Early Jurassic basal sauropodomorpha dominated tracks from Guizhou,China: Morphology,ethology, and paleoenvironment

The newly discovered large(350 m^2) Yantan dinosaur tracksite, in the Lower Jurassic Ziliujing Formation of Guizhou Province, China, reveals at least 250 footprints of which ~97 can be resolved into trackways of sauropodomorphs. All the trackways are sub parallel likely indicating gregarious behavior. One theropod track(cf. Grallator) was recorded. The sauropodomorph tracks predominantly represent quadrupedal progression(Morphotype A), and footprint morphology is similar to the ichnospecies Liujianpusshunan, characterized by outward pes rotation. Three trackways indicate bipedal progression, and two of these(Morphotype B) indicate inward pes rotation, accompanied by elongate pes digit scratch marks. For the latter phenomenon three possible scenarios are discussed:(1) significant rotation changes accompanying changes in gait,(2) swimming behavior,(3) formation of undertracks.Sedimentological evidence indicates the tracks were made on a linguloid rippled, muddy, immature sandstone substrate characterized by significant differences in substrate consistency across the trackbearing surface. Microbially induced sedimentary structures(MISS) characterized by distinctive wrinkle marks indicate a stressed, probably semi-arid, paleoenvironment that was not conducive to habitation by invertebrate organisms. This is consistent with other evidence that Lower Jurassic sauropodomorph tracks are often associated with semi-arid paleoenvironments.     

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.     

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.     

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 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.     

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.     

New dinosaur tracks from Korea, Ornithopodichnus masanensis ichnogen. et ichnosp. nov. (Jindong Formation, Lower Cretaceous): implications for polarities in ornithopod foot morphology

Twelve trackways of ornithopods from Lower Cretaceous lacustrine margin deposits of the Jindong Formation represent new dinosaur trackways described from Korea. The site, discovered during highway construction, was rescued by removing the most important trackways to the Korean Natural Heritage Center in Daejeon, where they are on permanent display. The new ichnotaxon Ornithopodichnus masanensis ichnogen. et ichnosp. nov. is recognized as a distinctive robust tridactyl track, slightly wider than long (l/w ratio = 0.91), with positive (inward) rotation. The toe prints are very thick, broad and U-shaped, resulting in a trefoil outline with a smoothly rounded hind margin. Digit III is short and projects anteriorly much less than digit II and IV (= weak mesaxony). Divarication of digits II-IV is about 70° with interdigital angle II-III larger than III-IV. Trackway width is narrow and the stride length/track length ratio is about 4.2–4.6. The Ornithopodichnus trackways evidently represent gregarious blunt-toed Iguanodon-like bipedal ornithopods, although poorly preserved manus traces are discerned in a few trackways. Ornithopodichnus is distinct from other well known iguanodontid tracks that display much stronger mesaxony and indicates a polarity in ornithopod foot morphology that can be verified by reference to known foot skeletons.     

基于恐龙足迹重建攀西地区白垩纪恐龙动物群

攀西地区没有白垩纪恐龙骨骼化石记录,对该区白垩纪恐龙动物群组合的认识尚属空白。1991年,攀西地区首次发现了恐龙足迹,此后陆续发现了8个足迹点。这些足迹点共发现9种不同的非鸟恐龙足迹(6种非鸟兽脚类、1种蜥脚类和2种鸟脚类足迹),以及翼龙和龟类足迹。共11种足迹形态类型由185道行迹(与孤立足迹)组成,可能代表着同等数量的造迹者。其中飞天山组的组合多样性最强,小坝组和雷打树组较弱。这个相对全面的足迹数据库,为该地区白垩纪四足类的古生态学普查提供了动物群的组成信息,这在该缺乏骨骼化石的地区显得尤为重要。     

记广东南雄盆地上白垩统主田组鸟脚类足迹及中国已命名的大型鸟脚类足迹的属级新观察

记述了广东省南雄盆地上白垩统主田组3组恐龙行迹,由杨梅坑足迹点的足迹命名了南雄鸭嘴龙足迹（Hadrosauropodus nanxiongensis ichnosp. nov.）一新种;古市足迹点的足迹归入鸭嘴龙足迹一未定种（Hadrosauropodus isp.）。这是中国鸭嘴龙足迹属的首次描述,将该足迹的分布从北美拓展至中国,并为南雄盆地存在大型的鸭嘴龙提供了新的化石证据。还整理了中国已描述的大型鸟脚类足迹：中国足迹（Sinoichnites）被定为非正式的塑模标本;扬子足印（Yangtzepus）被归入兽脚类足迹;禽龙足迹（Iguanodonopus）被认为是裸名,归入禽龙足迹（Iguanodontipus）;嘉荫足迹（Jiayinosauropus）增加了新材料并得以具体描述;内乡足迹（Neixiang footprint）得以重新描述。     

Cenomanian dinosaur tracks on the islet of Fenoliga in southern Istria, Croatia

One hundred and forty six dinosaur footprints occur on the Cenomanian outcrop of Fenoliga. They are arranged in six trackways and six footprint groups (a–f). Two basic footprint types can be distinguished: firstly, large circular-elliptical sauropod footprints, forming four trackways (I, III, V, VI) and five footprint groups (a–e), and secondly, smaller, tridactyl theropod footprints forming two trackways (II and IV) and one footprint group (f). Three type specimens, one sauropod and two theropod, were determined during the course of the work. Since no dinosaur bones have been discovered in the Cenomanian rocks of Istria so far, these findings are outstandingly important in the analysis of the regional dinosaur fauna.     

山东莒南地质公园发现小型兽脚类恐龙足迹化石MinisauripUS

在山东省莒南地质公园内发现6个小型恐龙足迹化石,被归入足迹属Minisauripus。化石产于下白垩统大盛群田家楼组．时代为早白垩世巴列姆（Barremian）-阿普特期（Apdan）。5个较大,长约6cm,产于下部“主”层面上,其中4个组成2节行迹：1个较小,长约3cm,位于“主”层面30cm之上的上部层面上。山东Minisauripus的特征是：足迹个体小（长3．1～5．6cm,宽2．0～3．7cm）,三趾型,略不对称;足迹纵长,各趾近平行,趾垫较清晰;趾末端较钝,但爪迹较尖。Ⅲ趾比Ⅳ趾略长,而Ⅳ趾比Ⅱ趾略长且窄。此外,步幅较长,足长与步长之比约为10：1。与四川、韩国Minisau却淞不同之处是：足迹个体较大,长约是它们的2倍。继中国四川和韩国之后,山东是Minisau而淞在全球的第三个发现点。     

The first dinosaur tracksite from Abruzzi (Monte Cagno,Central Apennines,Italy)

A new Lower Cretaceous (lower Aptian) dinosaur tracksite, from the eastern side of Monte Cagno (Abruzzi, Italy), is described. Different styles of track formation are represented on the site surface. Most of the footprints are preserved as deep tracks, produced by trackmakers sinking into soft mud. Some tracks, better preserved than the others, are characterized by metatarsal impressions and were interpreted as the resting traces of a crouching theropod (based on their orientation and three-dimensional morphology). The 135 cm length of the track with metatarsal impressions indicates huge pedal proportions and represents the largest theropod trackmaker ever documented from the Mesozoic peri-Adriatic platforms of Italy.     

An Important Ornithischian Tracksite in the Early Jurassic of the Shenmu Region,Shaanxi, China

A large track site with multiple,well-preserved trackways of an Early Jurassic quadrupedal ornithischian dinosaur is the first report of Moyenisauripus-like trackways from Asia,herein named Shenmuichnu...     

中国新沂马陵山白垩纪恐龙足迹——从虎到肉食龙及由传说至古生物学

江苏省新沂市马陵山地区的4个神秘的印记,传统上被解释为“李存孝打虎处”的“虎爪印”和“人足迹”,重新将其解释为大盛群田家楼组恐龙足迹。这些足迹中只有一个最深的“虎爪印”保存了可识别的趾印,表明它是一个有着后内侧拇趾印的兽脚类恐龙左足迹,是一道单步约50cm的直线行迹的一部分;另一种解释是该足迹为小型蜥脚类恐龙的右后足迹,形成行迹的右侧部分,其左侧部分没有保存。马陵山足迹点提供了恐龙足迹如何影响中国民间传说的另一个案例。     

New ornithopod footprints from the Lower Cretaceous Kitadani Formation,Fukui, Japan: Ichnotaxonomical implications

Two ichnogenus of “large-sized” ornithopods are found from the Lower Cretaceous (Aptian) Kitadani Formation in central Japan. Caririchnium isp. is characterized by the longer pes print than its width with strong mesaxony. Amblydactylus isp. is characterized by the wider pes print than its length with weak mesaxony. In the northern hemisphere, Caririchnium-type footprints are known from the Berriasian–Cenomanian strata, whereas Amblydactylus-type footprints are known from the Barremian–Maastrichtian strata. It is consistent with the temporal and geographic distribution of non-hadrosauroid iguanodontians and basal hadrosauroids. It suggests that footprint length-width ratio and mesaxony are important factors to indicate trackmakers (basal iguanodontian or hadrosauroid). Two “large-sized” ornithopod ichnogenus from the Kitadani Formation is also consistent with two iguanodontians from same site. It indicates a high diversity of the Kitadani ichnofauna and its importance to elucidate the ecosystem of the Kitadani Formation.     

藏东昌都地区首次发现中侏罗世兽脚类恐龙行迹

藏东昌都地区阿涅塘化石点中侏罗世达布卡组滨湖相紫红色中-细砂岩中,发现以凸形迹完好地保存的4层恐龙足迹,包括Grallator类、大型兽脚类、小型兽脚类和小型蜥脚类等足迹类型,并构成两列完美的行迹. 组成行迹T1的4个Grallator类足迹被命名为Grallator anietangensis isp. nov.,对其进行了详细的形态学分析,并讨论了造迹恐龙的大小和运动状态,这是Grallator类足迹在西藏地区的首次发现. 组成行迹T2的9个大型兽脚类足迹被归为Theropoda igen. et isp. indet.,对该类足迹进行了形态描述和功能分析;这类足迹在中国是首次记录,其连续且清晰的尾迹表明造迹者并非处于正常运动状态,而是应对某种特殊境况的特别行为方式,即其正以昂起的前躯一边观察,一边以极低的速度前进,因前躯昂起,尾部相应下垂,从而留下尾迹. 两列行迹均显示出造迹者正沿古昌都湖岸前进. 中国迄今所发现的恐龙骨骼和Grallator类足迹化石主要表现为以西南地区为中心,再向四周不断发散的扩散模式,但Grallator类足迹化石综合提供的信息显示恐龙动物的扩散速度比骨骼化石反映的速度更快.      

Minisauripus―the track of a diminutive dinosaur from the Cretaceous of China and South Korea: implications for stratigraphic correlation and theropod foot morphodynamics

The diminutive (2.5–3.0 cm long), Cretaceous dinosaur track ichnogenus Minisauripus, previously known only from the type ichnospecies, M. chuanzhuensis, from a single locality in Sichuan Province China, is here reported from two new localities in South Korea and one in China. Material from the new Chinese locality is assigned to the new ichnospecies M. zhenshuonani on the basis of its distinctive morphology. Most of the new material is well-preserved, revealing narrow asymmetric tracks with claw traces, long step and phalangeal formula (2-3-4 for digits II, III and IV, respectively), suggesting a theropod track maker rather than an ornithischian, as originally inferred for the Chinese type material.The South Korean samples (eight tracks), from two localities in the Haman Formation, are considered Early Cretaceous (Aptian-Albian) in age, whereas the Chinese type material (21 specimens) has been assigned both an Early and Late Cretaceous age. The former age is probably correct as suggested by a new Minisauripus locality (5 specimens) from the Early Cretaceous (Barremian-Albian) of Shandong Province, China.Other diminutive tracks from the Sichuan fauna include Aquatilavipes sinensis (2.5 cm long, a possible junior synonym of Koreanoris hamanensis), Grallator emeiensis (2 cm long) and Velociraptorichnus sichuanensis (10–11 cm), which occur, in various combinations, with Minisauripus at both the new Korean and Chinese localities.In Minisauripus, digit III is very short in comparison with other theropods and provides a striking contrast to G. emeinsis. This difference has significant implications for standard assumptions about theropod track allometry. Based on the classic Early Jurassic forms Grallator and Eubrontes, it has long been inferred that relative digit III length shrank with increasing size (up to foot lengths of 30–40 cm). The reiteration of reduction in relative length of digit III in specimens in the size range of 2–3 cm indicates that the allometric or morphodynamic ‘program’ that influenced development in large theropod clades reiterated fractally in theropod clades a full order of magnitude smaller. This shows that a given allometry can be size-dependent in one clade and size-independent in another. Thus, the developmental program appears ‘contracted’ or morphologically miniaturized by heterochrony to manifest paedomorphically in some clades and peramorphically in others. This strongly suggests that ‘formal’ developmental ‘programs operated’ along similar morphodynamic lines in quite different clades.     

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。