Patterns of correlated character evolution in flightless birds: a phylogenetic approach

Given a robust phylogeny for a particular higher taxon, it is possible to map the evolution of various character changes onto the phylogeny and study the extent to which they co-occur. Of particular interest are the questions of (a) whether particular morphological changes tend to accompany changes in ecology or behaviour to which they bear a functional relationship and (b) whether changes in those ‘primary’ morphological characters tend to be associated with correlated changes in other aspects of morphology, as would be expected given the high level of morphological integration that characterizes most organisms. Here we report a study of this kind, looking at morphological correlates of the evolution of flightlessness in birds, and using the concentrated changes test to determine whether associations are significant. We find that pectoral reduction, pelvic enlargement and changes in skull morphology significantly co-occur, and that these are usually achieved through heterochrony rather than other kinds of developmental reprogramming. This revised version was published online in November 2006 with corrections to the Cover Date.     

Nonheterochronic developmental changes underlie morphological heterochrony in the evolution of the Ardeidae

Evolutionary changes in developmental timing and rates (heterochrony) are a source of morphological variation. Here we explore a central issue in heterochronic analysis: are the alterations in developmental timing and rates the only factor underlying morphological heterochrony? Tarsometatarsal growth through endochondral ossification in Ardeidae evolution has been taken as a case study. Evolutionary changes in bone growth rate (morphological heterochrony) might be either (a) the result of alterations in the mitotic frequency of epiphyseal chondrocytes (process‐heterochrony hypothesis), or (b) the outcome of alterations in the number of proliferating cells or in the size of hypertrophic chondrocytes (structural hypothesis). No correlation was found between tarsometatarsal growth rates and the frequency of cell division. However, bone growth rates were significantly correlated with the number of proliferating cells. These results support the structural hypothesis: morphological acceleration and deceleration are the outcome of evolutionary changes in one structural variable, the number of proliferating cells.     

Patterns and processes of heterochrony in Mesozoic goniasterid sea-stars

Heterochronic processes are obvious in the most common family of Mesozoic asteroids, the Goniasteridae, whether studied from entire fossils or, as is more frequently the case, from isolated ossicles. When phylogeny is poorly known, comparison between species leads to such unsatisfactory interpretations as 'A is more paedomorphic than B' or 'C, compared with other species, has many peramorphic characters'. When the phylogeny is known, heterochronic trends appear amongst the lineages. The peramorphocline Metopaster parkinsoni - M. loirensis-M. trichilae-M. chilipora-M. hypertelicus is studied here from the Cenomanian to the Upper Campanian of France. Different morphological characters evolved at different speeds. Among each polymorphous population, some morphotypes foreshadowed the next step. Juveniles of one species look like the adults of one of the ascendants. The cline developed irrespective of the substrate, and proceeded together with a migration from the Anglo-Paris Basin to the Aquitanian Basin. Metopaster hunteri , during the Coniacian, and Metopaster meudonensis , during the Upper Campanian, both evolved from Metopaster parkinsoni by neoteny, and provide a good example of reiterated and canalized morphological evolution. Some non-heterochronic canalizations are described.     

The role of post-natal ontogeny in the evolution of phenotypic diversity in Podarcis lizards

Understanding the role of the developmental pathways in shaping phenotypic diversity allows appreciating in full the processes influencing and constraining morphological change. Podarcis lizards demonstrate extraordinary morphological variability that likely originated in short evolutionary time. Using geometric morphometrics and a broad suite of statistical tests, we explored the role of developmental mechanisms such as growth rate change, ontogenetic divergence/convergence/parallelism as well as morphological expression of heterochronic processes in mediating the formation of their phenotypic diversity during the post-natal ontogeny. We identified hypermorphosis - the prolongation of growth along the same trajectory - as the process responsible for both intersexual and interspecific morphological differentiation. Albeit the common allometric pattern observed in both sexes of any species constrains and canalizes their cephalic scales variation in a fixed portion of the phenotypic space, the extended growth experienced by males and some species allows them to achieve peramorphic morphologies. Conversely, the intrasexual phenotypic diversity is accounted for by non-allometric processes that drive the extensive morphological dispersion throughout their ontogenetic trajectories. This study suggests a model of how simple heterochronic perturbations can produce phenotypic variation, and thus potential for further evolutionary change, even within a strictly constrained developmental pathway.     

The genetic control of heterochrony: Evidence from developmental mutants of Pisum sativum L.

Heterochrony, as a means of evolution in which the rate or timing of developmental events of the descendant is altered compared with that of the ancestor, is of significance because it suggests that rapid and dramatic morphological changes are possible with few genetic changes. The putative origin of plant taxa by this means of evolution is becoming increasingly frequent in the literature but there is little evidence of the extent of the genetic change necessary to alter the timing of developmental events to produce such changes. This study shows that the onset of flowering can be altered independently from the vegetative transition in leaf form in at least one genotype of Pisum in response to different environments. Further, it identifies 9 mutations that act in a heterochronic manner to produce dramatic morphological changes that can be described as progenesis, neoteny, hypermorphosis or acceleration. In addition, it is demonstrated that the same heterochronic process (e.g. progenesis) may be caused by genes controlling distinctly different physiological processes. It is suggested that Pisum is an ideal model species for studies of heterochrony and that few genetic changes are necessary to bring about dramatic heterochronic changes.