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.     

Theropod assemblages and a new ichnotaxon Gigandipus chiappei ichnosp. nov. from the Jiaguan Formation,Lower Cretaceous of Guizhou Province,China

A newly discovered Jiaguan Formation(Lower Cretaceous) tracksite from the Linjiang region of Guizhou Province, China, reveals the first example of a Cretaceous track morphotype attributable to the non-avian theropod ichnogenus Gigandipus, here named Gigandipus chiappei ichnosp nov. The theropod dominated locality also reveals the second report of the avian theropod ichnogenus Wupus, one of the largest avian traces currently known from the Lower Cretaceous. The Linjiang site provides evidence to support previous interpretations of a distinctive Lower Cretaceous theropod-dominated ichnofauna that was widespread in China and East Asia and highlights the similarity between Lower Cretaceous theropod ichnotaxa in East Asia and those found in the Lower Jurassic both in East Asia and elsewhere. These similarities in turn create various ichnotaxonomic challenges familiar to researchers working on theropod tracks, and we recommend caution in the naming of new theropod ichnotaxa at the ichnogenus level.     

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.     

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.     

New dinosaur tracks from the Lower Cretaceous (Valanginian-Hauterivian) Saniri Formation of Yeongdong area,central Korea: Implications for quadrupedal ornithopod locomotion

Newly discovered tracks of a large quadrupedal ornithopod and a theropod with trail trace, and a bird track from the Cretaceous Saniri Formation of Yeongdong area are described. One ornithopod trackway is very unusual in having large (length width ∼15 and >15 cm respectively) clover-leaf-like manus tracks situated anteromedial to the pes tracks with atypical negative, inward rotation of 45°. Ornithopod pes tracks are quadripartite with three separated, elongate-oval, nearly parallel sided, wide digit impressions and a separate suboval heel impression (mean length and width about 41 and 36 cm respectively: l/w ratio 1.13). Manus track morphotypes are a clover-leaf-shaped configuration of three digit impressions, representing digits II–IV, in triangular configuration and registered just in front of (anteromedial to) pes track digits II and III. The pes morphotype is typical of Caririchnium, but the manus morphotype is quite distinct from previously described ichnotaxa, thus justifying a new ichnotaxon: Caririchnium yeondongensis ichnosp. nov., probably representing a facultatively-quadrupedal Iguanodon-like trackmaker.Theropod tracks are composed of three tapered pes digit impressions with interdigital angles between digits II and IV 45°. Length and width about 22.8 cm and 15.5 cm, respectively. Pace, stride, and pace angles are about 51 cm, 101 cm, and 170°, respectively. Theropod tracks are characteristically associated with a nearly continuous tail trace, which is up to 360 cm in length, 4.5 to 6 cm in width, and broad “U” shaped in cross section. Expulsion rims and dragging striae occur intermittently. In addition, dinosaur skin impressions, poorly preserved large sauropod tracks, a bird track, invertebrate and plant fossils are found from the lake margin deposits also containing rain drop impressions and desiccation cracks. Dinosaur tracks of the Yeongdong area represent the oldest (Valanginian-Hauterivian) dinosaur tracks of Korea.     

西南极利文斯顿岛晚三叠世遗迹化石及其环境意义

西南极利文斯顿岛晚三叠世迈尔斯陡崖组形成于海底浊积扇中扇的上、下部分,发育着许多海相遗迹化石.在采集到的样品中鉴定共有15个遗迹属、16个遗迹种,其中有10个可以鉴定到遗迹种、两个比较种,4个只鉴定到遗迹属,未鉴定到遗迹种;并建立了1个新遗迹属及新遗迹种是文献中从未发现的.除新遗迹属种外,其余14个遗迹属,15个遗迹种都曾经在深海相浊积岩内发现过,Belorhaphe、Glockerichnus、Lophoctenium、Rhabdoglyphus、Paleodictyon、Sublorenzinia、Spirophycus、Strobilorhaphe、Tuberculichnus、Cochlichnus等属于浊流前产生在深海相泥岩内的高度分异的雕画迹(Graphoglyptida),它们产于泥岩却保存为上覆砂岩底面的铸型凸起.Fucusopsis和Neonereites却产生在砂岩内代表浊流后形成的沉积后遗迹组合.这些遗迹化石属深水的Nereites遗迹相,为研究区的沉积环境增添了可靠依据.     

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.     

重庆綦江中白垩统夹关组恐龙足迹群的发现及其意义

记述了来自重庆綦江中白垩统夹关组恐龙足迹群，命名了甲龙亚目的中国綦江足迹（Qijiangpussinensis ichnogen．et ichnosp．nov．）、兽脚亚目的敏捷舞足迹（Wupusagilis ichnogen．etichnosp．nov．）和鸟脚亚目的炎热老瀛山足迹（Laoyingshanpus torridus ichnogen．etichnosp．nov．）和莲花卡利尔足迹（Caririchniumlotusichnosp．nov．）等新属种。中国綦江足迹是中国首次发现甲龙类的足迹。莲花卡利尔足迹提供了鸭嘴龙类在各发育阶段所留下不同的足迹。     

A new record and a new ichnospecies of Arachnostega from the Middle Miocene of Egypt

The ichnogenus Arachnostega Bertling, 1992 is recorded for the first time from a bioeroded, coral-rich, Middle Miocene limestone bed in the middle Siwa Escarpment Member (Marmarica Formation) at Siwa Oasis, northern Western Desert of Egypt. These burrowing traces are preserved on the surface of a few internal moulds of bivalves and gastropods. Until now, the ichnogenus Arachnostega included a single ichnospecies, A. gastrochaenae Bertling, 1992. In the material studied herein, two ichnospecies are identified, described, illustrated and compared with other, similar traces. One of these is a new ichnospecies of Arachnostega; this is here named A. siwaensis ichnosp. nov. It is distinguished from A. gastrochaenae in the possession of tunnels that usually branch in a V-pattern; intervals of ramification are longest in the main branch and decrease in second- and third-order branches. Arachnostega traces were produced in a warm, low-energy, shallow-marine (<10?m water depth) environment. Polychaetes are the most likely producers of marine Arachnostega, which are commonly occurred in the same studied bed. This new occurrence is significant in extending the known stratigraphical and geographical ranges of Arachnostega into the Middle Miocene of Egypt.     

First report of lacertiform (lizard) tracks from the Cretaceous of Asia

The well-preserved trackway of a lacertiform, lizard-like trackmaker from the Haman Formation (Cretaceous) of Korea is described as Neosauroides koreaensis ichnogen. et ichnosp. nov. This is the only example of a Cretaceous lacertiform or lizard-like trackway currently known in the global track record. Although lacertiform trackways, mostly assigned to the ichnogenus Rhynchosauroides, are common in the global Triassic, they are almost entirely absent in the Jurassic and Cretaceous. Moreover, ichnological classification criteria allow that Neosauroides is morphologically distinct from Rhynchosauroides at the genus level, and more like the tracks of the extant lizard Sceloporus. The reasons for the conspicuous lack of post-Triassic occurrences are not certain, but not due to a post-Triassic lack of potential lizard trackmakers. Thus, the preservation biases are likely due to paleobiological factors such as trackmaker ecology and paleoenvironmental preference.     

Fossil caddisflies (Insecta: Trichoptera) from the Early Cretaceous of southern England

The first fossil caddis cases from the Early Cretaceous (Wealden) of SE England are described and named Conchindusia rasnitsyni ichnosp. nov., Piscindusia sukachevae ichnogen. et isp. nov., ?Ostracindusia vyalovi ichnosp. nov. and Pelindusia percealleni ichnosp. nov. In addition, the necrotauliid caddisfly Paratrichopteridium purbeckianum (Handlirsch) comb. nov. from the earliest Cretaceous (Purbeck Group) of Dorset is redescribed.     

A new mammal track from the Laramie Formation (Maastrichtian) at the Fossil Trace locality,Golden, Colorado

A previously unknown and morphologically distinct pentadactyl mammal track was recovered from the Upper Cretaceous (Maastrichtian) Laramie Formation at the Fossil Trace site, a National Natural Landmark which is the type locality for Schadipes crypticus, the only named mammal track known from North America. The track is different, and larger than S. crypticus, and thus is evidence of a diversity of mammal trackmakers at this site. Although Cretaceous mammal tracks are very rare, preliminary indications are that those currently known are all morphologically distinct and therefore indicative of a global diversity of different trackmakers, as the body fossil record suggests. Lack of well-preserved mammal trackways with morphologically distinct manus and pes footprints hampers efforts to name diagnostic ichnotaxa.     

Zurupleuropholis gen. nov. (Teleostei,Albian, Argentina), first pleuropholids from the Cretaceous of South America

Although the Cretaceous is characterized by a rich fish diversity, Cretaceous continental fishes from Gondwana are poorly known and comparatively scarce. Among these fishes, the family Pleuropholidae is only known by a few species relatively poorly preserved, from the Middle Jurassic to Lower Cretaceous of Europe, Africa, North America, and South America. In this paper, two new species of the pleuropholid new genus Zurupleuropholis are described, Z. quijadensis gen. et sp. nov. and Z. decollavi gen. et sp. nov. The new fishes were recovered in the Lower Cretaceous lacustrine Lagarcito Formation of central-west Argentina. This taxon constitutes a relevant finding considering that the representation of the family Pleuropholidae is rare worldwide. Zurupleuropholis gen. nov. appears to be the youngest known member of Pleuropholidae, and it represents the second record of the family in South America and the first record in the Cretaceous of the continent.     

An Ichthyornis-like bird from the earliest Late Cretaceous (Cenomanian) of European Russia

Ornithuromorph birds (the clade which includes modern avian radiation) first appeared in the Early Cretaceous in Asia and achieved a great diversity during the latest ages of the Late Cretaceous (Campanian and Maastrichtian). The evolutionary history of orithuromorphs during the first 17 MYAs of the Late Cretaceous (Cenomanian to Santonian ages) remains very poorly known, as the fossil record for this time interval is largely restricted to several isolated finds of the classic avian genus Ichthyornis in North America. Here we describe an isolated distal tibiotarsus of an evolutionary advanced bird, morphologically similar to Ichthyornis, from the middle Cenomanian of Saratov Province, European Russia. This is the first documentation of an Ichthyornis-like bird in the Old World. The find further constitutes only the second pre-Campanian record of the Late Cretaceous Ornithuromorpha in Eurasia, the second record of Cenomanian birds in Russia. This discovery shows that Ichthyornis-like birds enjoyed a wide geographical distribution as early as the beginning of the Late Cretaceous. Given that the earliest and the most primitive ornithuromorph birds are known from Asia, the new find supports a Eurasian origin for Ichthyornithidae.     

云南下侏罗统禄丰组恐龙足迹的发现和张北足迹属新观察

记述了云南省禄丰县腰站乡下侏罗统禄丰组2个恐龙足迹,命名了棋盘张北足迹（Changpeipus pareschequier ichnosp. nov.）一新种。这是在禄丰组中首次发现恐龙足迹化石,重新观察并修订了张北足迹一属的属征。滦平张北足迹（Changpeipus luanpingeris）应为石炭张北足迹（Changpeipus carbonicus）的亚成年个体,为同物异名。此前被认为是石炭张北足迹（Changpeipus carbonicus）前足迹的IVPP V2472 2a,应为其未成年体的后足迹。张北足迹与卡岩塔足迹（Kayentapus）类似。综合种种特征,棋盘张北足迹（新种）的造迹恐龙很可能为禄丰组的腔骨龙类（Coelophysoidea）     

Trace Fossils from Lower Cambrian Wusongger Formation in Xiaoerbulake Outcrop,Kalpin Area,Xinjiang

Based on the analysis of trace fossils collected from the typical outcrop of the Lower Cambrian Wusongger Formation in the Kalpin area, ten ichnospecies of six ichnogenus were identified in the upper member of the Wusongger Formation, with most of them being found for the first time. The trace fossils are described seriatim. The ichnospecies are mainly represented by such common trace fossils as Ophiomorpha nodosa, Helminthopsis hieroglyphica, Helminthopsis ichnosp., Planolites beverleyensis, Planolites vulgaris, Planolites montanus, Palaeophycus striatus, Palaeophycus curvatus?, Cochlichnus anguineus and Rituichnus elongatum. The trace fossils are of high diversity and low abundance, and can be diagnosed as the Cruziana ichnofacies. They can be interpreted as having formed in a shallow water environment.     

Research on Dinoasur Footprints in Zizhou, Shaanxi Province, China

In recent years, the discoveries of dinosaur footprints have been successively reported from Dianshi Town, Zizhou City, Shaanxi Province. The footprints include the tracks of theropod, ornithopod and sauropod as well. Actually, the dinosaur footprints were found by local inhabitants much earlier in history, but not for science. The slabs bearing the dinosaur footprints were collected for domestic use, such as building stones, millstones, cellar covers, sheepfold fences, windlass holders, etc. This paper is to describe the dinosaur footprints on both sides of three slabs used for cellar covers, sheepfold fences and windlass holders by the local people. 24 dinosaur footprints and 4 trackways have been recognized and all of them belong to theropod. Four kinds of dinosaur footprints are identified, including 1 new ichnogenus and 2 new ichnospecies: (1) Shanbeipus caudatus ichnogen. et ichnosp. nov.; (2) Pengxianpus yulinensis ichnosp. nov.; (3) Eubrontes ichnosp; (4) Kayentapus ichnosp. Dinosaur-footprint-bearing beds were initially identified as the Fuxian Formation of the Lower Jurassic. Multipal dinosaur footprints associated with fragment plant fossils suggest a humid fluvio-lacustrine environment setting in the Ordos Basin during the early period of the Jurassic.     

New Early Cretaceous Pterosaur-Bird Track Assemblage from Xinjiang, China: Palaeoethology and Palaeoenvironment

A pterosaur-bird track assemblage from a sandstone-siltstone-mudstone sequence of the Lower Cretaceous Tugulu Group of Xinjiang comprises the first pterosaur track record from this province and the largest specimen thus far known from China.The pterosaur tracks are assigned to the ichnogenus Pteraichnus based on the triangular overall-shape,the four elongate digit traces and the robust manual digit trace Ⅲ.Supposed trackmakers were dsungaripterid pterodactyloids whose skeletal remains are well known from the Tugulu Group.The bird tracks that occur on the same surface,are those of typical shorebirds,known from different other localities in southeast Asia.The congruence with Koreanaornis dodsoni described from the same stratigraphic level justifies an assignment to this ichnospecies.This is a further evidence of the co-occurrence of pterosaurs and birds in a typical iakeshore environment with possible seasonal alteration of water supply and aerial exposure indicated by wave ripples,mudcracks and repeated cycles of coarse to fine sediment.Pterosaurs and birds frequented the shoreline and may have fed also on the numerous invertebrates such as the Scoyenia tracemaker that left abundant burrows.