Latitude or biogeographic breaks? Determinants of phenotypic (co)variation in fitness-related traits in Betaeus truncatus along the Chilean coast

Ectothermal organisms distributed along environmental gradients in a wide geographical distribution display extensive phenotypic variation. This is particularly pervasive along latitudinal clines, which are linked to gradual changes in environmental factors. Widespread species may also be distributed among biogeographic breaks, which in contrast to smooth clines, often show abrupt changes in phenotypic traits. In species with widespread latitudinal distribution that also encompass important biogeographical breaks, it is not clear which of those factors prevails on shaping the phenotypic variation or if some traits are particularly more sensitive to one or the other. To evaluate this, we measured 4 fitness-related traits in 6 populations of the intertidal snapping shrimp Betaeus truncatus, as its distribution along Chile expands over 40° in latitude and three major biogeographical provinces. Here, we statistically evaluated the role of both, latitude and biogeographic breaks, on mean population values of fitness-related traits but also on the variances and covariances (i.e., P-matrix) between them. Overall, our results (1) indicate that latitude is more important than breaks in shaping the phenotypic variation of most of these fitness-related traits, (2) show that the differences in the variance-covariance relationship among traits between the extremes of the gradient arises from gradual increases in variance and rather sharp changes in covariance at mid-latitudes and (3) show that at present, it is difficult to unambiguously determine whether natural selection or plasticity is responsible for the observed pattern in means, variances and covariances and only further work might disentangle these possibilities.