Mutualism between armoured scale insects and ants: new species and observations on a unique trophobiosis (Hemiptera: Diaspididae; Hymenoptera: Formicidae: Melissotarsus Emery)

The association between African armoured scale insects (Hemiptera: Coccoidea: Diaspididae) and ants belonging to Melissotarsus Emery (Hymenoptera: Formicidae: Myrmicinae) is the only trophobiosis known in which ants do not receive honeydew or nectar in exchange for protection and other services. Food reward for the ants in this mutualism remains unknown, despite repeated suggestions that diaspidids are consumed by the associated ants, thus serving as ‘domestic cattle’. We describe new observations on interactions between Melissotarsus emeryi Santschi and the diaspidid Morganella conspicua (Brain) from South Africa. Worker ants exhibited previously undescribed tending behaviours, most notably a ‘squeezing and licking’ performed on an adult female diaspidid and ‘culling’, in which a worker removed an adult female armoured scale from the host plant. These could represent the gathering of secretory products and the cultivation of an individual for consumption, respectively. An ant exclusion study over 12 days of isolation showed that adult female diaspidids and second-instar nymphs secreted no wax or exudates that attending ants would ordinarily collect. Workers of M. emeryi did not defend their nest against invading colonies of Crematogaster and other unidentified ants: we hypothesize that the primary mode of defence is maintenance of isolation within galleries. We describe three new ant-associated diaspidid species: Affirmaspis cederbergensis Schneider sp.n. from South Africa, Diaspis doumtsopi Schneider sp.n. from Cameroon, and Melissoaspis incola Schneider sp.n. from Madagascar. Melissoaspis formicaria (Ben-Dov) comb.n. is transferred from Morganella (Brain). Diagnostic characteristics for Melissoaspis Ben-Dov are revised, and additional taxonomic information defining this genus allows ease of identification. An updated identification key to the species of ant-associated diaspidids is provided