Copyedited by: SU The official journal of the Behavioral ISBE Ecology International Society for Behavioral Ecology Behavioral Ecology (2018), 00(00), 1–1. Invited Commentary Reticulitermes termites, the hydrocarbon heneicosane, identified by Queen pheromones, colony odors, and behavioral responses of workers (Funaro et al. 2018), turns out to better science: a comment on Holman also have effects on worker reproduction. Tristram D. Wyatt Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK RANDOMIZED, BLIND, TRIALS AND THE QUEST FOR RELIABLE SCIENCE The response of biomedical and psychological researchers to the QUEEN PHEROMONES “reproducibility crisis” seems to be well ahead of those in ecology and evolution in recognizing potential sources of bias in designing, With the identification of a growing number of queen pheromones running, analyzing, and interpreting experiments, as well as propos- in ants, wasps, bees, and termites in recent years, Holman’s (2018) ing remedies (see e.g. the Catalogue of Bias www.catalogofbias.org). analysis offers a timely stocktake of our current understanding. This Munafò et al. (2017) present a manifesto for more reproduc- paper shows the usefulness of systematic meta-analysis compared with ible behavioral science, starting with the design and conduct of “vote counting” or entirely qualitative reviews. The focus on effect experiments. Randomization and blinding are key requirements, sizes allows the assessment of evidence for recently proposed hypoth- as highlighted by Holman (2018), who observed that effect sizes eses, such as, for example, the interesting experiments which suggest were larger in queen pheromone experiments that were not run that queen pheromones in some ant species are not effective unless blind. Increasing scientific transparency is also important: pre- they are perceived alongside familiar colony odors (reviewed in Smith registration of experiments (to avoid p-Hacking for example) and and Liebig 2017). Holman observes that the small sample sizes in the incentives for open data sharing are starting to be encouraged in experiments do not allow firm conclusions to be drawn. It would repay evolutionary biology and ecology too (Parker et al. 2016; Fidler repetition, if possible, with larger samples designed to take account of et al. 2017). the likely effect sizes (for which Holman (2018) offers estimates). Queen pheromones can be identified because they are consist- Address correspondence to T.D. Wyatt. E-mail: email@example.com. ently carried by queens in different colonies (and have been dem- onstrated to have the effects on worker reproduction). Pheromones Received 23 April 2018; accepted 2 May 2018.; editorial decision 26 April 2018 superimposed on a colony-odor background that varies between col- onies are a feature of social insect pheromones (Wyatt 2010, 2014). doi:10.1093/beheco/ary074 The convergent evolution of queen pheromones in independently Editor-in-Chief: Leigh Simmons evolved eusocial lineages of ants, bees, and wasps is fascinating (see Holman 2018 for references) but the conservation of molecules as queen pheromones in many, but not all, taxa is not unprecedented. REFERENCES In the absence of selection, pheromones need not change. For Fidler F, Chee YE, Wintle BC, Burgman MA, McCarthy MA, Gordon A. example, in aphids, there is no selective advantage in species-specific 2017. Metaresearch for evaluating reproducibility in ecology and evolu- alarm pheromones (if anything, cross-species response is an advan- tion. Bioscience. 67:282–289. tage) and so aphid species across more than 30 genera share the Funaro CF, Boroczky K, Vargo EL, Schal C. 2018. Identification of a queen and king recognition pheromone in the subterranean termite same molecule, (E)-β-farnesene, as their alarm pheromone. Reticulitermes flavipes. Proc Natl Acad Sci USA. 115: 3888–3893. It is possible that queen pheromones in some species, currently Holman L (2018) Queen pheromones and reproductive division of labour: single molecules, will turn out to be multicomponent, even perhaps a meta-analysis. Behav Ecol. with components from a variety of glands. The complexity and Munafò MR, Nosek BA, Bishop DVM, Button KS, Chambers CD, Percie du Sert N, Simonsohn U, Wagenmakers E-J, Ware JJ, Ioannidis JPA. 2017. nonhydrocarbon nature of honeybee queen pheromones does not A manifesto for reproducible science. Nature Human Behaviour. 1:0021. fit with the pattern of other hymenopteran queen pheromones but Parker TH, Nakagawa S, Gurevitch J, IIEE (Improving Inference in examples from around the animal kingdom show that the evolu- Evolutionary Biology and Ecology) workshop participants. 2016. Promoting tion of new pheromones occurs commonly, as demonstrated by the transparency in evolutionary biology and ecology. Ecol Lett. 19:726–728. enormous variety of independently evolved male moth sex phero- Smith AA, Liebig J. 2017. The evolution of cuticular fertility signals in eusocial insects. Curr Opin Insect Sci. 22:79–84. mones (see Wyatt 2014). Wyatt TD. 2010. Pheromones and signature mixtures: defining species-wide A multitude of effects have been ascribed to queen pheromones. signals and variable cues for identity in both invertebrates and vertebrates. Blum noted that pheromone molecules often have multiple func- J Comp Physiol A Neuroethol Sens Neural Behav Physiol. 196:685–700. tions in social insects (“parsimony,” see Wyatt 2014). It will be Wyatt TD. 2014. Pheromones and animal behavior: chemical signals and signatures. 2nd ed. Cambridge: Cambridge University Press. interesting to see if the king and queen recognition pheromone in © The Author(s) 2018. Published by Oxford University Press on behalf of the International Society for Behavioral Ecology. Downloaded from https://academic.oup.com/beheco/advance-article-abstract/doi/10.1093/beheco/ary074/4995894 All rights reserved. For permissions, please e-mail: firstname.lastname@example.org by Ed 'DeepDyve' Gillespie user on 11 July 2018
Behavioral Ecology – Oxford University Press
Published: May 14, 2018
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