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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

An exceptionally well-preserved fossil Kalligrammatid from the Jehol Biota

A new species of Neuroptera, Sophogramma lii sp. nov. is described and illustrated in this paper. This new species is assigned to Sophogramma Ren of Kalligrammatidae, mainly based on its humeral recurrent veins, the basal cell present between posterior branch of media at base of wing, etc. The new species is distinguished from all other Sophogramma species on the difference of wing venation. This exceptionally well-preserved specimen is recovered from the Yixian Formation, Huangbanjigou, located near Beipiao City, in western Liaoning Province. The genus diagnosis has been emended based on new characters shown in this new species. Based on a summary of localities, distribution and geological ages of all genera and species of the Kalligrammatidae, we suggest that Kalligrammatids might have originated in Eastern Asia, especially in China. Function of the wing pigmentation is discussed briefly. Supported by National Natural Science Foundation of China (Grant Nos. 30430100 and 40872022), Beijing Natural Science Foundation Program (Grant No. 5082002), Scientific Research Key Program (Grant No. KZ200910028005) and PHR Project of of Beijing Municipal Commission of Education, Science-Technology Basic Condition Platform from the Ministry of Science and Technology of the People’s Republic of China (Grant No. 2005DKA21402)     

New dragonflies （Insecta： Odonata： Gomphaeschnidae） from the Yixian Formation in Inner Mongolia,China

Two fossil dragonflies from the Upper Jurassic to Lower Cretaceous Yixian Formation in Liutiaogou Village, Ningcheng County, Inner Mongolia, China are described and illustrated. They are assigned to two new genera and species, i.e., Sophoaeschna frigida gen. et sp. nov. and Falsisophoaeschna generalis gen. et sp. nov. within the family Gomphaeschnidae Tillyard ＆ Fraser, 1940. This is the first report of Odonata from Yixian Formation in Inner Mongolia and the second record of fossil Gomphaeschnidae from China.     

New dragonflies (Insecta: Odonata: Gomphaeschnidae) from the Yixian Formation in Inner Mongolia, China

Two fossil dragonflies from the Upper Jurassic to Lower Cretaceous Yixian Formation in Liutiaogou Village, Ningcheng County, Inner Mongolia, China are described and illustrated. They are assigned to two new genera and species, i.e., Sophoaeschna frigida gen. et sp. nov. and Falsisophoaeschna generalis gen. et sp. nov. within the family Gomphaeschnidae Tillyard & Fraser, 1940. This is the first report of Odonata from Yixian Formation in Inner Mongolia and the second record of fossil Gomphaeschnidae from China.     

New fossil elaterids （Insect： Coleoptera： Polyphaga： Elateridae） from the Middle Jurassic of Inner Mongolia, China

A new genus with a new species of fossil elaterid, Paraprotagrypnus superbus gen. et sp. nov., from the Middle Jurassic Jiulongshan Formation in Daohugou Village (N41°18.979′, E119°14.318′), Shantou Township, Ningcheng County, Inner Mongolia, China is described and illustrated. The genus Paraprotagrypnus belongs to the family of Elateridae, subfamily of Protagrypninae and tribe of Protagrypnini. Some primitive characters of the new genus and new species shed light on the ages between the Jiulongshan Formation in Inner Mongolia and the Yixian Formation in the western Liaoning Province of China. The habitat of the new species is briefly discussed.     

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.     

Mesozoic giant fleas from northeastern China (Siphonaptera): Taxonomy and implications for palaeodiversity

The recently discovered definite giant fleas from the Middle Jurassic Daohugou fauna and the Early Cretaceous Jehol fauna of northeastern China represent significant evidence for understanding ectoparasitism in the Mesozoic as well as the evolution of these giant blood feeders with their putative hosts (i.e. hairy or feathered vertebrates). On the basis of seven well-preserved specimens from Daohugou and Huangbanjigou we analyse the systematic classification of these primitive fleas, establishing two new genera and three new species as Pseudopulex wangi sp. nov., Hadropsylla sinica gen. et sp. nov., and Tyrannopsylla beipiaoensis gen. et sp. nov. All of them are assigned to the extinct siphonapteran family Pseudopulicidae, while the Early Cretaceous genus Tarwinia is transferred to Tarwiniidae fam. nov. The basal morphological disparities of Siphonaptera in the Mesozoic are evidenced by the occurrence of at least three distinct groups (pseudopulicids, tarwiniids, and saurophthirids). These disparate morphologies likely indicate adaptations to different hosts.     

New dragonflies (Insecta: Odonata: Gomphaeschnidae) from the Yixian Formation in Inner Mongolia, China

Two fossil dragonflies from the Upper Jurassic to Lower Cretaceous Yixian Formation in Liutiaogou Village, Ningcheng County,Inner Mongolia, China are described and illustrated. They are assigned to two new genera and species, I.e., Sophoaeschna frigida gen.et sp. Nov. And Falsisophoaeschna generalis gen. Et sp. Nov. Within the family Gomphaeschnidae Tillyard & Fraser, 1940. This is the first report of Odonata from Yixian Formation in Inner Mongolia and the second record of fossil Gomphaeschnidae from China.