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Abstract
The pterosaurs were a diverse group of Mesozoic flying reptiles that underwent a body
plan reorganization, adaptive radiation, and replacement of earlier forms midway through
their long history, resulting in the origin of the Pterodactyloidea, a highly specialized
clade containing the largest flying organisms. The sudden appearance and large suite
of morphological features of this group were suggested to be the result of it originating
in terrestrial environments, where the pterosaur fossil record has traditionally been
poor [1, 2], and its many features suggested to be adaptations to those environments
[1, 2]. However, little evidence has been available to test this hypothesis, and it
has not been supported by previous phylogenies or early pterodactyloid discoveries.
We report here the earliest pterosaur with the diagnostic elongate metacarpus of the
Pterodactyloidea, Kryptodrakon progenitor, gen. et sp. nov., from the terrestrial
Middle-Upper Jurassic boundary of Northwest China. Phylogenetic analysis confirms
this species as the basalmost pterodactyloid and reconstructs a terrestrial origin
and a predominantly terrestrial history for the Pterodactyloidea. Phylogenetic comparative
methods support this reconstruction by means of a significant correlation between
wing shape and environment also found in modern flying vertebrates, indicating that
pterosaurs lived in or were at least adapted to the environments in which they were
preserved.