A new bird track,Koreanaornis lii ichnosp. nov., from the Lower Cretaceous Hekou Group in the Lanzhou-Minhe Basin,Gansu, Northwest China,and implications for Early Cretaceous avian diversity

There are a growing number of Early Cretaceous avian tracks and trackways from around the world, with Asia (China and Korea) having the largest reported number and diversity of Mesozoic avian traces to date, and these new discoveries are increasing the Early Cretaceous avian ichnodivesrity of Laurasia. Here we report on a new Lower Cretaceous avian track locality in the Guanshan area, Yongjing County, Gansu Province, northwest China, and on a novel ichnospecies of Koreanaornis, Koreanaornis lii ichnosp. nov. Koreananornis lii is distinct from other Koreanaornipodidae in that it possesses a consistently wider digit divarication than previously described tridactyl tracks, and possess a short, small, posteromedially oriented hallux that displays a different orientation than that seen in Koreanaornis hamanensis. The lack of linear and angular data reported for digit I traces of many avian ichnotaxa has the potential to give misleading results in multivariate statistical analyses. Also, the wide divarication of Koreanaornis lii causes the ichnotaxon to not group with other Koreanornipodidae in multivariate analyses, but with Ignotornidae. Despite the results of the analyses, K. lii is morphologically distinct from these ichnotaxa. The results demonstrate that relying solely on multivariate statistical analyses without careful examination of footprint morphology will result in erroneous ichnospecies groupings. While new vertebrate ichnotaxa discoveries from Asia may support the hypotheses of the presence of a unique and endemic Asian vertebrate ichnofauna during the Cretaceous, the recent discovery of skeletal remains interpreted to be of a volant wading bird from the Early Cretaceous, and recent reports of tracks from volant avians, could suggest that flighted avians of the shore- and wading bird ecotypes could have had a Laurasian-wide distribution during the Early Cretaceous. However, strong convergence in foot morphology of shore- and wading birds suggests that avian ichnotaxa found in both present-day Asia and North America may have been made by birds endemic to eastern and western Laurasia during the Early Cretaceous.     

Saurischian track assemblages from the Lower Cretaceous Shenhuangshan Formation in the Yuanma Basin,Southern China

Cretaceous dinosaur tracks from Hunan Province are historically significant as the basis for three ichnotaxa: Xiangxipus chenxiensis, Xiangxipus youngi, and Hunanpus jiuquwanensis all representing theropodan tracks, described from a single site in 1982. Although the type locality has since been destroyed, the type specimen and replicas remained available for restudy in 2006, when paratype Hunanpus tracks and sauropod footprints were described from a second, nearby locality. Material from both localities is here re-described in detail. It is proposed that while Xiangxipus chenxiensis can be regarded as a distinct ichnospecies, probably representing an ornithomimid trackmaker, Xiangxipus youngi cannot be accommodated in the same ichnogenus. We consider it similar to the ichnogenus Wupus from the Lower Cretaceous of Sichuan Province, and therefore of possible avian affinity. We also find no justification for regarding Hunanpus jiuquwanensis as generically distinct from the widespread ichnogenus Grallator, and therefore rename it Grallator jiuquwanensis comb. nov. The Hunan track record is generally consistent with the sparse record of theropod skeletal remains in the province, but adds evidence of sauropods that was previously lacking.     

The longest theropod trackway from East Asia,and a diverse sauropod-, theropod-, and ornithopod-track assemblage from the Lower Cretaceous Jiaguan Formation,southwest China

Here we report a large dinosaur tracksite from an extensive fluvial sandstone surface in the Lower Cretaceous Jiaguan Formation of Sichuan Province, China. The site contains over 250 individual tracks comprising at least 18 recognizable trackways, including the longest theropod trackway (cf. Eubrontes) known from China. This exceptional theropod trackway consists of 81 successive footprints covering a distance of 69 m. The tracks are well-preserved and are expressed both as true tracks on the main “upper” surface and as transmitted undertracks on a locally exposed “lower” bed. Also recorded are six other theropod trackways, including small Grallator-like ichnites, eight sauropod trackways (cf. Brontopodus), and three small ornithopod (cf. Ornithopodichnus) trackways with a parallel orientation, which may indicate gregarious behavior. Several trackways of a larger theropod trackmaker show pes imprints with elongated traces of the metatarsals, suggesting extramorphological (substrate-controlled) variation and/or plantigrade posture, which is here interpreted as indicating a change in gait assumed in response to deep and soft sediment. The assemblage indicates a diverse dinosaur fauna in the Lower Cretaceous Sichuan Basin with variously sized theropods, sauropods, and ornithopods. The late occurrence of footprints of the Grallator-Eubrontes plexus in Lower Cretaceous strata is further evidence of the extended stratigraphic range of this morphotype and the distinct palaeobiogeographic distribution of these trackmakers in East Asia.     

白垩纪四足动物足印的生物地层学、生物年代学与遗迹相

从全球范围来看,白垩纪四足动物的足印多数是非鸟恐龙与鸟类留下的痕迹;少量足印来自翼龙、鳄鱼、龟、哺乳动物和其他四足动物。白垩纪的足迹化石以东亚(尤其是中国和朝鲜)和北美西部的最为人所知。南美(主要是阿根廷和巴西)也有一定数量广泛分布的足迹化石,欧洲、非洲与澳大利亚的白垩纪足迹组合则鲜为人知。以白垩纪四足动物的足印记录为基础,我们对两个全球足印生物年代重新进行了检查。早白垩世生物年代以蜥脚类与鸟脚类的足迹为特征。晚白垩世生物年代中的蜥脚类足迹较少,但是鸭嘴龙、暴龙和角龙的足迹增多了。另外,白垩纪足印化石的记录中记载了许多重要的生物地层学信息,如北美白垩纪中期蜥脚类恐龙的消失,以及白垩纪末恐龙的绝灭。越来越多来自东亚的白垩纪足印记录使我们对更精细的地方性白垩纪足印生物年代学有了初步印象。因此,以地方性四足恐龙(包括鸟类)遗迹属的地层分布为基础,可以识别出三个或四个足印生物年代。种类丰富并具有地方性特色的东亚的白垩纪鸟类动物的遗迹群,可能指示白垩纪时东亚存在着一个独特而繁盛的鸟类动物群。以足印化石为基础的这一假说有待进一步的验证。     

A Review of the Tetrapod Track Record in China,with Special Reference to Type Ichnospecies:Implications for Ichnotaxonomy and Paleobiology

<正>"Splitting" and "lumping" are perpetual problems in vertebrate,especially dinosaur, ichnotaxonomy.Chinese dinosaur ichnotaxonomy,which began in 1940,provides a series of interesting case studies,highlighting the dual problems of historical and dubious ichnotaxonomy. Chinese Mesozoic tetrapod track types have been placed into 63 ichnospecies(one Triassic,28 Jurassic,and 34 Cretaceous),exclusive of other,non-type ichnospecies or ichnotaxa identified from China.Fifty-two(～83%) of these 63 tetrapod ichnospecies were placed in monospecific ichnogenera. At the ichnogenus level,we prune—either by recognizing nomina dubia or by synonymy—17 from the list of 53 dinosaurian ichnogenera(a 32%reduction),leaving 36 ichnotaxa that we consider valid. Most of the cuts affect Jurassic theropod ichnotaxa,which are reduced from 23 to only nine because most ichnogenera are subjective junior synonyms of Grallator and Eubrontes.Fewer Chinese Cretaceous ichnotaxa(only six of 21 ichnogenera) are obvious nomina dubia or subjective synonyms, suggesting greater east Asian endemism during this time.Because ichnospecies differences are subtle, we provisionally retain ichnospecies as valid pending detailed comparative analyses of congeneric ichnospecies.This synthesis is long overdue and is necessary to address problems of historical and provincial ichnotaxonomy,which severely hamper comparisons of tetrapod ichnofaunas in space and time.     

First report of the small bird track Koreanaornis from the Cretaceous of North America: implications for avian ichnotaxonomy and paleoecology

Although body fossils of shorebirds and shorebird-like species are extremely rare from the Cretaceous, rapid increase in the discovery of bird footprint sites provides valuable alternate evidence to help fill gaps in the story of the early evolution of shorebird-like species. Newly discovered bird tracks from the Albian-Cenomanian Dakota Formation in northeastern Utah represent the first report of the ichnogenus Koreanaornis from North America and only the second report of bird tracks from this formation. These tracks are not attributable to Aquatilavipes as previously claimed. Three well-preserved trackways are described and provisionally referred to Koreanaornis cf. hamanensis (Kim). However, a review of the ichnotaxonomy of shorebird ichnites reveals that this ichnotaxon also closely resembles the Miocene ichnospecies Avipeda sirin (Vyalov). This latter comparison points to the need for a thorough evaluation of the similarity between Mesozoic and Cenozoic avian ichnotaxa, which may be over-split in some cases and under-differentiated in others.The new material helps distinguish ichnogenus Koreanaornis from the larger bird track Aquatilavipes, which is more abundant and widespread in North America. In some cases Aquatilavipes has been incorrectly used as a catch-all ichnogenus both in North America and Asia. The Dakota Formation stratigraphy at the tracksite indicates that the track makers lived in a marginal marine paleoenvironment. However, despite the widespread distribution of such facies, often replete with dinosaur tracks, the bird track record of the Dakota Formation, and the Cretaceous of the western USA remains relatively sparse in comparison with other areas such as east Asia.     

A Megatheropod Tooth from the Early Cretaceous of Fusui,Guangxi, Southern China

A nearly complete right maxillary or left dentary tooth（NHMG 10858） from the Lower Cretaceous Xinlong Formation of the Napai Basin, Fusui County, Guangxi, southern China, is described. The tooth is large in size, with a CBL of 37 mm. Given the geological age and its crown morphology, including the size, it is most likely that the tooth belongs to a carcharodontosaurid. The recovered specimen represents one of the largest theropod teeth hitherto reported from the Early Cretaceous of Asia.     

莱阳盆地莱阳群恐龙足迹化石的新发现

在山东莱阳盆地早白垩世莱阳群龙旺庄组发现15个足迹化石。化石产在灰绿色细砾粗砂岩(转石)的下层面,呈层底凸形(Convex hyporelief)保存。研究表明,足迹为二足行走、趾行式、三趾型,趾端具尖、长爪。足长约13 cm,长大于宽,近中趾对称。趾间角较大:平均值Ⅱ27°Ⅲ27.2°Ⅳ,Ⅱ、Ⅳ趾夹角平均54.2°。趾垫不清晰,无尾迹印痕。经研究,笔者认为这是一新的足迹类型,命名为Paragrallator ichnogen. nov.,并将其归入Coelurosauria Huene 1914次亚目,Anchisauripodidae(Lull 1904)足迹科。即该足迹系由小型兽脚类恐龙——虚骨龙类所形成。恐龙足迹形成于滨湖相—河流相环境中。     

陕西商洛地区下白垩统大型兽脚类恐龙足迹

记叙了中国陕西省商洛市邵涧村下白垩统东河群2个大型兽脚类恐龙足迹,并将其归于Megalosauripus isp.。 东河群的时代与热河生物群同期,邵涧Megalosauripus isp. 可能可与热河生物群的大型兽脚类恐龙之足部形态做简 单对比。此外,邵涧Megalosauripus isp.造迹者的体长约为7.4m,这意味着早白垩世的陕西中南部亦存在着大型的 兽脚类恐龙。邵涧Megalosauripus isp.的发现,不仅为中国早白垩世恐龙足迹增加了新的成员,而且为将来在该地 区下白垩统中发现大型兽脚类恐龙的骨骼化石提供了确凿的依据。     

A diversified vertebrate ichnite fauna from the Feitianshan Formation (Lower Cretaceous) of southwestern Sichuan,China

The Bajiu tracksite exposed on a fine sandstone surface at the top of the Feitianshan Formation, close the contact with the overlying Xiaoba Formation, reveals a saurischian dominated ichnofauna consisting of four sauropod and at least eight theropod trackways. The sauropod trackways display a medium-wide gauge pattern characteristic of titanosauriforms, while the theropod trackways can be subdivided into two distinct morphotypes. One is similar to the ichnogenera Eubrontes and Megalosauripus, the other (cf. Dromaeopodus) appears to represent a functionally didactyl dromaeosaur where pedal digit II is represented only by an oval basal pad. A single swim track is possibly attributable to a turtle or crocodylian. Such sauropod-theropod dominated ichnofaunas are consistently typical of the red-bed tracksites in the region and useful for characterizing a fauna that is otherwise poorly known from body fossils. The stratigraphic position and late occurrence of the (typical Jurassic) Eubrontes-Megalosauripus morphotype is well known from other Cretaceous localities in China and seems to reflect a peculiarity in the theropod faunas of East Asia.     

A theropod track assemblage including large deinonychosaur tracks from the Lower Cretaceous of Asia

A total of more than 40 tridactyl and didactyl tracks were preserved as natural casts on four fallen blocks of sandstone representing the Lower Cretaceous Jiaguan Formation of Gulin County in southeastern Sichuan Province, China. While several trackways can be distinctly followed, others are isolated imprints only. All have been flattened by overburden pressures. Tridactyl tracks are present with three size-classes being <10 cm, 10–20 cm and >20 cm in length. Morphologically they are similar to the ichnogenus Eubrontes, considering the relatively weak mesaxony. Eight of the tracks on one of the blocks are clearly didactyl and are here interpreted as representing large and medium sized dromaeosaurids. The largest track is about ∼30 cm long and comparable in size to the type of Dromaeopodus (∼28 cm), from the Lower Cretaceous of Shandong Province, which was the largest dromaeosaurid track previously reported. This report adds new data to the growing number of dromaeosaurid tracksites reports from China, and from the Jiaguan Formation, suggesting that this theropod group had a preference for fluvial paleoenvironments.     

A diverse saurischian (theropod–sauropod) dominated footprint assemblage from the Lower Cretaceous Jiaguan Formation in the Sichuan Basin,southwestern China: A new ornithischian ichnotaxon,pterosaur tracks and an unusual sauropod walking pattern

A sample of fallen blocks of fluvial sandstone from the Lower Cretaceous Jiaguan Formation of Sichuan Province yielded an assemblage of dinosaur and pterosaur tracks preserved as natural impressions and casts. Collectively the assemblage reveals 132 tracks representing at least 30 trackways of tridactyl and didactyl theropods, sauropods, ornithopods and pterosaurs. Ichnotaxonomically, the trackways of small tridactyl theropods (pes lengths 7–18 cm) are indeterminable, whereas the trackway of a small didactyl dromaeosaur (pes length up to 7.5 cm) is tentatively assigned to cf. Velociraptorichnus. The sauropod trackways are assigned to cf. Brontopodus based on the medium to nearly wide-gauge pattern. Other characteristics are the U-shaped manus and strong heteropody. One sauropod trackway shows a peculiar pattern with a lack of left manus imprints, and an unusual position and rotation of right manus imprints. Different scenarios and explanations for this phenomenon are discussed. Ornithopod trackways are the most abundant in the sample and characterized by pes imprints of a small biped that are assigned here to the new ichnospecies Caririchnium liucixini. It is characterized by an unusual broad shape and weak mesaxony. Bivariate analysis of different Caririchnium ichnospecies reveals increasing mesaxony toward the larger forms, a trend that is the reverse of typical theropod ichnotaxa, where large imprints have weak mesaxony. Three isolated, small pterosaur tracks (two manus, one pes) are visible on a single surface. They show strong similarities to the widespread ichnogenus Pteraichnus. This is the ninth report of tetrapod tracks from the Jiaguan Formation in recent years and represents one of the most diverse assemblages recorded to date. It is also rare evidence of typical didactyl dromaeosaur tracks and the co-occurrence of sauropod and ornithopod tracks in a fluvial depositional environment representing arid climate conditions.     

山东沂沭断裂带早白垩世晚期恐龙足迹特征差异性*

近些年来,在山东沂沭断裂带南部地区(莒南、临沭、郯城和东海）的下白垩统上部大盛群中发现了大量的恐龙足迹化石,而这些化石反映的足迹类型、形态大小、足迹踩踏深度等方面都存在有差异性。研究发现,在足迹类型方面,莒南地区以兽脚类和鸟脚类恐龙足迹为主,临沭、郯城和东海地区以蜥脚类恐龙足迹为主。足迹大小方面,莒南地区的兽脚类足迹可划分为类型Ⅰ和类型Ⅱ两种类型,而东海地区的恐龙足迹主要属于类型Ⅱ,但是莒南地区大多数足迹的趾间角要小于东海地区的,这可能是由于兽脚类恐龙属种的不同所造成的;莒南地区鸟脚类足迹绝大多数要小于临沭地区的鸟脚类足迹;郯城地区的蜥脚类足迹较小,而莒南、临沭和东海地区大多数蜥脚类足迹的大小较为相似,个别临沭和莒南地区的蜥脚类恐龙足迹较大。足迹踩踏深度方面,临沭、东海和郯城地区的足迹相比莒南地区的足迹要稍深,这与足迹形成在不同湿度、黏度、颗粒度的沉积物表面上有密切联系。基于上述研究认为,在早白垩世晚期,沂沭断裂带内生活着种类繁盛,数量庞大的恐龙动物群,联系到沂沭断裂带东部诸城地区早白垩世早期地层中也发现有大量的以兽脚类为主的恐龙足迹化石,推断在早白垩世,沂沭断裂带及胶莱盆地当时为一个理想的恐龙生存、繁衍的栖息地。同时,山东早白垩世大量恐龙足迹化石的发现也为整个华北甚至东北亚地区晚中生代恐龙属种的分布、演化以及古地理研究提供了丰富的材料和信息。     

Small scale scrapes suggest avian display behavior by diminutive Cretaceous theropods

A set of about 25 diminutive sub-parallel scrapes in the Cretaceous Haman Formation of South Korea fit the morphology of the small theropod track Minisauripus which is also known from this formation. The scrapes are interpreted as evidence of display behavior, and suggest that the trace maker was an adult engaged in avian-like courtship behavior. Although avian theropods are also known from this formation, the scrapes are inconsistent with their foot morphology. Although large theropod scrapes have been interpreted as evidence of display behavior they are only known from North America. Thus, the scrapes described here are the first reported from Asia, and the first interpreted as evidence of display behavior in such a diminutive species.     

Reassessment of coelurosaurian (Dinosauria,Theropoda) remains from the Upper Cretaceous Wangshi Group of Shandong Province,China

Non-avian theropods are well represented in the Cretaceous of Asia. However, the first theropod remains ever reported from China, from the Wangshi Group of Shandong Province, have not been reassessed since they were described by Carl Wiman in 1929, despite the great strides that have been made in theropod phylogenetics in the last eight decades. The remains redescribed herein consist of four vertebrae (two cervicals, one dorsal and one caudal) from one site, and a caudal vertebra and ungual phalanx from another. The vertebrae from the first site are assigned to tyrannosauroids and ornithomimosaurs, as is the caudal vertebra from the second site, whereas the manual ungual also bears resemblances to those of non-maniraptoran coelurosaurs (specifically compsognathids and ornithomimosaurs). Theropods remain relatively rare in the Wangshi Group; nevertheless, the remains redescribed herein highlight the potential for the discovery of more theropod remains in future excavations.     

中国早白垩世热河生物群中大型霸王龙类化石的发现

一件包括部分头骨、下颌、肠骨等的不完整兽脚类恐龙骨架发现于辽宁喀左早白垩世九佛堂组中,估计身体全长可达9~10m。其前颌骨主体部分高、肠骨外侧面具显著的直立的嵴等特征表明该化石应归霸王龙类。它以外鼻孔大、上颌骨背缘前部略内凹、上颌骨孔向前达眶前窝前缘、腹缘与眶前窝腹缘存在较大距离等特征不同于晚白垩世的霸王龙科分子,故被命名为一新属种——喀左中国暴龙(Sinotyrannus kazuoensis gen. et sp. nov.),或许代表了最早的霸王龙科类型。该属种是已知个体最大的前晚白垩世霸王龙类,也是辽西及周边地区热河生物群中个体最大的兽脚类恐龙。它的发现不仅表明东亚是霸王龙类最主要的演化地区之一,而且为探讨霸王龙科的起源、热河生物群的组成与生态系统等提供了重要依据。     

A new tyrannosaurine specimen (Theropoda: Tyrannosauroidea) with insect borings from the Upper Cretaceous Honglishan Formation of Northwestern China

A large theropod ilium was recently collected from the Upper Cretaceous Honglishan Formation in the Sangequan area of the northern Junggar Basin, Xinjiang, China, which represents the first officially reported dinosaur fossil from this formation. Several morphological features, including robust supracetabular ridge, reduced supracetabular crest, concave anterior margin of the pubic peduncle, ventral flange on the pubic peduncle, converging dorsal surface of the iliac blades, laterally visible cuppedicus shelf, and ventral flange on the posterior surface of pubic peduncle, suggest that this specimen can be referred to Tyrannosaurinae, and furthermore, a few differences between this specimen and other tyrannosaurines in particular the contemporary Asian tyrannosaurine Tarbosaurus suggest that IVPP V22757 may represent a new tyrannosaurine species. However, in the absence of extensive data that would make it possible to properly evaluate these differences, we refrain from naming a new taxon based on this specimen. Some insect borings are also identified in this specimen, and are referable to the ichnogenus Cubiculum, which is interpreted as the insect pupichnia. This new fossil documents the presence of a gigantic theropod in the Upper Cretaceous of Junggar Basin, adding new information on its poorly studied ecosystems.     

A proscoliine wasp (Insecta: Hymenoptera: Scoliidae) from Shandong peninsula, East Asia

A new species of wasp from the Laiyang Formation in Shandong, China is described and named Cretaproscolia asiatica sp. nov. It is the second record of the genus Cretaproscolia Rasnitsyn and Martínez-Delclòs, 1999, and extends the distribution of the genus from the Lower Cretaceous of South America into the upper Mesozoic of East Asia. The diagnosis of Cretaproscolia is emended based on the new material.     

Aturoidea (Nautilida) from the Upper Cretaceous Sada limestone in Shimanto City, Kochi Prefecture, Japan

The first specimen of Aturoidea to be recorded in East Asia has been found in the Upper Cretaceous Sada limestone in Shimanto City, Kochi Prefecture, Japan. The specimen is one of the few representatives from the Upper Cretaceous, along with species known from Libya, Angola, and India. The specimen is very similar to A. mathewsonni from the Paleocene deposits in California. However, we describe the specimen as A. cf. mathewsonni, as it slightly differs from A. mathewsonni in the shape of the lateral lobe of the suture. The finding reveals that Aturoidea had already lived in waters around East Asia in the Late Cretaceous.