Taphonomy and abundance of birds from the Lower Eocene Fur Formation of Denmark

The pattern, pace and extent of the evolutionary radiation of modern birds (Neornithes) by the end‐Cretaceous (65 Ma) has long been debated. Well‐dated, taphonomically understood and phylogenetically constrained fossil birds from both sides of the Cretaceous–Paleogene (K–Pg) boundary are required to quantify the shape of this radiation, but have largely been lacking. Here we report on a large collection of fossil birds from the Lower Eocene of Denmark (ca. 54 Ma) that includes three‐dimensionally preserved, articulated specimens from carbonate concretions as well as skeletal imprints and feathers. These birds are from a marine diatomite sequence (the Fur Formation), a low‐energy deep‐water preservational environment unique to the Cretaceous and Paleogene avian fossil record. We present taphonomic and palaeoecological information gleaned from these birds that in combination with phylogenetic data have implications for unravelling avian survivorship across the K–Pg boundary as well as for the pattern of the neornithine evolutionary radiation. Copyright © 2009 John Wiley & Sons, Ltd.     

New Schizolepis Fossils from the Early Cretaceous in Inner Mongolia, China and its Phylogenetic Position

Three Schizolepis species collected from the Lower Cretaceous layer of the Huolinhe Basin,Inner Mongolia,China are described.These fossils are Schizolepis longipetiolus Xu XH et Sun BN sp.nov.,which is a new species,Schizolepis cf.heilongjiangensis Zheng et Zhang,and Schizolepis neimengensis Deng.The new species is a well-preserved female cone,slender and cylindrical in shape.The seed–scale complexes have long petioles and are arranged on the cone axis loosely and helically.The seed scales are divided into two lobes from the base.Each lobe is semicircular or elongate ligulate in shape,widest at the middle or the lower middle part,with an obtuse or bluntly pointed apex.The inner margin is almost straight and the outer margin is strongly arched.On the surface of the lobe,there are longitudinal and somewhat radial striations from the base to the margin.The seed is borne on the adaxial surface at the base or middle of each lobe.Schizolepis was established in 1847,and,although more than twenty species have been discovered and reported,its phylogenetic position is controversial because of the imperfection of fossils.Most authors have considered there to be a close evolutionary relationship between Schizolepis and extant Pinaceae.Here,we analyze characteristics and compare Schizolepis with Picea crassifolia Kom,which is morphologically most similar to Schizolepis.The results indicate that the genus probably has a distant evolutionary relationship with extant Pinaceae.A detailed statistical analysis of the global paleogeographic distribution of Schizolepis showed that all the fossils of this genus appeared in strata ranging from the Upper Triassic to the Lower Cretaceous in the North Hemisphere,being rare in the Upper Triassic and Lower Jurassic,but being very common from the Middle Jurassic to the Lower Cretaceous,and particularly abundant in the Lower Cretaceous.According to the statistical results,we speculate that the genus originated in Europe in the Late Triassic then spread from Europe to Asia between the Late Triassic and the Late Jurassic.In the Early Cretaceous most species existed in China’s three northeastern Provinces and the Inner Mongolia Autonomous Region and adjacent areas.Combining the paleogeographic distribution of the genus with ancient climatic factors,we deduced that Schizolepis began to decline and became extinct in the Early Cretaceous,and the reason for its extinction is closely related to the icehouse climate during the Early Cretaceous.     

Two Early Cretaceous fishes discovered from the most northern area of China: implications for the palaeobiogeography of the Jehol Biota

From a newly discovered Early Cretaceous fossil site at the Sanlianhe Coal Mine in Tahe County, Heilongjiang Province, the most northern area in China, two freshwater fossil fish had been recovered. This paper reports their classifications and implications regarding the palaeobiogeography of the Jehol Biota. The fish assemblage consists of Yanosteus sp. (Acipenseriforms) and Sinamia sp. (Amiiformes). Both are important members of the well known Jehol Biota, and their occurrence supports the correlation of the Yiliekede Formation at Sanlianhe Coal Mine site with the Jiufotang Formation of West Liaoning. The remains of the Yanosteus sp. from this locality is the first known record of the genus other than in the Yan‐Liao area of China, and the specimens of Sinamia sp. is quite similar to what may be a new species of Sinamia from West Liaoning. The new fossil site represents the most northern site that is undoubtedly part of the extent of the Jehol Biota as currently understood. The northern Great Xing'an Range area is an important region that may link correlations between the Early Cretaceous continental biotas of China, Mongolia and Russia. Copyright © 2010 John Wiley & Sons, Ltd.     

Ginkgo Huolinhensis sp.nov.from the Lower Cretaceous of Huolinhe Coal Field,Inner Mongolia,China

A new species,Ginkgo huolinhensis sp.nov.,with well-preserved cuticle from the Lower Cretaceous,Huolinhe Formation of Huolinhe Coal Field in Inner Mongolia,China,was studied morphologically and anatomi...     

Late Cretaceous Aquatic Angiosperms from Jiayin,Heilongjiang, Northeast China

Three taxa of Late Cretaceous aquatic angiosperms, Queruexia angulata （Lesq.） Krysht., Cobbania corrugata （Lesq.） Stockey et al. and Nelumbites cf. extenuinervis Upchurch et al. from Jiayin of Heilongjiang, NE China, are described in detail. Among them, Cobbania and Nelumbites from the Upper Cretaceous in China are reported for the first time. The aquatic angiosperm assemblage of Queruexia-Cobbania-Nelumbites appears to imply a seasonal, warm and moist environment in the Jiayin area during the Santonian-Campanian time.     

A New Material of Lindera (Lauraceae) of the Late Pliocene from Tengchong, Yunnan and the Genus' Biogeography Significance

Lindera is a large genus of graceful, pleasantly scented and common native trees and shrubs of southern China and neighboring regions of SE Asia. There is a well-documented Cenozoic fossil record not only in these regions but also from elsewhere. A new fossil leaf record has been found in diatomite beds from the Upper Pliocene Mangbang Formation of Tuantian, Tengchong County, Yunnan. The leaves are identified and assigned to Lindera acuminatissima K. Q. Dao et B. N. Sun sp. nov., by comparing their leaf architecture and epidermal characteristics with those of 51 extant Lauraceae species and with 15 known fossil Lindera taxa. The specimens have well-preserved cuticles, with typical leaf architecture and epidermal characteristics of the Lauraceae, including entire leaf margin, intramarginal veins, basal ternate acrodromous primary veins, one-cell trichome base, paracytic stomatal apparatus, sunken guard cells, subsidiary hardly staining cells and presence of oil cells. These characteristics are consistent with Lindera sect. Daphnidium but are different from reported fossil and extant species of Lindera. The cuticles of Lindera are fragile and delicate with only three Lindera fossils reported based on this tissue. In terms of paleobiogeography, the fossil record indicates that Lindera is distributed in high- to mid-latitude regions of the Late Cretaceous to Paleocene northern hemisphere. Coincidentally, the records of Lindera located on both sides of the Bering Land Bridge possibly support the hypothesis that ancient plants extended via transcontinental exchanges through the Bering Corridor. In the Eocene, ancient Lindera spread to Europe through the Northern Degeer Route and the Southern Thulian Route. At the same time, ancient Lindera spread into Central Asia. Climatic changes and tectonization since the Neogene prevented the propagation of Lindera throughout Asia, North America and Europe, and hence the distribution areas have just regressed to the low-latitude regions in Asia and North America. From the Paleogene to the Neogene, Lindera has changed its distribution by surviving extreme climate changes. Quaternary glaciations ultimately led to Lindera becoming extinct in Europe. The new record from Tengchong, Yunnan, with its lower latitude located in tropical and subtropical regions, indicates that Lindera has lived in those regions since the late Pliocene.     

New Mesozoic Mesopsychidae(Mecoptera)from Northeastern China

<正>The Mesozoic family Mesopsychidae Tillyard,1917 presently consists of seven described genera and 13 species from the mid-Triassic to the Early Cretaceous of Australia,South Africa and Eurasia.In the present paper one new genus and three new species of fossil mesopsychids are described that add significant distributional and stratigraphic extensions to the family.This finding documents the first formal record of fossil Mesopsychidae in China.Both Lichnomesopsyche gloriae gen.et sp.nov.and L.daohugouensis gen.et sp.nov. were found from the Middle Jurassic Jiulongshan Formation of Inner Mongolia,whereas Vitimopsyche kozlovi sp. nov.,of mid Early Cretaceous age,was collected from the Jehol Biota of northern Hebei Province.This new,well-preserved material from China reveals complete and previously unknown body features,including head, antennae,mouthparts,legs and abdomen.The delicate and long proboscides of these new taxa indicate that they were feeding on externally exposed,nutrient-rich fluids of gymnospermous ovulate fructifications,and incapable of piercing surface epidermis-attributable principally to the absence of stylets.These proboscides originated, perhaps multiply,among basal Mecoptera and are functionally and structurally convergent with equivalent mouthparts borne by fossil and extant Diptera,Lepidoptera,Neuroptera and Coleoptera.     

A New Species of Cupressinocladus from the Lower Cretaceous of Guyang Basin, Inner Mongolia, China and Cluster Analysis

Cupressinocladus Seward is a fossil genus of conifers and conifer fossils with reproductive organs are very rare. In general, it is difficult to understand the natural affinities with other conifers. In this paper, a new species, Cupressinocladus guyangensis P.H. Jin et B.N. Sun sp. nov., is reported based on branches with immature female cones from the Lower Cretaceous Guyang Formation of the Guyang Basin in Inner Mongolia, northern China. The foliage shoots are decussate. Leaves are decussate, imbricate, scale-like, weakly dimorphic, and bear longitudinal glands on the abaxial view. Stomata complexes are haplocheilic, monocyclic, irregularly arranged, and spread along the leaf margin. Immature female cones are subglobose with 6–8 cone scales, and three subglobose ovules arranged in a row at the base of the cone scales. Moreover, we performed cluster analysis using a statistics and machine learning toolbox for 23 fossils and extant species based on 16 morphological characters. The result implies that the new species bears a close resemblance to the extant Cupressus funebris Endl. and might have nearest systematic affinities to it.     

Pelecinid wasps (Insecta, Hymenoptera, Proctotrupoidea) from the Cretaceous of Russia and Mongolia

Thirteen new species referable to four genera, of which one is new, from the Cretaceous of Russia and Mongolia are established herein and assigned to the family Pelecinidae. Among the four genera, Protopelecinus gen. nov., including four new species, is referred to the subfamily Pelecininae, while Iscopinus Kozlov, including three new species, Eopelecinus Zhang, Rasnitsyn and Zhang, including five new species, and Scorpiopelecinus laetus sp. nov. are assigned to the subfamily Iscopininae. Of these new taxa, eight, namely Protopelecinus regularis, P. furtivus, Iscopinus simplex, ?I. suspectus, Eopelecinus exquisitus, E. scorpioideus, E. rudis and Scorpiopelecinus laetus, are from the Lower Cretaceous Zaza Formation of Baissa, Transbaikalia, Russia; two, E. minutus and E. fragilis, are from the basal Lower Cretaceous Tsagan-Tsab Formation of Khutel-Khara, Mongolia; two, P. dubius and P. deformis, are from the Lower Cretaceous (Aptian?) of Bon Tsagan, Mongolia; and one, I. separatus, is from the Upper Cretaceous (Cenomanian) Ola Formation of Obeshchayushchiy, Russia. A key to fossil pelecinid wasps is provided and a morphological analysis shows that the Pelecinidae might be paraphyletic with respect to the Proctotrupidae. The Chinese insect fauna from both the Yixian and Laiyang formations is dominated by Eopelecinus and Sinopelecinus whereas the Siberian + Mongolian fauna from the Zaza and Tsagan-Tsab formations is dominated by Eopelecinus and Iscopinus. Hence, Eopelecinus is common to both. The differences between the two faunas are likely to be the result of geographical variation in populations.     

Solenites (Czekanowskiales) from the Late Mesozoic Jehol Biota of Southeastern Jilin, China and its Paleoclimatic Implications

Situated at the easternmost area of the Late Mesozoic Jehol Biota, China, abundant fossils, including invertebrates, vertebrates and plants, were collected from the Early Cretaceous Yingzuilazi Formation of the Baishan Basin, southeastern Jilin. The faunal remains from this formation distinctly belong to the Early Cretaceous Lycoptera–Ephemeropsis–Eosestheria assemblage of the Jehol Biota. Based on the gross leaf morphology and epidermal structures, two new species and one indeterminate species of genus Solenites(Czekanowskiales), S. baishanensis sp. nov., S. gracilis sp. nov. and S. sp. are described in this paper. This is the first fossil plant of Czekanowskiales reported from this new fossil locality of the easternmost area of the Late Mesozoic Jehol Biota, China. The new discovery contributes to improve our knowledge for understanding of leaf morphology, epidermal characters and diversity of this genus during the Early Cretaceous time, and extends geologic and geographic distribution of Solenites in northern China and Eurasia. Along with consideration of associated plant and faunal assemblages and sedimentology, we suggest that the regional climate was warm and moderately humid with seasonal temperature and precipitation variations in warm–temperate zone during the Early Cretaceous.     

A gigantic caenagnathid oviraptorosaurian (Dinosauria: Theropoda) from the Upper Cretaceous of the Gobi Desert,Mongolia

A large, isolated symphyseal region of fused dentaries of an oviraptorosaurian was found in the Upper Cretaceous Bayn Shire Formation at Tsagaan Teg in the Mongolian Gobi Desert. A phylogenetic analysis places this specimen within Caenagnathidae. This specimen is comparable in size and morphology to the gigantic caenagnathid Gigantoraptor erlianensis from the Iren Dabasu Formation in China and is likely closely related to it. The occurrence of the specimen with possible affinities to G. erlianensis in the Bayn Shire Formation is consistent with the hypothesized correlation between the Bayn Shire and Iren Dabasu formations based on the co-occurrences of vertebrate fossils, especially turtles.     

Glacial forest refugium in Howard Valley,South Island,New Zealand

Fossils of forest habitat beetles and leaves of Nothofagus menziesii provide evidence of a forest refugium at times between ca. 34 000 and ca. 18 500 cal. a BP at an upland site in Howard Valley, located adjacent to glaciated valleys in South Island, New Zealand. The stratigraphy of the glacial‐aged terrace sequence of organic‐rich silts and fluvial sand/gravels indicates that soil development occurred episodically for around 15 000 a. Fifty‐four beetle taxa represent seven habitat types: forest, forest or scrub, riparian and aquatic, litter, grass/tussock, marshland and moss habitats. Leaf and beetle fossils indicate that forest dominated by N. menziesii persisted at the site for most of the time period represented, and tree line taxa such as Taenarthrus sp. 1 (Carabidae) and Podocarpus sp. (Podocarpaceae) indicate that the site may represent the upper tree limit for full‐glacial time. The finding of forest at this elevated site adds to the growing fossil evidence for multiple forest refugia in New Zealand during the last glaciation and is consistent with the pollen records, which have consistently indicated the presence of forest species during the last glaciations. Copyright © 2009 John Wiley & Sons, Ltd.     

Fossil Liverworts from the Lower Cretaceous Huolinhe Formation in Inner Mongolia, China

A new genus and species, Rebouliothallus huolinhensis sp. nov., and two uncertain species, Ricciopsis sp. and Hepaticites sp., are described. The fossils were collected from the Lower Cretaceous Huolinhe Formation of Huolinhe Basin, northeastern China. The new genus Rebouliothallus was established in the Aytoniaceae family of Marchantiales. Rebouliothallus huolinhensis appears to be quite similar to species of the extant genus Reboulia Raddi. The species is characterized by the relatively large ventral scales. Ventral scales are large, imbricate and arranged in two rows on the ventral surface. Rhizoids are either pegged or smooth. Ricciopsis sp. is characterized by the rosette-forming thallus. Hepaticites sp. shows some similarities to liverworts. Of the forty-nine Early Cretaceous floras of China, only two floras, one from the Huolinhe basin, Inner Mongolia and the other from the Jixi Basin, Heilongjiang Province, contain fossil liverworts. The rare fossil liverwort records in the Early Cretaceous floras of China may be the result of taphonomic bias.     

Revision of Cenomanian regular echinoids in collections at the Cairo Geological Museum,Egypt

Cenomanian rocks of Egypt are rich in moderately to well-preserved echinoids. From the large Cretaceous echinoid collection at the Cairo Geological Museum, a total of 27 regular species, in 12 genera, eight families, six orders and two subclasses, are revised and systematically described from these strata as exposed at several localities in Egypt. One new family (Pedinopsidae), one new subfamily (Lorioliinae), one new genus (Alternocidaris) and two new species of Goniopygus, G. subaequalis (Gebel Ekma) and G. macrotuberculatus (Gebel El Minsherah), are erected. Additionally, two species of Diplopodia, D. micropyga Fourtau, 1921 and D. halperti Fourtau, 1921, are transferred to the genus Pedinopsis.     

Reassessment of Prejanopterus curvirostris,a Basal Pterodactyloid Pterosaur from the Early Cretaceous of Spain

Prejanopterus curvirostra, from the Early Cretaceous of La Rioja province, was the first pterosaur genus and species described from Spain. The material comprises disarticulated cranial and postcranial remains from several individuals. The fossil-bearing bed is assigned to the lacustrine Leza Formation (eastern Cameros Basin, NW margin of the Iberian Range). This unit is regarded as either Berriasian-Valanginian or Barremian-Aptian. Prejanopterus curvirostris (specific name emended) was originally diagnosed on the basis of several characters of which the most significant was a lateral curvature of the rostrum. Re-examination of the holotype (rostrum) and paratype (partial rostrum with teeth) indicates that there is no genuine sideways bend of the preserved premaxilla-maxilla segments, but a slight dorsal curvature. Prejanopterus is characterized by a unique combination of characters: an emended diagnosis is provided. In contrast with previous estimates, the wing span of Prejanopterus was probably not much (if ever) in excess of 2 m. A phylogenetic analysis suggests that Prejanopterus is a basal pterodactyloid positioned between Pterodactylus and Cycnorhamphus-Gallodactylus. Prejanopterus represents the first evidence of Pterodactylidae in the Early Cretaceous of the Iberian Peninsula.     

An Unusual, Three-Dimensionally Preserved, Large Hadrosauriform Pes Track from “Mid”-Cretaceous Jiaguan Formation of Chongqing, China

Three-dimensional tracks provide unique insights into the locomotor mechanics of their track makers. An isolated, large hadrosauriform print attributable to Caririchnium lotus from the “mid”-Cretaceous Lotus track site (Jiaguan Formation) in China permits reconstruction of the footfall, weight-bearing, and kick-off phases of the step cycle. Large-scale modifications of the pes during the step cycle indicate C. lotus trackmakers were capable of locomotory modifications in response to substrate consistency beyond the “expected” shift between bipedal and quadrupedal postures. An unusual curvature to the trace of one of the outer digits indicates substantial transverse mobility. The remaining digits demonstrate lesser degrees of transverse movement accompanied by extension of the digits during footfall. The absence of overprinted scale-scratch marks and toe drags are consistent with a vertical kick-off of the pes and concomitant flexion of the digits. This track suggests that pedal mobility in C. lotus track makers was greater than previously suspected and has implications for reconstructions of hadrosauriform locomotion.     

Revision of the Clam Shrimp Genus Magumbonia from the Upper Jurassic of the Luanping Basin,Hebei, Northern China

The diverse clam shrimp Nestoria-Keratestheria fauna is widely distributed in the Dabeigou Formation in northern Hebei and eastern Inner Mongolia of China. Its important component genus Magumbonia from the Dabeigou Formation in the Luanping Basin, northern Hebei, China, is revised on the basis of a scanning electron microscope (SEM) examination of the type species M. jingshangensis Wang, 1984, which revealed morphological features not recognized previously. These include coarse reticulation on the umbo and prominent growth lines ornamented with densely spaced, small tubercles and fine lirae.     

The first lizard fossil (Reptilia: Squamata) from the Mesozoic of South Korea

Upper Cretaceous deposits in Mongolia, Chinese Inner Mongolia, and, more recently, southern China, have yielded individually rich and taxonomically diverser lizard assemblages. Here we describe the remains of a new terrestrial lizard, Asprosaurus bibongriensis gen. et sp. nov., from the Upper Cretaceous of South Korea. It represents the first record of a Mesozoic lizard from the Korean Peninsula and, although incomplete, is exceptional in its very large size. Characters of the mandible support attribution to crown-group Anguimorpha, with the closest similarities being to monstersaurs, the group represented today by the venomous North American Beaded lizard and Gila monster, genus Heloderma. This group is well-represented in the Upper Cretaceous fossil record in of eastern Asia, and the remains of large monstersaurs have been recovered from several dinosaur egg localities, suggesting dietary preferences similar to those of the living genus. The new Korean lizard, recovered from the Boseong Bibong-ri Dinosaur Egg Site, fits the same pattern.     

Clypeorbis? ultima n. sp. from the uppermost Maastrichtian of Austria: The youngest representative of the Clypeorbinae Sigal, 1952 (calcareous benthic foraminifera)?

A second possible species of the genus Clypeorbis Douvillé [type-species C. mammillata (Schlumberger)] is described from the topmost Maastrichtian of the Kambühel Formation of the Northern Calcareous Alps, Austria. This comparatively small-sized, asymmetric calcareous benthic foraminifer is characterized by its rather large biloculine embryonic apparatus and a reduced ventral umbo. It occurs in a mixed calcilithic–bioclastic littoral facies together with orbitoids, rotaliids, Siderolites, and other mainly calcareous benthic foraminifera as well as red algae. The stratigraphy is constrained by means of associated planktonic foraminifera, indicating the latest Maastrichtian age (CF-3 hariaensis zone). Clypeorbis? ultima n. sp. might represent the youngest representative of the Clypeorbinae Sigal that became extinct at the Cretaceous–Paleogene boundary.     

A New Mesopsychid (Mecoptera) from the Middle Jurassic of Northeastern China

The family Mesopsychidae Tillyard,1917 presently consists of ten described genera from the Early Permian to the Early Cretaceous of Australia,China,Kyrgyzstan,Russia,South Africa,Tajikistan,and Ukraine.Herein,a new genus and a new species of fossil mesopsychid,Epicharmesopsyche pentavenulosa gen.et sp.nov.,is described from the Middle Jurassic Jiulongshan Formation of Inner Mongolia,China,supplementing the family-level diversity.The well-preserved,new material reveals many characters,including antennae,wing venation,shape and genitalia that increase our morphological understanding of the family,although unfortunately the mouthparts and most of the legs are not preserved.Diagnosis of the Mesopsychidae is emended to include a four-or five-branched MP in both the fore-and hind wings.This is the first documentation of a wing coupling structure in the Mesopsychidae,consisting of three to four frenula bristles on the humeral lobe at the base of the costal margin of both hind wings.Asymmetrical shape and size of the left and right wings on a specimen of E.pentavenulosa gen.et sp.nov.,seems to be a common condition for mesopsychid taxa from northeastern China.