A new pterosaur (Pterodactyloidea,Tapejaridae) from the Early Cretaceous Jiufotang Formation of western Liaoning,China and its implications for biostratigraphy

In this article we describe a new and exceptionally weel-preserved pterodactyloid pterosaur,Sinopterus dong gen.et sp.nov.from the Jiufotang Formation in western Liaoning Province of northeast China,The new species is referred to the family Tapejaridae,representing its first record outside Brazil.It also represents the earliest occurrence as well as the most complete sketeton of the famil.Some revisions are made about the family according to the mophological observations of the postcranial bones of Sinopterus.Two pterosaur assemblages appear to have existed in the Jehol Group,represented by the lower Yixian Formation and upper Jiufotang Formation,respectively,The lower pterosaur assemblage shows some resembalance to that of the Late Jurassic in Solnhofen (Tithonian)by sharing members of the Pterodactylidae and Anurognathidae.The upper one shows more resemblance to that to the Early Cretaceous Santana Formation (Aptian/Albian) by comprising only pterodactyloids such as the Tapejaridae,The age of the Yixian Formation is younger than that of the Solnhofen lihographic limestone ,and the age of the Jiufotang Formation (Aptian )is slightly older than the Santana Formation.     

U-Pb zircon age for the Daohugou Biota at Ningcheng of Inner Mongolia and comments on related issues

SHRIMP U-Pb zircon dating was carried out for the Daohugou Biota near Ningcheng of Inner Mongolia and for lavas overlying or underlying sala-mander-bearing strata at Reshuitang in Lingyuan of West Liaoning. The results suggest that the Dao- hugou Biota occurred at an interval from 168 Ma to 164―152 Ma. Both the Daohugou Biota and the salamander-bearing fossil assemblage are the same biota and thus developed from 168 to 152 Ma, i.e. from late Middle Jurassic to the early Late Jurassic. The Daohugou Biota-bearing rocks, resting on the Jiulongshan Formation in disconformity and being overlain in unconformity by Late Jurassic Tuchengzi Formation and Early Cretaceous rocks containing the Jehol Biota, are mainly composed of volcanic-sedi- mentary rocks in a normal sequence. It is recom- mended that the Daohugou Biota and the related stratigraphy should be correlated with the Tiaojishan Formation (Lanqi Formation in West Liaoning) or its synchronous rocks. It is suggested that the Dao- hugou Biota and the Jehol Biota would be neither taken into one biota nor considered as the earliest elements of the Jehol Biota. The Daohugou Biota and the related rocks and the Yixian Formation were respectively formed in different periods of volcanic-sedimentary tectonics.     

Stratigraphy and age of the Daohugou Bed in Ningcheng, Inner Mongolia

Recent fieldwork has extended the distribution of the Daohugou Bed deposits from the Daohugou Village to its several neighboring areas. The fossil-bearing Daohugou deposits uncomformably overlie complex bedrocks, and comprise three major parts. The red shales in the lower part were misidentified as belonging to the Tuchengzi Formation. Field excavation has indicated that the shales of upper part of the bed are the major fossil-bearing horizon. Due to strong tectonic activities, sediments were often folded with the sequences inverted in the region. Some newly recognized contacts between the Daohugou Bed and the volcanic rocks showed that the ignimbrite of the Tiaojishan Formation （159-164 Ma） underlies the Daohugou deposits, rather than overlying the latter as previously proposed. Thus, the age of the Daohugou deposits should be younger than the age of the ignimbrite, and thus it was incorrect to correlate the Daohugou Bed with the Middle Jurassic Jiulongshan Formation. Although biostratigraphic studies based on conchostracans and insects support a Middle Jurassic-early Late Jurassic age for the Daohugou deposits, vertebrate fossils such as Liaoxitriton, Jeholopterus and feathered maniraptorans show much resemblance to those of the Yixian Formation. In other words, despite the absence of Lycoptera, a typical fish of the Jehol Biota, the Daohugou vertebrate assemblage is closer to that of the Early Cretaceous Jehol Biota than to any other biota. We propose that the Daohugou fossil assemblage probably represents the earliest evolutionary stage of the Jehol Biota based on both vertebrate biostratigraphy and the sedimentological and volcanic features which suggest the Daohugou deposit belongs to the same cycle of volcanism and sedimentation as the Yixian Formation of the Jehol Group.     

Sinobaatar gen. nov.: First multituberculate from the Jehol Biota of Liaoning, Northeast China

A multituberculate skeleton from the Lower Cretaceous Yixian Formation at Dawangzhangzi, Lingyuan City, Liaoning Province, Northeast China, provides new morphological information for early multituberculates. The specimen is the holotype of Sinobaatar lingyuanensis gen. et sp. nov. It has a narrow skull that lacks the superorbital crest or postorbital process. The dental formula is 3·?·5·2/1·0·3·2. The dental morphology, especially that of cheek teeth, of S. lingyuanensis is similar to that of Eobaatar, which places it in the family Eobaataridae. The postcranial skeleton of Sinobaataris similar to that of other multituberculates. As in Holotheria, Metatarsal V of Sinobaatar articulates only with the cuboid and has no contact with the calcaneus, which probably represents the primitive condition of multituberculates. Nine carpals of Sinobaatar resemble those of Zhangheotherium, except the centrale being larger than the trapezoid. The dental features of Sinobaatar show again that eobaatarids are obviously intermediate between Late Jurassic multituberculates and the later forms. Because eobaatarids are only known from the Early Cretaceous, the finding of Sinobaatar, therefore, supports that the age of the Jehol Biota is most likely Early Cretaceous.     

热河生物群--探索中生代生命演化的世界级化石宝库

最近十多年来,我国在热河生物群各门类生物研究方面取得了一系列重要的发现和成果。我国辽西地区已经成为世界上研究鸟类起源,恐龙的进化,鸟类、哺乳动物以及被子植物的早期演化和辐射等最重要的地区之一。同位素测年技术的广泛应用,表明热河生物群属于早白垩世,在地球上延续了大约2千万年。热河生物群的繁盛和东亚地区当时特有的温暖的淡水和陆地环境具有密切的联系。而频繁的火山活动又是化石得以大量完美保存的重要因素之一。热河生物群的研究还具有很大的潜力,而对化石的科学发掘和保护的任务还十分艰巨。     

The discovery of aeschnidiid nymphs (Aeschnidiidae, Odonata, Insecta)

Aeschnidiid nymphs are here first identified and described, and its phylogenetic relationship and bioecological characteristics are deduced. The nymphs known from China, Mongolia, Russia and Brazil and arranged in the family Aeschnidiidae are, in fact, unrelated to this group. All the known five Chinese species in four genera which were erected based on fossil nymphs and placed in the family Gomphidae may be transferred in Aeschnidiidae and merged into one alone, theSinaeschnidia cancellosa. It is a geographically widespread species from the “Jehol biota” of East Asia and appeared in the latest Late Jurassic age.     

Preliminary elemental analysis of fossil insects from the Middle Jurassic of Daohugou, Inner Mongolia and its taphonomic implications

Fossil insects from the Middle Jurassic of Daohugou, Inner Mongolia were investigated using Scanning Electron Microscope (SEM) for the first time, and portions and distribution of some elements in compression and pyrited fossils were also revealed by X-ray Energy Dispersive Spectroscopy (EDS) attached to SEM. Most of compression fossil insects from the Daohugou Biota are preserved in organic remains (diagenetic products of the original organic components). A small part of compression fossils retain a comparatively high Fe concentration which probably resulted from the absorption of Fe by biopolymers during the decaying period. Pyritized insect fossils suggest that the “fossil envelop” model found in the Early Cretaceous Jehol Biota probably also occurs in the Daohugou Biota. Different preservation modes show various mechanisms of fossilization, and also suggest that several different microenvironments are present in Daohugou palaeolakes. Supported by National Basic Research Program of China (Grant No. 2006CB806400), National Natural Science Foundation of China (Grant Nos. 40872015, 40523004, 40632010), and Special Fund for Graduates in the Graduate University of the Chinese Academy of Sciences     

Earliest Asian discoglossid frog from western Liaoning

Based on a superbly preserved skeleton, a new anuran taxon has been named and described from the Yixian Formation (Upper Jurassic/Lower Cretaceous), Liaoning Province, northeastern China. This discovery documents the first discoglossid fossil from China, the earliest record of the group in Asia, and the only second Mesozoic discoglossid known from fully articulated material in the world. The family Discoglossidae is widely perceived as a primitive anuran group, but has no definite fossils found in collaboration with their recent distribution in Asia. The new discovery from Liaoning extends the temporal range of the group in Asia back at least 120 million years, and provides valuable fossil material for study of anatomical details of early discoglossids, as well as of historical distribution of this primitive anuran group.     

A new species of Gobiconodon （Triconodonta, Mammalia） and its implication for the age of Jehol Biota

A new species of Gobiconodon is found from the Yixian Formation of western Liaonlng, China. The new taxa,G zofiae sp. nov., has a confluent opening for branches Ⅱand Ⅲ of the trigeminal nerve on the anterior lamina of the petrosal. G zofiae sp. nov. is similar to Repenomamus in having an ossified Meckel‘s cartilage connecting the lower jaws and ear region. The new species, with enlarged I^1/I1, posteriorly located infraorbital foramen and four mental foramina,distinctly differs from the other species of Gobiconodon. The new material indicates that Gobiconodon has four, not five,upper molariforms. The presence of Gobiconodon in Jchol Biota makes it possible to correlate Jehol Biota with faunas in eastern Asia and North America, and suggests the age of the Yixian Formation to be Early Cretaceous.     

Earliest known crown-group salamanders

Salamanders are a model system for studying the rates and patterns of the evolution of new anatomical structures. Recent discoveries of abundant Late Jurassic and Early Cretaceous salamanders are helping to address these issues. Here we report the discovery of well-preserved Middle Jurassic salamanders from China, which constitutes the earliest known record of crown-group urodeles (living salamanders and their closest relatives). The new specimens are from the volcanic deposits of the Jiulongshan Formation (Bathonian), Inner Mongolia, China, and represent basal members of the Cryptobranchidae, a family that includes the endangered Asian giant salamander (Andrias) and the North American hellbender (Cryptobranchus). These fossils document a Mesozoic record of the Cryptobranchidae, predating the previous record of the group by some 100 million years. This discovery provides evidence to support the hypothesis that the divergence of the Cryptobranchidae from the Hynobiidae had taken place in Asia before the Middle Jurassic period.     

A new iguanodontid (Jinzhousaurus yangi gen. et sp. nov.) from the Yixian Formation of western Liaoning, China

A new iguanodontid,Jinzhousaurus yangi gen. et sp. nov., is erected based on the cranial and dental morphology of a specimen from the Yixian Formation of western Liaoning. Although a few primitive characteristics remain,Jinzhousaurus yangi display a number of characteristics similar to those seen in derived iguanodontids. Other interesting characteristics include the absence of antorbital fenestra and the frontal excluded from the participation in orbit, which are closer to the condition in hadrosaurids. The unusual combination of the characteristics provides important data for assessing morphological evolution toward hadrosaurs. The new discovery also represents the first reported large-sized dinosaur from Jehol fauna in western Liaoning, and increases the diversity of this famous fauna. Furthermore, the discovery of a derived iguanodontid from the Yixian Formation provides further evidence supporting a Cretaceous age of the Jehol fauna, which is consistent with an isotopic age.     

Highly specialized mammalian skulls from the Late Cretaceous of South America

Dryolestoids are an extinct mammalian group belonging to the lineage leading to modern marsupials and placentals. Dryolestoids are known by teeth and jaws from the Jurassic period of North America and Europe, but they thrived in South America up to the end of the Mesozoic era and survived to the beginnings of the Cenozoic. Isolated teeth and jaws from the latest Cretaceous of South America provide mounting evidence that, at least in western Gondwana, dryolestoids developed into strongly endemic groups by the Late Cretaceous. However, the lack of pre-Late Cretaceous dryolestoid remains made study of their origin and early diversification intractable. Here we describe the first mammalian remains from the early Late Cretaceous of South America, including two partial skulls and jaws of a derived dryolestoid showing dental and cranial features unknown among any other group of Mesozoic mammals, such as single-rooted molars preceded by double-rooted premolars, combined with a very long muzzle, exceedingly long canines and evidence of highly specialized masticatory musculature. On one hand, the new mammal shares derived features of dryolestoids with forms from the Jurassic of Laurasia, whereas on the other hand, it is very specialized and highlights the endemic, diverse dryolestoid fauna from the Cretaceous of South America. Our specimens include only the second mammalian skull known for the Cretaceous of Gondwana, bridging a previous 60-million-year gap in the fossil record, and document the whole cranial morphology of a dryolestoid, revealing an unsuspected morphological and ecological diversity for non-tribosphenic mammals.     

On the horizon of Protopteryx and the early vertebrate fossil assemblages of the Jehol Biota

Protopteryx, a monotypic fossil bird discovered from the Sichakou basin in Fengning, Hebei, is the most primitive enantiornithine currently known. The bird-bearing strata do not contain the index fossils of the Yixian Formation in western Liaoning; the fish and bird fossils have more primitive features than the related forms found in the Yixian Formation, and the conchostracans are those usually distributed in the Dabeigou and Dadianzi formations in northern Hebei. Besides, the Protopteryx-bearing strata underlie the deposits bearing the index fossils of the Yixian Formation in the neighboring basin. Thus, it could be confirmed that the horizon of Protopteryx should be lower than the Yixian Formation, and Is approximately equivalent to the Dadianzi Formation in northern Hebei. This is the lowest horizon of the known fossil birds in China and Mesozoic enantiornithine birds in the world. Accompanying Protopteryx, there are other birds, acipenseriform fishes, salamanders, and mammals, which compose the Peipiaosteus fengningensis-Protopteryx fengningensis assemblage. This new assemblage traces the vertebrate evolution history of the Jehol Biota back to 130.7 Ma before. It is suggested that the demarcation of the Jehol Biota should be based on the large-scale tectonic-sedimentary cycles, and Peipiaosteus, instead of Lycoptera, could be taken as the vertebrate representative of the Jehol Biota.     

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.     

大港探区中生代原型盆地恢复及成盆模式

对大港探区中生代残留盆地进行原型盆地恢复．原型盆地可分为：早中侏罗世坳陷型盆地,晚侏罗至早白垩世断坳共存盆地及晚白垩世热坳陷盆地．中生代原型盆地分布方向为ＮＥ－ＮＮＥ向,分布范围有限,且沉积厚度较小,这与中生代中国东部受伊泽奈崎板块斜向俯冲和华北与扬子板块碰撞所致的挤压环境有关     

The earliest known sauropod dinosaur

Sauropods were a very successful group of dinosaurs during the Jurassic and Cretaceous periods, but their earlier history is poorly known. Until now, the earliest reported sauropod bones were from the Early Jurassic, and the only tentative evidence of earlier sauropods was in the form of controversial footprints. Here we report the discovery of an incomplete sauropod skeleton from the Late Triassic period of Thailand, which provides the first osteological evidence of pre-Jurassic sauropods. This dinosaur is markedly different from prosauropods and substantiates theoretical predictions that there was a fairly long period of sauropod evolution during the Triassic.     

中国的蜥脚类恐龙化石及其层位

文章对中国的蜥脚类恐龙化石及其层位、产地进行了较为系统的总结,划分出５个相关的蜥脚类恐龙动物群：（１）早侏罗世资中龙动物群（ＺｉｚｈｏｎｇｏｓａｕｒｕｓＦａｕｎａ）;（２）中侏罗世蜀龙－峨眉龙动物群（Ｓｈｕｎｏｓａｕｒｕｓ－ＯｍｅｉｓａｕｒｕｓＦａｕｎａ）;（３）晚侏罗世马门溪龙动物群（ＭａｍｅｎｃｈｉｓａｕｒｕｓＦａｕｎａ）;（４）早白垩世亚洲龙动物群（ＡｓｉａｔｏｓａｕｒｕｓＦａｕｎａ）;（５）晚白垩世纳摩盖吐龙动物群（ＮｅｍｅｇｔｏｓａｕｒｕｓＦａｕｎａ）。文章还对中国的蜥脚类恐龙的地理分布特征进行了探讨。     

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.     

A gigantic feathered dinosaur from the lower cretaceous of China

Numerous feathered dinosaur specimens have recently been recovered from the Middle-Upper Jurassic and Lower Cretaceous deposits of northeastern China, but most of them represent small animals. Here we report the discovery of a gigantic new basal tyrannosauroid, Yutyrannus huali gen. et sp. nov., based on three nearly complete skeletons representing two distinct ontogenetic stages from the Lower Cretaceous Yixian Formation of Liaoning Province, China. Y. huali shares some features, particularly of the cranium, with derived tyrannosauroids, but is similar to other basal tyrannosauroids in possessing a three-fingered manus and a typical theropod pes. Morphometric analysis suggests that Y. huali differed from tyrannosaurids in its growth strategy. Most significantly, Y. huali bears long filamentous feathers, thus providing direct evidence for the presence of extensively feathered gigantic dinosaurs and offering new insights into early feather evolution.     

Transitional mammalian middle ear from a new Cretaceous Jehol eutriconodont

The transference of post-dentary jaw elements to the cranium of mammals as auditory ossicles is one of the central topics in evolutionary biology of vertebrates. Homologies of these bones among jawed vertebrates have long been demonstrated by developmental studies; but fossils illuminating this critical transference are sparse and often ambiguous. Here we report the first unambiguous ectotympanic (angular), malleus (articular and prearticular) and incus (quadrate) of an Early Cretaceous eutriconodont mammal from the Jehol Biota, Liaoning, China. The ectotympanic and malleus have lost their direct contact with the dentary bone but still connect the ossified Meckel's cartilage (OMC); we hypothesize that the OMC serves as a stabilizing mechanism bridging the dentary and the detached ossicles during mammalian evolution. This transitional mammalian middle ear narrows the morphological gap between the mandibular middle ear in basal mammaliaforms and the definitive mammalian middle ear (DMME) of extant mammals; it reveals complex changes contributing to the detachment of ear ossicles during mammalian evolution.