Sapeornis and the flight modes of birds

Sapeornis chaoyangensis (DNHM-3078) showing well-preserved primary (P) and secondary (S) feathers. From Serrano and Chiappe, 2017

Birds originated from a theropod lineage more than 150 million years ago. By the Early Cretaceous, they diversified, evolving into a number of groups of varying anatomy and ecology. Most of these fossils, like Sapeornis chaoyangensis (125 to 120 Ma), are from the Jehol Biota of northeastern China. Sapeornis shows a combination of derived and primitive features, like a short, robust non-strut-like coracoid and a fibula reaching the distal end of the tarsal joint, a pygostyle, reduced manual digits, and a well-fused carpometacarpus. All of these features indicates a mosaic pattern in the early evolution of birds and confirm the basal position of Sapeornis near Archaeopteryx and Jeholornis in the phylogeny of early birds.

The evolution of flight involved a series of adaptive changes at the morphological and molecular levels, that included the fusion and elimination of some bones and the pneumatization of the remaining ones. Archaeopteryx lacked a bony sternum and a compensatory specialized gastral basket for anchoring large flight muscles, while Jelohornis had several derived flight-related features of modern birds like fused sacral vertebrae, an elongated coracoid with a procoracoid process, a complex sternum, a narrow furcula, and curved scapula. In Enantiornithines, their robust pygostyle appears to have been unable to support the muscles that control the flight feathers on the tail in modern birds.

Morphofunctional fitness of the wing shape for soaring as depicted by the relation between the lift surface and the wingspan in modern soaring birds and Sapeornis (From Serrano and Chiappe, 2017)

The flight modes of modern birds are a reflection of their different strategies to reduce the energetic costs of a highly demanding style of locomotion. Among these features are wing shape, and the use of thermals and tail winds. Flapping flight is energetically more costly  than gliding and soaring flight, consequently, large birds have either elongated wingspans that allow them to gain height through air currents and to glide for long distances with much lower transit costs than flapping.

Fossil evidence suggests that S. chaoyangensis was a specialized flier that used continental soaring as its main flight mode. Computational models of S. chaoyangensis are also congruent with other morphological similarities between S. chaoyangensis and modern soaring birds including the shape of the furcula and the proportions of the forelimbs. Modern soaring birds include dynamic soarers that exploit air velocity gradients over sea waves, and thermal soarers that use ascending air currents mainly generated in continental areas. Because, exceptionally well preserved fossils of S. chaoyangensis have revealed seeds and/or fruits in its intestinal tract, this interpretation of the flight capabilities of S. chaoyangensis is consistent with the energetic disadvantages from a herbivorous diet, because soaring is a less demanding flight mode than continuous flapping.


Serrano FJ, Chiappe LM. 2017 Aerodynamic modelling of a Cretaceous bird reveals thermal soaring capabilities during early avian evolution. J. R. Soc. Interface 14: 20170182.

Butler PJ. 2016 The physiological basis of bird flight. Phil. Trans. R. Soc. B 371, 20150384 doi:10. 1098/rstb.2015.0384

Zhou, Zhonghe & Zhang, Fucheng (2003): Anatomy of the primitive bird Sapeornis chaoyangensis from the Early Cretaceous of Liaoning, China. Canadian Journal of Earth Sciences 40(5): 731–747. doi 10.1139/E03-011


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