Zhonghe Zhou

An exceptionally preserved Lower Cretaceous ecosystem

Fieldwork in the Early Cretaceous Jehol Group, northeastern China has revealed a plethora of extraordinarily well-preserved fossils that are shaping some of the most contentious debates in palaeontology and evolutionary biology. These discoveries include feathered theropod dinosaurs and early birds, which provide additional, indisputable support for the dinosaurian ancestry of birds, and much new evidence on the evolution of feathers and flight. Specimens of putative basal angiosperms and primitive mammals are clarifying details of the early radiations of these major clades. Detailed soft-tissue preservation of the organisms from the Jehol Biota is providing palaeobiological insights that would not normally be accessible from the fossil record.

Four-winged dinosaurs from China

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.

A long-tailed, seed-eating bird from the Early Cretaceous of China

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.

Timing of the Jiufotang Formation (Jehol Group) in Liaoning, northeastern China, and its implications

The timing of the Jiufotang Formation remains speculative despite recent progress in the study of the Jehol Biota. In this paper we contribute to this topic with Ar-40/Ar-39 dating on K-feldspar (sanidine and orthoclase) from tuffs interbedded within the fossil-bearing shales of the Jiufotang Formation, from the upper part of the Jehol Group in Chaoyang, Liaoning, northeastern China. Ar-40/(39) Ar step heating analyses of K-feldspar and the SHRIMP U-Pb zircon data indicate that tuffs at the Shangheshou section erupted at 120.3 +/- 0.7 million years ago. This result confirms an Aptian age for the Jiufotang Formation that was mainly based on biostratigraphic evidence. It also places stringent controls on the age of the fossils from the formation, providing a minimum age (120 Ma) for the four-winged dinosaur, Microraptor, and the seed-eating bird, Jeholornis.

Branched integumental structures in Sinornithosaurus and the origin of feathers

The evolutionary origin of feathers has long been obscured because no morphological antecedents were known to the earliest, structurally modern feathers of Archaeopteryx. It has been proposed that the filamentous integumental appendages on several theropod dinosaurs are primitive feathers; but the homology between these filamentous structures and feathers has been disputed, and two taxa with true feathers (Caudipteryx and Protarchaeopteryx) have been proposed to be flightless birds. Confirmation of the theropod origin of feathers requires documentation of unambiguously feather-like structures in a clearly non-avian theropod. Here we describe our observations of the filamentous integumental appendages of the basal dromaeosaurid dinosaur Sinornithosaurus millenii, which indicate that they are compound structures composed of multiple filaments. Furthermore, these appendages exhibit two types of branching structure that are unique to avian feathers: filaments joined in a basal tuft, and filaments joined at their bases in series along a central filament. Combined with the independent phylogenetic evidence supporting the theropod ancestry of birds, these observations strongly corroborate the hypothesis that the integumental appendages of Sinornithosaurus are homologous with avian feathers. The plesiomorphic feathers of Sinornithosaurus also conform to the predictions of an independent, developmental model of the evolutionary origin of feathers.

Fossilized melanosomes and the colour of Cretaceous dinosaurs and 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.

A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers

Recent coelurosaurian discoveries have greatly enriched our knowledge of the transition from dinosaurs to birds, but all reported taxa close to this transition are from relatively well known coelurosaurian groups. Here we report a new basal avialan, Epidexipteryx hui gen. et sp. nov., from the Middle to Late Jurassic of Inner Mongolia, China. This new species is characterized by an unexpected combination of characters seen in several different theropod groups, particularly the Oviraptorosauria. Phylogenetic analysis shows it to be the sister taxon to Epidendrosaurus, forming a new clade at the base of Avialae. Epidexipteryx also possesses two pairs of elongate ribbon-like tail feathers, and its limbs lack contour feathers for flight. This finding shows that a member of the avialan lineage experimented with integumentary ornamentation as early as the Middle to Late Jurassic, and provides further evidence relating to this aspect of the transition from non-avian theropods to birds.

Pterosaur diversity and faunal turnover in Cretaceous terrestrial ecosystems in China.

New specimens and an analysis of the Jehol pterosaur faunae of northeastern China show an unexpected diversity of flying reptile groups in terrestrial Cretaceous ecosystems. Here we report two new pterosaurs that are referred to European groups previously unknown in deposits of northeastern China. Feilongus youngi, from the Yixian Formation, is closely related to the Gallodactylidae and is distinguished by the presence of two independent sagittal crests and a protruding upper jaw. Nurhachius ignaciobritoi, from the Jiufotang Formation, has teeth formed by labiolingually compressed triangular crowns, only previously reported in Istiodactylus latidens from England. With these new discoveries, the Jehol pterosaurs show a wide range of groups including both primitive and derived forms that are not matched by any other deposit in the world. The discoveries also document the turnover of pterosaur faunae, with the primitive Anurognathidae and early archaeopterodactyloids being replaced by derived pterodactyloids. Furthermore, these deposits offer an opportunity to examine the interaction and competition between birds and pterosaurs—it indicates that the avian fauna during the Lower Cretaceous (and possibly most of the Mesozoic) dominated terrestrial, inland regions, whereas pterosaurs were more abundant in coastal areas.

Early Adaptive Radiation of Birds: Evidence from Fossils from Northeastern China

Late Jurassic and Early Cretaceous birds from northeastern China, including many complete skeletons of Confuciusornis, provide evidence for a fundamental dichotomy in the class Aves that may antedate the temporal occurrence of the Late Jurassic Archaeopteryx. The abundance of Confuciusornis may provide evidence of avian social behavior. Jurassic skeletal remains of an ornithurine bird lend further support to the idea of an early separation of the line that gave rise to modern birds. Chaoyangia, an ornithurine bird from the Early Cretaceous of China, has premaxillary teeth.

Insight into the evolution of avian flight from a new clade of Early Cretaceous ornithurines from China and the morphology of Yixianornis grabaui

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.