An Unusual Sauropod Turning Trackway from the Early Cretaceous of Shandong Province, China

Palaeogeography, Palaeoclimatology, Palaeoecology 437 (2015) 74–84 Contents lists available at ScienceDirect Palaeogeography, Palaeoclimatology, Palaeoecology journal homepage: www.elsevier.com/locate/palaeo An unusual sauropod turning trackway from the Early Cretaceous of Shandong Province, China Lida Xing a,⁎,DanielMartyb,KebaiWangc, Martin G. Lockley d, Shuqing Chen c, Xing Xu e,YongqingLiuf, Hongwei Kuang f, Jianping Zhang a, Hao Ran g, W. Scott Persons IV h a School of the Earth Sciences and Resources, China University of Geosciences, Beijing 100083, China b Naturhistorisches Museum Basel, Augustinergasse 2, CH-4001 Basel, Switzerland c Dinosaur Research Center of Zhucheng, Zhucheng 262200, China d Dinosaur Trackers Research Group, University of Colorado Denver, P.O. Box 173364, Denver, CO 80217, USA e Key Laboratory of Evolutionary Systematics of Vertebrates, Chinese Academy of Sciences, Beijing 100044, China f Institute of Geology, Chinese Academy of Geological Science, Beijing 100037, China g Key Laboratory of Ecology of Rare and Endangered Species and Environmental Protection, Ministry of Education, Guilin 541004, China h Department of Biological Sciences, University of Alberta, 11455 Saskatchewan Drive, Edmonton, Alberta T6G 2E9, Canada article info abstract Article history: An unusual turning and a regular, straight sauropod trackway both left by small trackmakers, as well as additional Received 21 April 2015 isolated medium-sized to large sauropod tracks are described from the Early Cretaceous Dasheng Group of the Received in revised form 14 July 2015 Zhucheng Basin, Shandong Province, China. Based mainly on well-preserved tracks exhibiting three forwardly- Accepted 21 July 2015 directed digit/claw impressions and a pronounced heteropody, and to a lesser degree due to a predominantly Available online 31 July 2015 narrow to medium trackway gauge, the two small trackways are assigned to the Parabrontopodus ichnotaxon. As there is no clear trackway and no well-preserved tracks amongst the medium-sized to large sauropod tracks, Keywords: fi Dinosaur track these tracks can only be identi ed as of sauropod origin but they cannot be assigned to an ichnotaxon. The Sauropoda unusual turning trackway is characterized by a highly variable trackway configuration (pes and manus outward Turning trackway rotation, gauge, pace, stride) and pattern (different degree of manus overprinting by pes tracks) along its course, Trackway configuration & pattern evidently related to the narrow, semicircular turn to the left that the animal made. This is also associated with a Locomotion pronounced change from a narrow–medium (in the straight part at the beginning) to a (very) wide (within the Gait turn) gauge. This demonstrates that these two stances could have been used by one and the same sauropod Parabrontopodus trackmaker, even if in the present case associated with turning and not simply during straight progression, as it was already reported from a Late Jurassic tracksite from NW Switzerland and an Early Cretaceous tracksite from Spain. Such ‘untypical’ trackways provide important constraints for the reconstruction of locomotor charac- teristics of sauropods such as unsteady locomotion and changes in locomotor behavior, and they will be of particular interest in the future to model and understand the different ‘locomotor styles/capabilities’ sauropods were engaged in. © 2015 Elsevier B.V. All rights reserved. 1. Introduction (theropod, sauropod, and turtle tracks, Li et al., 2011; Lockley et al., 2015) from the Laiyang Group. At the Wenxiyuan tracksite (also from Shandong Province is one of the most important Mesozoic verte- the Laiyang Group), pterosaur tracks are preserved (Xing et al., 2012). brate fossil producing areas in China. Body fossils (Hu, 1973), tracks The tracksites of the Dasheng Group are primarily found along the (Young, 1960), and eggs (Chow, 1951) of dinosaurs are all known Yishu Fault zone, which extends (north–northeast) from northwest from Shandong's Mesozoic strata for a long time. The dinosaur track re- Jiangsu Province into central Shandong Province (Zhucheng–Junan– cord includes tracks from the Upper Jurassic Santai Formation (Li et al., Linshu–Tancheng). The so far described tracksites of the Dasheng 2002), the Lower Cretaceous Laiyang Group (~130–120 Ma) and the Group (see Xing et al., 2015a: Fig. 1) include the Zhangzhuhewan upper Lower Cretaceous Dasheng Group (~110–100 Ma) (Kuang et al., (Xing et al., 2010a) and Tangligezhuang tracksites (Wang et al., 2013). The largest known tracksite is the Huanglonggou tracksite 2013a) of Zhucheng; the Houzuoshan Dinosaur Park tracksite (Li et al., 2005a, 2005b, 2008; Lockley et al., 2007, 2008) of Junan; the Jishan tracksites I–VIII (Xing et al., 2013) of Linshu; the Beilin (Wang et al., ⁎ Corresponding author. Tel./fax: +86 1 82321827. 2013b; Xing et al., 2015a) and Qingquan tracksites (Xing et al., in E-mail address: [email protected] (L. Xing). press) of Tancheng; and the Nanguzhai tracksite of Donghai (Xing http://dx.doi.org/10.1016/j.palaeo.2015.07.036 0031-0182/© 2015 Elsevier B.V. All rights reserved. L. Xing et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 437 (2015) 74–84 75 Fig. 1. Geographic position (A) and stratigraphic sections (B, Modified from Wang et al., 2013a) of the Tangdigezhuang dinosaur tracksite. et al., 2010b). Except for the Houzuoshan Dinosaur Park tracksite, all T = track (single footprint); TDGZ = Tangdigezhuang tracksite, these tracksites contain sauropod tracks, most of which were identified Shandong Province, China as small Parabrontopodus and large Brontopodus tracks (Xing et al., 2013). The largest trackway with a pes length of 84 cm is found at the Qingquan tracksite within the Dasheng Group, it is of the 3. Geological setting Parabrontopodus-/Breviparopus-type (Xing et al., in press). Xing et al. (2010a) described the Zhangzhuhewan tracksite, which is Yanlu Fault is a large regional fault and significant geological charac- close to the Tangdigezhuang tracksite, and reported tracks of sauropod, teristic of Eastern China. The portion of the Yanlu Fault within Shandong birds, and possibly ornithopods. Initially, the Zhangzhuhewan tracksite Province is called Yishu Fault zone, spanning 300 km in length and 20 to was thought to belong to the Laiyang Group. However, the latest geolog- 60 km in width and consisting of four major, parallel faults all with ical survey showed that it to belongs to the Tianjialou Formation of the approximately the same direction. The strata exposed in the Yishu Dasheng Group (Kuang et al., 2013), and therefore has approximately Fault zone is divided into the Lower Cretaceous Laiyang, Qingshan and the same age as the Tangdigezhuang tracksite. Dasheng groups, and the Upper Cretaceous Wangshi Group (T'an, Since 2000, Chen S. Q. from the Dinosaur Research Center of 1923). Liu et al. (2003) divided the Dasheng Group into the Malanggou, Zhucheng has discovered several new tracksites (Wang et al., 2013a), Tianjialou, Siqiancun, and Mengtuan formations. However, Kuang et al. including the Tangdigezhuang tracksite, but lacking any detailed de- (2013) recently argued that the Malanggou and Siqiancun formations, scriptions. Wang et al. (2013a) briefly described the Tangdigezhuang and the Tianjialou and Mengtuan formations, respectively, are probably tracksite, but did not provide a morphological discussion. The first contemporaneous but developed in different facies. The Tianjialou/ author inspected the Tangdigezhuang tracksite during 2013 and 2014, Mengtuan formations are composed of lacustrine facies deposits, and here a detailed description, notably of the remarkable turning dominated by detrital rocks, and with a thickness exceeding 500 m. sauropod trackway, is provided for the first time. The Tangdigezhuang tracksite, situated 2 km to the north of the Zhangzhuhewan tracksite (Xing et al., 2010a)(Fig. 1A), belongs to Early Cretaceous Tianjialou Formation of the Dasheng Group, and is 2. Institutional and other abbreviations slightly younger than the strata exposed in the Zhangzhuhewan area. At the Tangdigezhuang tracksite, yellow–green and grayish green I = Isolated tracks; L and R = left and right; NGZ = Nanguzhai siltstones, fine sandstones and pelitic siltstones are exposed. These tracksite, Jiangsu Province, China; P = pes impression; S = Sauropoda; frequently exhibit small climbing ripple bedding, cross-bedding and 76 L. Xing et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 437 (2015) 74–84 parallel bedding, indicating that these are shallow lacustrine deposits be narrow-gauge (see also Farlow, 1992). Accordingly, a value of 1.0 (Wang et al., 2013a). separates narrow-gauge from medium-gauge trackways, whereas the value 1.2 is arbitrarily fixed between medium-gauge and wide-gauge 4. Material and methods trackways, and trackways with a value higher than 2.0 are considered to be very wide gauge (Marty, 2008). In 2014, due to the pronounced inclination (~45°) of the bedding Terminology for trackway patterns (e.g., quadrupedal, pes/ planes at the Tangdigezhuang tracksite, the site was studied using a manus-dominated) is used according to Marty et al. (2006) and system of safety ropes, which were set up by technicians from Dinosaur Marty (2008, fig. 2.16 on p. 39). Sauropod size classes are used ac- Research Center of Zhucheng. cording to Marty (2008, table 2.2. on p. 40):tiny:P′L b 25 cm; Wang et al. (2013a) have measured the geologic sections in small: 25 cm b P′L b 50 cm; medium-sized: 50 b P′Lcmb

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