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      Hints of the Early Jehol Biota: Important Dinosaur Footprint Assemblages from the Jurassic-Cretaceous Boundary Tuchengzi Formation in Beijing, China

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          Abstract

          New reports of dinosaur tracksites in the Tuchengzi Formation in the newly established Yanqing Global Geopark, Beijing, China, support previous inferences that the track assemblages from this formation are saurischian-dominated. More specifically, the assemblages appear theropod-dominated, with the majority of well-preserved tracks conforming to the Grallator type (sensus lato), thus representing relatively small trackmakers. Such ichnofaunas supplement the skeletal record from this unit that lacks theropods thus far, proving a larger diversity of dinosaur faunas in that region. Sauropods are represented by medium to large sized and narrow and wide-gauge groups, respectively. The latter correspond with earlier discoveries of titanosauriform skeletons in the same unit. Previous records of ornithischian tracks cannot be positively confirmed. Purported occurrences are re-evaluated here, the trackways and imprints, except of a single possible specimen, re-assigned to theropods. Palecologically the Tuchengzi ichnofauna is characteristic of semi-arid fluvio-lacustrine inland basins with Upper Jurassic-Lower Cretaceous deposits in northern China that all show assemblages with abundant theropod and sauropod tracks and minor components of ornithopod, pterosaur and bird tracks.

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          A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus.

          The early evolution of the major groups of derived non-avialan theropods is still not well understood, mainly because of their poor fossil record in the Jurassic. A well-known result of this problem is the 'temporal paradox' argument that is sometimes made against the theropod hypothesis of avian origins. Here we report on an exceptionally well-preserved small theropod specimen collected from the earliest Late Jurassic Tiaojishan Formation of western Liaoning, China. The specimen is referable to the Troodontidae, which are among the theropods most closely related to birds. This new find refutes the 'temporal paradox'1 and provides significant information on the temporal framework of theropod divergence. Furthermore, the extensive feathering of this specimen, particularly the attachment of long pennaceous feathers to the pes, sheds new light on the early evolution of feathers and demonstrates the complex distribution of skeletal and integumentary features close to the dinosaur-bird transition.
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            A ‘Terror of Tyrannosaurs’: The First Trackways of Tyrannosaurids and Evidence of Gregariousness and Pathology in Tyrannosauridae

            The skeletal record of tyrannosaurids is well-documented, whereas their footprint record is surprisingly sparse. There are only a few isolated footprints attributed to tyrannosaurids and, hitherto, no reported trackways. We report the world’s first trackways attributable to tyrannosaurids, and describe a new ichnotaxon attributable to tyrannosaurids. These trackways are from the Upper Cretaceous (Campanian - Maastrichtian) of northeastern British Columbia, Canada. One trackway consists of three tridactyl footprints, and two adjacent trackways consist of two footprints each. All three trackways show animals bearing southeast within an 8.5 meter-wide corridor. Similarities in depth and preservation of the tyrannosaurid tracks indicate that these three trackways were made by track-makers walking concurrently in the same direction. These trackways add significantly to previous osteology-based hypotheses of locomotion and behavior in Tyrannosauridae by providing ichnologic support for gregariousness in tyrannosaurids, and the first record of the walking gait of tyrannosaurids.
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              Discriminating between Medium-Sized Tridactyl Trackmakers: Tracking Ornithopod Tracks in the Base of the Cretaceous (Berriasian, Spain)

              Background Recent work on the Jurassic-Cretaceous transition of the Iberian Range (Spain) has opened a new window onto the interpretation of the trackmakers of some medium-sized tridactyl tracks. The ichnotaxon Therangospodus oncalensis has been described in the Huérteles Formation (Berriasian) and is one of the classical tracks from the area assigned to medium-sized theropods. Methodology/Principal Findings A review of the type locality of Therangospodus oncalensis (Fuentesalvo tracksite) and other tracksites from the Huérteles Formation (Berriasian) has yielded new information on the morphology, gait and trackmaker identity of the aforementioned ichnospecies. The new data suggest that the trackmaker is an ornithopod rather than a theropod on the basis of the length/width ratio, the anterior triangle length-width ratio, the short steps, the round to quadrangular heel pad impression and the probable manus impressions. Conclusions/Significance T. oncalensis shows similarities with various tracks from the Berriasian of Europe assigned to Iguanodontipus. The ichnotaxonomical status of this ichnospecies is here considered as Iguanodontipus? oncalensis due to the current state of knowledge of the ichnotaxonomy of medium-sized ornithopod tracks. This reassessment of I? oncalensis also has two significant implications for the palaeoecology of the faunas during the deposition of the Huérteles Formation: 1- the high number and percentage of theropod tracks would be lower than previous papers have suggested. 2- the gregarious behaviour described in the type locality (Fuentesalvo) would be among ornithopods instead of theropods.
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                Author and article information

                Contributors
                Role: Academic Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                22 April 2015
                2015
                : 10
                : 4
                : e0122715
                Affiliations
                [1 ]School of the Earth Sciences and Resources, China University of Geosciences, Beijing 100083, China
                [2 ]Dinosaur Trackers Research Group, University of Colorado Denver, PO Box 173364, Denver, CO 80217, United States of America
                [3 ]Peace Region Palaeontology Research Centre, Box 1540, Tumbler Ridge, British Columbia V0C 2W0, Canada
                [4 ]Saurierwelt Paläontologisches Museum, Alte Richt 7, D-92318 Neumarkt, Germany
                [5 ]Fundación Conjunto Paleontológico de Teruel–Dinópolis, Teruel, Aragón E-44002, Spain
                [6 ]Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9, Canada
                [7 ]Institute of Evolutionary Biology, University Witten/Herdecke, Stockumerstr. 10–12, 58454 Witten, Germany
                University of Pennsylvania, UNITED STATES
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: LX ML JZ. Performed the experiments: LX ML RM HK LB SK QH. Analyzed the data: LX ML RM LB HK QH. Contributed reagents/materials/analysis tools: LX ML RM. Wrote the paper: LX JZ ML RM HK LA LB MB SK QH.

                Article
                PONE-D-14-40274
                10.1371/journal.pone.0122715
                4406591
                25901363
                d7c501de-269b-4bc7-acdc-40c4546ea542
                Copyright @ 2015

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited

                History
                : 8 September 2014
                : 12 February 2015
                Page count
                Figures: 13, Tables: 4, Pages: 34
                Funding
                This research project was supported by Yanqing Global Geopark (LX, JZ), and the 2013 support fund for graduate student's science and technology innovation from China University of Geosciences (Beijing), China (LX). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
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