Investigating the role of non-helpers in group living thrips.

Behavioural variation among individuals is a hallmark of cooperative societies, which commonly contain breeders and non-breeders, helpers and non-helpers. In some cases, labour is divided with non-breeders "helping." Conversely, in some societies, subordinate non-breeders may not help. These individuals may be (i) an insurance workforce ensuring continuity of help for breeders when other helpers are lost, (ii) conserving energy while waiting to breed themselves or (iii) simply of too poor physiological quality either to help or breed. In the Australian Outback, Acacia thrips Dunatothrips aneurae (Thysanoptera) glue Acacia phyllodes into "domiciles" using silk-like secretions, either alone or cooperatively. Domicile maintenance is important for humidity, so repair can be interpreted as helping. I found that not all females helped to repair damage; some repaired partially or not at all ("non-helpers"). At the same time, some co-foundresses are non- or partially reproductive ("non-breeders"), and their role is currently unknown. I first tested the possibility that helping and breeding are divided, with non-helping females breeding, and non-breeders helping. In a lab experiment, I rejected this idea. Experimentally damaged domiciles were typically repaired by reproductive females, and less so by non- or partially reproductive individuals. To test whether non-helpers are an insurance workforce, I successively removed repairing females and found that non-helping females from the same domicile did not increase effort as a result, rejecting this hypothesis. Then I tested whether non-helpers were conserving energy waiting to breed. In a field experiment, I removed all other females, allowing either a helpful female or a non-helper to "inherit" her domicile. Isolated like this, non-helpers laid very few eggs compared to helpers or naturally occurring single foundresses, despite similar ovarian development. My findings show that labour was not divided: reproduction and helping covaried positively, probably depending on individual variation in female quality and intra-domicile competition. Non-helping females were neither an insurance workforce nor conserving energy waiting to breed. They are likely simply of poor quality, freeloading by benefiting from domicile maintenance by others. I hypothesize they are tolerated because of selection for indiscriminate communal brood care in the form of domicile repair.