Search

The Lower Cretaceous Huajiying Formation of the Sichakou Basin in northern Hebei Province, northern China contains key vertebrate taxa of the early Jehol Biota, e.g., Protopteryx fengningensis , Archaeornithura meemannae , Peipiaosteus fengningensis , and Eoconfuciusornis zhengi This formation arguably documents the second-oldest bird-bearing horizon, producing the oldest fossil records of the two major Mesozoic avian groups Enantiornithes and Ornithuromorpha. Hence, precisely determining the depositional ages of the Huajiying Formation would advance our understanding of the evolutionary history of the Jehol Biota. Here we present secondary ion mass spectrometry (SIMS) U-Pb zircon analysis results of eight interbedded tuff/tuffaceous sandstone samples from the Huajiying Formation. Our findings, combined with previous radiometric dates, suggest that the oldest enantiornithine and ornithuromorph birds in the Jehol Biota are ∼129-131 Ma, and that the Jehol Biota most likely first appeared at ∼135 Ma. This expands the biota's temporal distribution from late Valanginian to middle Aptian with a time span of about 15 My., Competing Interests: The authors declare no competing interest., (Copyright © 2020 the Author(s). Published by PNAS.)

Birds have a highly specialized and efficient digestive system, but when this system originated remains uncertain. Here we report six gastric pellets attributable to the recently discovered 160-million-year-old troodontid dinosaur Anchiornis, which is among the key taxa for understanding the transition to birds. The gastric pellets contain lightly acid-etched lizard bones or fish scales, and some are associated with Anchiornis skeletons or even situated within the oesophagus. Anchiornis is the earliest and most basal theropod known to have produced gastric pellets. In combination with other lines of evidence, the pellets suggest that a digestive system resembling that of modern birds was already present in basal members of the Paraves, a clade including troodontids, dromaeosaurids, and birds, and that the evolution of modern avian digestion may have been related to the appearance of aerial locomotion in this lineage.

Feathers are remarkable evolutionary innovations that are associated with complex adaptations of the skin in modern birds. Fossilised feathers in non-avian dinosaurs and basal birds provide insights into feather evolution, but how associated integumentary adaptations evolved is unclear. Here we report the discovery of fossil skin, preserved with remarkable nanoscale fidelity, in three non-avian maniraptoran dinosaurs and a basal bird from the Cretaceous Jehol biota (China). The skin comprises patches of desquamating epidermal corneocytes that preserve a cytoskeletal array of helically coiled α-keratin tonofibrils. This structure confirms that basal birds and non-avian dinosaurs shed small epidermal flakes as in modern mammals and birds, but structural differences imply that these Cretaceous taxa had lower body heat production than modern birds. Feathered epidermis acquired many, but not all, anatomically modern attributes close to the base of the Maniraptora by the Middle Jurassic.

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.

The crop is characteristic of seed-eating birds today, yet little is known about its early history despite remarkable discoveries of many Mesozoic seed-eating birds in the past decade. Here we report the discovery of some early fossil evidence for the presence of a crop in birds. Two Early Cretaceous birds, the basal ornithurine Hongshanornis and a basal avian Sapeornis, demonstrate that an essentially modern avian digestive system formed early in avian evolution. The discovery of a crop in two phylogenetically remote lineages of Early Cretaceous birds and its absence in most intervening forms indicates that it was independently acquired as a specialized seed-eating adaptation. Finally, the reduction or loss of teeth in the forms showing seed-filled crops suggests that granivory was possibly one of the factors that resulted in the reduction of teeth in early birds.

Nudds and Dyke (Reports, 14 May 2010, p. 887) compared the rachis diameters of the primary feathers of Archaeopteryx and Confuciusornis to those of modern birds and found that the primary feathers of these two basal birds were too weak to support sustained flight. Our measurements of Confuciusornis specimens suggest that their conclusions need to be further evaluated.

In extant birds, the hand is permanently abducted towards the ulna, and the wrist joint can bend extensively in this direction to fold the wing when not in use. Anatomically, this asymmetric mobility of the wrist results from the wedge-like shape of one carpal bone, the radiale, and from the well-developed convexity of the trochlea at the proximal end of the carpometacarpus. Among the theropod precursors of birds, a strongly convex trochlea is characteristic of Coelurosauria, a clade including the highly derived Maniraptora in addition to tyrannosaurs and compsognathids. The shape of the radiale can be quantified using a 'radiale angle' between the proximal and distal articular surfaces. Measurement of the radiale angle and reconstruction of ancestral states using squared-change parsimony shows that the angle was small (15 degrees) in primitive coelurosaurs but considerably larger (25 degrees) in primitive maniraptorans, indicating that the radiale was more wedge-shaped and the carpal joint more asymmetric. The radiale angle progressively increased still further within Maniraptora, with concurrent elongation of the forelimb feathers and the forelimb itself. Carpal asymmetry would have permitted avian-like folding of the forelimb in order to protect the plumage, an early advantage of the flexible, asymmetric wrist inherited by birds.

Spectacular fossils from the Early Cretaceous Jehol Group of northeastern China have greatly expanded our knowledge of the diversity and palaeobiology of dinosaurs and early birds, and contributed to our understanding of the origin of birds, of flight, and of feathers. Pennaceous (vaned) feathers and integumentary filaments are preserved in birds and non-avian theropod dinosaurs, but little is known of their microstructure. Here we report that melanosomes (colour-bearing organelles) are not only preserved in the pennaceous feathers of early birds, but also in an identical manner in integumentary filaments of non-avian dinosaurs, thus refuting recent claims that the filaments are partially decayed dermal collagen fibres. Examples of both eumelanosomes and phaeomelanosomes have been identified, and they are often preserved in life position within the structure of partially degraded feathers and filaments. Furthermore, the data here provide empirical evidence for reconstructing the colours and colour patterning of these extinct birds and theropod dinosaurs: for example, the dark-coloured stripes on the tail of the theropod dinosaur Sinosauropteryx can reasonably be inferred to have exhibited chestnut to reddish-brown tones.

Most of Mesozoic bird diversity comprises species that are part of one of two major lineages, namely Ornithurae, including living birds, and Enantiornithes, a major radiation traditionally referred to as 'opposite birds'. Here we report the largest Early Cretaceous enantiornithine bird from north-east China, which provides evidence that basal members of Enantiornithes share more morphologies with ornithurine birds than previously recognized. Morphological evolution in these two groups has been thought to be largely parallel, with derived members of Enantiornithes convergent on the 'advanced' flight capabilities of ornithurine birds. The presence of an array of morphologies previously thought to be derived within ornithurine and enantiornithine birds in a basal enantiornithine species provides evidence of the complex character evolution in these two major lineages. The cranial morphology of the new specimen is among the best preserved for Mesozoic avians. The new species extends the size range known for Early Cretaceous Enantiornithes significantly and provides evidence of forelimb to hind limb proportions distinct from all other known members of the clade. As such, it sheds new light on avian body size evolution and diversity, and allows a re-evaluation of a previously proposed hypothesis of competitive exclusion among Early Cretaceous avian clades.

In studies of the evolution of avian flight there has been a singular preoccupation with unravelling its origin. By contrast, the complex changes in morphology that occurred between the earliest form of avian flapping flight and the emergence of the flight capabilities of extant birds remain comparatively little explored. Any such work has been limited by a comparative paucity of fossils illuminating bird evolution near the origin of the clade of extant (i.e. 'modern') birds (Aves). Here we recognize three species from the Early Cretaceous of China as comprising a new lineage of basal ornithurine birds. Ornithurae is a clade that includes, approximately, comparatively close relatives of crown clade Aves (extant birds) and that crown clade. The morphology of the best-preserved specimen from this newly recognized Asian diversity, the holotype specimen of Yixianornis grabaui Zhou and Zhang 2001, complete with finely preserved wing and tail feather impressions, is used to illustrate the new insights offered by recognition of this lineage. Hypotheses of avian morphological evolution and specifically proposed patterns of change in different avian locomotor modules after the origin of flight are impacted by recognition of the new lineage. The complete articulated holotype specimen of Yixianornis grabaui, from the Early Cretaceous Jiufotang Formation of Liaoning Province, in north-eastern China, arguably the best-preserved basal ornithurine specimen yet discovered, provides the earliest evidence consistent with the presence of extant avian tail feather fanning.

An ornithurine bird, Hongshanornis longicresta gen. et sp. nov., represented by a nearly complete and articulated skeleton in full plumage, has been recovered from the lacustrine deposits of the Lower Cretaceous Jehol Group in Inner Mongolia, northeast China. The bird had completely reduced teeth and possessed a beak in both the upper and lower jaws, representing the earliest known beaked ornithurine. The preservation of a predentary bone confirms that this structure is not unique to ornithischian dinosaurs but was common in early ornithurine birds. This small bird had a strong flying capability with a low aspect ratio wing. It was probably a wader, feeding in shallow water or marshes. This find confirms that the aquatic environment had played a key role in the origin and early radiation of ornithurines, one branch of which eventually gave rise to extant birds near the Cretaceous/Tertiary boundary. This discovery provides important information not only for studying the origin and early evolution of ornithurines but also for understanding the differentiation in morphology, body size, and diet of the Early Cretaceous birds.

Yanornis martini is an Early Cretaceous basal ornithurine bird. Its fish-eating diet was previously recognized from a discrete mass of disarticulated fish remains discovered in its abdominal region. A new complete and articulated specimen of Yanornis martini preserves abundant in-situ gastroliths such as have been associated with a herbivorous diet. We suggest that the occurrence of gastroliths in this specimen, fish remains in a second, and the lack of gastroliths in three others, is consistent with diet-switching in Yanornis martini. Incompatibility of the preserved data with explanations of the grit as an artifact of preservation or result of accidental ingestion is discussed. This discovery indicates the earliest presence of intermittent diet change (and associated gizzard plasticity) observed in extant birds seasonally and in response to changes in available food sources.

An embryo of an enantiornithine bird has been recovered from the Lower Cretaceous rocks of Liaoning, in northeast China. The bird has a nearly complete articulated skeleton with feather sheet impressions and is enclosed in egg-shaped confines. The tucking posture of the skeleton suggests that the embryo had attained the final stage of development. The presence of well-developed wing and tail feather sheets indicates a precocial developmental mode, supporting the hypothesis that precocial birds appeared before altricial birds.

Here we describe a fossil of an enantiornithine bird from the Early Cretaceous period in China that has substantial plumage feathers attached to its upper leg (tibiotarsus). The discovery could be important in view of the relative length and aerodynamic features of these leg feathers compared with those of the small 'four-winged' gliding dinosaur Microraptor and of the earliest known bird, Archaeopteryx. They may be remnants of earlier long, aerodynamic leg feathers, in keeping with the hypothesis that birds went through a four-winged stage during the evolution of flight.

We report on a new Mesozoic bird, Longirostravis hani, from the Early Cretaceous Jehol Biota of northeastern China. The new taxon has a long, slender rostrum and mandible, and a small number of rostralmost teeth. Postcranial characters such as a furcular ramus wider ventrally than dorsally, a centrally concave proximal margin of the humeral head, and a minor metacarpal that projects distally more than the major metacarpal, support the placement of Longirostravis within euenantiornithine Enantiornithes, the most diverse clade of Mesozoic birds. The morphology of the skull, however, suggests that Longirostravis had a probing feeding behavior, a specialization previously unknown for Enantiornithes. Indeed, this discovery provides the first evidence in support of the existence of such a foraging behavior among basal lineages of Mesozoic birds.

The recently reported Jeholornis represents the only known bird with a complete long skeletal tail except for Archaeopteryx. Two newly discovered specimens referable to Jeholornis provide some important new information about its anatomy. The tail of Jeholornis is much longer than that of Archaeopteryx and comprises a maximum of 27 caudal vertebrae compared with only 23 in Archaeopteryx. More interestingly, the tail feathers are shaped more like those of dromaeosaurs than those of Archaeopteryx. We conclude that the common ancestor of birds must have a more primitive tail than that in Archaeopteryx, confirming the side branch position of Archaeopteryx in the early avian evolution. The synsacrum is composed of six sacrals, representing a transitional stage between Archaeopteryx and more advanced birds. The scapula of Jeholornis has a dorso-laterally exposed glenoid facet, and the coracoid has a supracoracoid foramen. The presence of a pair of fenestrae in the sternum of Jeholornis has further implications for the air-sac system in early birds.

Although the dinosaurian hypothesis of bird origins is widely accepted, debate remains about how the ancestor of birds first learned to fly. Here we provide new evidence suggesting that basal dromaeosaurid dinosaurs were four-winged animals and probably could glide, representing an intermediate stage towards the active, flapping-flight stage. The new discovery conforms to the predictions of early hypotheses that proavians passed through a tetrapteryx stage.

Here we report an unequivocal arboreal coelurosaur, Epidendrosaurus ningchengensis gen. et sp. nov. This juvenile coelurosaur's third manual digit is extremely elongated, distinctively different from that of other known dinosaurs and birds. It represents certainly a type of adaptation previously unreported from the Mesozoic although the exact function of the third manual digit is unclear. The relatively long forelimb, penultimate phalanx of manual digit II, and pedal penultimate phalanges, are interpreted as evidence for the arboreal habit of Epidendrosaurus. Because Epidendrosaurus is more similar to advanced birds in some arboreal features than to Archaeopteryx, we suggest that the initial appearance of tree-adaptation in theropods was probably not directly related to flight but to other functions, such as seeking food or escaping from predators. Electronic Supplementary Material is available if you access this article at http://dx.doi.org/10.1007/s00114-002-0353-8. On that page (frame on the left side), a link takes you directly to the supplementary material.

The lacustrine deposits of the Yixian and Jiufotang Formations in the Early Cretaceous Jehol Group in the western Liaoning area of northeast China are well known for preserving feathered dinosaurs, primitive birds and mammals. Here we report a large basal bird, Jeholornis prima gen. et sp. nov., from the Jiufotang Formation. This bird is distinctively different from other known birds of the Early Cretaceous period in retaining a long skeletal tail with unexpected elongated prezygopophyses and chevrons, resembling that of dromaeosaurids, providing a further link between birds and non-avian theropods. Despite its basal position in early avian evolution, the advanced features of the pectoral girdle and the carpal trochlea of the carpometacarpus of Jeholornis indicate the capability of powerful flight. The dozens of beautifully preserved ovules of unknown plant taxa in the stomach represents direct evidence for seed-eating adaptation in birds of the Mesozoic era.

With only one known exception, early Cretaceous birds were smaller than their closest theropod dinosaur relatives. Here we report on a new bird from the Early Cretaceous feathered-dinosaur-bearing continental deposits of Liaoning, northeast China, which is not only larger than Archaeopteryx but is nearly twice as large as the basal dromaeosaur Microraptor. The new taxon, Sapeornis chaoyangensis gen. et sp. nov., has a more basal phylogenetic position than all other birds except for Archaeopteryx. Its exceptionally long forelimbs, well-developed deltoid crest of the humerus, proximally fused metacarpals, relatively short hindlimbs and short pygo-style indicate powerful soaring capability and further suggest that by the Early Cretaceous ecological diversification of early birds was greater than previously assumed. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00114-001-0276-9.

Presents information on the ornithurine birds Yanornis martini and Yixianornis grabaui from the early Cretaceous of western Liaoning, China. Description of the birds; Differences between the birds; Information on enantiornithine and ornithurine birds.

Deals with a study which described an enantiornithine bird Longipteryx chaoyangensis from the Early Cretaceous Jiufotang Formation in Chaoyang, western Liaoning Province, China. Systematics; Discussion; Indication of its discovery.