Forelimb shortening of Carcharodontosauria (Dinosauria: Theropoda): an update on evolutionary anterior micromelias in non-avian theropods.

Evolutionary teratology recognises certain anatomical modifications as developmental anomalies. Within non avian-theropod dinosaurs, the strong forelimb shortening of Tyrannosauridae, Carnotaurinae and Limusaurus - associated with a reduction or loss of autonomy - have been previously diagnosed as evolutionary anterior micromelias. The feature is here examined with Acrocanthosaurus atokensis (Carcharodontosauridae) and Gualicho shinyae (Neovenatoridae). The micromelic diagnosis is confirmed for Acrocanthosaurus, without supplementary malformations. Gualicho is considered as a borderline case, outside of the micromelic spectrum, but shows a total phalangeal loss on digit III. The reduction in the biomechanical range of Acrocanthosaurus' forelimbs was compensated by the skull and jaws as main predatory organs. The same is assumed for Gualicho, but its robust first digit and raptorial claw are to be underlined. Other gigantic-sized and derived representatives of Carcharodontosauridae probably shared the anterior micromelia condition, potentially due to developmental modifications involving differential forelimbs/hindlimbs embryological growth rates, secondarily associated with post-natal growth rates leading to large and gigantic sizes; a converging state with Tyrannosauridae. Nevertheless, whereas developmental growth rates are also considered in the shortened condition of Gualicho, there is no association with post-natal gigantism. Finally, the digit III reduction likely followed the same evolutionary pathways as Tyrannosauridae, potentially involving BMPs, Fgfs and Shh signalling.