Selection on bristle length has the ability to drive the evolution of male abdominal appendages in the sepsid fly Themira biloba.

Many exaggerated and novel traits are strongly influenced by sexual selection. Although sexual selection is a powerful evolutionary force, underlying genetic interactions can constrain evolutionary outcomes. The relative strength of selection vs. constraint has been a matter of debate for the evolution of male abdominal appendages in sepsid flies. These abdominal appendages are involved in courtship and mating, but their function has not been directly tested. We performed mate choice experiments to determine whether sexual selection acts on abdominal appendages in the sepsid Themira biloba. We tested whether appendage bristle length influenced successful insemination by surgically trimming the bristles. Females paired with males that had shortened bristles laid only unfertilized eggs, indicating that long bristles are necessary for successful insemination. We also tested whether the evolution of bristle length was constrained by phenotypic correlations with other traits. Analyses of phenotypic covariation indicated that bristle length was highly correlated with other abdominal appendage traits, but was not correlated with abdominal sternite size. Thus, abdominal appendages are not exaggerated traits like many sexual ornaments, but vary independently from body size. At the same time, strong correlations between bristle length and appendage length suggest that selection on bristle length is likely to result in a correlated increase in appendage length. Bristle length is under sexual selection in T. biloba and has the potential to evolve independently from abdomen size.

Testing hypotheses in ecoimmunology using mixed models: disentangling hierarchical correlations.

Considerable research in ecoimmunology focuses on investigating variation in immune responses and linking this variation to physiological trade-offs, ecological traits, and environmental conditions. Variation in immune responses exists within and among individuals, among populations, and among taxonomic groupings. Understanding how variation and covariation are distributed and how they differ across these levels is necessary for drawing appropriate ecological and evolutionary inferences. Moreover, variation at the among-individual level directly connects to underlying quantitative genetic parameters. In order to fully understand immune responses in evolutionary and ecological contexts and to reveal phylogenetic constraints on evolution, statistical approaches must allow (co)variance to be partitioned among levels of individual, population, and phylogenetic organization (e.g., population, species, genera, and various higher taxa). Herein, we describe how multi-response mixed-effects models can be used to partition variation in immune responses among different hierarchical levels, specifically within-individuals, among-individuals, and among-species. We use simulated data to demonstrate that mixed models allow for proper partitioning of (co)variances. Importantly, these simulations also demonstrate that conventional statistical tools grossly misestimate relevant parameters, which urges caution in relating ecoimmunological hypotheses to existing empirical research. We conclude by discussing the advantages and caveats of a mixed-effects modeling approach.