Exceptional Three-Dimensional Preservation and Coloration of an Originally Iridescent Fossil Feather from the Middle Eocene Messel Oil Shale

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Exceptional Three-Dimensional Preservation and Coloration of an Originally Iridescent Fossil Feather from the Middle Eocene Messel Oil Shale Exceptional three-dimensional preservation and coloration of an originally iridescent fossil feather from the Middle Eocene Messel Oil Shale Natasha S. Vitek, Jakob Vinther, James D. Schiffbauer, Derek E. G. Briggs & Richard O. Prum Paläontologische Zeitschrift Scientific Contributions to Palaeontology ISSN 0031-0220 Volume 87 Number 4 Paläontol Z (2013) 87:493-503 DOI 10.1007/s12542-013-0173-5 1 23 Your article is protected by copyright and all rights are held exclusively by Springer- Verlag Berlin Heidelberg. This e-offprint is for personal use only and shall not be self- archived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 1 23 Author's personal copy Pala¨ontol Z (2013) 87:493–503 DOI 10.1007/s12542-013-0173-5 RESEARCH PAPER Exceptional three-dimensional preservation and coloration of an originally iridescent fossil feather from the Middle Eocene Messel Oil Shale Natasha S. Vitek • Jakob Vinther • James D. Schiffbauer • Derek E. G. Briggs • Richard O. Prum Received: 3 September 2012 / Accepted: 12 February 2013 / Published online: 16 March 2013 Ó Springer-Verlag Berlin Heidelberg 2013 Abstract A feather from the Eocene Messel Formation, generated by incoherent light diffraction between this Germany, has been demonstrated to have been originally sediment layer and melanosomes and, although related to structurally colored by densely packed sheets of melano- the original iridescence of the feather, is not a feature of the somes similar to modern iridescent feathers exhibiting thin- feather itself. The reddish and greenish hues frequently film diffraction. The fossil itself currently exhibits a silvery exhibited by fossil feathers from the Messel Formation sheen, but the mechanism for generating this optical effect appear to be due to precipitates on the surface of individual was not fully understood. Here we use scanning electron melanosomes. microscopy, electron probe microanalysis, and dual-beam focused ion beam scanning electron microscopy to inves- Keywords Exceptional preservation Á Color Á Melanin Á tigate the source of the silvery sheen that occurs in the Iridescence Á Fossil bird apical feather barbules. Focused ion beam scanning elec- tron microscopy provides a powerful tool for studying Kurzfassung Bei einer Feder aus der eoza¨nen Messel three-dimensionality of nanostructures in fossils. Use of the Formation von Deutschland wurde eine originale strukturelle method reveals that the flattened apical barbules are pre- Fa¨rbung aus dicht gepackten Melanosomen nachgewiesen, die served almost perfectly, including smooth structural mel- modernen irridisierenden Federn a¨hnlich ist, die Du¨nns- anosome sheets on the obverse surface of the fossil feather chichtdiffraktion zeigen. Das Fossil selbst zeigt in seinem that are identical to those that cause iridescence in modern derzeitigen Zustand einen silbrigen Glanz, aber es ist unklar, bird feathers. Most of each apical barbule is preserved durch welchen Mechanismus dieser entsteht. Mit Hilfe von beneath a thin layer of sediment. The silvery sheen is Rasterelektronenmikroskopie, Elektronenstrahlmikroanalyse N. S. Vitek (&) Á J. Vinther Á D. E. G. Briggs D. E. G. Briggs Á R. O. Prum Department of Geology and Geophysics, Yale University, Peabody Museum of Natural History, Yale University, 210 Whitney Avenue, New Haven, CT 06520, USA 170 Whitney Avenue, New Haven, CT 06511, USA e-mail: [email protected] e-mail: [email protected] N. S. Vitek Á J. Vinther R. O. Prum Jackson School of Geosciences, The University of Texas Department of Ecology and Evolutionary Biology, at Austin, Austin, TX 78712, USA Yale University, 21 Sachem Street, New Haven, CT 06520, USA J. Vinther e-mail: [email protected] Department of Earth Sciences and Biological Sciences, University of Bristol, Bristol BS8 1UG, UK e-mail: [email protected] J. D. Schiffbauer Department of Geological Sciences, University of Missouri, Columbia, MO 65211, USA e-mail: [email protected] 123 Author's personal copy 494 N. S. Vitek et al. und doppelt fokussierter Ionenfeinstrahlmikroskopie untersu- presence of iron in different oxidation states. The silvery chen wir die Herkunft der gla¨nzenden Fa¨rbung, die an den sheen was hypothesized to be a direct result of the apikalen Hakenstrahlen auftritt. Ionenfeinstrahlmikroskopie arrangement of the fossil’s originally iridescent melano- stellt eine leistungsfa¨hige Methode zur Untersuchung dreidi- some sheets. The latter hypothesis was based on the spatial mensionaler Nanostrukturen in Fossilien dar. Ergebnisse dieser correlation between the sheets and the silvery sheen in the Untersuchungsmethode zeigen, dass sowohl die abgeflachten, barbules of the distal part of the feather vane. In this case, apikalen Hakenstrahlen, als auch die einzelnen Schichten aus such a silvery sheen would provide an easily detectable Strukturmelanosomen, die mit denen heutiger Vogelfedern indication of iridescence in other fossil birds and in nahezu identisch sind, anna¨hernd unversehrt u¨berliefert sind. feathered dinosaurs. Ein Großteil der apikalen Hakenstrahlen ist unter einer du¨nnen Melanosomes are known to absorb metal ions in other Sedimentschicht erhalten, die aufgrund der Interferenz mit den contexts. Melanin granules in the squid Sepia have been Melanosomenzudemsilbergla¨nzenden Farbeffekt fu¨hrt. shown to absorb metals such as iron, calcium, and mag- Dieser Effekt wird durch die irridisierende Fa¨rbung der Feder nesium without alteration of the melanosome macrostruc- versta¨rkt, trat allerdings nicht im Originalzustand auf. Dies ture (Liu et al. 2004). Extant birds may accumulate metals deutet darauf hin, dass die ro¨tliche und gru¨nliche Fa¨rbung, die such as calcium, zinc, copper, iron, and magnesium in their typisch fu¨r Federn aus Messel ist, aufgrund von Ablagerungen feather melanin (Niecke et al. 1999; McGraw 2008). Those auf der Oberfla¨che der einzelnen Melanosome zustande metals are released in reducing conditions (Liu et al. 2004), kommt. and presumably also during diagenesis, with the removal of hydroxyl (–OH) and carboxyl (–COOH) functional groups Schlu¨sselwo¨rter Außergewo¨hnliche Erhaltung Á that chelated the metals in the macromolecule (Riley Farberhaltung Á Melanin Á Iridisieren Á Fossile Vo¨gel 1997). Here we investigate the mechanism giving rise to the silvery sheen in the Messel feather and test the hypothesis Introduction that the red hue is the result of diagenetic precipitation of metal ions, likely sourced from the melanosomes (Vinther Distinct arrangements and morphologies of melanosomes et al. 2010). Investigating the mechanism of color pro- can generate structural coloration and metallic iridescence duction in the fossil, and any correlation between the in modern bird feathers by creating coherently scattered preserved color and that of the original feather, requires a light (Prum 2006). Melanin is highly decay resistant and clear understanding of which components of the fossil can survive with little alteration in fossils at least as old as remain on the part and counterpart when the shale is split. the Jurassic (Glass et al. 2012). Many fossil feathers are Furthermore, as compared with modern feathers, only the preserved as remains of melanosomes, which are melanin- highly melanized components of the fossil feather are containing organelles (Vinther et al. 2008, 2010; Zhang likely to survive decay (Goldstein et al. 2004). The et al. 2010). Additionally, melanosome shape and distri- appearance of the fossil may be affected by both the bution offer a means of inferring ancient coloration and associated sediment matrix and the loss of the nonmelan- color patterns (Li et al. 2010, 2012; Clarke et al. 2010). ized feather parts. Fossil feathers may preserve smooth sheets of melano- somes in arrangements similar to those that generate iri- descence in modern birds (Vinther et al. 2010). The Methods iridescent sheens in such feathers are created by so-called thin-film diffraction of a smooth layer of keratin above an We investigated the Messel feather fossil SMF ME 3850 equally smooth melanosome surface (Shawkey et al. using a dual-beam focused ion beam scanning electron 2006). The fossil feather examined here, from the Senc- microscope (FIB-SEM) to image surface topography kenberg Museum Frankfurt, SMF ME 3850, was collected and concurrently ion-mill site-specific cross-sections from the famous Eocene Messel locality (*47 Ma) in (Schiffbauer and Xiao 2009, 2011) in order to document Germany. It exhibits a conspicuous red hue that alternates the three-dimensional nature of the exceptional preserva- in places with stripes of a silvery sheen in the barbules of tion and elucidate the origin of the silvery sheen. We used the apical region (Fig. 1). Initially, the red hue was inter- FIB-SEM analysis and electron probe microanalysis preted as an effect of the absorption of metal ions onto the (EPMA) to determine the chemical composition of the surface of the eumelanosomes, which are rod-shaped and feather and surrounding sediment, and to investigate the originally produce black and gray colors (Vinther et al. origin of the reddish and greenish hues exhibited by some 2010). The fossil feathers in Messel vary from black to fossil feathers from the Messel Formation (Vinther et al. reddish and greenish in hue, which may indicate the 2010, suppl. Figs. 1 and 2). 123 Author's personal copy Eocene Messel Oil Shale 495 Fig. 1 Color photo of fossil SMF ME 3850 (main slab) showing the Inset shows a detail of the apical barbs shown in Figs. 1d and 4. FIB- locations of each FIB-SEM cross-section. Labeled boxes correspond SEM cuts are marked in blue. Arrows indicate the direction of view.
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