Social benefits require a community: the influence of colony size on behavioral immunity in honey bees

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Apidologie * The Author(s), 2020 DOI: 10.1007/s13592-020-00754-5

Social benefits require a community: the influence of colony size on behavioral immunity in honey bees Rachael E. BONOAN1 , Paola M. IGLESIAS FELICIANO1,2 , Joanna CHANG1,3 , Philip T. STARKS1 1 Department of Biology, Tufts University, 200 College Ave, Medford, MA, USA College of Natural Sciences, Department of Biology, University of Puerto Rico, San Juan, PR 00931, USA 3 Department of Biology, Pomona College, 333 North College Way, Claremont, CA, USA

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Received 22 March 2019 – Revised 14 February 2020 – Accepted 25 February 2020

Abstract – Emergent properties of eusocial insect colonies (e.g., nest architecture and defense) highlight benefits of group living. Such emergent properties, however, may only function as a benefit if the group is large enough. We tested the effect of group size on colony-level fever in honey bees. When a colony is infected with Ascosphaera apis , a heat-sensitive brood pathogen, adult bees raise the temperature to kill the pathogen and keep brood disease free. In relatively large colonies, we show a rhythm to honey bee fever: colonies inoculated with A . apis generated a fever in the afternoon and at night but not in the morning. In comparison, relatively small colonies did not generate a fever following inoculation, although they invested more in thermoregulation on a “per bee” basis than control colonies. Thus, in small colonies, honey bee fever could be regarded as a cost of group living: individuals futilely exerted valuable energy towards fighting a pathogen. group size constraints / behavioral immunity / thermoregulation / sociality

1. INTRODUCTION Group living comes with costs and benefits (Wilson 1975). Some of the most successful group-living organisms on earth are social insects, which exhibit a range of social organization (Wilson 1975). At the pinnacle of social evolution and cooperation are the eusocial insects: individuals work so well together en masse ; they can be called a superorganism (Wheeler 1910). In order to successfully function as a superorganism, however, the colony likely needs to host a large enough Electronic supplementary material The online version of this article (https://doi.org/10.1007/s13592-020-00754-5) contains supplementary material, which is available to authorized users.

Corresponding author: R. Bonoan, [email protected] Manuscript editor: James Nieh

workforce. This is likely the case when it comes to group-level defense, such as social immunity. For our purposes, we use Meunier (2015)’s definition of social immunity: any collective or individual mechanism that has emerged and/or is maintained at least partly due to the anti-parasite defense it provides to other group members. Many studies have examined the effect of group size on survival and social immunity in ants and termites. For example, following fungal infection, dampwood termite survival is lower in isolated nymphs than those in groups (Rosengaus et al., 1998). Accordingly, infected dampwood termites show increa