Blouch: Bayesian linear Ornstein-Uhlenbeck models for comparative hypotheses

Relationships among species in the tree of life can complicate comparative methods and testing adaptive hypotheses. Models based on the Ornstein-Uhlenbeck process permit hypotheses about adaptation to be tested by allowing traits to either evolve towards fixed adaptive optima (e.g., regimes or niches) or track continuously changing optima that can be influenced by other traits. These models allow estimation of the effects of both adaptation and phylogenetic inertia – resistance to adaptation due to any source – on trait evolution, an approach known as the “adaptation-inertia” framework. However, previous applications of this framework, and most approaches suggested to deal with the issue of species non-independence, are based on a maximum likelihood approach and thus it is difficult to include information based on prior biological knowledge in the analysis, which can affect resulting inferences. Here I present Blouch, (Bayesian Linear Ornstein-Uhlenbeck Models for Comparative Hypotheses), which fits allometric and adaptive models of continuous trait evolution in a Bayesian framework based on fixed or continuous predictors and incorporates measurement error. I first briefly discuss the models implemented in Blouch, and then the new applications for these models provided by a Bayesian framework. This includes the advantages of assigning biologically meaningful priors when compared to non-Bayesian approaches, allowing for varying effects (intercepts and slopes), and multilevel modeling. Validations on simulated data show good performance in recovering the true evolutionary parameters for all models. To demonstrate the workflow of Blouch on an empirical dataset, I test the hypothesis that the relatively larger antlers of larger bodied deer are the result of more intense sexual selection that comes along with their tendency to live in larger breeding groups. While results show that larger bodied deer that live in larger breeding groups have relatively larger antlers, deer living in the smallest groups appear to have a different and steeper scaling pattern of antler size to body size than other groups. These results are contrary to previous findings and may argue that a different type of sexual selection or other selective pressures govern optimum antler size in the smallest breeding groups.

Methods
Tsuboi et al. (2024) collected an expansive dataset on antler size and body size and generated a new Cervidae phylogeny. I supplemented this dataset with data on Cervidae breeding group size from Plard et al. (2011) and Clutton-Brock et al. (1980).