Fossil Feathers Research Articles

Bio-molecular analyses enable new insights into the taphonomy of feathers.

Exceptionally preserved feathers from the Mesozoic era have provided valuable insights into the early evolution of feathers and enabled color reconstruction of extinct dinosaurs, including early birds. Mounting chemical evidence for the two key components of feathers-keratins and melanins-in fossil feathers has demonstrated that exceptional preservation can be traced down to the molecular level. However, the chemical changes that keratin and eumelanin undergo during fossilization are still not fully understood, introducing uncertainty in the identification of these two molecules in fossil feathers. To address this issue, we need to examine their taphonomic process. In this study, we analyzed the structural and chemical composition of fossil feathers from the Jehol Biota and compared them with the structural and chemical changes observed in modern feathers during the process of biodegradation and thermal degradation, as well as the structural and chemical characteristics of a Cenozoic fossil feather. Our results suggest that the taphonomic process of feathers from the Cretaceous Jehol Biota is mainly controlled by the process of thermal degradation. The Cretaceous fossil feathers studied exhibited minimal keratin preservation but retained strong melanin signals, attributed to melanin's higher thermal stability. Low-maturity carbonaceous fossils can indeed preserve biosignals, especially signals from molecules with high resistance to thermal degradation. These findings provide clues about the preservation potential of keratin and melanin, and serve as a reference for searching for those two biomolecules in different geological periods and environments.

Cellular structure of dinosaur scales reveals retention of reptile-type skin during the evolutionary transition to feathers

Fossil feathers have transformed our understanding of integumentary evolution in vertebrates. The evolution of feathers is associated with novel skin ultrastructures, but the fossil record of these changes is poor and thus the critical transition from scaled to feathered skin is poorly understood. Here we shed light on this issue using preserved skin in the non-avian feathered dinosaur Psittacosaurus. Skin in the non-feathered, scaled torso is three-dimensionally replicated in silica and preserves epidermal layers, corneocytes and melanosomes. The morphology of the preserved stratum corneum is consistent with an original composition rich in corneous beta proteins, rather than (alpha-) keratins as in the feathered skin of birds. The stratum corneum is relatively thin in the ventral torso compared to extant quadrupedal reptiles, reflecting a reduced demand for mechanical protection in an elevated bipedal stance. The distribution of the melanosomes in the fossil skin is consistent with melanin-based colouration in extant crocodilians. Collectively, the fossil evidence supports partitioning of skin development in Psittacosaurus: a reptile-type condition in non-feathered regions and an avian-like condition in feathered regions. Retention of reptile-type skin in non-feathered regions would have ensured essential skin functions during the early, experimental stages of feather evolution.

Preservation of corneous β-proteins in Mesozoic feathers.

Fossil proteins are valuable tools in evolutionary biology. Recent technological advances and better integration of experimental methods have confirmed the feasibility of biomolecular preservation in deep time, yielding new insights into the timing of key evolutionary transitions. Keratins (formerly α-keratins) and corneous β-proteins (CBPs, formerly β-keratins) are of particular interest as they define tissue structures that underpin fundamental physiological and ecological strategies and have the potential to inform on the molecular evolution of the vertebrate integument. Reports of CBPs in Mesozoic fossils, however, appear to conflict with experimental evidence for CBP degradation during fossilization. Further, the recent model for molecular modification of feather chemistry during the dinosaur-bird transition does not consider the relative preservation potential of different feather proteins. Here we use controlled taphonomic experiments coupled with infrared and sulfur X-ray spectroscopy to show that the dominant β-sheet structure of CBPs is progressively altered to α-helices with increasing temperature, suggesting that (α-)keratins and α-helices in fossil feathers are most likely artefacts of fossilization. Our analyses of fossil feathers shows that this process is independent of geological age, as even Cenozoic feathers can comprise primarily α-helices and disordered structures. Critically, our experiments show that feather CBPs can survive moderate thermal maturation. As predicted by our experiments, analyses of Mesozoic feathers confirm that evidence of feather CBPs can persist through deep time.

Downy feather fossils from the Lower Cretaceous of the Jiuquan Basin, Gansu Province, Northwest China

Mesozoic feather fossils are well known for their exceptional preservation. However, the scarcity of quantities and the lack of fine structural details preserved in carbonaceous fossils, most studies have focused on the macro-morphology of such feather fossils, neglecting the significance of the microscopic morphology. A group of 28 downy feather fossils preserved in the Lower Cretaceous strata of the Jiuquan Basin are similar to the modern feather, including basic structures such as calamus, barb, and barbules, and are considered to be early downy feathers. Based on well-preserved and quantified feather characteristics, multivariate statistical analysis (PCA and CVA) was used to compare fossil groups with modern down feathers. The results revealed that the microscopic morphology of fossil feathers was noticeably different from living birds, that feathers from the Early Cretaceous may not yet be true down feathers. And the species specificity between different specimens within the fossil taxa suggests that they may have come from different coelurosaurs. This gives us further information about the evolution of Early Cretaceous feathers.

Sediment-encased pressure–temperature maturation experiments elucidate the impact of diagenesis on melanin-based fossil color and its paleobiological implications

AbstractMelanin pigments are central to colors and patterns in modern vertebrate integuments, which inform upon ecological and behavioral strategies like crypsis, aposematism, and sociosexual selection. Over the last decade, melanin has emerged as a valuable tool for predicting color in exceptionally preserved fossil feathers and subsequent testing of paleobiological hypotheses in long-extinct dinosaurs and birds. Yet much remains to be learned about melanin stability, diagenetic alterations to melanin chemistry, and their implications for “paleocolor reconstruction.” Pressure–temperature maturation experiments with modern feathers offer a way to examine these topics but have mostly been conducted in closed-system capsules or open-system aluminum foil. Both methods have operational limitations and do not consider the filtering effect of porous sediment matrices on thermally labile chemical groups versus stable ones during natural fossilization. We use sediment-encased maturation to resolve this issue and demonstrate replication of organic preservation of melanin highly comparable to compression fossils. Our experiments, coupled with time-of-flight secondary ion mass spectrometry, show predictable volatilization of N/S-bearing molecules and increased melanin cross-linking with elevated temperatures. We also suggest that eumelanin is more stable compared with pheomelanin at higher temperatures, explaining why eumelanic colors (black, dark brown, iridescent) are better preserved in fossils than pheomelanic ones (reddish brown). Furthermore, we propose that proteins preferentially undergo hydrolysis more so than forming N-heterocycles in selectively open systems analogous to natural matrices. Thus, we conclude that melanin pigments and not diagenetically altered protein remnants are the key players in promoting fossilization of soft tissues like feathers.

Immature feathers preserved in Burmite provide evidence of rapid molting in enantiornithines

Fossil feathers are recorded in the greatest fidelity in amber. A new specimen of Cenomanian aged Burmese amber provides the first unequivocal evidence of immature feathers in the Mesozoic fossil record. The small, clear piece, identified as aerial amber, preserves a cluster of body feathers with their basal portions encased in keratinous sheaths. The feathers are identified as belonging to a young enantiornithine individual based on the combination of preserved feather morphotypes, degree of feather asymmetry in the single preserved remix, their diminutive size, and because only enantiornithines have so far been recovered from these amber deposits. The body feathers are all at approximately the same stage of development suggesting that immature enantiornithines rapidly molted body feathers. This interpretation is supported by evidence from lithic specimens of juveniles from the Lower Cretaceous Jehol group that preserve immature feathers throughout the body. This specimen contributes to evidence that indicates enantiornithines hatched flight ready, with fully developed remiges, but with sparse body plumage that was simultaneously replaced during the prejuvenal molt. Rapid molts indicate periodic increases in thermostatic costs due to the temporary loss of insulation. This information leads to an intriguing new hypothesis regarding the extinction of Enantiornithes: that periods of increased thermostatic cost during early ontogeny created untenable energetic budgets during the asteroid induced impact winter, which resulted in both decreased global temperature and resource scarcity. High energetic budgets are associated with extinction vulnerability and may have been a major factor underpinning the extinction of this clade.

Silicification of feathers in a modern hot spring in New Zealand

ABSTRACT Fossil feathers have greatly improved our understanding of the evolutionary transition from non-avian dinosaurs to birds and the evolution of feathers, and may be the only evidence for their source animals in the fossil record. Hot spring environments have been demonstrated to be conducive to the preservation of fossils, but internal silicification of feathers was not observed in the only avian carcass so far discovered in ancient hot spring deposits. To determine whether feathers can be internally silicified, here we analyse feathers sampled from a modern hot spring vent pool – Champagne Pool – in New Zealand. Our results of scanning electron microscopy (SEM)-energy dispersive X-ray spectrometry elemental mapping show that the sampled feathers are silicified to different degrees, and one of them is pervasively silicified. SEM observations show that feathers can be silicified at the cellular level. Degradation is involved in the silicification of feathers, as indicated by the reduction of the abundance of carbon and the loss of keratin fibrils. Our findings suggest that ancient deposits of hot spring vent pools are promising targets in search for fossil feathers.

Exceptional Preservation of Feather Micro-Structures in Amber from the Middle-Cretaceous of Myanmar

Feathers are under-represented in the fossil record because soft tissues do not usually preserve well in sedimentary sequences. Fossil feathers are nevertheless extremely important in resolving pattern and process related to the origin of dinosaur flight. In recent years, a number of feathers have been discovered which have been mummified in amber; these feathers are preserved in three dimensions with remarkable sub-microscopic details and are especially important for our understanding of the early development of feathers. In this paper, we describe a diverse assemblage of mid-Cretaceous feathers contained within seven pieces of amber that have been recovered from northern Myanmar (Burma). These pieces include pennaceous primary feathers, contour feathers, and rachis-dominated feathers, and also a plumulaceous (downy) feather. Subcomponents of these feathers, such as barbs, barbules, and nodes are immediately recognizable. One extraordinary piece contains the distal remains of the first four primary flight feathers and a small number of possible hooklets. These pieces are discussed in terms of evolutionary development and comments are made on flight ability where appropriate. These feather-types are classified and compared with similar structures seen in Mesozoic and extant birds. We consider that integumentary feathers and ‘feather-like’ structures fall within two major structural categories (‘shafted’ and ‘non-shafted’).

Evidence corroborates identity of isolated fossil feather as a wing covert of Archaeopteryx

The historic fossil feather from the Jurassic Solnhofen has played a pivotal but controversial role in our evolutionary understanding of dinosaurs and birds. Recently, a study confirmed the diagnostic morphology of the feather’s original calamus, but nonetheless challenged the proposed identity as an Archaeopteryx covert. However, there are errors in the results and interpretations presented. Here we show that the feather is most likely an upper major primary covert, based on its long calamus (23.3% total length) and eight other anatomical attributes. Critically, this hypothesis is independently supported by evidence of similar primary coverts in multiple specimens of Archaeopteryx–including from the same fossil site and horizon as the isolated feather. We also provide additional insights, such as an updated colour reconstruction of the entire feather as matte black, with 90% probability. Given the isolated nature of the fossil feather, we can never know the anatomical and taxonomic provenance with 100% certainty. However, based on all available evidence, the most empirical and parsimonious conclusion is that this feather represents a primary covert from the ancient wing of Archaeopteryx.

Chemical preservation of tail feathers fromAnchiornis huxleyi, a theropod dinosaur from the Tiaojishan Formation (Upper Jurassic, China)

AbstractA panel of geochemical techniques is used here to investigate the taphonomy of fossil feathers preserved in association with the skeleton of the Jurassic theropodAnchiornis huxleyi. Extant feathers were analysed in parallel to test whether the soft tissues morphologically preserved in the fossil also exhibit a high degree of chemical preservation. Scanning electron microscopy (SEM) and energy dispersive spectroscopy (EDS) indicate that clays and iron oxide pseudomorphs occur in the surrounding sediment and also reveal the preservation of melanosome‐like microbodies in the fossil. Carbon gradient along a depth profile and co‐occurrence of carbon and sulphur are shown in the fossil by elastic backscattering (EBS) and particle‐induced x‐ray emission (PIXE), which are promising techniques for the elemental analysis of fossil soft tissues. The molecular composition of modern and fossil soft tissues was assessed from micro‐attenuated total reflectance fourier transform infrared spectroscopy (micro‐ATR FTIR), solid‐state13C nuclear magnetic resonance (CP‐MAS13C NMR) and pyrolysis gas chromatography mass spectrometry in the presence of TMAH (TMAH‐Py‐GC‐MS). Results indicate that the proteinaceous material that comprises the modern feathers is not present in the fossil feathers. The fossil feathers and the embedding sediment exhibit a highly aliphatic character. However, substantial differences exist between these samples, revealing that the organic matter of the fossil feathers is, at least partially, derived from original constituents of the feathers. Our results suggest that, despite the morphological preservation ofAnchiornisfeathers, original proteins, that is keratin, were probably not preserved in the 160‐myr‐old feathers.

Mimicry in Cretaceous Bugs.

SummaryMimicry is ubiquitous in nature, yet understanding its origin and evolution is complicated by the scarcity of exceptional fossils that enable behavioral inferences about extinct animals. Here we report bizarre true bugs (Hemiptera) that closely resemble beetles (Coleoptera) from mid-Cretaceous amber. The unusual fossil bugs are described as Bersta vampirica gen. et sp. nov. and Bersta coleopteromorpha gen. et sp. nov. and are placed into a new family, Berstidae fam. nov. The specialized mouthparts of berstids indicate that they were predaceous on small arthropods. Their striking beetle-like appearance implies that they were either involved in defensive mimicry or mimicked beetles to attack unsuspecting prey. The latter would represent the first case of aggressive mimicry in the invertebrate fossil record. These findings enrich our understanding of the paleoecological associations and extinct behavioral strategies of Mesozoic insects.

EXPERIMENTAL MATURATION OF FEATHERS: IMPLICATIONS FOR INTERPRETATIONS OF FOSSIL FEATHERS

ABSTRACT Exceptionally preserved fossil feathers and feather-like integumentary structures provide valuable insights into the early evolution of feathers and flight, but taphonomic biases often make interpretations at the microstructural and ultrastructural levels ambiguous. Maturation experiments have been demonstrated to be useful for investigating the taphonomic alterations of soft tissues, including feathers, during diagenesis. However, experimentally matured feathers resembling fossil feathers preserving keratinous matrix have not yet been obtained. Here we employ experimental maturation to obtain feathers corresponding to different degradation stages, and compare these matured feathers with untreated feathers and fossil feathers at the macroscopic, microstructural, and ultrastructural levels. Results show that several features of thermally matured feathers are similar to those found in fossil feathers. The fusion of barbules that occurred in thermally matured feathers suggests that such a process could occur during diagenesis, making barbules difficult to identify in fossil feathers. Under the most extreme experimental condition, the keratinous matrix can partially survive when the whole feather is turned into ash-like remains and many melanosomes are exposed. Moreover, our results show that the keratinous matrix immediately surrounding melanosomes appears to be more resistant to degradation than the unpigmented keratinous matrix, supporting the hypothesis that melanin can act as a fixative agent to prevent the degradation of keratin.

A polar dinosaur feather assemblage from Australia

Exceptionally preserved Mesozoic feathered dinosaur fossils (including birds) are famous, but recognized from only very few localities worldwide, and are especially rare in the Southern Hemisphere. Here we report an assemblage of non-avian and avian dinosaur feathers from an Early Cretaceous polar (around 70°S) environment in what is now southeastern Australia. The recovered remains incorporate small (10–30 mm long) basal paravian-like tufted body feathers, open-vaned contour feathers, and asymmetrical bird-like wing feathers that possess high-angled barbs with possible remnants of barbicels — amongst the geologically oldest observed to date. Such morphological diversity augments scant skeletal evidence for a range of insulated non-avian theropods and birds inhabiting extreme southern high-latitude settings during the Mesozoic. Although some of these fossil feathers exhibit what may be residual patterning, most are uniformly toned and preserve rod-shaped microbodies, as well as densely-packed microbody imprints on the barbules that are structurally consistent with eumelanosomes. Geochemical analysis detected no identifiable residual biomolecules, which we suspect were lost via hydrolysis and oxidization during diagenesis and weathering. Nevertheless, an originally dark pigmentation can be reasonably inferred from these melanic traces, which like the coloured feathers of modern birds, might have facilitated crypsis, visual communication and/or thermoregulation in a cold polar habitat.

Taphonomic experiments resolve controls on the preservation of melanosomes and keratinous tissues in feathers.

Fossils are a key source of data on the evolution of feather structure and function through deep time, but their ability to resolve macroevolutionary questions is compromised by an incomplete understanding of their taphonomy. Critically, the relative preservation potential of two key feather components, melanosomes and keratinous tissue, is not fully resolved. Recent studies suggesting that melanosomes are preferentially preserved conflict with observations that melanosomes preserve in fossil feathers as external moulds in an organic matrix. To date, there is no model to explain the latter mode of melanosome preservation. We addressed these issues by degrading feathers in systematic taphonomic experiments incorporating decay, maturation and oxidation in isolation and combination. Our results reveal that the production of mouldic melanosomes requires interactions with an oxidant and is most likely to occur prior to substantial maturation. This constrains the taphonomic conditions under which melanosomes are likely to be fossilized. Critically, our experiments also confirm that keratinous feather structures have a higher preservation potential than melanosomes under a range of diagenetic conditions, supporting hitherto controversial hypotheses that fossil feathers can retain degraded keratinous structures.

Ancient amino acids from fossil feathers in amber

Ancient protein analysis is a rapidly developing field of research. Proteins ranging in age from the Quaternary to Jurassic are being used to answer questions about phylogeny, evolution, and extinction. However, these analyses are sometimes contentious, and focus primarily on large vertebrates in sedimentary fossilisation environments; there are few studies of protein preservation in fossils in amber. Here we show exceptionally slow racemisation rates during thermal degradation experiments of resin enclosed feathers, relative to previous thermal degradation experiments of ostrich eggshell, coral skeleton, and limpet shell. We also recover amino acids from two specimens of fossil feathers in amber. The amino acid compositions are broadly similar to those of degraded feathers, but concentrations are very low, suggesting that much of the original protein has been degraded and lost. High levels of racemisation in more apolar, slowly racemising amino acids suggest that some of the amino acids were ancient and therefore original. Our findings indicate that the unique fossilisation environment inside amber shows potential for the recovery of ancient amino acids and proteins.

Detection of lost calamus challenges identity of isolated Archaeopteryx feather

In 1862, a fossil feather from the Solnhofen quarries was described as the holotype of the iconic Archaeopteryx lithographica. The isolated feather’s identification has been problematic, and the fossil was considered either a primary, secondary or, most recently, a primary covert. The specimen is surrounded by the ‘mystery of the missing quill’. The calamus described in the original paper is unseen today, even under x-ray fluorescence and UV imaging, challenging its original existence. We answer this question using Laser-Stimulated Fluorescence (LSF) through the recovery of the geochemical halo from the original calamus matching the published description. Our study therefore shows that new techniques applied to well-studied iconic fossils can still provide valuable insights. The morphology of the complete feather excludes it as a primary, secondary or tail feather of Archaeopteryx. However, it could be a covert or a contour feather, especially since the latter are not well known in Archaeopteryx. The possibility remains that it stems from a different feathered dinosaur that lived in the Solnhofen Archipelago. The most recent analysis of the isolated feather considers it to be a primary covert. If this is the case, it lacks a distinct s-shaped centerline found in modern primary coverts that appears to be documented here for the first time.

Combined microscopy and spectroscopy techniques to characterize a fossilized feather with minimal damage to the specimen

The study of fossil feathers has been revitalized in the last few decades and has contributed significantly to paleontological studies of dinosaurs and birds. Specific morphological and physicochemical characteristics of the microscale structures of feathers and the protein keratin are key targets when preserved during the fossilization process. Keratin is a fibrous protein that composes some hard tissues such as hair, nails and feathers. It is part of the so called intermediate filaments inside keratinocyte cells and is rich in sulfur containing amino acid cysteine. To date, different microscopy and analytical methods have been used for the analysis and detailed characterization and classification of feathers. However, in this work we showed that analytical optical and electron microscopies can be quick and precise methods with minimal effects on the sample during analysis. This association of different approaches on the same sample results in correlative data albeit in different length scales. Intracellular bodies called melanosomes originally present in melanocyte cells were identified with Scanning Electron Microscopy (SEM) and Transmission Electron Microscopy (TEM), and had well-defined orientation and a mean aspect ratio comparable to melanosomes extant in dark feathers. The detection of sulphur in melanosomes via Energy Dispersive Spectroscopy both in SEM and TEM shows that, along the fossilization process, sulphur from the degraded keratin matrix could have been trapped inside the melanosomes. Chemical groups that make up keratin and melanin in the fossil sample were detected via FT-IR Spectroscopy and Confocal Laser Scanning Microscopy (CLSM). The use of combined analytical microscopy techniques can contribute significantly to the study of fossils generating precise results with minimum damage to the original sample.

Fossil feathers reveal how dinosaurs took flight

Fossil feathers reveal how dinosaurs took flight

Preservation potential of keratin in deep time.

Multiple fossil discoveries and taphonomic experiments have established the durability of keratin. The utility and specificity of antibodies to identify keratin peptides has also been established, both in extant feathers under varying treatment conditions, and in feathers from extinct organisms. Here, we show localization of feather-keratin antibodies to control and heat-treated feathers, testifying to the repeatability of initial data supporting the preservation potential of keratin. We then show new data at higher resolution that demonstrates the specific response of these antibodies to the feather matrix, we support the presence of protein in heat-treated feathers using ToF-SIMS, and we apply these methods to a fossil feather preserved in the unusual environment of sinter hot springs. We stress the importance of employing realistic conditions such as sediment burial when designing experiments intended as proxies for taphonomic processes occurring in the fossil record. Our data support the hypothesis that keratin, particularly the β-keratin that comprises feathers, has potential to preserve in fossil remains.

Experimental subaqueous burial of a bird carcass and compaction of plumage

‘Exceptional fossils’ of dinosaurs preserving feathers have radically changed the way we view their paleobiology and the evolution of birds. Understanding how such soft tissues preserve is imperative to accurately interpreting the morphology of fossil feathers. Experimental taphonomy has been integral to such investigations. One such experiment used a printing press to mimic compaction, done subaerially and without sediment burial, and concluded that the leaking of bodily fluid could lead to the clumping of feathers by causing barbs to stick together such that they superficially resemble simpler, less derived, filamentous structures. Here we use a novel, custom-built experimental setup to more accurately mimic subaqueous burial and compaction under low-energy, fine-grain depositional environments applicable to the taphonomic settings most plumage-preserving ‘exceptional fossils’ are found in. We find that when submerged and subsequently buried and compacted, feathers do not clump together and they maintain their original arrangement. Submersion in fluid in and of itself does not lead to clumping of barbs; this would only occur upon pulling feathers out from water into air. Furthermore, sediment encases the feathers, fixing them in place during compaction. Thus, feather clumping that leads to erroneously plesiomorphic morphological interpretations may not be a taphonomic factor of concern when examining fossil feathers. Our current methodology is amenable to further improvements that will continue to more accurately mimic subaqueous burial and compaction, allowing for various hypothesis testing.