Abstract
Although an increasing number of studies are combining skeletal and neural morphology data in a phylogenetic context, most studies do not include extinct taxa due to the rarity of preserved endocasts. The early Eocene avifauna of the Fur Formation of Denmark presents an excellent opportunity for further study of extinct osteological and endocranial morphology as fossils are often exceptionally preserved in three dimensions. Here, we use X-ray computed tomography to present additional material of the previously described taxon Scandiavis mikkelseni and reassess its phylogenetic placement using a previously published dataset. The new specimen provides novel insights into the osteological morphology and brain anatomy of Scandiavis. The virtual endocast exhibits a morphology comparable to that of modern avian species. Endocranial evaluation shows that it was remarkably similar to that of certain extant Charadriiformes, yet also possessed a novel combination of traits. This may mean that traits previously proposed to be the result of shifts in ecology later in the evolutionary history of Charadriiformes may instead show a more complex distribution in stem Charadriiformes and/or Gruiformes depending on the interrelationships of these important clades. Evaluation of skeletal and endocranial character state changes within a previously published phylogeny confirms both S. mikkelseni and a putative extinct charadriiform, Nahmavis grandei, as charadriiform. Results bolster the likelihood that both taxa are critical fossils for divergence dating and highlight a biogeographic pattern similar to that of Gruiformes.
Highlights
The early Eocene Fur Formation of north-western Jutland, Denmark, is well-known for its rich fossil bird fauna that includes three-dimensionally preserved, articulated specimens, as well as occasional soft tissues [1,2,3]
The diatomaceous sediments of the Fur Formation are unusual in the Paleogene fossil record of birds in deriving from a marine offshore environment [3]
Recovery of additional material and new phylogenetic analysis of Scandiavis mikkelseni more robustly establishes both S. mikkelseni and Nahmavis grandei as charadriiform, and most consistently places them as stem-charadriiform taxa. This is critical to reconstructing phylogenetic relationships of Paleogene fossils, as previous studies produced conflicting results as to whether either taxon was charadriiform or gruiform [17,18]
Summary
The early Eocene Fur Formation of north-western Jutland, Denmark, is well-known for its rich fossil bird fauna that includes three-dimensionally preserved, articulated specimens, as well as occasional soft tissues [1,2,3]. We employ high-resolution computed tomography (μCT) to describe new remains of Scandiavis mikkelseni from the Fur Formation This specimen, NHMD 625345, consists of a fully articulated skull preserved in three dimensions. Phylogenetic analysis of both osteological and endocranial characters further constrains the affinities of this taxon by more robustly placing both S. mikkelseni and N. grandei within the Charadriiformes, primarily as stem-Charadriiformes This new specimen is another example of the excellent preservation of early Eocene avifauna and testifies to the potential of the Fur Formation in providing material that is crucial to better understanding avian evolution. This software was used for the linear and volumetric measurements
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