Feathered dinosaur is possible that feathers first developed in an even earlier group, in light of the discovery of pycnofibers of pterosaurs. Crocodilians also possess beta keratin similar to those of birds, which suggests that they evolved from common ancestral genes. Shortly after the 1859 publication of Charles Darwin's "On the Origin of Species", British biologist Thomas Henry Huxley proposed that birds were descendants of dinosaurs. He compared the skeletal structure of "Compsognathus", a small theropod dinosaur, and the 'first bird' "Archaeopteryx lithographica" (both of which were found in the Upper Jurassic Bavarian limestone of Solnhofen). He showed that, apart from its
Archaeopteryx be related particularly closely to "Anchiornis" and "Xiaotingia", which were recovered as basal troodontids instead. Agnolín and Novas (2013) found "Archaeopteryx" and (possibly synonymous) "Wellnhoferia" to be the basalmost avialans (Avialae being defined by the authors as including "Archaeopteryx lithographica" and "Passer", their most recent common ancestor and all of its descendants), with Microraptoria, Unenlagiinae, and the clade containing "Anchiornis" and "Xiaotingia" being successively closer outgroups to the Avialae. Another phylogenetic study by Godefroit, "et al.", using a more inclusive matrix than the one from the analysis in the description of "Eosinopteryx brevipenna", also found "Archaeopteryx" to be a member