When populations experience suboptimal conditions, the mechanisms involved in the regulation of phenotypic variation can be challenged, resulting in increased phenotypic variance. This kind of disturbance can be diagnosed by using morphometric tools to study morphological patterns at different hierarchical levels and evaluate canalization, developmental stability, integration, modularity, and allometry. We assess the effect of urbanization on phenotypic variation in the common wall lizard (Podarcis muralis) by using geometric morphometrics to assess disturbance to head shape development. The head shapes of urban lizards were more variable and less symmetric, suggesting that urban living is more likely to disturb development. Head shape variation was congruent within and across individuals, which indicated that canalization and developmental stability are two related phenomena in these organisms. Furthermore, urban lizards exhibited smaller mean head sizes, divergent size-shape allometries, and increased deviation from within-group allometric lines. This suggests that mechanisms regulating head shape allometry may also be disrupted. The integrated evaluation of several measures of developmental instability at different hierarchical levels, which provided in this case congruent results, can be a powerful methodological guide for future studies, as it enhances the detection of environmental disturbances on phenotypic variation and aids biological interpretation of the results.

The increase in human activities that leads to wildlife decline and species extinction poses an urgent need for simple indicators of environmental stress in animal populations. Several studies have suggested that fluctuating asymmetry (FA) can be an easy, direct measure of developmental instability because it is associated to environmental stress and, as such, it can be a useful indicator of population disturbance. We examined three different morphological traits in urban and rural populations of the common wall lizard (Podarcis muralis) to test whether anthropogenic disturbance causes an increase in FA. Compared to rural populations, urban ones showed higher levels of FA in all analyzed traits, thus providing evidence that FA can respond to anthropogenic disturbance. However, we also found significant differences in FA among traits, where femoral pores and subdigital lamellae, traits with a functional relevance, were more stable developmentally compared to supracilliar granules which have no evident function. Unsigned FA [abs(right-left)] exhibited significant, but weak, positive correlations among traits, indicating that developmental noise does not have a uniform effect across characters and thus questioning the view of developmental stability as an organism-wide property. The degree of signed FA (right-left) was more similar between structurally associated traits, possibly as an outcome of morphological integration. In conclusion, our results demonstrate that FA can be a reliable indicator of disturbance provided that it is analyzed on multiple traits simultaneously and examined at the population level.

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Canalization and developmental stability (DS) are important organismal properties involved in determining the level of phenotypic variation. Ontogenetic patterns of phenotypic variance components can shed light on the mechanistic basis of developmental buffering (DB). Here, we analyze how individual FA and among-individual variation in head shape change in ontogenetic series of three lizard species raised in laboratory. The degree of asymmetry increased slightly with size, suggesting that developmental mechanisms hypothesized to correct for deviations either do not exist, or that their efficiency is truncated with increasing size. Alternatively, they may need the disturbance as a trigger. The relationship between asymmetry and age was complex, with asymmetry being stable across the age range in two species but increased with age in the third. Lack of congruence in ontogenetic patterns of asymmetry might be due to intrinsic differences in buffering mechanisms or a result of species-specific growth patterns. Head shape was shown to be equally canalized across both size and age range in all species, probably as a result of a balance between the buffering mechanisms and mechanisms generating variance. The patterns of symmetric and asymmetric head shape variation were highly correlated across species meaning that DS and canalization may rely on similar mechanisms.

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