects the interaction between a plant-specialist ant and its myrmecophyte host

Abstract

By estimating relative costs and bene¢ts, we explored the role of the homopteran partner in the protection mutualism between the myrmecophyte Leonardoxa africana T3, the ant Aphomomyrmex afer, and sap-sucking homopterans tended by ants in the tree's swollen hollow twigs. The ants obtain nest sites and food from their host-plant (food is obtained either directly by extra£oral nectar or indirectly via homopterans). Aphomomyrmex workers patrol the young leaves of L. africana T3 and protect them against phytophagous insects. Because ants tended, either solely or primarily, coccids in some trees and pseudococcids in others, we were able to study whether the nature of the interaction was dependent on the identity of the third partner. First, the type of homopteran a¡ects the bene¢ts to the tree of maintaining a large ant colony. Larger colony size (relative to tree size) confers greater protection against herbivory; this relationship is more pronounced for trees whose ants tend pseudococcids than for those in which ants tend coccids. Second, for trees (and associated ant colonies) of comparable size, homopteran biomass was much larger in trees harbouring coccids than in trees with pseudococcids. Thus, the cost to the tree of maintaining ants may be greater when ants are associated with coccids. The net bene¢ts to the plant of maintaining ants appear to be much greater with pseudococcids as the third partner. To explore how the type of homopteran a¡ects functioning of the system, we attempted to determine which of the resources (nest sites, extra£oral nectar, and homopterans) is likely to limit ant colony size. In trees where ants tended coccids, ant-colony biomass was strongly dependent on the number of extra£oral nectaries. In contrast, in trees whose ants tended only pseudococcids, colony biomass was not related to the number of nectaries and was most strongly determined by the volume of available nest sites. We present hypotheses to explain how the type of homopteran a¡ects functioning of this symbiosis, and discuss the implications of our study for the evolutionary ecology of antp lant^homopteran relationships.

Plant-ants feed their host plant, but above all a fungal symbiont to recycle nitrogen

“…Fungi are not the only neglected actors in these systems. Bacteria [40,41] and domatia inhabitants [42] are also important components of these interaction webs. Our study provides a new framework, demonstrating interactions that should stimulate more detailed analyses of food webs in ant -plant mutualisms.…”

The evolution of mutualisms: exploring the paths between conflict and cooperation

“…Several studies have documented that net costs and benefits can vary over relatively short timescales 4,36 resulting from: (1) changes in the presence or abundance of influential third parties 36,37 ; (2) variation in host densities that results in shifts in patterns of transmission 24 ; (3) variation in resource availablility 3,36 ; or (4) variation in physical conditions (Box 1). Furthermore, such studies raise questions concerning the degree of local adaptation in host and symbiont populations, such as whether hosts generally benefit most from local, presumably more highly coadapted symbionts.…”

Extrafloral nectaries indirectly protect small aphid colonies via ant-mediated interactions

“…Ant-tended aphids frequently parasitize plants with EFNs (Buckley, 1983;Gaume et al, 1998;Offenberg, 2000;Engel et al, 2001;Katayama and Suzuki, 2003a;Suzuki et al, 2004), and ant-mediated interactions between aphids and plants are important for plant fitness . With increased aphid density, Vicia plants (V. faba L. and V. angustifolia L.) show decreased ability to attract ants via their EFNs (Katayama and Suzuki, 2003a;Suzuki et al, 2004) because the amount of aphid honeydew is much greater than that of extrafloral nectar (Katayama, unpublished data).…”