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.     

河北赤城地区土城子组中一例极可能的兽脚类恐龙蹲伏迹(英文)

描述了河北省赤城县寺梁山土城子(后城)组一例极可能的兽脚类恐龙蹲伏迹及相关的足迹,该蹲伏迹也是该组地层目前发现的最大的兽脚类足迹。虽然缺乏前足迹和尾迹,但因保存了左跖骨印、关联的坐骨及可能的耻骨胼胝印被定为蹲伏迹。该蹲伏迹是世界上第三例非对称的兽脚类恐龙蹲伏迹。     

河北赤城地区土城子组中一例极可能的兽脚类恐龙蹲伏迹

描述了河北省赤城县寺梁山土城子（后城）组一例极可能的兽脚类恐龙蹲伏迹及相关的足迹,该蹲伏迹也是该组地层目前发现的最大的兽脚类足迹。虽然缺乏前足迹和尾迹。但因保存了左跖骨印、关联的坐骨及可能的耻骨胼胝印被定为蹲伏迹。该蹲伏迹是世界上第三例非对称的兽脚类恐龙蹲伏迹。     

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。     

安徽齐云山晚白垩世兽脚类恐龙足迹点沉积环境

安徽齐云山兽脚类足迹群是目前中国晚白垩世保存最好的恐龙足迹群之一,对其沉积环境的研究为恢复白垩纪末期恐龙动物群的生活环境提供了证据。研究区足迹全部为兽脚类凸型足迹,与无脊椎动物遗迹Palaeophycus tubularis和Scoyenia sp.保存在一起,指示陆相沉积中的浅水沉积。通过薄片鉴定,足迹层位主要为岩屑砂岩,其平均粒径、标准偏差、偏度和峰度4个粒度参数特征与典型的河相砂一致;粒级分布直方图、概率累积曲线图及散点图表明,足迹层位从下到上颗粒逐渐变细,水动力条件变弱,沉积环境由河床沉积转变为边滩沉积。综上,齐云山兽脚类足迹点沉积环境为大型曲流河沉积,足迹保存在边滩沉积的下部,是白垩纪末期干旱气候条件下兽脚类恐龙动物群的水源地。     

A new Cenomanian vertebrate tracksite at Tamajón (Guadalajara,Spain): Palaeoichnology and palaeoenvironmental implications

A new Upper Cretaceous vertebrate tracksite has been discovered at Tamajón (Iberian Ranges, Guadalajara, Spain). The track level is a relatively smooth and slightly undulating sandy ferruginous crust, corresponding to an erosive surface at the base of a small meandering channel. It is incised into the underlying planar cross-bedded sandstones of coastal bars located at the middle-upper part of the Utrillas Formation (middle-upper Cenomanian). The site shows an extraordinary concentration of vertebrate tracks, among which numerous sets of two to five isolated digit impressions (“swim tracks”) and, at least, two trackways referred to crocodyliforms, and a single tridactyl footprint probably produced by a theropod dinosaur can be recognized. There are also several long traces (epichnial grooves) revealing sharp direction changes (up to 90°) which seem to be fish fin traces (Undichna unisulca), although crocodyliforms (tail marks) and/or fish invertebrates cannot be rejected as possible tracemakers. Some crocodyliform tracks reveal a thin raised rim, due to the displacement of the sediment by the pressure produced by the feet. Several impressions are moderately deformed by small sediment slides, only preserving their deepest part (claw marks). This is clearly indicative of a soft substrate with a high degree of plasticity and water content at the time of the track registration. Nevertheless, the sediment was hard enough to preserve manus and pes print morphologies and also possible crocodyliform tail and/or fish fin traces. Small rhizoliths can also be recognized and may belong to herbaceous wetland vegetation. The morphology of the palaeochannel, the sedimentary context and the track preservation seem to indicate that the tracks were impressed in a shallow channel located near the coast, under wet conditions and in different moments of time. This discovery represents the first occurrence of vertebrate ichnites in the Utrillas Formation, a stratigraphic unit where osteological and ichnological remains are relatively scarce, and it confirms that some crocodyliforms lived in near coast channels during the deposition of this unit.     

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.     

Dinosaur footprint assemblage from the Lower Cretaceous Khok Kruat Formation,Khorat Group,northeastern Thailand

The Khok Kruat Formation is the upper part of the Khorat Group, which consists of upper Lower Cretaceous non-marine sedimentary rocks in northeastern Thailand. Many dinosaur footprints have been known from the upper Lower Cretaceous (Aptian–Albian) Khok Kruat Formation at the Huai Dam Chum (Tha Uthen) site, northeastern Thailand. Approximately 600 tracks occur in thin mudstone layer of the northern part of the outcrop at the Huai Dam Chum track site. Two types of footprints, small-sized theropod and crocodylomorph are imprinted with mud cracks and ripple marks on the thin mud layer. Most of footprints are referred to cf. Asianopodus, and are imprinted by small-sized theropoda, probably ornithomimosauria. Theropod tracks are mainly separated into two groups, Group A and Group B. From ichnological viewpoints, the small-sized theropod track assemblage indicates the herd behaviour and its idiosyncratic group composition. In particular, the histogram of size-frequency measurements of Group A shows the anomalous bimodal distribution. We consider that there are two hypotheses; the first one is due to the male-female difference, and the second is a result of the different growing stage.     

Mid-Cretaceous dinosaur track assemblage from the Tongfosi Formation of China: Comparison with the track assemblage of South Korea

A Yanji Basin tracksite in the Cretaceous Tongfosi Formation in the Yanbian Korean Autonomous Prefecture, Jilin Province was reinvestigated twenty years after the original study.The re-examination confirms the presence of tracks tentatively referred to ornithopods and theropod tracks including several not previously reported. In addition to the main track-bearing level, a second footprint horizon was identified from which a probable ornithopod track was collected. Some of the formerly identified tracks were documented using photogrammetry, whereas others that had been collected, eroded, buried or obscured by dense vegetation. The age of the track-bearing beds is inferred to be late Albian to early Cenomanian. Comparison with track-bearing units of the same age in Shandong, China, and Korea indicated quite different compositions of track assemblages. In the case of the Korean sites the differences may be facies related. While the Korean ichnofauna is characterized by the co-occurrence of ornithopod, sauropod and tridactyl theropod tracks with didactyl dromaeosaurid, Minisauripus (small theropod), pterosaur and avian tracks, the latter four are absent in the Tongfosi assemblage. This could reflect a restricted environment being not favourable for activities of these trackmakers or simply a preservational bias.     

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。     

Didactyl raptor tracks from the Cretaceous,Plainview Sandstone at Dinosaur Ridge

Two natural casts of two-toed (didactyl) tracks from the Cretaceous (Albian) Plainview Sandstone (Plainview Member) of the South Platte Formation (Dakota Group) at Dinosaur Ridge, Colorado are attributed to deinonychosaurian theropod dinosaurs and placed in the ichnogenus Dromaeosauripus. This is both the first report of tracks from this unit in the Dinosaur Ridge area and the first report of deinonychosaurian tracks from Colorado. It is also only the third report of this track type from North America. The rarity of tracks from the Albian-aged, Plainview Sandstone (Dakota Group Sequence 2) contrasts with their abundance in the upper (Cenomanian) part of the overlying South Platte Formation (Dakota Group Sequence 3), which has yielded more than 120 sites mostly in Colorado, giving rise to the “Dinosaur Freeway” concept. As no deinonychosaurid tracks are known from the sequence 3 part of the South Platte Formation, despite the large vertebrate and invertebrate ichnological database available, it is evident that the sparse vertebrate ichnofauna from the Plainview Member (Sequence 2) is inherently different. This striking difference in both track abundance and track type reflects differences in both age and depositional environment. Based on the Albian age, and track type, the Plainview tracks invite comparison with the ichnofaunas of the Cedar Mountain Formation and not with those well-known from the upper part of the South Platte Formation known as the Dinosaur Freeway.     

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。     

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。     

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。     

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.     

山东诸城早白垩世中期超大规模恐龙足迹群及其古地理与古生态*

山东诸城黄龙沟恐龙足迹产于下白垩统中部莱阳群杨家庄组,其中以兽脚类为主,同时有蜥脚类和疑似鸟脚类足迹;兽脚类足迹又可进一步区分为大型兽脚类、似跷脚龙类(Paragrallator）和东方强壮百合龙类(Corpulentapus lilasia）。足迹共组成至少63条行迹,作者系统实测了其中50条,并据足迹大小指数—复步长和GAE(跷脚龙—安琪龙—实雷龙）图解给出了识别3类兽脚类足迹的经验性数值。足迹点沉积特征显示,浅湖相粉砂质泥岩和粉砂岩逐渐过渡为滨湖相砂岩。足迹层及其邻近层位波痕、泥裂发育,根据波脊线恢复的古岸线方向为东西向,波浪运动方向呈现出北至北西周期性变化,表明可能存在季节性风向变化。泥裂以及恐龙分布特征显示足迹点以S/WS为湖心方向,以N/EN为湖岸方向。似跷脚龙类和东方强壮百合龙类足迹分布区域的差异性表明东方强壮百合龙类生活区域更靠近湖岸。恐龙足迹中存在的明显叠盖现象表明足迹不是同时,而是多次活动遗迹。小型兽脚类行迹中较多近于平行的行迹体现了集群性生活习性。行进行为研究又显示兽脚类的绝大多数为快速奔跑状态,但蜥脚类等为慢行状态。兽脚类与蜥脚类、疑似鸟脚类足迹数量之比远大于正常捕食者和被捕食者之比也表明,早白垩世中期的胶莱盆地生存着以兽脚类为主、蜥脚类和鸟脚类为辅的恐龙动物群。早白垩世晚期,胶莱盆地以及沂沭断裂带内蜥脚类、鸟脚类恐龙以及鸟类逐渐繁盛,这与华北北部恐龙足迹所恢复的造迹恐龙动物群特征较为一致。中国北方、韩国、日本白垩纪恐龙足迹以及骨骼化石发育情况暗示,晚中生代中国北方以及东亚地区发育一个以恐龙等脊椎动物为主的大型陆地动物群。     

山东诸城盆地下白垩统莱阳群的非鸟恐龙与鸟足迹化石组合

中国山东省诸城盆地张祝河湾村下白垩统杨家庄组发现了非鸟恐龙(蜥脚类和鸟脚类)与古鸟类足迹。该足迹点的非鸟恐龙足迹深受水浸沉积物的影响,发生了强烈变形。蜥脚类足迹的前后足迹面积相近,造迹者行进缓慢。鸟脚类足迹属于禽龙-鸭嘴龙类型的足迹。古鸟类足迹并不类似于中国以往发现的中生代鸟类足迹,而类似于韩国滨水鸟类足迹的金东鸟足迹(Jindongornipes)。讨论了莱阳足迹(Laiyangpus)和拟跷脚龙足迹(Paragrallator,亦译为拟似鹬龙足迹),前者因模式标本遗失而不明确,后者可归入安琪龙足迹未定种(Anchisauripus isp)。这批足迹的发现大大增加了山东省东部恐龙的多样性。     

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.     

山东诸城盆地下白垩统莱阳群的非鸟恐龙与鸟足迹化石组合（英文）

中国山东省诸城盆地张祝河湾村下白垩统杨家庄组发现了非鸟恐龙（蜥脚类和鸟脚类）与古鸟类足迹。该足迹点的非鸟恐龙足迹深受水浸沉积物的影响,发生了强烈变形。蜥脚类足迹的前后足迹面积相近,造迹者行进缓慢。鸟脚类足迹属于禽龙-鸭嘴龙类型的足迹。古鸟类足迹并不类似于中国以往发现的中生代鸟类足迹,而类似于韩国滨水鸟类足迹的金东鸟足迹（Jin-dongornipes）。讨论了莱阳足迹（Laiyangpus）和拟跷脚龙足迹（Paragrallator,亦译为拟似鹬龙足迹）,前者因模式标本遗失而不明确,后者可归入安琪龙足迹未定种（Anchisauripus isp）。这批足迹的发现大大增加了山东省东部恐龙的多样性。