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      Recent advances in amniote palaeocolour reconstruction and a framework for future research

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          ABSTRACT

          Preserved melanin pigments have been discovered in fossilised integumentary appendages of several amniote lineages (fishes, frogs, snakes, marine reptiles, non‐avialan dinosaurs, birds, and mammals) excavated from lagerstätten across the globe. Melanisation is a leading factor in organic integument preservation in these fossils. Melanin in extant vertebrates is typically stored in rod‐ to sphere‐shaped, lysosome‐derived, membrane‐bound vesicles called melanosomes. Black, dark brown, and grey colours are produced by eumelanin, and reddish‐brown colours are produced by phaeomelanin. Specific morphotypes and nanostructural arrangements of melanosomes and their relation to the keratin matrix in integumentary appendages create the so‐called 'structural colours'. Reconstruction of colour patterns in ancient animals has opened an exciting new avenue for studying their life, behaviour and ecology. Modern relationships between the shape, arrangement, and size of avian melanosomes, melanin chemistry, and feather colour have been applied to reconstruct the hues and colour patterns of isolated feathers and plumages of the dinosaurs Anchiornis, Sinosauropteryx, and Microraptor in seminal papers that initiated the field of palaeocolour reconstruction. Since then, further research has identified countershading camouflage patterns, and informed subsequent predictions on the ecology and behaviour of these extinct animals. However, palaeocolour reconstruction remains a nascent field, and current approaches have considerable potential for further refinement, standardisation, and expansion. This includes detailed study of non‐melanic pigments that might be preserved in fossilised integuments. A common issue among existing palaeocolour studies is the lack of contextualisation of different lines of evidence and the wide variety of techniques currently employed. To that end, this review focused on fossil amniotes: ( i) produces an overarching framework that appropriately reconstructs palaeocolour by accounting for the chemical signatures of various pigments, morphology and local arrangement of pigment‐bearing vesicles, pigment concentration, macroscopic colour patterns, and taphonomy; ( ii) provides background context for the evolution of colour‐producing mechanisms; and ( iii) encourages future efforts in palaeocolour reconstructions particularly of less‐studied groups such as non‐dinosaur archosaurs and non‐archosaur amniotes.

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          The biology of color.

          Coloration mediates the relationship between an organism and its environment in important ways, including social signaling, antipredator defenses, parasitic exploitation, thermoregulation, and protection from ultraviolet light, microbes, and abrasion. Methodological breakthroughs are accelerating knowledge of the processes underlying both the production of animal coloration and its perception, experiments are advancing understanding of mechanism and function, and measurements of color collected noninvasively and at a global scale are opening windows to evolutionary dynamics more generally. Here we provide a roadmap of these advances and identify hitherto unrecognized challenges for this multi- and interdisciplinary field.
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                Author and article information

                Contributors
                arindam.roy@connect.hku.hk
                mpittman@hku.hk
                Journal
                Biol Rev Camb Philos Soc
                Biol Rev Camb Philos Soc
                10.1111/(ISSN)1469-185X
                BRV
                Biological Reviews of the Cambridge Philosophical Society
                Blackwell Publishing Ltd (Oxford, UK )
                1464-7931
                1469-185X
                19 September 2019
                February 2020
                : 95
                : 1 ( doiID: 10.1111/brv.v95.1 )
                : 22-50
                Affiliations
                [ 1 ] Vertebrate Palaeontology Laboratory, Department of Earth Sciences The University of Hong Kong Pokfulam Hong Kong SAR China
                [ 2 ] Integrative Research Center, Section of Earth Sciences Field Museum of Natural History 1400 S. Lake Shore Drive, Chicago IL 60605 U.S.A.
                [ 3 ] Foundation for Scientific Advancement 7023 Alhambra Drive, Sierra Vista AZ 85650 U.S.A.
                [ 4 ] Institute of Vertebrate Paleontology and Paleoanthropology Chinese Academy of Sciences 142 Xizhimenwai Street. Beijing 100044 China
                Author notes
                [*] [* ]Authors for correspondence (Tel: +852 66549715; E‐mail: arindam.roy@ 123456connect.hku.hk ); (Tel: +852 3917 7840; E‐mail: mpittman@ 123456hku.hk )
                Author information
                https://orcid.org/0000-0002-4890-6851
                https://orcid.org/0000-0002-6149-3078
                https://orcid.org/0000-0002-9306-9060
                https://orcid.org/0000-0001-7996-618X
                Article
                BRV12552
                10.1111/brv.12552
                7004074
                31538399
                00e909b0-f7a3-4ada-958d-3e2b6725335c
                © 2019 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.

                This is an open access article under the terms of the http://creativecommons.org/licenses/by-nc/4.0/ License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited and is not used for commercial purposes.

                History
                : 26 February 2019
                : 12 August 2019
                : 15 August 2019
                Page count
                Figures: 6, Tables: 3, Pages: 29, Words: 23089
                Funding
                Funded by: Faculty of Science , open-funder-registry 10.13039/501100003803;
                Funded by: HKU MOOC course on Dinosaur Ecosystems , open-funder-registry 10.13039/501100003803;
                Funded by: Hong Kong PhD Fellowship
                Award ID: HKPF PF16‐09281
                Categories
                Original Article
                Original Articles
                Custom metadata
                2.0
                February 2020
                Converter:WILEY_ML3GV2_TO_JATSPMC version:5.7.5 mode:remove_FC converted:06.02.2020

                Ecology
                palaeocolour,melanin,melanosomes,exceptional preservation,amniotes,taphonomy
                Ecology
                palaeocolour, melanin, melanosomes, exceptional preservation, amniotes, taphonomy

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