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Concept: Feathered dinosaurs


Ticks are currently among the most prevalent blood-feeding ectoparasites, but their feeding habits and hosts in deep time have long remained speculative. Here, we report direct and indirect evidence in 99 million-year-old Cretaceous amber showing that hard ticks and ticks of the extinct new family Deinocrotonidae fed on blood from feathered dinosaurs, non-avialan or avialan excluding crown-group birds. A †Cornupalpatum burmanicum hard tick is entangled in a pennaceous feather. Two deinocrotonids described as †Deinocroton draculi gen. et sp. nov. have specialised setae from dermestid beetle larvae (hastisetae) attached to their bodies, likely indicating cohabitation in a feathered dinosaur nest. A third conspecific specimen is blood-engorged, its anatomical features suggesting that deinocrotonids fed rapidly to engorgement and had multiple gonotrophic cycles. These findings provide insight into early tick evolution and ecology, and shed light on poorly known arthropod-vertebrate interactions and potential disease transmission during the Mesozoic.

Concepts: Bird, Extinction, Dinosaur, Feathered dinosaurs, Archaeopteryx, Feather, Avialae, Microraptor


Two skeletons of the large compsognathid Sinocalliopteryx gigas include intact abdominal contents. Both specimens come from the Jianshangou Beds of the lower Yixian Formation (Neocomian), Liaoning, China. The holotype of S. gigas preserves a partial dromaeosaurid leg in the abdominal cavity, here attributed to Sinornithosaurus. A second, newly-discovered specimen preserves the remains of at least two individuals of the primitive avialan, Confuciusornis sanctus, in addition to acid-etched bones from a possible ornithischian. Although it cannot be stated whether such prey items were scavenged or actively hunted, the presence of two Confuciusornis in a grossly similar state of digestion suggests they were consumed in rapid succession. Given the lack of clear arboreal adaptations in Sinocalliopteryx, we suggest it may have been an adept stealth hunter.

Concepts: Cretaceous, Liaoning, Dinosaur, Feathered dinosaurs, Theropoda, Dromaeosauridae, Compsognathidae, Jiufotang Formation


The famous ‘feathered dinosaurs’ from the Early Cretaceous of Liaoning Province, northeastern China, include several dromaeosaurids, which are among the closest relatives of birds. Most of these are small-bodied taxa with long arms and broad wings comprised of vaned feathers, but a single specimen (the holotype of Tianyuraptor) belongs to a much larger individual with reduced forelimbs, which unfortunately lacks any preserved integument. We describe a new specimen of large-bodied, short-armed Liaoning dromaeosaurid, which we designate as a new genus and species, Zhenyuanlong suni. The integument is well preserved and provides the first evidence of feather morphologies and distribution in a short-armed (and probably non-volant) dromaeosaurid, indicating that these rare and aberrant taxa had large wings consisting of pennaceous feathers on the arms and long pennaceous feathers on the tail very similar to their smaller and longer-armed relatives, but potentially lacked vaned feathers on the legs. Zhenyuanlong adds yet more diversity to the Liaoning dromaeosaurid fauna, helps further reveal a distinct short-armed bauplan among dromaeosaurids, and illuminates previously-unrecognized homoplasy that complicates dromaeosaurid phylogeny and suggests that the Liaoning taxa may not have formed their own clade.

Concepts: Bird, Cretaceous, Dinosaur, Feathered dinosaurs, Archaeopteryx, Feather, Dromaeosauridae, Sinosauropteryx


In the two decades since the discovery of feathered dinosaurs [1-3], the range of plumage known from non-avialan theropods has expanded significantly, confirming several features predicted by developmentally informed models of feather evolution [4-10]. However, three-dimensional feather morphology and evolutionary patterns remain difficult to interpret, due to compression in sedimentary rocks [9, 11]. Recent discoveries in Cretaceous amber from Canada, France, Japan, Lebanon, Myanmar, and the United States [12-18] reveal much finer levels of structural detail, but taxonomic placement is uncertain because plumage is rarely associated with identifiable skeletal material [14]. Here we describe the feathered tail of a non-avialan theropod preserved in mid-Cretaceous (∼99 Ma) amber from Kachin State, Myanmar [17], with plumage structure that directly informs the evolutionary developmental pathway of feathers. This specimen provides an opportunity to document pristine feathers in direct association with a putative juvenile coelurosaur, preserving fine morphological details, including the spatial arrangement of follicles and feathers on the body, and micrometer-scale features of the plumage. Many feathers exhibit a short, slender rachis with alternating barbs and a uniform series of contiguous barbules, supporting the developmental hypothesis that barbs already possessed barbules when they fused to form the rachis [19]. Beneath the feathers, carbonized soft tissues offer a glimpse of preservational potential and history for the inclusion; abundant Fe(2+) suggests that vestiges of primary hemoglobin and ferritin remain trapped within the tail. The new finding highlights the unique preservation potential of amber for understanding the morphology and evolution of coelurosaurian integumentary structures.

Concepts: Bird, Dinosaur, Feathered dinosaurs, Archaeopteryx, Feather, Coelurosauria, Dromaeosauridae, Sinosauropteryx


Feathers are amongst the most complex epidermal structures known, and they have a well-documented evolutionary trajectory across non-avian dinosaurs and basal birds. Moreover, melanosome-like microbodies preserved in association with fossil plumage have been used to reconstruct original colour, behaviour and physiology. However, these putative ancient melanosomes might alternatively represent microorganismal residues, a conflicting interpretation compounded by a lack of unambiguous chemical data. We therefore used sensitive molecular imaging, supported by multiple independent analytical tests, to demonstrate that the filamentous epidermal appendages in a new specimen of the Jurassic paravian Anchiornis comprise remnant eumelanosomes and fibril-like microstructures, preserved as endogenous eumelanin and authigenic calcium phosphate. These results provide novel insights into the early evolution of feathers at the sub-cellular level, and unequivocally determine that melanosomes can be preserved in fossil feathers.

Concepts: DNA, Evolution, Bird, Paleontology, Dinosaur, Feathered dinosaurs, Archaeopteryx, Troodontidae


Features that were once considered exclusive to modern birds, such as feathers and a furcula, are now known to have first appeared in non-avian dinosaurs. However, relatively little is known of the early evolutionary history of the hyperinflated brain that distinguishes birds from other living reptiles and provides the important neurological capablities required by flight. Here we use high-resolution computed tomography to estimate and compare cranial volumes of extant birds, the early avialan Archaeopteryx lithographica, and a number of non-avian maniraptoran dinosaurs that are phylogenetically close to the origins of both Avialae and avian flight. Previous work established that avian cerebral expansion began early in theropod history and that the cranial cavity of Archaeopteryx was volumetrically intermediate between these early forms and modern birds. Our new data indicate that the relative size of the cranial cavity of Archaeopteryx is reflective of a more generalized maniraptoran volumetric signature and in several instances is actually smaller than that of other non-avian dinosaurs. Thus, bird-like encephalization indices evolved multiple times, supporting the conclusion that if Archaeopteryx had the neurological capabilities required of flight, so did at least some other non-avian maniraptorans. This is congruent with recent findings that avialans were not unique among maniraptorans in their ability to fly in some form.

Concepts: Evolution, Bird, Dinosaur, Feathered dinosaurs, Archaeopteryx, Theropoda, Maniraptora, Avialae


Recent discoveries of large leg feathers in some theropods have implications for our understanding of the evolution of integumentary features on the avialan leg, and particularly of their relevance for the origin of avialan flight. Here we report 11 basal avialan specimens that will greatly improve our knowledge of leg integumentary features among early birds. In particular, they provide solid evidence for the existence of enlarged leg feathers on a variety of basal birds, suggest that extensively scaled feet might have appeared secondarily at an early stage in ornithuromorph evolution, and demonstrate a distal-to-proximal reduction pattern for leg feathers in avialan evolution.

Concepts: Bird, Metaphysics, Dinosaur, Feathered dinosaurs, Archaeopteryx, Wing, Theropoda, Avialae


Discoveries of bird-like theropod dinosaurs and basal avialans in recent decades have helped to put the iconic ‘Urvogel’ Archaeopteryx into context and have yielded important new data on the origin and early evolution of feathers. However, the biological context under which pennaceous feathers evolved is still debated. Here we describe a new specimen of Archaeopteryx with extensive feather preservation, not only on the wings and tail, but also on the body and legs. The new specimen shows that the entire body was covered in pennaceous feathers, and that the hindlimbs had long, symmetrical feathers along the tibiotarsus but short feathers on the tarsometatarsus. Furthermore, the wing plumage demonstrates that several recent interpretations are problematic. An analysis of the phylogenetic distribution of pennaceous feathers on the tail, hindlimb and arms of advanced maniraptorans and basal avialans strongly indicates that these structures evolved in a functional context other than flight, most probably in relation to display, as suggested by some previous studies. Pennaceous feathers thus represented an exaptation and were later, in several lineages and following different patterns, recruited for aerodynamic functions. This indicates that the origin of flight in avialans was more complex than previously thought and might have involved several convergent achievements of aerial abilities.

Concepts: Bird, Dinosaur, Feathered dinosaurs, Archaeopteryx, Feather, Dromaeosauridae, Maniraptora, Bird anatomy


Microbodies associated with feathers of both nonavian dinosaurs and early birds were first identified as bacteria but have been reinterpreted as melanosomes. Whereas melanosomes in modern feathers are always surrounded by and embedded in keratin, melanosomes embedded in keratin in fossils has not been demonstrated. Here we provide multiple independent molecular analyses of both microbodies and the associated matrix recovered from feathers of a new specimen of the basal bird Eoconfuciusornis from the Early Cretaceous Jehol Biota of China. Our work represents the oldest ultrastructural and immunological recognition of avian beta-keratin from an Early Cretaceous (∼130-Ma) bird. We apply immunogold to identify protein epitopes at high resolution, by localizing antibody-antigen complexes to specific fossil ultrastructures. Retention of original keratinous proteins in the matrix surrounding electron-opaque microbodies supports their assignment as melanosomes and adds to the criteria employable to distinguish melanosomes from microbial bodies. Our work sheds new light on molecular preservation within normally labile tissues preserved in fossils.

Concepts: Protein, Bird, Cretaceous, Keratin, Paleontology, Dinosaur, Feathered dinosaurs, Feather


Understanding the aerodynamic performance of feathered, non-avialan dinosaurs is critical to reconstructing the evolution of bird flight. Here we show that the Early Cretaceous five-winged paravian Microraptor is most stable when gliding at high-lift coefficients (low lift/drag ratios). Wind tunnel experiments and flight simulations show that sustaining a high-lift coefficient at the expense of high drag would have been the most efficient strategy for Microraptor when gliding from, and between, low elevations. Analyses also demonstrate that anatomically plausible changes in wing configuration and leg position would have made little difference to aerodynamic performance. Significant to the evolution of flight, we show that Microraptor did not require a sophisticated, ‘modern’ wing morphology to undertake effective glides. This is congruent with the fossil record and also with the hypothesis that symmetric ‘flight’ feathers first evolved in dinosaurs for non-aerodynamic functions, later being adapted to form lifting surfaces.

Concepts: Evolution, Bird, Aerodynamics, Fossil, Paleontology, Dinosaur, Feathered dinosaurs, Dromaeosauridae