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 Protop- teryx, 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 de- marcation 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.     

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.     

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.     

Discovery of a pterodactylid pterosaur from the Yixian Formation of western Liaoning, China

A well-preserved pterosaur with nearly complete skull is described from the Lower Cretaceous Yixian Formation at Sihetun in western Liaoning. It is characterized by a low and long crestless skull, slender and pointed teeth, long metacarpal, nearly equal length of metatarsals I–III and short pedal digit V. It is referred to a new genus and species of the family Pterodactylidae:Haopterus gracilis gen. et sp. nov. This is the first pterosaur with a nearly complete skull from the Jehol Biota; it also represents the first non-controversial fossil record of Pterodactylidae in Asia.Haopterus is more derived thanPterodactylus from the Late Jurassic Solnhofen in Germany. This discovery extends the distribution of the family Pterodactylidae from Europe and Africa to Asia and its latest occurrence from the Late Jurassic to the Early Cretaceous. The discovery ofHaopterus gracilis provides further evidence for the study of the origin and radiation of the Jehol Biota; it also sheds new light on the evolution and distribution of pterosaurs in the late Mesozoic.     

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     

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。     

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.     

Liaoxiornis delicatus gen. et sp. nov., the smallest Mesozoic bird

Recently, a very small and complete bird specimen was collected from theConfuciusornis-bearing Yixian. Formation. This finding represents a new kind of Mesozoic bird, namedLiaoxiornis delicatus. It is characterized by a short skull with teeth, a long neck and a very small sternum with only posterior process; the humeral head is not internally curved, and the hindlimbs are longer than the forelimbs.Liaoxiornis delicatus is not only much smaller thanConfuciusomis, but also smaller thanCathayomis and other early Cretaceous birds. It is the smallest bird known from Mesozoic deposit.Liaoxiornis provides new evidence indicating that the early adaptive radiation of birds is complicated.     

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.     

Pyrite framboids associated with the Mesozoic Jehol Biota in northeastern China: Implications for microenvironment during early fossilization

Despite of the recent influx of reports describing important fossil specimens from the Mesozoic Jehol Biota, detailed information on the mechanism of fossilization and paleoenvironment in which these fossils were deposited is scanty. We present an analysis of microenvironment based upon scanning electron microscope observations of in situ pyrite framboids and microcrystallines of plant and vertebrate feather fossils in the Jehol Biota. Pyrite microcrystallines and framboids occur extensively inside and on surface of plant fossils.Framboids found on feathers and in sedimentary matrix were in a lower abundance. These framboids have diameters ranging from 6 μm to 31 μm with an average of 20 μm, indicating a dysoxic aqueous condition with free oxygen level less than 30 μmol/L for the microenvironment where these framboids were formed. The outgrowth of framboids inside plant tissues suggests the presence of water molecules and free oxygen at the cellular level during pyritization; the relative timing between tissue decay and framboid formation implies a rapid tissue degradation occurred during the very early stage of fossilization. This line of reasoning is consistent with the observation that cell level structure of plant fossils from these deposits is rarely preserved. We propose a "fossil envelop" model to accommodate the different geochemical conditions between the microenvironment surrounding the fossil material and the macroenvironment of background lake bottom water.     

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.     

A new family of primitive mammal from the Mesozoic of western Liaoning, China

A fairly well-preserved specimen of a new primitive mammal, Repenomamus robustus gen. et sp. nov., has been described from the Yixian Formation (Lower Cretaceous), western Liaoning, China. This animal has several derived characteristics, such as well-developed den-tary/squamosal articulation, reduced number of tooth, differentiation of premolars and molars in postcanine teeth, presence of a dorsal process of the premaxilla that is not in contact with the nasal, closed medial wall of the orbit, and presence of fingerlike promontorium on the petrosal. It also retains some primitive reptile-like features. It is the most primitive taxon among the three mammals known from the Jehol Biota, and also represents the largest mammal of Mesozoic age over the world.     

A new discovery of macroscopic fossils from the Ediacaran Doushantuo Formation in the Yangtze Gorges area

The Ediacaran section in the Yangtze Gorges area, South China, is one of the most important in the worldand contains abundant fossils[1-4]. The earliest discov- ery of macrofossils in 1984, including macroscopic algal fossils from the uppermost Doushantuo…     

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.     

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.     

A new species from an ornithuromorph（Aves： Ornithothoraces）dominated locality of the Jehol Biota

We report on a new species of ornithuromorph bird, Iteravis huchzermeyeri gen. et sp. nov., from the previously unreported Sihedang locality of the Lower Cretaceous Yixian Formation, the oldest ornithuromorph bearing deposit in the world. Unlike most other Cretaceous localities, specimens from this new quarry are largely referable to Ornithuromorpha, similar to the Lower Cretaceous Aptian Xiagou Formation in Gansu Province. Also similar to the Xiagou avifauna, the fauna at Sihedang is largely dominated by a single taxon（described here）.Differences in faunal dominance may suggest the Sihedang records a unique ecological habitat. This may also explain the dominance of Gansus in the younger Xiagou Formation locality and suggests that previous hypotheses regarding the shift in dominance between Enantiornithes and Ornithuromorpha need to be reassessed in terms of potential ecological biases due to limited sampling. Furthermore, the recognition of an ornithuromorph dominated locality in the Sihedang significantly weakens the signal of such an inferred trend. Compared to most Jehol birds, the new specimen is relatively better preserved in three dimensions revealing morphological details of the skeleton, as well as preserves feather impressions including a rectricial morphology previously unknown among Mesozoic birds.     

辽西早白垩世义县组大康堡层纹泥的古环境意义

辽西地区义县组湖泊沉积层中产出著名的“热河生物群”化石,对于此时期古环境的研究一直是诸多学者关注的焦点.本文对大康堡层湖相纹泥展开研究,重建此时期古环境.通过野外及室内对纹泥组成、类型、厚度等方面的观测,对此时期古气候有了新的认识.纹泥厚度特征变化表明沉积期降水量和沉积速率逐渐减小,统计洪水沉积间纹层数量可指示沉积期洪水事件的频率和规模.结合近现代纹泥形成和保存条件,可知该地区沉积期湖泊水体较深,湖水分层,存在一个缺氧环境,具有较小的表面积/深度比,湖盆形态适合,未受到风的侵扰.     

A new eutriconodont mammal from the early Cretaceous Jehol Biota of Liaoning,China

A new genus and species of eutriconodont mammal, Chaoyangodens lii gen. et sp. nov., from the Dawangzhangzi bed of Yixian Formation, Lingyuan, Liaoning, is reported. The new species has a tooth formula I5-C1-P1-M3/i4-c1-p1-m4, unique among eutriconodonts in having only one premolar in lower and upper jaws, respectively, and a distinctive diastema between the canine and the premolar. Its simple incisors and reduced premolars show a mosaic combination of primitive and derived features. This new taxon adds to the diverse group of Jehol eutriconodonts. Among the known species from at least three horizons, there seems no evolutionary trend in morphology that is recognizable, except for that larger species are all from the Lujiatun bed of the Yixian Fm. A thorough and systematic analysis involving all the Jehol eutriconodonts is needed to understand their phylogenetic relationships.     

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.     

Constraint on the late Mesozoic regional angular unconformity in West Liaoning--North Hebei by LA-ICP-MS dating

The angular unconformity between the Tuchengzi Formation and the Zhangjiakou Formation in the Chengde area is commonly regarded as the direct evidence of boundary between Jurassic and Cretaceous systems, and as the evidence of tectonic transformation interfaces in Mesozoic Era in the West Liaoning-North Hebei. Moreover, the lower boundary age of the Zhangjiakou Formation in the Chengde was to be thought about 135 Myr in the past. Based on the detailed study and systematic sampling in Yixian-Beipiao, Lingy- uan, Chengde and Luanping areas in the field, the authors obtained the following isotopic ages by means of LA-ICP-MS dating： In the Lingyuan area, the Zhangjiakou Formation and the Yixian Formation are in angular unconformable contact; the age range of the Zhangjiakou Formation volcanic rock is from 133 to 130 Myr （seven samples）, and that of the Yixian Formation volcanic rock is from 126 to 121 Myr （eleven samples）. In the Luotuoshan of Chengde, the Tuchengzi Formation and the Zhangjiakou Formation are in angu- lar unconformable contact; the age at the lower Zhangjiakou Formation is about 124 Myr （one sample）, which reflects that the age range of the lower Zhangjiakou Formation is from 129 to 124 Myr combined with the previous study. In the Luanping basin, there is the larg- est sediment gap in upper Mesozoic, whose lower boundary age is confined in 131 to 130 Myr. In the Yixian-Beipiao area, the second Member of the Yixian Formation, making up more than 3/4 thickness of the Yixian Formation, was formed in 126 to 119 Myr, and the age of the Yixian Formation at the bottom is about 132 Myr. Moreover, the upper boundary age of the Tuchengzi Formation is about 136 Myr in the West Liaoning-North Hebei region. Based on the above isotopic data combined with the geological characteristics in the field in the West Liaoning-North Hebei region, it reflects that there was a widespread tectonic event in the period of 136 to 126 Myr （mainly in 130-126 Myr） in the West Liaoning-North Hebei, and it formed th