The forelimbs of Alvarezsauroidea (Dinosauria: Theropoda): Insight from evolutionary teratology.

Alvarezsauroidea (Tetanurae) are nonavian theropod dinosaurs whose forelimb evolution is characterised by the overdevelopment of digit I, at the expense of the other two digits, complemented by a drastic forelimb shortening in derived species (Parvicursorinae). These variations are recognised as evolutionary developmental anomalies. Evolutionary teratology hence leads to a double diagnosis with (1) macrodactyly of digit I and microdactyly of digits II and III, plus (2) anterior micromelia. The teratological macrodactyly/microdactyly coupling evolved first. Developmental mechanisms disturbing limb proportion are thought to be convergent with those of other Tetanurae (Tyrannosauridae, Carcharodontosauridae). As for the manual anomalies, both are specific to Alvarezsauroidea (macrodactyly/microdactyly) and inherited (digit loss/reduction). While considering the frame-shift theory, posterior digits develop before the most anterior ones. There would therefore be a decrease in the area devoted to digits II (condensation 3) and III (condensation 4), in connection with the Shh signalling pathway, interacting with other molecular players such as the GLI3 protein and the Hox system. Developmental independence of digit I (condensation 2) would contribute to generating a particular morphology. Macrodactyly would be linked to a variation in Hoxd-13, impacting Gli3 activity, and increasing cell proliferation. The loss/reduction of digital ray/phalanges (digits II and III), would be associated with Shh activity, a mechanism inherited from the theropodan ancestry. The macrodactyly/microdactyly coupling, and then anterior micromelia, fundamentally changed the forelimb mechanical function, compared to the 'classical' grasping structure of basal representatives and other theropods. The distal ossification of the macrodactylian digit has been identified as physiological, implying the use of the structure. However, the debate on a particular 'adaptive' use is pointless as the ecology of an organism is interactively complex, being both at the scale of the individual and dependent on circumstances. Other anatomical features also allow for compensation and different predation (cursorial hindlimbs).