First Discovery of Middle Jurassic Theropod Trackways in Changdu Region, Eastern Tibet of China
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摘要: 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.Abstract: A new dinosaur track site, Anietang site, was discovered in Changdu region of eastern Tibet. The lithostratigraphy of this track site is mainly lakeside purplish red medium-fine sandstone of the Middle Jurassic Dabuka Formation. Four exquisitely convex track layers (footprint types including Grallator, large theropods, small theropods and small sauropods) and two perfect track-ways were found. Four sequential Grallator footprints that make up the T1 trackway are named Grallator anietangensis isp nov., and a detailed morphological analysis was performed on it. It is doubtless that the Grallator was found in Tibet for the first time, and the size and gait of track-maker are discussed. Nine large theropod footprints that make up T2 trackway are classified as Theropoda igen et isp. indet.. The current study has described the morphology of Theropoda igen et isp. indet., and it is determined that this kind of footprint is found for the first time in China. The consecutive and clear trail tracks of T2 indicate that the trackmaker was not in a normal gait, but a special behavior to deal with a particular situation. It is speculated that the trackmaker was detecting situation with its raised body while walking at a very low speed. The forequarters were raised, resulted in the tail to sag correspondingly, and left trail tracks. Both trackways show that the trackmakers were walking along the bank of ancient Changdu Lake. At present, the dinosaur skeletons and the Grallator footprints found in China all show a migration pattern with the Southwest China as the center and then spreading around. But the speed of dinosaur migration reflected by the Grallator footprints is faster than that of skeletons.
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Key words:
- theropod track /
- dinosaur /
- Middle Jurassic /
- Dabuka Formation /
- Changdu region
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图 1 研究区及阿涅塘化石点地理位置图
a.研究区构造位置（KL=阿尼玛卿‒昆仑结合带，GL=甘孜‒理塘结合带，JS=金沙江结合带，AL=哀牢山结合带，NL=北澜沧江结合带，SL=南澜沧江结合带，LS=龙木错‒双湖结合带，CM=昌宁‒孟连结合带，BN=班公湖‒怒江结合带，YL=印度河‒雅鲁藏布江结合带）；b. 西藏地区行政区划图；c. 阿涅塘化石点位置卫星图；d. 研究区地质图. 数据来源：图a据安显银等（2021），图d据王全立等（2013），略修改
Fig. 1. Geographic location map of the study region and Anietang site
图 2 阿涅塘剖面岩性分布特征及足迹位置
a. 非对称浪成波痕；b. 砾屑沉积；c. 平行层理；d. 不均一的铁质砂岩沉积；e. 交错层理；f. 砂岩与泥岩互层状沉积（ANT1、ANT2、ANT3、ANT4分别标示各足迹层的位置；M=泥岩，S=砂岩，G=砾岩）
Fig. 2. Lithostratigraphy and footprint layer position of Anietang Section
图 3 阿涅塘剖面和ANT2、ANT3足迹层照片
a. 剖面照片（1=ANT1足迹层，2=ANT2足迹层，3=ANT3足迹层，4=ANT4足迹层）；b. ANT2和ANT3足迹层照片；c. ANT2和ANT3足迹层之间被清理掉的紫红色中层状中到细砂岩见发育良好的波痕；d. ANT2和ANT3足迹层素描图（PA=复步角，γ=行迹偏角，We=行迹外宽，PL=步长，SL=复步长；足迹编号简写：15=ANT3-15，16=ANT3-16，17=ANT3-17，18=ANT3-18，19=ANT3-19，20=ANT3-20，21=ANT3-21，22=ANT3-22，23=ANT3-23，25=ANT3-25，26=ANT3-26，27=ANT3-27，28=ANT3-28，24=ANT2-24）
Fig. 3. Photos of Anietang Section, ANT2 and ANT3 footprint layers
图 4 ANT3和ANT2足迹层的兽脚类足迹照片及素描图
a~d. Grallator anietangensis isp. nov.（足迹编号，a. ANT3-15；b. ANT3-16；c. ANT3-17；d. ANT3-18）；e~f. Theropoda igen. et isp. indet.（足迹编号，e. ANT3-23；f. ANT2-24）（FL=足迹长，FW=足迹宽，DL=趾长，DW=趾宽，Ⅱ=Ⅱ趾，Ⅲ=Ⅲ趾，Ⅳ=Ⅳ趾，α=Ⅱ趾和Ⅲ趾趾间角，β=Ⅲ趾和Ⅳ趾趾间角；X/Y=中趾并齐率，a/b=中趾前凸值）
Fig. 4. Photos and sketches of theropod footprints on ANT3 and ANT2 footprint layers
图 5 中国所发现的Grallator类足迹和典型的Grallator类足迹
a. Grallator anietangensis isp. nov.，本文；b. Grallator ssatoi，辽宁阳山晚侏罗世土城子组（Shikama，1942）；c. Grallator ssatoi，河北承德晚侏罗世土城子组（Young，1960）；d. Grallator类足迹，四川自贡早侏罗世自流井组（李日辉等，2021），本文素描；e. Grallator wuhuangensis，四川自贡中侏罗世新田沟组（Lockley et al., 2013）；f. Grallator wuhuangensis，四川自贡中侏罗世新田沟组（Lockley et al., 2013；Xing et al., 2017）；g. Grallator emeiensis，四川峨眉早白垩世夹关组（Zhen et al., 1994）；h. Grallator yangi，山东莱阳早白垩世龙旺庄组（Li et al., 2011）；i. Grallator yangi，山东莱阳早白垩世龙旺庄组（李日辉和张光威，2000；Li et al., 2011）；j. Grallator yangi，山东莱阳早白垩世水南组（李日辉等，2021），本文素描；k. Grallator yangi，山东诸城早白垩世杨家庄组（李日辉等，2021）；l. Grallator pingchuanensis，甘肃白银中侏罗世王家山组（Li et al., 2019），m. Grallator microiscus，四川资中中侏罗世新田沟组（杨兴隆和杨代环，1987；Lockley et al., 2013；Xing et al., 2017）；n. Grallator yemiaoxiensis，重庆南岸中侏罗世下沙溪庙组（杨兴隆和杨代环，1987；Lockley et al., 2013；Xing et al., 2017）；o. Grallator limnosus，云南晋宁早侏罗世冯家河组（Lockley et al., 2013；Xing et al., 2017）；p. cf. Grallator，四川喜德早白垩世小坝组（Xing et al., 2019；李日辉等，2021）；q. cf. Grallator，山东新泰中侏罗世三台组（李日辉等, 2002, 2021）；r. cf. Grallator，山东新泰中侏罗世三台组（李日辉等，2021）；s. Grallator，辽宁北票晚侏罗世土城子组（李日辉等，2021）；t. Grallator，辽宁北票晚侏罗世土城子组（李日辉等，2021）；u. Grallator，辽宁北票早白垩世义县组（Xing et al., 2009）；v. Grallator，云南双柏中‒晚侏罗世蛇店组（Xing et al., 2016b）；w. Grallator，内蒙古鄂托克旗早白垩世泾川组（Li et al., 2009；李建军等，2009；李日辉等，2021）；x. Grallator？，陕西中鸡早白垩世洛河组（李日辉等，2021），Xing et al.（2018）将其归为Eubrontes更合理；y. Grallator？，甘肃临洮早白垩世河口组（Xing et al., 2014a）；z. Grallator，四川古蔺早侏罗世自流井组（Xing et al., 2017）；aa. Grallator，辽宁阳山晚侏罗世土城子组（李日辉等，2021）；ab. Grallator，辽宁北票早白垩世义县组（Xing et al., 2009）；ac. Grallator，西班牙早侏罗世（Castanera et al., 2016）；ad. Grallator parallelus（Olsen et al., 1998）
Fig. 5. Sketches of typical Grallator found in Jurassic and abundant Grallator found in China
图 6 中国各足迹点Grallator类足迹测量数据平均值综合统计结果
a. 各足迹点Grallator类足迹FL与FW平均值散点图（西藏地区的数据由此次调查所获取，Anchisauripus类足迹的数据来自于李日辉等（2021），该足迹被鉴定为Grallator类足迹）；b. Grallator类足迹FL/FW值不同区间足迹点数量；c. Grallator类足迹α+β值不同区间足迹点数量. 数据来源：附表2
Fig. 6. Comprehensive statistical results of Grallator average measurement data of all footprint sites in China
图 7 ANT3兽脚类足迹造迹情形模拟
吉姣（E-mail：1182748971@qq.com），北海艺术设计学院（广西北海536000）；邵迪（E-mail：785142043@qq.com），贵阳建筑勘察设计有限公司（贵州贵阳550081）
Fig. 7. Taphonomic scenario for the footprint preservation of ANT3
图 8 中国恐龙骨骼化石点和Grallator类足迹化石点分布图
数据来源：骨骼化石点分布图方晓思等（2008），略修改
Fig. 8. Distribution map of dinosaur skeleton and Grallator footprint sites in China
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