title: Non-additive benefit or cost? Disentangling the indirect effects that occur when plants bearing extrafloral nectaries and honeydew-producing insects share exotic ant mutualists.
abstract: OBJECTIVE
  In complex communities, organisms often form mutualisms with multiple different partners simultaneously. Non-additive effects may emerge among species linked by these positive interactions. Ants commonly participate in mutualisms with both honeydew-producing insects (HPI) and their extrafloral nectary (EFN)-bearing host plants. Consequently, HPI and EFN-bearing plants may experience non-additive benefits or costs when these groups co-occur. The outcomes of these interactions are likely to be influenced by variation in preferences among ants for honeydew vs. nectar. In this study, a test was made for non-additive effects on HPI and EFN-bearing plants resulting from sharing exotic ant guards. Preferences of the dominant exotic ant species for nectar vs. honeydew resources were also examined.
  
  
  METHODS
  Ant access, HPI and nectar availability were manipulated on the EFN-bearing shrub, Morinda citrifolia, and ant and HPI abundances, herbivory and plant growth were assessed. Ant-tending behaviours toward HPI across an experimental gradient of nectar availability were also tracked in order to investigate mechanisms underlying ant responses.
  
  
  RESULTS
  The dominant ant species, Anoplolepis gracilipes, differed from less invasive ants in response to multiple mutualists, with reductions in plot-wide abundances when nectar was reduced, but no response to HPI reduction. Conversely, at sites where A. gracilipes was absent or rare, abundances of less invasive ants increased when nectar was reduced, but declined when HPI were reduced. Non-additive benefits were found at sites dominated by A. gracilipes, but only for M. citrifolia plants. Responses of HPI at these sites supported predictions of the non-additive cost model. Interestingly, the opposite non-additive patterns emerged at sites dominated by other ants.
  
  
  CONCLUSIONS
  It was demonstrated that strong non-additive benefits and costs can both occur when a plant and herbivore share mutualist partners. These findings suggest that broadening the community context of mutualism studies can reveal important non-additive effects and increase understanding of the dynamics of species interactions.
authors.affiliation: Department of Biology, North Carolina State University, Raleigh, NC 27607, USA. amy_savage@ncsu.edu
authors.forename: Amy M
authors.initials: AM
authors.lastname: Savage
authors.forename: Jennifer A
authors.initials: JA
authors.lastname: Rudgers
mesh_terms: D000818:Animals; D001522:Behavior, Animal; D017753:Ecosystem; D007313:Insecta / Q000502:physiology*; D058865:Introduced Species; D032066:Morinda / Q000502:physiology*; D057048:Plant Nectar / Q000378:metabolism*; D011156:Population Density; D018947:Samoa; D013559:Symbiosis
chemical_list: D057048:Plant Nectar
keywords: Additive effects; Anoplolepis gracilipes; EFN; Morinda citrifolia; community context; extrafloral nectar; honeydew; indirect effects; invasive ants; mutualism; non-additive benefit; non-additive cost; simultaneous interactions