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Concept: Archaeopteryx


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


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


The ‘Solnhofen Limestone’ beds of the Southern Franconian Alb, Bavaria, southern Germany, have for centuries yielded important pterosaur specimens, most notably of the genera Pterodactylus and Rhamphorhynchus. Here we describe a new genus of non-pterodactyloid pterosaur based on an extremely well preserved fossil of a young juvenile: Bellubrunnus rothgaengeri (gen. et sp. nov.).

Concepts: Biology, Pterosaur, Dinosaur, Archaeopteryx, Jurassic, Late Jurassic, Solnhofen limestone, Pterodactylus


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


Archaeopteryx is an iconic fossil that has long been pivotal for our understanding of the origin of birds. Remains of this important taxon have only been found in the Late Jurassic lithographic limestones of Bavaria, Germany. Twelve skeletal specimens are reported so far. Archaeopteryx was long the only pre-Cretaceous paravian theropod known, but recent discoveries from the Tiaojishan Formation, China, yielded a remarkable diversity of this clade, including the possibly oldest and most basal known clade of avialan, here named Anchiornithidae. However, Archaeopteryx remains the only Jurassic paravian theropod based on diagnostic material reported outside China.

Concepts: Bird, Dinosaur, Archaeopteryx, Theropoda, Dromaeosauridae, Maniraptora, Jurassic, Avialae


Theropod dinosaurs were the dominant predators in most Mesozoic era terrestrial ecosystems. Early theropod evolution is currently interpreted as the diversification of various carnivorous and cursorial taxa, whereas the acquisition of herbivorism, together with the secondary loss of cursorial adaptations, occurred much later among advanced coelurosaurian theropods. A new, bizarre herbivorous basal tetanuran from the Upper Jurassic of Chile challenges this conception. The new dinosaur was discovered at Aysén, a fossil locality in the Upper Jurassic Toqui Formation of southern Chile (General Carrera Lake). The site yielded abundant and exquisitely preserved three-dimensional skeletons of small archosaurs. Several articulated individuals of Chilesaurus at different ontogenetic stages have been collected, as well as less abundant basal crocodyliforms, and fragmentary remains of sauropod dinosaurs (diplodocids and titanosaurians).

Concepts: Bird, Reptile, Cretaceous, Dinosaur, Saurischia, Archaeopteryx, Theropoda, Jurassic


The wings of birds and their closest theropod relatives share a uniform fundamental architecture, with pinnate flight feathers as the key component. Here we report a new scansoriopterygid theropod, Yi qi gen. et sp. nov., based on a new specimen from the Middle-Upper Jurassic period Tiaojishan Formation of Hebei Province, China. Yi is nested phylogenetically among winged theropods but has large stiff filamentous feathers of an unusual type on both the forelimb and hindlimb. However, the filamentous feathers of Yi resemble pinnate feathers in bearing morphologically diverse melanosomes. Most surprisingly, Yi has a long rod-like bone extending from each wrist, and patches of membranous tissue preserved between the rod-like bones and the manual digits. Analogous features are unknown in any dinosaur but occur in various flying and gliding tetrapods, suggesting the intriguing possibility that Yi had membranous aerodynamic surfaces totally different from the archetypal feathered wings of birds and their closest relatives. Documentation of the unique forelimbs of Yi greatly increases the morphological disparity known to exist among dinosaurs, and highlights the extraordinary breadth and richness of the evolutionary experimentation that took place close to the origin of birds.

Concepts: Bird, Reptile, Dinosaur, Archaeopteryx, Wing, Theropoda, Coelurosauria, Maniraptora


The Late Jurassic Lourinhã Formation is known for its abundant remains of dinosaurs, crocodylomorphs and other vertebrates. Among this record are nine localities that have produced either dinosaur embryos, eggs or eggshell fragments. Herein, we describe and identify the first crocodiloid morphotype eggs and eggshells from the Lourinhã Formation, from five occurrences. One clutch from Cambelas, composed of 13 eggs, eggshell fragments from Casal da Rola and Peralta, one crushed egg and eggshells from Paimogo North, and four crushed eggs as well as eggshell fragments from Paimogo South. We observed and confirmed diagnostic morphological characters for crocodiloid eggshells and which are consistent with a crocodylomorph affinity, such as the ellipsoidal shape, wedge-shaped shell units, triangular extinction under cross-polarized light, and tabular ultrastructure. This material is distinctive enough to propose two new ootaxa within the oofamily Krokolithidae, Suchoolithus portucalensis, oogen. and oosp. nov., for the material from Cambelas, the most complete clutch known for crocodiloid eggs, and Krokolithes dinophilus, oosp. nov., for the remaining material. These are the oldest crocodylomorph eggs known, extending the fossil record for this group to the Late Jurassic. Furthermore, except for the clutch from Cambelas, the material was found with theropod eggs and nests, in the other four occurrences, which seem to suggest some form of biological relationship, still unclear at this point.

Concepts: Egg, Bird, Reptile, Chicken, Fossil, Paleontology, Dinosaur, Archaeopteryx


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


Maniraptora includes birds and their closest relatives among theropod dinosaurs. During the Cretaceous period, several maniraptoran lineages diverged from the ancestral coelurosaurian bauplan and evolved novel ecomorphologies, including active flight, gigantism, cursoriality and herbivory. Propagation X-ray phase-contrast synchrotron microtomography of a well-preserved maniraptoran from Mongolia, still partially embedded in the rock matrix, revealed a mosaic of features, most of them absent among non-avian maniraptorans but shared by reptilian and avian groups with aquatic or semiaquatic ecologies. This new theropod, Halszkaraptor escuilliei gen. et sp. nov., is related to other enigmatic Late Cretaceous maniraptorans from Mongolia in a novel clade at the root of Dromaeosauridae. This lineage adds an amphibious ecomorphology to those evolved by maniraptorans: it acquired a predatory mode that relied mainly on neck hyperelongation for food procurement, it coupled the obligatory bipedalism of theropods with forelimb proportions that may support a swimming function, and it developed postural adaptations convergent with short-tailed birds.

Concepts: Bird, Dinosaur, Saurischia, Archaeopteryx, Theropoda, Coelurosauria, Dromaeosauridae, Maniraptora