SPASOS 1.1: a program for the inference of ancestral shape ontogenies.

We recently published a method to infer ancestral landmark-based shape ontogenies that takes into account the possible existence of changes in developmental timing. Here we describe SPASOS, a software to perform that analysis. SPASOS is an open-source Windows program written in C. Input data include landmark coordinates for each specimen -with the corresponding information about developmental timing- and a phylogenetic tree showing the relationships among the species sampled. As output, the program produces image files for an easy visualization of the results and data files useful for post-processing. The program incorporates an interpolating function, based on weighting moving averages, which allows analysis of data with scarce information along the ontogenetic trajectory. An empirical evaluation of this function showed its suitability to fill in incomplete ontogenetic trajectories. Finally, we present the results of a reanalysis in SPASOS of a published dataset, where changes in developmental timing were originally inferred by considering PCA scores as shape variables. Both approaches retrieved the same four largest changes in developmental timing, but differed in the ancestral shapes inferred.