Long-term storage shapes ejaculate traits in a monogamous stingless bee ( Scaptotrigona aff. depilis )

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Original article

Long-term storage shapes ejaculate traits in a monogamous stingless bee (Scaptotrigona aff. depilis ) Sheina KOFFLER1,2 , Andreas HOPPE3 , Kaspar BIENEFELD3 , Astrid de Matos Peixoto KLEINERT1 , Rodolfo JAFFÉ1,4 1 Department of Ecology, University of São Paulo, Rua do Matão, 321, São Paulo, SP 05508-090, Brazil Institute of Advanced Studies, University of São Paulo, Rua Praça do Relógio, 109, São Paulo, SP 05508-970, Brazil 3 lnstitute of Bee Research, Friedrich Engels Str. 32, 16540, Hohen Neuendorf, Germany 4 Instituto Tecnológico Vale, Rua Boaventura da Silva, 955, Belém, PA 66055-090, Brazil

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Received 8 February 2020 – Revised 3 June 2020 – Accepted 29 September 2020

Abstract – The influence of long-term storage on the evolution of social insect male ejaculates remains a puzzle. Here, we studied the influence of sperm storage on male ejaculates in the stingless bee Scaptotrigona aff. depilis , a species where queens mate with a single male and thus do not trigger sperm competition. We estimated quantitative genetic parameters by sampling males from six colonies and two consecutive generations (n = 172). Heritability for sperm length was lower than heritability estimates for other traits. Coefficients of additive genetic variance (CVa) suggest sperm viability and sperm counts are under directional selection, sperm length under stabilizing selection, and male size under weak selection. Both genetic and phenotypic correlations were weak and not significant, indicating independent trait evolution. Overall, our findings provide evidence of selection acting on social insect ejaculates and bring insights for stingless bee breeding programs. Breeding / Heritability / Sexual selection / Hymenoptera

1. INTRODUCTION The evolution of social insect male traits remains poorly understood, as few studies have assessed how sexual selection operates in this group (Abell et al. 1999; Boomsma et al. 2005; Couvillon et al. 2010; Jaffé and Moritz 2010; Koffler et al. 2016). Earlier attempts have been made to understand how competition for mating shapes male morphology (Abell et al. 1999, Couvillon et al. 2010). However, ejaculate Electronic supplementary material The online version of this article (https://doi.org/10.1007/s13592-020-00813-x) contains supplementary material, which is available to authorized users.

Corresponding author: S. Koffler, [email protected] Handling Editor: Mathieu Lihoreau

evolution is expected to be under more specific selective pressures. Males of the haplodiploid social Hymenoptera (ants, bees, and wasps) originate from unfertilized eggs, so to achieve fitness gains, these haploid males must sire reproductive females (queens). This implies that most sperm are used for the production of sterile worker offspring until colonies are large enough to start producing queens (Oster and Wilson 1978). Male fitness gains can also be realized indirectly through the production of males by unmated workers, albeit worker reproduction is rare or absent in some groups (Wenseleers and Ratnieks