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IntroductionThe ‘social brain hypothesis’ proposes that brain development (particularly primates) is driven by social complexity, more than group size. Yet, small insects with minute brains are capable of the most complex social organization in animals - which warrants further attention. Research has focused on highly eusocial hymenopterans with extreme caste specialization and very large colony sizes that have passed social evolutionary points of no return. However, facultatively social insects that form small colonies (< 20 individuals) are likely to provide greater insight on brain selection at the origin-point of social group living. MethodsWe undertake the first neurobiological investigation of the facultatively social allodapine bees (Apidae: Xylocopinae: Allodapini), an exploratory study comparing single- and multi-female colonies ofExoneura angophorae. Using volume as a proxy for neural investment, we measured mushroom body calyces, optic lobes, antennal lobes and whole brains of queens, workers, and single-females to test three theories associating brain development with behavior: social brain hypothesis; distributed cognition hypothesis; sensory environment hypothesis. ResultsMushroom bodies were reduced in subordinate workers, but did not differ between queens and single-females. Workers had larger optic lobes than queens, but did not differ from single-females. There were no differences in antennal lobes or whole brain volume. DiscussionSocial caste, rather than multi-female versus single-female nesting, influenced mushroom body volume in this allodapine bee – counter to both social brain and distributed cognition theories and in alignment with halictine and ceratinine bees that also form small facultatively social colonies. Optic lobe enhancement is likely a response to dietary niche requirements for extra-nidal foraging behavior – which may be a highly plastic trait capable of rapid transition among allodapine and ceratinine bees that conforms with ecological intelligence hypotheses. These broad volumetric trends require further investigations on the functional neural circuitry involved in the aforementioned environmental contexts. 

Abstract Many insects show plasticity in the area of the brain called the mushroom bodies (MB) with foraging and social experience. MBs are paired neuropils associated with learning and memory. MB volume is typically greater in mature foragers relative to young and/or inexperienced individuals. Long-term studies show that extended experience may further increase MB volume, but long-term studies have only been performed on non-reproductive social insect workers. Here we use the subsocial beeCeratina calcaratato test the effect of extended foraging experience on MB volume among reproductive females.Ceratina calcaratafemales forage to provision their immature offspring in the spring, and then again to provision their adult daughters in the late summer. We measured the volume of the MB calyces and peduncle, antennal lobes (AL), optic lobes (OL), central complex (CX), and whole brains of three groups of bees: newly emerged females, reproductive females in spring (foundresses), and post-reproductive mothers feeding their adult daughters in late summer. Post-reproductive late summer mothers had smaller MB calyces and ALs than foundresses. Moreover, among late mothers (but not other bees), wing wear, which is a measure of foraging experience, negatively correlated with both MB and OL volume. This is contrary to previously studied non-reproductive social insect workers in which foraging experience correlates postiviely with MB volume, and suggests that post-reproductive bees may reduce neural investment near the end of their lives. 

Abstract Social behavior has been predicted to select for increased neural investment (the social brain hypothesis) and also to select for decreased neural investment (the distributed cognition hypothesis). Here, we use two related bees, the social Augochlorella aurata (Smith) (Hymenoptera: Halictidae) and the related Augochlora pura (Say), which has lost social behavior, to test the contrasting predictions of these two hypotheses in these taxa. We measured the volumes of the mushroom body (MB) calyces, a brain area shown to be important for cognition in previous studies, as well as the optic lobes and antennal lobes. We compared females at the nest foundress stage when both species are solitary so that brain development would not be influenced by social interactions. We show that the loss of sociality was accompanied by a loss in relative neural investment in the MB calyces. This is consistent with the predictions of the social brain hypothesis. Ovary size did not correlate with MB calyx volume. This is the first study to demonstrate changes in mosaic brain evolution in response to the loss of sociality.