Morphological integration during postnatal ontogeny: implications for evolutionary biology.

How covariance patterns of phenotypes change during development is fundamental for a broader understanding of evolution. There is compelling evidence that mammalian cranium covariance patterns change during ontogeny. However, it is unclear to what extent variation in covariance patterns during ontogeny can impact the response to selection. To tackle this question, we explored: (a) the extent to which covariance patterns change during postnatal ontogeny; (b) in which ontogenetic stages covariance patterns differ the most; and (c) the extent to which the phenotypic covariance pattern at different ontogenetic stages can be explained by the same processes determining additive genetic covariance. We sampled the postnatal ontogenetic series for both marsupials and placentals. Within each ontogenetic series, we compared covariance matrices (P-matrices) at different ontogenetic stages. Furthermore, we compared these P-matrices to two target matrices [adult P-matrix and an additive genetic covariance matrix (G-matrix)]. Our results show that for all ontogenetic series, covariance patterns from weaning onward are conserved and probably shaped by the same processes determining the G-matrix. We conclude that irrespective of eventual differences in how selection operates during most of the postnatal ontogeny, the net response to such pressures will probably not be affected by ontogenetic differences in the covariance pattern.

The pre-eminent role of directional selection in generating extreme morphological change in glyptodonts (Cingulata; Xenarthra).

The prevalence of stasis on macroevolution has been classically taken as evidence of the strong role of stabilizing selection in constraining morphological change. Rates of evolution calculated over longer timescales tend to fall below the expected under genetic drift, suggesting that directional selection signals are erased at longer timescales. Here, we investigated the rates of morphological evolution of the skull in a fossil lineage that underwent extreme morphological modification, the glyptodonts. Contrary to what was expected, we show here that directional selection was the primary process during the evolution of glyptodonts. Furthermore, the reconstruction of selection patterns shows that traits selected to generate a glyptodont morphology are markedly different from those operating on extant armadillos. Changes in both direction and magnitude of selection are probably tied to glyptodonts' invasion of a specialist-herbivore adaptive zone. These results suggest that directional selection might have played a more critical role in the evolution of extreme morphologies than previously imagined.

Measuring the magnitude of morphological integration: The effect of differences in morphometric representations and the inclusion of size.

The magnitude of morphological integration is a major aspect of multivariate evolution, providing a simple measure of the intensity of association between morphological traits. Studies concerned with morphological integration usually translate phenotypes into morphometric representations to quantify how different morphological elements covary. Geometric and classic morphometric representations translate biological form in different ways, raising the question if magnitudes of morphological integration estimates obtained from different morphometric representations are compatible. Here we sought to answer this question using the relative eigenvalue variance of the covariance matrix obtained for both geometric and classical representations of empirical and simulated datasets. We quantified the magnitude of morphological integration for both shape and form and compared results between representations. Furthermore, we compared integration values between shape and form to evaluate the effect of the inclusion or not of size on the quantification of the magnitude of morphological integration. Results show that the choice of morphological representation has significant impact on the integration magnitude estimate, either for shape or form. Despite this, ordination of the integration values within representations is relatively the same, allowing for similar conclusions to be reached using different methods. However, the inclusion of size in the dataset significantly changes the estimates of magnitude of morphological integration, hindering the comparison of this statistic obtained from different spaces. Morphometricians should be aware of these differences and must consider how biological hypothesis translate into predictions about integration in each particular choice of representation.

Early South Americans Cranial Morphological Variation and the Origin of American Biological Diversity.

Recent South Americans have been described as presenting high regional cranial morphological diversity when compared to other regions of the world. This high diversity is in accordance with linguistic and some of the molecular data currently available for the continent, but the origin of this diversity has not been satisfactorily explained yet. Here we explore if this high morphological variation was already present among early groups in South America, in order to refine our knowledge about the timing and origins of the modern morphological diversity. Between-group (Fst estimates) and within-group variances (trace of within-group covariance matrix) of the only two early American population samples available to date (Lagoa Santa and Sabana de Bogotá) were estimated based on linear craniometric measurements and compared to modern human cranial series representing six regions of the world, including the Americas. The results show that early Americans present moderate within-group diversity, falling well within the range of modern human groups, despite representing almost three thousand years of human occupation. The between-group variance apportionment is very low between early Americans, but is high among recent South American groups, who show values similar to the ones observed on a global scale. Although limited to only two early South American series, these results suggest that the high morphological diversity of native South Americans was not present among the first human groups arriving in the continent and must have originated during the Middle Holocene, possibly due to the arrival of new morphological diversity coming from Asia during the Holocene.

EvolQG - An R package for evolutionary quantitative genetics.

We present an open source package for performing evolutionary quantitative genetics analyses in the R environment for statistical computing. Evolutionary theory shows that evolution depends critically on the available variation in a given population. When dealing with many quantitative traits this variation is expressed in the form of a covariance matrix, particularly the additive genetic covariance matrix or sometimes the phenotypic matrix, when the genetic matrix is unavailable and there is evidence the phenotypic matrix is sufficiently similar to the genetic matrix. Given this mathematical representation of available variation, the \textbf{EvolQG} package provides functions for calculation of relevant evolutionary statistics; estimation of sampling error; corrections for this error; matrix comparison via correlations, distances and matrix decomposition; analysis of modularity patterns; and functions for testing evolutionary hypotheses on taxa diversification.