Patterns of sexual dimorphism in the armoured tardigrades

Sexual dimorphism is widespread among animals, with diverse patterns and proposed explanations observed across the Tree of Life. Here we present the first formal analysis of the patterns of sexual dimorphism in body size and cephalic sensory appendages across 40 species (from 10 genera) of armoured tardigrades (Echiniscidae). Phylogenetic signal was found for body size traits and the cephalic papilla relative size, indicating that the association between these traits between the sexes has high evolutionary persistence. The Echiniscidae body size dimorphism is generally female-biased, which would be in accordance with the fecundity hypothesis. No strong evidence of allometric patterns of body size sexual dimorphism was found. In contrast, some of the cephalic appendages show male-biased sexual dimorphism, particularly those that, by being more innervated, are thought to function as chemodetection organs used by males during mate search. The latter is consistent with the sexual selection hypothesis. As the first systematic quantification and analysis of the patterns of sexual dimorphism in the phylum Tardigrada, this study provides important insights into their ecology and evolution, such as corroborating the suggestion that cephalic appendages evolved for mate searching.