Head shape evolution in Tropidurinae lizards: does locomotion constrain diet?

Different components of complex integrated systems may be specialized for different functions, and thus the selective pressures acting on the system as a whole may be conflicting and can ultimately constrain organismal performance and evolution. The vertebrate cranial system is one of the most striking examples of a complex system with several possible functions, being associated to activities as different as locomotion, prey capture, display and defensive behaviours. Therefore, selective pressures on the cranial system as a whole are possibly complex and may be conflicting. The present study focuses on the influence of potentially conflicting selective pressures (diet vs. locomotion) on the evolution of head shape in Tropidurinae lizards. For example, the expected adaptations leading to flat heads and bodies in species living on vertical structures may conflict with the need for improved bite performance associated with the inclusion of hard or tough prey into the diet, a common phenomenon in Tropidurinae lizards. Body size and six variables describing head shape were quantified in preserved specimens of 23 species, and information on diet and substrate usage was obtained from the literature. No phylogenetic signal was observed in the morphological data at any branch length tested, suggesting adaptive evolution of head shape in Tropidurinae. This pattern was confirmed by both factor analysis and independent contrast analysis, which suggested adaptive co-variation between the head shape and the inclusion of hard prey into the diet. In contrast to our expectations, habitat use did not constrain or drive head shape evolution in the group.     

Molecular tests of phylogenetic taxonomies: a general procedure and example using four subfamilies of the lizard family Iguanidae

A general procedure is described for examining when results of molecular phylogenetic analyses warrant formal revision of taxonomies constructed using morphological characters. We illustrate this procedure with tests of monophyly for four subfamilies in the lizard family Iguanidae using 1561 aligned base positions (838 phylogenetically informative) of mitochondrial DNA sequences, representing coding regions for eight tRNAs, ND2, and portions of ND1 and COI. Ten new sequences ranging in length from 1732 to 1751 bases are compared with 12 previously reported sequences and 67 morphological characters (54 phylogenetically informative) from the literature. New morphological character states are provided for Sator. Phylogenies derived from the molecular and combined data are in agreement but both conflict with phylogenetic inferences from the morphological data alone. Strong support is found for the monophyly of the subfamilies Crotaphytinae and Phrynosomatinae. Monophyly of the Iguaninae is weakly supported in each analysis. All analyses suggest that the Tropidurinae is not monophyletic but the hypothesis of monophyly cannot be rejected. A phylogenetic taxonomy is proposed in which the Tropidurinae* is maintained as a metataxon (denoted with an asterisk), for which monophyly has not been demonstrated. Within the Phrynosomatinae, the close relationship of Sator and Sceloporus is questioned and an alternative hypothesis in which Sator is the sister taxon to a clade comprising Petrosaurus, Sceloporus, and Urosaurus is presented. Statistical tests of monophyly provide a powerful way to evaluate support for taxonomic groupings. Use of the metataxon prevents premature taxonomic rearrangements where support is lacking.     

Ecological constraints on the evolutionary association between field and preferred temperatures in Tropidurinae lizards

Ectothermic body temperatures affect organismal performances and presumably fitness, and are strongly influenced by the thermal environment. Therefore, the processes of colonization of novel thermal habitats by lizards might involve changes in thermal preferences, performance curves (reaction norms) and field activity temperatures. According to theory based on optimality analysis, diverse aspects of the thermal biology of vertebrate ectotherms should co-evolve as to maximize performance at the temperature range more often experienced by animals in the field. One corollary of this premise is that derived lizard clades that experienced a significant shift in thermal ecology, in comparison with the ancestral condition, should prefer and select temperatures in a thermal gradient similar to those experienced in nature. Here we report an analysis of the premise stated before. Specifically, we verify whether or not Tropidurinae species from three major Brazilian habitats (the Rainforests, the semi-arid Caatingas and the Cerrados, a Savannah-like biome) differ in thermal ecology and thermoregulatory behavior. The Caatinga is believed to be the ancestral habitat of this sub-family, and differences are expected because species from semi-arid habitats usually exhibit high body temperatures for lizards, whereas forest specialists might be thermoconformers and active at low temperatures. We also compared selected temperatures in the laboratory by species from the two open habitats (Caatingas and Cerrados). Data were analyzed using both conventional and phylogenetic analysis tools. Although species from Caatingas exhibited higher activity temperatures in nature than those from Cerrados, mean selected temperatures were similar between ecological groups. Phylogenetic analyses confirmed these findings and evidenced large␣evolutionary divergence in field activity temperatures between sister species from different␣open habitats without coupled divergence in selected temperatures. Therefore, thermoregulatory behavior and ecological parameters did not evolve similarly during the colonization of contrasting open habitats by Tropidurinae.