Evolutionary novelties in the skeleton are usually expressed as changes in the timing of growth of features intrinsically integrated at different hierarchical levels of development1. As a consequence, most of the shape-traits observed across species do vary quantitatively rather than qualitatively2, in a multivariate space3 and in a modularized way4,5. Because most phylogenetic analyses normally use discrete, hypothetically independent characters6, previous attempts have disregarded the phylogenetic signals potentially enclosed in the shape of morphological structures. When analysing low taxonomic levels, where most variation is quantitative in nature, solving basic requirements like the choice of characters and the capacity of using continuous, integrated traits is of crucial importance in recovering wider phylogenetic information. This is particularly relevant when analysing extinct lineages, where available data are limited to fossilized structures. Here we show that when continuous, multivariant and modularized characters are treated as such, cladistic analysis successfully solves relationships among main Homo taxa. Our attempt is based on a combination of cladistics, evolutionary-development-derived selection of characters, and geometric morphometrics methods. In contrast with previous cladistic analyses of hominid phylogeny, our method accounts for the quantitative nature of the traits, and respects their morphological integration patterns. Because complex phenotypes are observable across different taxonomic groups and are potentially informative about phylogenetic relationships, future analyses should point strongly to the incorporation of these types of trait.
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We thank P. Goloboff, M. Hernández, C. Lalueza, N. Martínez-Abadías, D. Pol and F. Ramírez-Rozzi for reading and discussing previous versions of this paper. We also thank R. E. Ambrosetto, R. Nicoletti and B. Nicoletti for their assistance during this work. The program TNT is freely available, thanks to a subsidy from the Willi Hennig Society.
The file contains Supplementary Figure 1 and Supplementary Data including the synapomorphies for the MP tree (pages 6-12), SI text file corresponding to Morphologika files for the traits: FCB_Morphologika file (pages 13-16), FR_Morphologika file (pages 17-22), NG_Morphologika file (pages 23-30), MA_Morphologika file (pages 31-41).
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Re-evaluation of temporal changes in alveolar protrusion in Japan according to two newly introduced craniometric angles
Anthropological Science (2016)