A Lower Cretaceous dinosaur track assemblage from the Taoqihe Formation in Central Heilongjiang, China

Palaeogeography, Palaeoclimatology, Palaeoecology 532 (2019) 109275

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A Lower Cretaceous dinosaur track assemblage from the Taoqihe Formation in Central Heilongjiang, China

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Lida Xinga,b,c, , Martin G. Lockleyd, Hendrik Kleine, Liang Qiub, Chunyong Choub, Donghao Wangb, W. Scott Persons IVf a

State Key Laboratory of Biogeology and Environmental Geology, China University of Geosciences, Beijing 100083, China School of the Earth Sciences and Resources, China University of Geosciences, Beijing 100083, China c State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China d Dinosaur Tracks Museum, University of Colorado Denver, PO Box 173364, Denver, CO 80217, USA e Saurierwelt Paläontologisches Museum, Alte Richt 7, D-92318 Neumarkt, Germany f Department of Biological Sciences, University of Alberta, 11455 Saskatchewan Drive, Edmonton, Alberta T6G 2E9, Canada b

A R T I C LE I N FO

A B S T R A C T

Keywords: Ankylosaur Sauropod Theropod Ornithopod Swim tracks

An Early Cretaceous dinosaur ichnofauna is reported from the Taoqihe Formation (Aptian–Albian) in Heilongjiang Province, China. The assemblage, which comprises > 70 tracks in 5 trackways, includes parallel sauropod trackways, a toe-trace dominated tridactyl ornithopod trackway and elongate tridactyl theropod tracks, the latter tentatively be attributed to swimming activity. The combination of possible swim and walking trackways suggest that they were formed at different times and in different environments. This is the first Lower Cretaceous tracksite reported from Heilongjiang Province with the exception of an historic record of a large pentadactyl track of probable thyreophoran (ankylosaurian) affinity. Ankylosaurian tracks are rare in China, and reported tentatively from only one other location. The dinosaur ichnofauna from the Taoqihe Formation is similar in composition to the skeletal assemblages of the Early Cretaceous Jehol Biota in the nearby Liaoning Province. The new track records are important for better understanding the palaeobiogeography and faunal developments of northeastern China and adjacent Russia.

1. Introduction The Jehol Biota (Barremian-Aptian) constitutes the most famous Early Cretaceous dinosaur record in Northeast China, with a majority of its specimens being discovered in the western part of Liaoning, the western part of Inner Mongolia, and the northern part of Hebei. The dinosaur components of the fauna include sauropods, theropods/birds, ornithopods, ceratopsids, and ankylosaurids (Zhou and Wang, 2010). Jilin Province, which borders Liaoning to the north and Heilongjiang to the south, has a limited dinosaur record. The ornithischian Changchunsaurus (Zan et al., 2005), the ceratopsian Helioceratops (Jin et al., 2009), and the titanosaur Jiutaisaurus (Wu et al., 2006) have been found in the mid-Cretaceous Quantou Formation (primarily Albian), in central Jilin and are the only dinosaur fossils. In contrast, Early Cretaceous dinosaurs are almost nowhere to be found in Heilongjiang Province, the northernmost part of Northeast China, although other Early Cretaceous vertebrate remains have been unearthed, including the soft-shelled turtle “Trionyx” jixiensis from the

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Lower Cretaceous Chengzihe Formation, in the Jixi area (Li et al., 2015). Possible dinosaur and fish fossils have been reported from this area as well (Zhao et al., 2005), but without detailed description. However, Heilongjiang Province has yielded a well-known Late Cretaceous dinosaur fauna. So far, most Late Cretaceous dinosaur fossils in this area have been discovered from the Yuliangzi Formation (lowermiddle Maastrichtian) of the Jiayin area. These include the Jiayin and Wulaga bonebeds, both located on the Chinese side of the Heilongjiang River (Amur River) (Riabinin, 1930). The adjacent Blagoveschensk and Kundur bonebeds are located in the Udurchukan Formation, on the north (Russian) side of the Heilongjiang River (Riabinin, 1930). These bonebeds are dominated by hadrosaurs (Godefroit et al., 2008). Until recently, Heilongjiang Province was thought to be devoid of dinosaur tracks (Zhen et al., 1996), until hadrosaur tracks named Jiayinosauropus johnsoni (Dong et al., 2003; Xing et al., 2009) were reported from the Upper Cretaceous Yong'ancun Formation in the Jiayin area (Xing et al., 2016a). However, when sorting through old literature, the senior author (LX) found a significant report of a Lower

Corresponding author at: State Key Laboratory of Biogeology and Environmental Geology, China University of Geosciences, Beijing 100083, China. E-mail address: [email protected] (L. Xing).

https://doi.org/10.1016/j.palaeo.2019.109275 Received 20 April 2019; Received in revised form 14 July 2019; Accepted 14 July 2019 Available online 17 July 2019 0031-0182/ © 2019 Elsevier B.V. All rights reserved.

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Fig. 1. Geographic map indicating the location of the Jiayin, Yilan and Guanghui dinosaur tracksite localities in Yilan County, Heilongjiang Province, China.

mudstone and siltstone, interbedded with medium-grained and finegrained sandstone. The middle member is dominated by coarse- to medium- grained sandstone, interlayered with siltstone and mudstone. The lower member is dominated by glutenite, interbedded with sandstone. The Guanghui tracksite is located in the upper member of the Taoqihe Formation (BGMRH, 1972). Based on the biostratigraphic evidence, such as spore-pollen record (e.g. Appendicisporites jansonii), Tian (2008) suggested the lower member of Taoqihe Formation is Hauterivian–Barremian, the upper member of Taoqihe Formation is late Early Cretaceous Aptian–Albian. The well-exposed outcrop hosting the Guanghui tracksite is ~200 m-long and ~10 m high, situated along a paved road. The tilted bedding dips to southwest at an dip angle of ~30°. The outcrop, from the bottom to top, is dominated by coarse- to medium-grained lithic sandstone (thicker than 30 cm), siltstone (~20–30 cm), and mudstone (~10 cm), indicating a fining- upwards sequence. The various clasts show poor sorting and angular to subangular sphericity, suggesting a proximal source for these materials. The horizontal lamination is well developed and contains freshwater bivalve fossils (Trigonioides, Plicatounio and Cyrena), suggesting a lacustrine facies, such as a lake in a terrestrial intermontane freshwater basin (BGMRH, 1972). In addition, there are abundant plant fossils, most of which are ferns and ginkgos, with minor conifers and bennettites, which represent a temperate or tropical flora (BGMRH, 1972).

Cretaceous dinosaur track from Yilan County by Binglin Chen, from Daqing Petroleum Institute, in the Journal of Daqing Petroleum Institute (Chen, 1984). Though the paper is only one page, with only one photo and corresponding outline illustration, without English abstract, it represents the first documentation and earliest geological record of a dinosaur track in Heilongjiang. The Yilan County tracksite was probably destroyed by road reconstruction. In August 2018, amber hunter Zhu Li went to Heilongjiang and informed the lead author (LX) of another dinosaur track site (Fig. 1)., which forms the focus of this paper.

2. Geological setting The eastern North China Craton underwent a large-scale lithospheric thinning during the Early Cretaceous as a consequence of the roll back of the subducted Paleo-pacific plate (e.g., Qiu et al., 2018; Wu et al., 2019). A series of metamorphic core complexes and grabens or supradetachment basins developed during the lithospheric thinning and crustal extension (e.g., Liu et al., 2013). The recently discovered tracksites in this study come from one of these basins that were filled with Early Cretaceous continental clastic rocks. The Guanghui tracksite is located in an Early Cretaceous basin in the southern margin of Yilan County (GPS: 46°12′13.36”N, 129°34′10.12″E) (Fig. 2). The Cretaceous strata of the Yilan region belong to the Lower Cretaceous Taoqihe Formation and Upper Cretaceous Songmuhe Formation (Yilan Geological map L-52-16, scale 1: 20, 000; BGMRH, 1972; Fig. 2). The Taoqihe Formation in the Yilan area unconformably overlies Jurassic biotite granite and plagiogranite, and is overlain by the Upper Cretaceous Songmuhe Formation that consists of albitophyre and intermediate volcanic rocks. The Taoqihe Formation in the Yilan area is an ~860 m thick continental clastic deposit composed of glutenite, sandstone, and mudstone. It can be divided into three lithologic members (Fig. 2). The upper member is mainly

3. Materials and methods No new tracks were found at the Yilan tracksite, which we infer to have been lost to road construction since it was reported (Chen, 1984). However, 11 km south of the Yilan tracksite, is the Guanghui tracksite, which may have been first exposed during road construction, about 20 years ago. The track-bearing surface has a dip of about 45–50°, and 2

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Fig. 2. Stratigraphic sections of Guanghui dinosaur tracksite. The wavy lines indicated unconformity.

pace length (PL), stride length (SL), pace angulation (PA) and rotation (R) of the tracks were measured. The methods of Olsen (1980), Weems (1992), and Lockley (2009) were used to measure the mesaxony of tridactyl tracks. Mesaxony refers to the degree to which the central digit (III) protrudes anteriorly beyond the medial (II) and lateral (IV) digits. Using the ratio between the width of the angulation pattern of the pes (WAP) and the pes length (PL), gauge (trackway width) was quantified for pes and manus tracks in the trackways of quadrupeds (Marty, 2008; Marty et al., 2010). Gauge was calculated with pace and stride length, assuming that the width of the trackway can be measured perpendicular to stride at the approximate midpoint of the stride (Marty, 2008). When the (WAP/P'ML)-ratio is 1.0, the inner margin of the pes tracks may register on the trackway midline. When the ratio is smaller, the tracks would cross the trackway midline: i.e., registering a narrowgauge configuration (see Farlow, 1992). Accordingly, a ratio of 1.0 distinguishes narrow-gauge from medium-gauge trackways, whereas the value 1.2 arbitrarily divides medium-gauge and wide-gauge

overlying layers that consist of alternations of siltstone and mudstone. This rock surface is about 200 m2 in extent and contains three sauropod trackways, one ornithopod trackway, and two theropod trackways. The tracks usually penetrate the yellow siltstone on the surface to expose underlying black mudstone. Despite a certain degree of weathering, the track outlines are still clearly identifiable, and repeat consistently in trackway sequences. Due to the steepness of the bedding planes at the Guanghui tracksite, it was necessary to use safety ropes during the study of the trackbearing surfaces. In order to make accurate maps, tracks were photographed, outlined with chalk, and traced on large sheets of transparent plastic. Videotaping was employed to convert the full-size tracing maps into a digital format. On a lift truck, provided by the City Highway Bureau, the senior author photographed the track-bearing surface and trackway close-ups from 10 m above the ground. According to the standard methods of Leonardi (1987) and Lockley and Hunt (1995a), the maximum length (ML), maximum width (MW), 3

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Fig. 3. Photograph (A) and interpretative outline drawing (B) by Chen (1984), and (C,) an interpretative sketch by the authors, based on the original photo, of possible Tetrapodosaurus track from the Yilan dinosaur tracksite locality in Yilan County, Heilongjiang Province, China.

Fig. 4. Photograph (A) and interpretative outline drawing (B) with overview of dinosaur track surface at the Guanghui dinosaur tracksite locality in Yilan County, Heilongjiang Province, China. The tracksite surface include three sauropod trackways (GH-S1–S3), one ornithopod trackway (GH-O1), and two theropod (? swim) trackways (GH-T1 and T2).

4.1. Material and description

trackways, and trackways with a value higher than 2.0 are considered very wide-gauge (Marty, 2008). To calculate hip heights and speed estimates of the sauropod trackways, the methods of Alexander (1976), Thulborn (1990) and González Riga (2011) were adopted. The following abbreviations are used in the description: GH, Guanghui tracksite, Heilongjiang Province, China. JDGP, Jiayin Dinosaur National Geological Park, Heilongjiang Province, China. YL = Yilan tracksite, Heilongjiang Province, China.

The track was never catalogued or collected, but its measurements were given. The track was 40 cm long and 49 cm wide. Digits IeV were 18, 34, 32, 40, and 32 cm long. The tip distances were 18, 13, 18 and 16.5 cm, and the angulations were 40°, 65°, 50° and 40°. The center of the track was reported as 4.5 cm deep, and digit III as 6 cm deep (Chen, 1984). The track was named as “Mudanjiangpus yilanensis”, but is here considered a probable synonym of Tetrapodosaurus (see below). The photographs provided by Chen (1984) are black and white (Fig. 3A), and the outline (Fig. 3B) appears distorted. The author redrew the outline based on the photograph (Fig. 3C). We catalogued this specimen as YL-T1 for comparison. Morphologically, the impression is pentadactyl and fits the outline of a rough pentagon; it is wider than long, being 49 cm and 40 cm, respectively (ML/MW = 0.82). The left digits seems to be more developed than the right digits. All digits are relatively short and blunt. The digit total divarication is 110° (digits IeV).

4. The Yilan tracksite The Yilan tracksite yielded only one isolated track, which was discovered on August 4, 1983 by Binglin Chen, Qipeng Che, Xinhua Bai and Shangming Shi (Fig. 3A-C). The Yilan tracksite is located along the west bank of the Mudanjiang River, just outside the south gate of Yilan County.

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The YL-T1 track is morphologically consistent with the manus skeletons of some thyreophorans (e.g., Apesteguía and Gallina, 2011). Pentadactyl manus tracks with terminally round and blunt digits are present in the ichnogenera Tetrapodosaurus (Sternberg, 1932; McCrea et al., 2001; Lockley et al., 2014) and Ceratopsipes (Lockley and Hunt, 1995b). Ceratopsipes has been interpreted as a ceratopsian track (Lockley and Hunt, 1995b), and Tetrapodosaurus as an ankylosaurian track due to their morphologies being similar to the pedal osteology of these respective groups (McCrea et al., 2001). Morphologically, YL-T1 is quite similar to manus impressions of Tetrapodosaurus type, except that the heel trace appears posteriorly convex rather than concave as in type Tetrapodosaurus. However, with only one track for comparison the significance of this difference is uncertain. YL-T1 is also different from the manus impressions of Ceratopsipes, whose digit I and V show proximolateral and proximomedial orientations (Lockley and Hunt, 1995b), as well as a concave posterior margin. In addition to this pattern, some specimens of Tetrapodosaurus type show anteromedial and posterolateral orientations of digit I and V respectively (e.g., Apesteguía and Gallina, 2011). Xing et al. (2016b) described an isolated occurrence of Tetrapodosaurus isp. from the Early Cretaceous Jiaguan Formation, Sichuan Province, and suggested a possible ankylosaurian identity, which is the first Tetrapodosaurus record in China. However, the specimens are lost with only one incomplete pes track left. The Jiaguan Tetrapodosaurus isp only has pes impressions, and thus cannot be compared to YL-T1. The only abundant and well-documented thyreoporan tracks so far reported from the Lower Cretaceous of China are Deltapodus from Xinjiang Province (Xing et al., 2013a) which have been attributed to stegosaurians, not ankylosaurs. The Xinjiang trackmakers registered blunt manus digit traces similar to those described here and illustrated from Yilan (Fig. 3), however, in the former the manus is rather kidney-shaped, with short digits, whereas in the latter it is fan-shaped with longer digits. 5. The Guanghui tracksite 5.1. Theropod swim tracks 5.1.1. Materials and description Five complete natural molds of pes prints were catalogued as GHT1-R1–L1, and GH-T2-L1–L2, representing two trackways (Figs. 46, Table 1). GH-T1 is composed of two tracks, and GH-T2 is composed of three tracks. GH-T1 and GH-T2 may be from the same trackway, with the space between lacking registration of comparable traces. Both have a similar orientation (directed west and west-northwest) along similar directions of progression, and all the individual prints are roughly the same size. The GH-T1 and GH-T2 tracks consist of long slender, parallel and tapering digit impressions, and lack any impressions made by the metatarsophalangeal regions. The absence of metatarsophalangeal region impressions is common in swim tracks, interpreted as scratch marks made on the sediment by the distal ends (claws or toe tips) of the trackmaker's hindfeet (Coombs, 1980; Milner et al., 2006; Ezquerra et al., 2007; Xing et al., 2013b, 2016c). The digit III marks are longer and deeper, while the digits II and IV marks are shorter and shallower. The digit II marks are always shorter and deeper than the digit IV marks. There is a bivalve cast medial to the trace of digit IV of GH-T2R1. We note that there are no elongated sand mounds at the posterior ends of the GH-T1 and GH-T2 tracks (Fig. 6). Such sand mounds are a common feature of swim tracks (Swanson and Carlson, 2002), including those of theropods (such as Ezquerra et al., 2007; Xing et al., 2013b, 2016c) showing that substrate sediments were raked by the digits and pulled posteriorly. Therefore we cannot exclude that the absence of this feature in GH-T1 and GH-T2 could be the result of later weathering or the presence of undertracks, or penetrative tracks registered on a sediment layer below the surface the trackmaker was originally moving

Fig. 5. Interpretative outline drawing of theropod (? swim) trackways (GH-T1, and T2, R = right pes, L = left pes) from the Guanghui site (A) and one theropod swim trackway from the Zhaojue site, Sichuan Province (B) for comparison (Xing et al., 2013b).

4.2. Comparisons and discussion YL-T1 is most likely referable to a thyreophoran (possibly an ankylosaurian). Such tracks are common in the earliest Cretaceous to early Late Cretaceous of Canada (McCrea et al., 2001, 2014) and the western USA (Lockley et al., 2014) and are also known from Europe (Nopcsa, 1923; Ensom, 1987; Petti et al., 2008; Sacchi et al., 2009; Hornung and Reich, 2014; Shillito and Davies, 2019) and Australia (Lockley et al., 2012) but are rarely reported from Asia (Fujita et al., 2003). 5

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Fig. 6. Photograph (A, C) and interpretative outline drawing (B, D) of well-preserved theropod (? swim) tracks from the Guanghui site. The arrow indicated a bivalve near the theropod digit traces.

tracks from the Upper Jurassic-Lower Cretaceous Tuchengzi Formation of Chicheng County, Hebei Province, China. In overall morphology, these tracks are similar to Characichnos, but the tracks were isolated and poorly preserved. Theropod swim tracks from the Zhaojue site of Sichuan Province described by Xing et al. (2013b) (Fig. 5B) are the most definitive Characichnos record in China. Similar, is also a theropod swim track report from the Jurassic-Cretaceous boundary, Anning Formation, Konglongshan, Yunnan Province, but it is only a single isolated track (Xing et al., 2016c).

Table. 1 Measurements (in cm) of theropod and ornithopod tracks from Guanghui tracksite, Heilongjiang Province, China. Number

ML

MW

PL

SL

PA

ML/MW

M

GH-T1-R1 GH-T1-L1 Mean

21.4 16.5 18.9

18.2 – 18.2

205.1 – 205.1

– – –

– – –

1.2 – 1.2

– – –

GH-T2-L1 GH-T2-R1 GH-T2-L2 Mean

16.4 25.6 16.7 19.6

– 15.4 – 15.4

164.3 107.7 – 136.0

271.5 – – 271.5

173 – – 173.0

– 1.7 – 1.7

– – – –

GH-O1-R1 GH-O1-L1 GH-O1-R2 GH-O1-L2 Mean

12.0 14.2 19.6 12.9 14.7

33.8 32.4 33.5 27.4 31.8

122.2 134.0 122.0 – 126.1

221.0 220.3 – – 220.7

119 119 – – 119

0.4 0.4 0.6 0.5 0.5

0.27 0.35 0.47 0.34 0.36

5.2. Sauropod tracks 5.2.1. Materials and description The Guanghui Tracksite preserves three large trackways: GH-S1–S3 (Figs. 4, 79, Table 2, Supplementary material). All the tracks remain in situ. Trackway GH-S1 is a typical trackway of a quadruped with 12 pes impressions and 11 corresponding manus impressions (Fig. 7). The pes tracks are 48.4 cm long on average. The manus tracks are 13.2 cm long on average. The average length/width ratios of the manus and pes impressions are 0.5 and 1.3 respectively. The manus impressions of GHS1 lie slightly anteromedially to the pes impressions. In the best-preserved manus-pes association: RP3-RM3, the manus imprints show oval digit impressions, while the digit trace area and the metacarpophalangeal region are distinct and the pes impression is oval, the trace of digit I is extremely anteriorly oriented, digit II–IV traces have recognizable claw marks, and the metatarsophalangeal region is smoothly curved. The manus impression is rotated approximately 26° outward/ inward from the trackway axis, which is more than the outward rotation of the pes impressions (approximately 20°). The average manus PA is 110°, while the average pes PA is 97°. There are three unusual aspects to trackway GH-S1:1) An extremely antero-medially located digit I trace occurs in all right pes impressions, but only occasionally in left tracks (such as LP3 and LP4). This is consistent with the fact that (2) the pace length (PL) of the right tracks are significantly larger than that of the left track (153.3 cm > 116.9 cm). This may indicate limping, or an unusual gait. 3) GH-S1-LP2 appears to have two corresponding manus impressions, here labelled as LM2a and LM2b, which are basically the same in shape and area and may be the result of repeat stepping by the track maker. This is consistent with the fact that single step of LP2 (i.e. LP2-RP2 = 125.9 cm) is slightly longer than the preceding pace (LP1-RP1 = 112.5 cm) and following pace (LP3- RP3 = 116.5 cm). The LP2-LP3 stride (244.5 cm) is also the longest in the trackway (Table 2).

Abbreviations: ML: Maximum length; MW: Maximum width (measured as the distance between the tips of digits II and IV); PL: Pace length; SL: Stride length; PA: Pace angulation; ML/MW is dimensionless; M mesaxony (length/width ratio for the anterior triangle).

on. The formation of undertracks has been analyzed in detail for example in Gatesy and Falkingham (2017) and Marchetti et al. (2019).

5.1.2. Comparisons and discussion The ichnotaxon Characichnos, from the Middle Jurassic Saltwick Formation of England, represents dinosaur swim tracks of likely theropod affinities (Whyte and Romano, 2001). Characichnos tracks attributed to theropod producers have also been identified in the Moenave Formation, at the St. George Dinosaur Discovery Site at Johnson Farms in southwestern Utah, USA (Milner et al., 2006). Tridactyl Characichnos is clearly distinguished from the often tetradactyl morphologies of Hatcherichnus, which is attributed to crocodylomorphs or other tetrapods (Lockley et al., 2010). [Some Hatcherichnus maybe incomplete: i.e., tridactyl, but most are preserved as natural casts unlike the natural Characichnos impressions from England and those described here]. The characteristics of the GH-T1 and GH-T2 tracks are consistent with Characichnos in having three elongate and parallel epichnial grooves, the terminations of which are straight or sharply reflexed (Whyte and Romano, 2001; Figs. 5A, 6). Therefore, GH-T1 and GH-T2 tracks can assigned to Characichnos. Previously, Xing et al. (2011) reported five possible theropod swim 6

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suggesting acceleration. The pes impressions are rotated approximately 18° outward from the trackway axis. The average pes PA is 103°. Trackway GH-S3 has 15 pes impressions, without any manus impression (Figs. 4, 8–9). The pes impressions are 29.7 cm long on average, with an average length/width ratio of 0.8. Among all the pesonly tracks, the best preserved are GH-S3-LP2, LP3, RP3, and RP5 (Fig. 9). In GH-S3-LP2, for example, digit I, II, III and IV have recognizable claw marks. The digit III impression is the most developed (deepest) and is oval in shape (length greater than width). The impressions of digit II and IV are also oval in shape, slightly larger than the digit I impression and runs deep into the sediment proximolaterally. Digit I impression is relatively shallow. The left pes impression is rotated approximately 11° inward from the trackway axis, which is a little different from the outward rotation of the right pes impressions (approximately 12°). Some tracks only show three digits traces, such as LP3 (Fig. 9). Of note, the width of the right tracks are 40.6 cm, larger than the 35.3 cm widths of the left tracks. Digit I traces in the right pes impression are more developed than the left counterparts. Lingulate sand mounds are preserved at the posterior end of the almost all the pes traces, showing that the substrate was pushed up posteriorly behind the deeply impressed digits (Fig. 9). 5.2.2. Comparisons and discussion The pes and manus morphology and trackway configuration of the Guanghui large quadruped trackways is typical of sauropods (Lockley, 1999, 2001; Lockley and Hunt, 1995a). Most sauropod trackways in China are wide- (or medium-) gauge and are therefore referred to the ichnogenus Brontopodus (Lockley et al., 2002). The Guanghui sauropod trackways are between medium-gauge and wide-gauge trackways, with a WAP/P'ML ratio of 1.8, 2.5 and 3.8 (Marty, 2008). The Guanghui sauropod trackway configurations are also consistent with the characteristics of Brontopodus type tracks from the Lower Cretaceous of the USA (Farlow et al., 1989; Lockley et al., 1994a) and the Upper Jurassic of Portugal and Switzerland (Meyer and Pittman, 1994; Santos et al., 2009). These characteristics include 1) wide-gauge; 2) pes tracks that are longer than wide, and large and outwardly directed; 3) U-shaped manus prints; and 4) a high degree of heteropody (ratio of manus to pes size). The mean heteropody of the well-preserved Guanghui sauropod tracks from GH-S1 is 1: 3.6. This is close to Brontopodus birdi (1:3) and Breviparopus (1:3.6) but significantly less than in the Parabrontopodus (1:4 or 1:5) (Lockley et al., 1994a. We refer the Guanghui sauropod trackway GH-S1 to Brontopodus isp. The widegauge of the Brontopodus-type trackways suggests that the tracks were left by titanosaurian sauropods (Wilson and Carrano, 1999; Lockley et al., 2002). Lingulate sand mounds are preserved at the posterior end of the GHS3 tracks (Fig. 9). This configuration is a common feature of swim tracks (Swanson et al., 2002), including those of theropods (Ezquerra et al., 2007; Xing et al., 2013b) and turtle tracks (Xing et al., 2014a), but typically occurs in irregular configuration and not in regular walking sequences. There are at least two trackways potentially related to swimming sauropods in China: the Yanguoxia digit-only sauropod pes tracks, from the Cretaceous of Gansu Province (Xing et al., 2016d), and Yantan basal sauropodomorpha tracks with slender, tapering, parallel pes digit imprints. However, their patterns are typical and diagnostic for walking. One of the main arguments against swimming sauropod scenarios based on tracks is that trackway spacing patterns (step length and gauge) in purported swim traces are often the same as in trackways made by normal walking animals (Xing et al., 2016d, 2019). Despite the possibility that the GH-S3 trackway could indicate swimming behavior, the tracks are similar to the Yanguoxia digit-only sauropod pes tracks, though all the toe traces are wider and the digits do not taper from I to IV. More likely, if the tracks were left on slippery sediment, in which the claws of the trackmaker dig in deeper for stability. In such cases, claw penetration may serve a substrate gripping

Fig. 7. Photograph and interpretative outline drawing of sauropod trackway GH-S1 from the Guanghui site.

Trackway GH-S2 has 16 pes impressions without any manus impression (Figs. 4, 8). The pes impressions are 26.6 cm long on average, with an average length/width ratio of 1.2. Though different in size, the GH-S2 tracks are quite similar to the GH-S1 pes impressions. Digits I–IV of some of the best-preserved pes traces have well-developed claw marks, such as GH-S2-RP3. However, the step lengths of right and left tracks are not clearly different (mean 141.4 cm vs. 130.2 cm). The single steps of the pes impressions tend to increase gradually; 7

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Fig. 8. Photograph and interpretative outline drawing of best preserved portion of sauropod trackways GH-S2 and GH-S3 from the Guanghui site.

h = 4.586 × foot length. The relative stride length (SL/h) may be used to determine whether the animal is walking (SL/h ≤ 2.0), trotting (2 < SL/h < 2.9), or running (SL/h ≥ 2.9) (Alexander, 1976; Thulborn, 1990). The SL/h ratios of the GH-S1, S2, and S3 are 0.92, 1.46 and 1.29, and accordingly suggest walking. Using the equation to estimate speed from trackways (Alexander, 1976), the mean locomotion speed of the trackmakers are 1.02 m/s (3.67 km/h), 1.62 m/s (5.83 km/h) and 1.4 m/s (5.04 km/h).

function. Changpeipus from the Shanshan site, Middle Jurassic of Xingjiang, also has wider digit traces due to slippery sediment (Xing et al., 2014b). It has long been known that many incomplete sauropod tracks represent undertracks or partly penetrative tracks (toe traces) made by trackmakers that walked on a surface above that now exposed (Lockley and Rice, 1990; Lockley et al., 1994b, and that many examples of purported swim tracks have been reinterpreted, especially where the pattern of incomplete trackways is no different from that made during normal walking progression. Thus, it is particularly important that the role of preservation, and the degree to which trackways configurations conform to normal walking patterns, should be considered in the case of sites where sauropod trackways appear incomplete.

5.3. Ornithopod tracks 5.3.1. Materials and description The ornithopod trackway from Guanghui site is catalogued as GHO1 and consists of a sequence of twelve tridactyl tracks (Figs. 4, 10, Table 1). The GH-O1 tracks are only preserved as digit traces, showing three separate toe traces without heel impressions (Fig. 10). Thus, track lengths are partial measurements, and track widths more reliable as a measure of trackmaker size. The tracks have lengths of ~14.7 cm,

5.2.3. Speed estimates For sauropods, Alexander (1976) first suggested that hip height h = 4 × foot length, whereas, later, Thulborn (1990) estimated h = 5.9 × foot length. González Riga (2011), proposed a formula for titanosaurid tracks based on anatomical and ichnological evidences as 8

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Fig. 9. Photograph and interpretative outline drawings of well-preserved sauropod tracks GH-S3 from the Guanghui site. Table 2 Measurements (in cm) of the sauropod trackways from Guanghui tracksite, Heilongjiang Province, China. Number

ML

MW

R

PL

SL

PA

ML/MW

WAP

WAP/P'ML

GH-S1 GH-S1 GH-S2 GH-S3

48.4 13.2 26.6 29.7

40.1 24.4 23.6 38.1

|20| |26| |18| |12|

133.5 126.6 111.4 135.8

204.5 209.5 177.9 175.6

97 110 103 79

1.3 0.5 1.2 0.8

86.9 – 64.3 103.2

1.8 – 2.5 3.8

pes manus pes pes

Abbreviations: ML: Maximum length; MW: Maximum; R: Rotation; PL: Pace length; SL: Stride length; PA: Pace angulation; WAP: Width of the angulation pattern of the pes (calculated value); ML/MW, WAP/P'ML and are dimensionless.

Fig. 10. Photograph and interpretative outline drawing of well-preserved ornithopod tracks (four specimens from GH-O1 trackway) from the Guanghui site.

Falkingham, 2017; Marchetti et al., 2019). Similar findings can be seen in Late Cretaceous Yangmeikeng specimens from the southern part of Nanxiong Basin, China. GH-O1 tracks from the Yanguoxia assemblage and Yangmeikeng specimens are especially similar, suggesting that the former are also undertracks. The mesaxony of the GH-O1 tracks is only 0.36, smaller than the 0.45 in the Yangmeikeng specimens, but still in the common range for China's Caririchnium type (~0.22–0.52) (Xing et al., 2017)

widths of ~31.8 cm, and are separated by a step length of 126.1 cm (Table 1). GH-O1-R1–L2 is perfectly preserved. Digit III is the most developed. The ungual traces of digits II and IV are nearly triangular, with a blunt anterior edge (Fig. 10). The L/W ratio of the anterior triangle is 0.36.

5.3.2. Comparisons and discussion Similar digit-only tracks from the Yanguoxia sites, Gansu Province, have been interpreted as swim traces (Fujita et al., 2012). However, recent research interprets such prints as the result of animals walking on a soft mud-silt substrate, projecting their claws deeply to register their traces on an underlying sand layer, where they naturally gained more grip during progression (Xing et al., 2016d; Gatesy and

6. Discussion As mentioned above, the hip height of the theropod trackmaker of the swimming traces is estimated at 74 cm–100.8 cm, reflecting the 9

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groups appear to have persisted in the Heilongjiang area throughout the Cretaceous.

probable water depth of this area at the time the tracks were made and therefore giving some clues to the paleoenvironment. The theropod swimming traces go west and west-northwest while the ornithopod tracks go northeast, and the sauropod tracks go north-northwest. The three parallel sauropod trackways may be a sign of gregarious behavior, but it is interesting to note that the mode of preservation of the tracks changes from west to east (Fig. 4) in a zone only ~7.0 m wide: i.e., trackway S1, representing the largest animals (mean pes length 48.4 cm) shows the best preserved manus pes sets (Fig. 7), whereas trackway S2, the smallest (mean pes length 26.6 cm) is a pes-only trackway, and trackway S3 (mean pes length 29.7 cm) is also a pes-only trackway with deep anterior toe traces, and a notable asymmetry in pes track width between right (40.6 cm) and left (35.3 cm) tracks (Figs. 4, 8–9, Table 2, Supplementary material). The length/width ratio (0.78) of S3 is also very anomalous for sauropods in comparison with the pes length/width ratios of S1 and S2 (1.21 and 1.13, respectively). It is not possible to offer a compelling interpretation for this variation in mode of preservation between trackways S1, S2 and S3. However, it is reasonable to suggest that such variation strengthens the case for the animals passing over the area (possibly shore parallel) at different times, when different substrate conditions prevailed, or in a zone where the substrate consistency varied significant across an east-west zone only about 7 m wide. These three sauropod trackways generally do not increase in measurable depth, across this small east-west zone, and may be parallel to the lakeshore, while the theropod tracks are perpendicular to the lakeshore. (Milner et al., 2006). Some traces in GH-T1 are overlapped by GH-S2, reflecting the sequence of formation (Fig. 4). We infer that the theropod traces were left first, followed by ornithopod and sauropod tracks. Tetrapodosaurus discovered from the Yilan site reflects a potential ankylosaur trackmaker, in addition to ornithopod, sauropod, and theropod trackmakers. This is the first time that a diverse Early Cretaceous dinosaur fauna has been discovered in Heilongjiang, the northernmost part of China. The Taoqihe dinosaur trackway assemblage is comparable in composition to the vertebrate fauna of the Jehol Biota. According to the new definition the Jehol Biota (Barremian to Aptian) is discovered in deposits of the Yixian, Jiufotang and Huajiying formations in western Liaoning, adjacent Inner Mongolia, and northern Hebei (Pan et al., 2013). Although this biota is famous for its small-sized feathered dinosaurs, it also contained large theropod (e.g. Tyrannosauroidea Yutyrannus Xu et al., 2012), sauropod (e.g., Titanosauriformes Dongbeititan Wang et al., 2007), ankylosaur (e.g. Ankylosauridae Crichtonsaurus Dong, 2002) and ornithopod (e.g. Hadrosauroidea Jinzhousaurus Wang and Xu, 2001) fossils. All these latter groups are also represented in the Taoqihe ichnofauna. Because small-sized feathered dinosaurs are mostly arboreal species, it is unlikely they would leave an ichnological signature. This indicates a higher consistency and larger distribution of these dinosaur associations in northeastern China, than previously documented. The Late Cretaceous dinosaur fossil records from the Jiayin and Wulaga bonebeds, Blagoveschensk and Kundur bonebeds in the Heilong River region of Russia are dominated by hadrosaurs, including Mandschurosaurus amurensis (Riabinin, 1930); Amurosaurus riabinini (Bolotsky and Kurzanov, 1991); Charonosaurus jiayinensis (Godefroit et al., 2000); Olorotitan arharensis (Godefroit et al., 2003); Kerberosaurus manakini (Bolotsky and Godefroit, 2004); Sahaliyania elunchunorum (Godefroit et al., 2008); Wulagasaurus dongi (Godefroit et al., 2008) and Kundurosaurus nagornyi (Godefroit et al., 2012). Theropoda indet. are also found, including ornithomimosaurs, troodontids, dromaeosaurs (Li, 2004) and an albertosaurine tooth (Lü and Han, 2012). Sauropoda are present with the titanosaur Arkharavia heterocoelica and indeterminate forms (Alifanov and Bolotsky, 2010), furthermore Ankylosauridae indet. (Tumanova et al., 2004). Tracks from Yilan and Guanghui sites suggest that ankylosaurs, sauropods, theropods and ornithopods can be traced to the Early Cretaceous, and these dinosaur

7. Conclusions Apart from the report of a Late Cretaceous hadrosaur track Jiayinosauropus, from the Upper Cretaceous Yong'ancun Formation, Heilongjiang Province was thought to be devoid of other Cretaceous dinosaur tracks. Our new study has brought to light evidence of an almost-forgotten historic record of a single large, Lower Cretaceous pentadactyl track of probable thyreophoran (ankylosaurian) affinity. More importantly, the new discovery of a diverse Lower Cretaceous dinosaur tracks ichnofauna (> 70 tracks in 5 trackways) from the Taoqihe Formation reveals the presence of parallel sauropod trackways, a toe-trace dominated tridactyl ornithopod trackway and elongate tridactyl theropod (? swim) tracks. This is the first Lower Cretaceous tracksite reported from the province with multiple trackways. Palaebiogeographically, the new record is important, because it reveals that faunas evidenced from Late Cretaceous skeletons of northeastern China and adjacent Russia, were already present in this area in the Early Cretaceous. Supplementary data to this article can be found online at https:// doi.org/10.1016/j.palaeo.2019.109275. Acknowledgments The authors thank Spencer G. Lucas and an anonymous reviewer for their constructive comments. Li Zhu is thanked for providing local support. This research was funded by the National Natural Science Foundation of China (No. 41888101, 41790455, 41772008); the Fundamental Research Funds for the Central Universities (No. 2652017215), the State Key Laboratory of Palaeobiology and Stratigraphy (Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences) (No. 173127). References Alexander, R.M., 1976. Estimates of speeds of dinosaurs. Nature 26, 129–130. Alifanov, V.R., Bolotsky, Y.L., 2010. Arkharavia heterocoelica gen. et sp. nov., a new sauropod dinosaur from the Upper Cretaceous of far eastern Russia. Paleontol. Z. (in Russian) 1, 76–83. Apesteguía, S., Gallina, P.A., 2011. Tunasniyoj, a dinosaur tracksite from the JurassicCretaceous boundary of Bolivia: Ann. Brazilian Acad. Sci. 83, 267–277. BGMRH (The Fifth Geological Team of Heilongjiang, Provincial Bureau of Geology Regional Geological Survey team), 1972. Geological Map of the People’s Republic of China, Yilan Map Sheet 1:200000 (L-52-16) 84. Bolotsky, Y.L., Godefroit, P., 2004. A new hadrosaurine dinosaur from the Late Cretaceous of Far Eastern Russia. J. Vert. Paleontol. 24 (2), 351–365. Bolotsky, Y.L., Kurzanov, S.K., 1991. The hadrosaurs of the Amur Region. In: Geology of the Pacific Ocean Border. Blagoveschensk: Amur KNII, 94–103. Chen, B.L., 1984. The discovery and significance of dinosaur footprints in Yilan County. Heilongjiang Province. J. Daqing Petrol. Inst. 4, 195. Coombs, W.P., 1980. Swimming ability of carnivorous dinosaurs. In: Science. 207. pp. 1198–1200. Dong, Z.M., 2002. A new armored dinosaur (Ankylosauria) from Beipiao Basin, Liaoning Province, northeastern China. Vert. PalAsiat. 40 (4), 276–285. Dong, Z.M., Zhou, Z.L., Wu, S.Y., 2003. Note on a hadrosaur footprint from Heilongjiang River area of China. Vert. PalAsiat. 41 (4), 324–326. Ensom, P., 1987. Dinosaur tracks in Dorset. Geol. Today 31, 182–183. Ezquerra, R., Doublet, S., Costeur, L., et al., 2007. Were non-avian theropod dinosaurs able to swim? Supportive evidence from an Early Cretaceous trackway, Cameros Basin (La Rioja, Spain). Geology 35, 507–510. Farlow, J.O., 1992. Sauropod tracks and trackmakers: integrating the ichnological and skeletal record. Zubia 10, 89–138. Farlow, J.O., Pittman, J.G., Hawthorne, J.M., 1989. Brontopodus birdi. Lower Cretaceous sauropod footprints from the US Gulf coastal plain. In: Gillette, D.D., Lockley, M.G. (Eds.), Dinosaur Tracks and Traces, 371–394. Fujita, M., Azuma, Y., Goto, M., Tomida, Y., Hayashi, S., Arakawa, Y., 2003. First ankylosaur footprints from Japan and their significance: J. Vert. Paleontol. 23, 52A. Fujita, M., Lee, Y.N., Azuma, Y., Li, D., 2012. Unusual tridactyl trackways with tail traces from the Lower Cretaceous Hekou Group, Gansu Province, China. PALAIOS 27, 560–570. Gatesy, S.M., Falkingham, P.L., 2017. Neither bones nor feet: track morphological variation and ‘preservation quality’. J. Vert. Paleontol. 37 (3), e1314298.

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