Dinosaur Plumage Coloration and appearance provide important behavioral and evolutionary information in animals. However, for the most part, we do not know the coloration of fossil terrestrial animals. Li et al. (p. 1369 , published online 4 February) have reconstructed the appearance of a theropod dinosaur by mapping features of its well-preserved feathers and comparing them with modern samples from birds. Feather color is partly determined by melanosome density and shape, and this information is preserved in a recently discovered fossil from China. The dinosaur was gray with white limbs and had a reddish crest and a speckled face.

Iridescent feather colors involved in displays of many extant birds are produced by nanoscale arrays of melanin-containing organelles (melanosomes). Data relevant to the evolution of these colors and the properties of melanosomes involved in their generation have been limited. A data set sampling variables of extant avian melanosomes reveals that those forming most iridescent arrays are distinctly narrow. Quantitative comparison of these data with melanosome imprints densely sampled from a previously unknown specimen of the Early Cretaceous feathered Microraptor predicts that its plumage was predominantly iridescent. The capacity for simple iridescent arrays is thus minimally inferred in paravian dinosaurs. This finding and estimation of Microraptor feathering consistent with an ornamental function for the tail suggest a centrality for signaling in early evolution of plumage and feather color.

Melanosomes are organelles that produce and store melanin, a widespread biological pigment with a unique suite of properties including high refractive index, semiconducting capabilities, material stiffness, and high fossilization potential. They are involved in numerous critical biological functions in organisms across the tree of life. Individual components such as melanin chemistry and melanosome development have recently been addressed, but a broad synthesis is needed. Here, we review the hierarchical structure, development, functions, and evolution of melanosomes. We highlight variation in melanin chemistry and melanosome morphology and how these may relate to function. For example, we review what is known of the chemical differences between different melanin types (eumelanin, pheomelanin, allomelanin) and whether/how melanosome morphology relates to chemistry and color. We integrate the distribution of melanin across living organisms with what is known from the fossil record and produce hypotheses on its evolution. We suggest that melanin was present in life forms early in evolutionary history and that melanosomes evolved at the origin of organelles. Throughout, we discuss the (sometimes gaping) holes in our knowledge and suggest areas that need particular attention as we move forward in our understanding of these still-mysterious organelles and the materials that they contain.

Penguin feathers are highly modified in form and function, but there have been no fossils to inform their evolution. A giant penguin with feathers was recovered from the late Eocene (~36 million years ago) of Peru. The fossil reveals that key feathering features, including undifferentiated primary wing feathers and broad body contour feather shafts, evolved early in the penguin lineage. Analyses of fossilized color-imparting melanosomes reveal that their dimensions were similar to those of non-penguin avian taxa and that the feathering may have been predominantly gray and reddish-brown. In contrast, the dark black-brown color of extant penguin feathers is generated by large, ellipsoidal melanosomes previously unknown for birds. The nanostructure of penguin feathers was thus modified after earlier macrostructural modifications of feather shape linked to aquatic flight.

Inference of colour patterning in extinct dinosaurs has been based on the relationship between the morphology of melanin-containing organelles (melanosomes) and colour in extant bird feathers. When this relationship evolved relative to the origin of feathers and other novel integumentary structures, such as hair and filamentous body covering in extinct archosaurs, has not been evaluated. Here we sample melanosomes from the integument of 181 extant amniote taxa and 13 lizard, turtle, dinosaur and pterosaur fossils from the Upper-Jurassic and Lower-Cretaceous of China. We find that in the lineage leading to birds, the observed increase in the diversity of melanosome morphologies appears abruptly, near the origin of pinnate feathers in maniraptoran dinosaurs. Similarly, mammals show an increased diversity of melanosome form compared to all ectothermic amniotes. In these two clades, mammals and maniraptoran dinosaurs including birds, melanosome form and colour are linked and colour reconstruction may be possible. By contrast, melanosomes in lizard, turtle and crocodilian skin, as well as the archosaurian filamentous body coverings (dinosaur 'protofeathers' and pterosaur 'pycnofibres'), show a limited diversity of form that is uncorrelated with colour in extant taxa. These patterns may be explained by convergent changes in the key melanocortin system of mammals and birds, which is known to affect pleiotropically both melanin-based colouration and energetic processes such as metabolic rate in vertebrates, and may therefore support a significant physiological shift in maniraptoran dinosaurs.

Animal integumentary coloration plays a crucial role in visual communication and camouflage, and varies extensively among and within species and populations. To understand the pressures underlying such diversity, it is essential to elucidate the mechanisms by which animals have created novel integumentary coloration. Colours can be produced by selective absorption of light by skin pigments, through light scattering by structured or unstructured tissues, or by a combination of pigments and nanostructures. In this review, we highlight our current understanding of the interactions between pigments and structural integumentary tissues and molecules. We analyse the available evidence suggesting that these combined mechanisms are capable of creating colours and optical properties unachievable by either mechanism alone, thereby effectively expanding the animal colour palette. Moreover, structural and pigmentary colour mechanisms frequently interact in unexpected and overlooked ways, suggesting that classification of colours as being of any particular type may be difficult. Finally, we discuss how these mixtures are useful for investigating the largely unknown genetic, developmental and physical processes generating phenotypic diversity.This article is part of the themed issue 'Animal coloration: production, perception, function and application'.

Archaeopteryx has been regarded as an icon of evolution ever since its discovery from the Late Jurassic limestone deposits of solnhofen, Germany in 1861. Here we report the first evidence of colour from Archaeopteryx based on fossilized colour-imparting melanosomes discovered in this isolated feather specimen. using a phylogenetically diverse database of extant bird feathers, statistical analysis of melanosome morphology predicts that the original colour of this Archaeopteryx feather was black, with 95% probability. Furthermore, reexamination of the feather's morphology leads us to interpret it as an upper major primary covert, contrary to previous interpretations. Additional findings reveal that the specimen is preserved as an organosulphur residue, and that barbule microstructure identical to that of modern bird feathers had evolved as early as the Jurassic. As in extant birds, the extensive melanization would have provided structural advantages to the Archaeopteryx wing feather during this early evolutionary stage of dinosaur flight.

The Jurassic Yanliao theropods have offered rare glimpses of the early paravian evolution and particularly of bird origins, but, with the exception of the bizarre scansoriopterygids, they have shown similar skeletal and integumentary morphologies. Here we report a distinctive new Yanliao theropod species bearing prominent lacrimal crests, bony ornaments previously known from more basal theropods. It shows longer arm and leg feathers than Anchiornis and tail feathers with asymmetrical vanes forming a tail surface area even larger than that in Archaeopteryx. Nanostructures, interpreted as melanosomes, are morphologically similar to organized, platelet-shaped organelles that produce bright iridescent colours in extant birds. The new species indicates the presence of bony ornaments, feather colour and flight-related features consistent with proposed rapid character evolution and significant diversity in signalling and locomotor strategies near bird origins.