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Exquisitely-Preserved, High-Definition Skin Traces in Diminutive Theropod

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Exquisitely-Preserved, High-Definition Skin Traces in Diminutive Theropod www.nature.com/scientificreports OPEN Exquisitely-preserved, high- defnition skin traces in diminutive theropod tracks from the Received: 28 September 2018 Accepted: 3 January 2019 Cretaceous of Korea Published: xx xx xxxx Kyung Soo Kim 1, Martin G. Lockley2, Jong Deock Lim3 & Lida Xing 4 Small theropod tracks, ichnogenus Minisauripus, from the Jinju Formation (Cretaceous) of Korea reveal exquisitely preserved skin texture impressions. This is the frst report for any dinosaur of skin traces that cover entire footprints, and every footprint in a trackway. Special sedimentological conditions allowed footprint registration without smearing of skin texture patterns which consist of densely- packed, reticulate arrays of small (<0.5 mm) polygons, preserved as both impressions and casts, the latter essentially foot replicas. The skin texture resembles that reported for two Lower Cretaceous avian theropods (birds) from China which had quite diferent foot morphologies. This is also the oldest report of Minisauripus from Korea predating fve reports from the Haman Formation of inferred Albian age. Minisauripus is now known from six Korean and three Chinese localities, all from the Lower Cretaceous. This gives a total sample of ~95 tracks representing ~54 trackways. With >80% of tracks <3.0 cm long, Minisauripus is pivotal in debates over whether small tracks represent small species, as the database suggests, or juveniles of large species. Te diminutive theropod track ichnogenus Minisauripus is represented by two ichnospecies. It was frst described from Sichuan Province China1, as ichnospecies M. chuanzhuensis, from the Upper Cretaceous Daergun Formation of the Chiating Group, referred to as Late Cretaceous in age2, but elsewhere considered part of the Lower Cretaceous Jiaguan Formation3. Minisauripus was subsequently discovered at two sites in the Early Cretaceous Haman Formation (Hayang Group) of Korea and another site in the Early Cretaceous of Tianjialou Formation of Shandong Province, China4. Finds in the 2000s indicated that the Minisauripus track was not made by a miniature blunt toed ornithopod as had initially inferred1 but was instead made by a theropod which ofen registered sharp claw traces, and ofen clearly showed two phalangeal pads on digit II and three on digit III, diagnostic of theropods. Clearly defned phalangeal pad traces were not observed on digit IV. Due to these diagnostic morphological features the new ich- nospecies M. zhenshounani was erected on the basis of the Shandong material4, which included tracks somewhat longer than those reported from other sites: i.e., with footprint length (FL) up to 6.1 cm, including distinctive fne claw traces, whereas >80% of the global sample was in the FL range of 1.1–2.9 cm. Since 2008, Minisauripus isp. has been discovered at a number of other Korean localities, all in the Haman Formation5, and at a new site in the Feitianshan Formation of Sichuan Province China6. Te former, 2012 paper summarized the total Minisauripus database as consisting of “more than 80 well-preserved tracks comprising a minimum of 50 trackways from two localities in China and fve in Korea”5: i.e., a total of seven localities. One of the localities, Buyun-ri consisted of “Five separate localities, in very close proximity”5. Te addition of the new Chinese site increased the database so that “a total of ~92 Minisauripus tracks, representing at least 55 trackways, are now recorded from a total of eight localities”6 exclusive of sites in very close proximity which fall geographically within single key site locations. Clearly this constitutes a large multi-site sample of diminutive tracks, which bear only slight resemblance to larger theropod tracks assigned to quite diferent ichnotaxa (Suppl. Info). 1Department of Science Education, Chinju National University of Education, 3 Jinnyangho-ro 369beon-gil, Jinju-si, Gyeongnam, 52673, Korea. 2Dinosaur Trackers Research Group, University of Colorado Denver, P.O. Box 173364, Denver, CO, 80217, USA. 3Cultural Heritage Administration, Government Complex-Daejeon, 189, Cheongsa-ro, Seo-gu, Daejon, 35208, Korea. 4School of the Earth Sciences and Resources, China University of Geosciences, Beijing, 100083, China. Correspondence and requests for materials should be addressed to M.G.L. (email: Martin. [email protected]) SCIENTIFIC REPORTS | (2019) 9:2039 | https://doi.org/10.1038/s41598-019-38633-4 1 www.nature.com/scientificreports/ Figure 1. (A) Geological map of Gyeongsang Basin, in southeast sector of Korean peninsula (inset), showing outcrop of group-level units. (B) Geological map of formations around Jinju City showing fossil locality ~5 km south of the city center. Maps made by K-S K in Adobe photoshop (version CS6 www.adobe.com/Photoshop) and Canvas X (version, 2017 Build 160, http://www.canvasgfx.com/). Te present study, adds signifcantly to the growing data on Minisauripus morphology and distribution in space and time by adding well-preserved material, with exquisitely-preserved high-defnition skin traces, from a ninth site, from the Early Cretaceous Jinju Formation in the Jinju City area of Korea (Fig. 1). Te Jinju Formation in this area has recently attracted global ichnological attention for having produced large volumes of new track specimens 7 including a number of important new ichnotaxa, attributable to major groups such as mammals7–11, frogs8 and lizards11 which had not previously been reported from the Cretaceous of Korea. In addition diminutive tracks of purported microraptorine afnity have been described9. Tis new Minisauripus occurrence described here is the frst Korean site not associated with the younger Haman Formation (Fig. 1), and it is also the frst from any global site to reveal Minisauripus skin impressions. Moreover, these skin traces are extraordinarily well preserved, and the only example known for such a small dinosaur. Tus, the material extends the known stratigraphic range of Minisauripus in Korea and gives us a high defnition snapshot revealing the texture and ornamentation of the foot integument, which resembles a living foot replica, and can be compared with that reported from other theropod tracks. SCIENTIFIC REPORTS | (2019) 9:2039 | https://doi.org/10.1038/s41598-019-38633-4 2 www.nature.com/scientificreports/ Figure 2. (A) Counterpart slab CUE JJ_M01 showing trackway with four consecutive Minisauripus track casts TL1-TR2. (B) Natural impression slab (CUE JJ_M02) showing tracks TL2 and TR2. (C) Isolated track specimen (CUE JJ_M03). Compare with Fig. 3. Photographs by K-S Kim and layout created in Canvas X (version, 2017 Build 160, http://www.canvasgfx.com/). Material and Methods The Minisauripus material described here, originates from major excavations in the Jinju Innovation City area (Fig. 1), at the Ppuri Industrial Complex, not far from the highly productive outcrops associated with the Korea National Natural Monument Number 534 project (named Te Pterosaur, Bird and Dinosaur Tracksite of Hotan-dong), and the newly built Jinju Pterosaur Footprint Museum which already contains a number of recently described type and fgured specimens7–11 from the Jinju Formation. Te specimens are Chinju National University of Education (CUE) JJ_M01-M03 (Figs 2–3) and consist of a part and counterpart (M01 and M02) comprising a thin (~2.0 cm thick) slab of very fne grey sandstone with a dark grey to black mudstone drape less than 1.0 mm thick (Fig. 2B), with a four track-trackway (Figs 2A,B and 3B), as well as an isolated cast (M03), with a single track, from the same surface which, based on size and mode of preservation, likely belongs to the same trackway. Te slab with the natural impressions (Fig. 2B) reveals two Minisauripus tracks from the same trackway and one pterosaur manus track in concave epirelief. Tere are also a few faint invertebrate traces, including very small Cochlichnus trails, and many small raindrop impressions between 2.0 and 5.0 mm in diameter. Replicas of casts TR1-TR2, impression TL2 and the isolated M03 track were made for comparative study as University of Colorado Museum of Natural History (UCM) replicas UCM 214.323, 324 and 325 respectively. Te 7.0 cm-thick counterpart slab (CUE JJ_M01) which originally overlay the clay-draped unit shows another ~1.0 cm of dark grey to black mudstone overlain by ~6.0 cm of fne-grained sand. Te whole counterpart slab, with track casts, is larger (~50 × 50 cm) than the aforementioned slab with impressions, and reveals a sequence of four Minisauripus tracks comprising a single narrow trackway (Figs 2A,3B,C) in negative (convex hyporelief), and an additional pterosaur manus track. Te density of raindrop impressions is estimated at 1000/m2. Te rain- drop impressions and the Minisauripus are very shallow and fat bottomed. Tese physical features of both the biogenic and non-biogenic traces give clues as to how the substrate conditions facilitated superior preservation. SCIENTIFIC REPORTS | (2019) 9:2039 | https://doi.org/10.1038/s41598-019-38633-4 3 www.nature.com/scientificreports/ Figure 3. (A) Map of specimens CUE JJ_M01-3 showing four-track Minisauripus trackway, and additional isolated ffh track on small unconnected slab. Map based on counterpart cast of track-bearing surface. Red outline shows part of surface preserved as natural impressions. Pterosaur manus tracks, desiccation cracks (stippled areas), raindrop impressions and invertebrate traces also shown. Compare with Fig. 2 and text for details. (B) Shows microstratigraphy of part and counterpart of track-bearing slab. (C) Shows four track-trackway with dashed line to highlight steps and pace angulations. See Table 1 for measurements. Map made by M. G. Lockley and K-S Kim with layout created in Canvas X (version, 2017 Build 160, http://www.canvasgfx.com/). Track # Length Width L/W Step Stride T width Pace angle TL1 2.4 2.0 1.20 — — — — TR1 2.4 2.0 1.20 20.8 40.4 2.9 172° TL2 2.4 1.9 1.26 19.6 38.8 2.7 174° TR2 2.3 1.8 1.28 19.7 — — — means 2.38 1.93 1.23 20.03 39.6 2.8 173° Isolated t 2.3 1.3 1.28 — — — — Table 1.
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