Apsaravis

Abstract: Abstract The avialan taxon Apsaravis ukhaana from the Late Cretaceous of southern Mongolia is completely described and its phylogenetic position is evaluated. Apsaravis ukhaana is from continental sandstones exposed at the locality of Ukhaa Tolgod, Omnogov Aimag, Mongolia. The holotype specimen consists of the nearly complete, articulated skeleton of a small volant avialan. Apsaravis ukhaana is unambiguously differentiated from other avialans based on the presence of several unique morphologies: a strong tubercle on the proximal humerus, a hypertrophied trochanteric crest on the femur, and extremely well-projected posterior wings of a surface of the distal tibiotarsus that in Aves articulates with the tibial cartilage. Ten other homoplastic characters optimize as autapomorphies of Apsaravis ukhaana in the phylogenetic analysis. They are as follows: ossified mandibular symphysis; dentary strongly forked posteriorly; hooked acromion process on scapula; highly angled dorsal condyle of humerus; humeral condyles weakly defined; distal edge of humerus angling strongly ventrally; humerus flared dorsoventrally at its distal terminus; lateral condyle of tibiotarsus wider than medial one; neither condyle of tibiotarsus tapering toward the midline; and metatarsal II trochlea rounded rather than ginglymoid. Phylogenetic placement of Apsaravis ukhaana as the sister taxon of Hesperornithes + Aves resulted from analysis of 202 characters scored for 17 avialan ingroup taxa. The implications of Apsaravis ukhaana, and the results of the phylogenetic analysis, for the evolution of flight after its origin and character support for enantiornithine monophyly are extensively discussed.

Abstract: Despite the discoveries of well-preserved Mesozoic birds, a key part of avian evolution, close to the radiation of all living birds (Aves), remains poorly represented. Here we report on a new taxon from the Late Cretaceous locality of Ukhaa Tolgod, Mongolia, that offers insight into this critically unsampled period. Apsaravis and the controversial alvarezsaurids are the only avialan taxa known from the continental deposits at Ukhaa Tolgod, which have produced hundreds of fossil mammals, lizards and other small dinosaurs. The new taxon, Apsaravis ukhaana, is the best-preserved specimen of a Mesozoic ornithurine bird discovered in over a century. It provides data important for assessing morphological evolution across Avialae, with implications for, first, the monophyly of Enantiornithes and Sauriurae; second, the proposition that the Mesozoic sister taxa of extant birds, as part of an 'ecological bottleneck', inhabited exclusively near-shore and marine environments; and third, the evolution of flight after its origin.