First Turtle Remains from the Middle-Late Jurassic Yanliao Biota, NE China
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Vol.7, No.1 pages 1-11 Science Technology and Engineering Journal (STEJ) Research Article First Turtle Remains from the Middle-Late Jurassic Yanliao Biota, NE China Lu Li1,2,3, Jialiang Zhang4, Xiaolin Wang1,2,3, Yuan Wang1,2,3 and Haiyan Tong1,5* 1 Laboratory of Vertebrate Evolution and Human Origins of the Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China 2 CAS Center for Excellence in Life and Paleoenvironment, Beijing 100044, China 3 College of Earth and Planetary Sciences, University of China Academy of Sciences, Beijing 100044, China 4 State Key Laboratory of Biogeology and Environmental Geology, China University of Geosciences, Beijing 100083, China 5 Palaeontological Research and Education Centre, Mahasarakham University, Kantarawichai, Maha Sarakham 44150, Thailand * Corresponding author: [email protected] (Received: 14th August 2020, Revised: 11th January 2021, Accepted: 9th March 2021) Abstract - The Middle-Late Jurassic Yanliao Biota, preceding the Early Cretaceous Jehol Biota in NE China has yielded a rich collection of plant, invertebrate and vertebrate fossils. But contrary to the Jehol Biota which is rich in freshwater vertebrates, in the Yanliao Biota the aquatic reptiles are absent, and turtles have not been reported so far. In this paper, we report on the first turtle remains from the Yanliao Biota. The material consists of a partial skeleton from the Upper Jurassic Tiaojishan Formation of Bawanggou site (Qinglong, Hebei Province, China). Characterized by a broad skull with a pair of sulci carotici and a remnant of an interpterygoid vacuity, a well-developed anterior lobe of the plastron with mesiolaterally elongated epiplastra and a relatively large oval entoplastron; it is assigned to Annemys sp. (Xinjiangchelyidae). This discovery adds a new element to the poorly known freshwater vertebrate fauna of the Yanliao Biota and extends the geographical distribution of Annemys to NE China. Keywords: Testudines, Xinjiangchelyidae, Yanliao Biota, NE China, Tiaojishan formation, late jurassic 2 Lu Li, Jialiang Zhang, Xiaolin Wang, Science Technology and Engineering Journal (STEJ) Yuan Wang & Haiyan Tong 1. Introduction (see Xu et al., 2016 for a thorough review). But contrary to the Jehol Biota that is The Middle-Late Jurassic non-marine rich in freshwater animals, in the Yanliao deposits in northeastern China (Western Biota the freshwater reptiles are absent; Liaoning, northern Hebei and southeastern and turtles and crocodiles have hitherto Inner Mongolia) have yielded a rich collection not been reported (Xu et al., 2016, 2019 ; of fossil assemblages, termed Yanliao Biota. Zhou & Wang, 2017). In this paper we Like the famous Early Cretaceous Jehol report on the first turtle remains from Biota that covers the same area, remains the Yanliao Biota. The material consists of plants, invertebrates and vertebrates of of a partial skeleton of xinjiangchelyid the Yanliao Biota are mainly preserved in turtle from the Upper Jurassic Tiaojishan lacustrine sediments rich in volcanic ash. Formation of Bawanggou, Qinglong, Hebei Vertebrates include fishes, amphibians, Province, China. pterosaurs, dinosaurs and mammaliaforms Figure 1A. Map showing the location of Bawanggou site, Qinglong Manchu Autonomous County, Hebei Province, China. B. Stratigraphical distribution of the Middle-Late Jurassic Yanliao Biota (shading). 2. Geological setting the salamander Chunerpeton tianyiensis (Sullivan et al., 2014), the pterosaur The Bawanggou site (also called Mutoudeng Qinglongopterus guoi and Dendrorhynchoides site) is located north of Mutoudeng Town, mutoudengensis (Jiang et al., 2015 ; Lü Qinglong Manchu Autonomous County, et al., 2012), the dinosaur Yi qi (Xu et al., in the northern part of Hebei Province 2015) and the mammal Arboroharamiya (Figure 1). It has yielded some important jenkinsi (Zheng et al., 2013). vertebrates of the Yanliao Biota, such as Volume 7, Number 1, January-June 2021 First turtle remains from the Middle-Late Jurassic 3 Yanliao Biota, NE China The Yanliao Biota straddled the laminography system (160-Micro-CL) at Middle and Late Jurassic for about 10 the Key Laboratory of Vertebrate Evolution million years. It is divided in two phases: and Human Origin of Chinese Academy of the Bathonian-Callovian Daohugou phase Sciences, IVPP, Beijing. (early Yanliao Biota, Jiulongshan/ Haifengshan formations, about 168-164 million years ago) and the Oxfordian 4. Systematic palaeontology Linglongta phase (late Yanliao Biota, Tiaojishan/Lanqi formations, 164-159 Testudines Linnaeus (1758) million years ago). The fossil-bearing beds Cryptodira Cope (1868) of Bawanggou site pertain to the Tiaojishan Formation, which is regarded as Middle Xinjiangchelyidae Nessov in Kaznyshkin Jurassic (Daohugou Phase) (Sullivan et al., et al. (1990) 2014) or Late Jurassic in age (Linglongta Annemys Sukhanov and Narmandakh. (2006) Phase) (Liao & Huang, 2014) based on palaeontological and biostratigraphic Annemys sp. studies. A recent study on SIMS zircon (Figure 2-3) U-Pb yielded an age of 159 .0-159.8 Ma, which places the Tiaojishan Formation of Bawanggou site in the Oxfordian (Upper Referred material: IVPP V16730, a Jurassic) (Yu et al., 2020), providing a partial skeleton presented in ventral view, direct isotopic age to support the opinion including skull with attached lower jaw; of Liao et Huang (2014). disarticulated cervicals, complete right and left humeri, right forearm, right and left 3. Material and method scapula, anterior portion of the carapace and anterior lobe of the plastron with both The specimen studied in the present work right and left axillary buttresses, as well (IVPP V16730, anterior portion of a skeleton) as other shell fragments. The specimen is is housed in the Institute of Vertebrate preserved on a slab and further included Palaeontology and Palaeoanthropology in plaster. It is dorsoventrally crushed. (IVPP), Chinese Academy of Sciences, Locality and horizon: Bawanggou China. Preserved on a slab, the ventral surface site, Xinglongtaizi Village, Mutoudeng of the specimen is prepared mechanically. Township, Qinglong Manchu Autonomous To reveal the hidden features, the X-ray County, Qinhuangdao City, Hebei scanning technology is used. The Province, China. Upper Jurassic (Oxofordian) scanning is carried out using a micro-computed Tiaojishan Foramation. 4 Lu Li, Jialiang Zhang, Xiaolin Wang, Science Technology and Engineering Journal (STEJ) Yuan Wang & Haiyan Tong Fig. 2.Figure Annemys 2. Annemys sp. (Xinjiangchelyidae) sp. (Xinjiangchelyidae) from the fromUpper the Jurassic Upper Tiaojishan Jurassic Tiaojishan Formation of FormationBawamgou, of Bawamgou, Qinglong Qinglong Manchu Manchu Autonomous Autonomous County, County Hebei, Hebei Province, Province, China. A-B, partial China.skeleton A-B, partial(IVPP skeletonV16730) ( IVPPin ventral V16730 view;) in C-D, ventral detail view; of skull.C-D, detail Scale of bar skull. = 5 cm. Abbreviations: Scalebo, basioccipital;bar = 5 cm. Abbreviations: bs, basisphenoid; bo, basioccipital; co, costal; coc, bs, condylus basisphenoid; occipitalis; co, condylus cso, crista supraoccipitalis; occipitalis;ento, cso, entoplastron; crista supraoccipitalis; epi, epiplastron; ento, entoplastron; hyo, hyoplastron; epi, epiplastron; ipv, interpterygoid hyo, vacuity; lj, hyoplastron;lower jaw; lj, lower mx, maxilla;jaw; mx, n, maxilla; neural; nu, nu, nuchal; nuchal; pe, pe, peripheral; peripheral; pm, pm, premaxilla; premaxilla; pt, pt, pterygoid; qu, quadrate; sc, sulcus caroticus; so, supraoccipital; sq, squamosal. pterygoid; qu, quadrate; sc, sulcus caroticus; so, supraoccipital; sq, squamosal. Volume 7, Number 1, January-June 2021 First turtle remains from the Middle-Late Jurassic 5 Yanliao Biota, NE China 5. Description ventral exposure is roughly as long as wide, with the lateral borders slightly convergent Skull The skull, measuring 47 mm. from the posteriorly. The tubercula basioccipitale snout to the posterior end of the squamosal are reduced and set close one to another. and 38 mm. at the level of the quadrates, A strong occipital condyle is preserved. is elongate, with almost parallel lateral The supraoccipital bears a relatively long borders and a blunt snout. The ventral crista supraoccipitalis. The lower jaw is surface of the skull is somewhat crushed somewhat disarticulated, allowing the and moreover covered by the lower jaw, right condylus mandibularis to be exposed. the hyoid apparatus, a cervical vertebra The articular surface is concave and Descriptionand other :bone fragments, so that many widermandibularis than long. to be exposed. The articular Skull The skull, measuring 47 mm. from the surface is concave and wider than long. structures and sutures are not clearly visible. snout to the posterior end of the squamosal Lower jaw The lower jaw is robust, andThe 38 triturating mm. at the surface, level of as the partly quadrates, exposed is withLower a roughjaw The outer lower surface jaw is similar robust, towith that a elongate,on the left with side, almost appears parallel to be lateral narrow borders with ofrough the outershell. surfaceThe left similar ramus to is thatseparated of the anda sharp a blunt labial snout. ridge. The ventralThe pterygoids surface of theare fromshell. theThe rightleft ramusone by is crushing,separated itfrom is notthe skullpartly is exposed; somewhat the processuscrushed and pterygoideus moreover clearright onewhether by crushing, they are it unitedis not clearby