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Review
. 2022 Jun;97(3):960-1004.
doi: 10.1111/brv.12829. Epub 2022 Jan 6.

Morphology and distribution of scales, dermal ossifications, and other non-feather integumentary structures in non-avialan theropod dinosaurs

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Review

Morphology and distribution of scales, dermal ossifications, and other non-feather integumentary structures in non-avialan theropod dinosaurs

Christophe Hendrickx et al. Biol Rev Camb Philos Soc. 2022 Jun.

Abstract

Modern birds are typified by the presence of feathers, complex evolutionary innovations that were already widespread in the group of theropod dinosaurs (Maniraptoriformes) that include crown Aves. Squamous or scaly reptilian-like skin is, however, considered the plesiomorphic condition for theropods and dinosaurs more broadly. Here, we review the morphology and distribution of non-feathered integumentary structures in non-avialan theropods, covering squamous skin and naked skin as well as dermal ossifications. The integumentary record of non-averostran theropods is limited to tracks, which ubiquitously show a covering of tiny reticulate scales on the plantar surface of the pes. This is consistent also with younger averostran body fossils, which confirm an arthral arrangement of the digital pads. Among averostrans, squamous skin is confirmed in Ceratosauria (Carnotaurus), Allosauroidea (Allosaurus, Concavenator, Lourinhanosaurus), Compsognathidae (Juravenator), and Tyrannosauroidea (Santanaraptor, Albertosaurus, Daspletosaurus, Gorgosaurus, Tarbosaurus, Tyrannosaurus), whereas dermal ossifications consisting of sagittate and mosaic osteoderms are restricted to Ceratosaurus. Naked, non-scale bearing skin is found in the contentious tetanuran Sciurumimus, ornithomimosaurians (Ornithomimus) and possibly tyrannosauroids (Santanaraptor), and also on the patagia of scansoriopterygids (Ambopteryx, Yi). Scales are surprisingly conservative among non-avialan theropods compared to some dinosaurian groups (e.g. hadrosaurids); however, the limited preservation of tegument on most specimens hinders further interrogation. Scale patterns vary among and/or within body regions in Carnotaurus, Concavenator and Juravenator, and include polarised, snake-like ventral scales on the tail of the latter two genera. Unusual but more uniformly distributed patterning also occurs in Tyrannosaurus, whereas feature scales are present only in Albertosaurus and Carnotaurus. Few theropods currently show compelling evidence for the co-occurrence of scales and feathers (e.g. Juravenator, Sinornithosaurus), although reticulate scales were probably retained on the mani and pedes of many theropods with a heavy plumage. Feathers and filamentous structures appear to have replaced widespread scaly integuments in maniraptorans. Theropod skin, and that of dinosaurs more broadly, remains a virtually untapped area of study and the appropriation of commonly used techniques in other palaeontological fields to the study of skin holds great promise for future insights into the biology, taphonomy and relationships of these extinct animals.

Keywords: Theropoda; dinosaur; feathers; footprints; integument; scales; skin; trace fossils.

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REFERENCES

    1. Abourachid, A., Herrel, A., Decamps, T., Pages, F., Fabre, A.-C., Hoorebeke, L. V., Adriaens, D. & Amado, M. A. G. (2019). Hoatzin nestling locomotion: acquisition of quadrupedal limb coordination in birds. Science Advances 5, eaat0787.
    1. Acosta Hospitaleche, C., De Los Reyes, M., Santillana, S. & Reguero, M. (2020). First fossilized skin of a giant penguin from the Eocene of Antarctica. Lethaia 53, 409-420.
    1. Agnolín, F. L. & Novas, F. E. (2013). Avian Ancestors - A Review of the Phylogenetic Relationships of the Theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae. SpringerBriefs South America and the Southern Hemisphere, Springer Verlag, Dordrecht, Heidelberg, New York, London.
    1. Alcober, O. A. & Martínez, R. N. (2010). A new herrerasaurid (Dinosauria, Saurischia) from the Upper Triassic Ischigualasto Formation of northwestern Argentina. ZooKeys 63, s55-s81.
    1. Alexander, D. E., Gong, E., Martin, L. D., Burnham, D. A. & Falk, A. R. (2010). Model tests of gliding with different hindwing configurations in the four-winged dromaeosaurid Microraptor gui. Proceedings of the National Academy of Sciences of the United States of America 107, 2972-2976.

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