DOI: 10.5061/dryad.bs41q
title: Data from: Evolution of a complex phenotype with biphasic ontogeny: contribution of development versus function and climatic variation to skull modularity in toads

description: The theory of morphological integration and modularity predicts that if
 functional correlations among traits are relevant to mean population
 fitness, the genetic basis of development will be molded by stabilizing
 selection to match functional patterns. Yet, how much functional
 interactions actually shape the fitness landscape is still an open
 question. We used the anuran skull as a model of a complex phenotype for
 which we can separate developmental and functional modularity. We
 hypothesized that functional modularity associated to functional demands
 of the adult skull would overcome developmental modularity associated to
 bone origin at the larval phase because metamorphosis would erase the
 developmental signal. We tested this hypothesis in toad species of the
 Rhinella granulosa complex using species phenotypic correlation pattern
 (P-matrices). Given that the toad species are distributed in very distinct
 habitats and the skull has important functions related to climatic
 conditions, we also hypothesized that differences in skull trait
 covariance pattern are associated to differences in climatic variables
 among species. Functional and hormonal-regulated modules are more
 conspicuous than developmental modules only when size variation is
 retained on species P-matrices. Without size variation, there is a clear
 modularity signal of developmental units, but most species have the
 functional model as the best supported by empirical data without
 allometric size variation. Closely related toad species have more similar
 climatic niches and P-matrices than distantly related species, suggesting
 phylogenetic niche conservatism. We infer that the modularity signal due
 to embryonic origin of bones, which happens early in ontogeny, is blurred
 by the process of growth that occurs later in ontogeny. We suggest that
 the species differing in the preferred modularity model have different
 demands on the orbital functional unit and that species contrasting in
 climate are subjected to divergent patterns of natural selection
 associated to neurocranial allometry and T3 hormone regulation.
description: Phenotypic matrices with
 size variationPearson product-moment
 correlation matrices of all toad species for the 21 linear
 distances.P_matrices_ECE-2017-06-00822.csvList of covariance matrices of all toad speciesList of residual covariance matrices for the 21 linear distances extracted from toad skull. The matrices were constructed using residuals of multivariate linear models controlling for sex and locality. This list can be read in R using load('cov.list_data'). To construct correlation matrices, use lapply(cov.list, cov2cor).cov.list_dataDefinition of each developmental, hormonal and functional unitData frame describing which distances belong to each developmental, hormonal or functional unit. 1 for distances that belong to the unit and 0 for distances that do not belong to the unit. May be used as the argument 'modularity.hypot' in the function 'TestModularity' of the 'evolqg' R package.mod_hypo.csvModularity models for P-matrices with size variationData frame describing the modularity models used to test P-matrices with size variation. May be used as argument 'mod' in the 'EMMLi' function in R.mod_models_size.csvModularity models for P-matrices without allometryDescription of modularity models used to test P-matrices without allometric size variation. May be used as argument 'mod' in 'EMMLi' R function.mod_models_nosize.csvModularity models for P-matrices without isometryDescription of modularity models used to test P-matrices without isometric size variation. May be used as argument 'mod' in 'EMMLi' R function.mod_models_noiso.csv