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Home , Chunerpeton, Shishugou Formation, Tiaojishan Formation, Xinjiangchelyidae, Yanliao Biota

Vol.7, No.1 pages 1-11 Science Technology and Engineering Journal (STEJ) Research Article

First Remains from the Middle-Late , NE

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 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- Yanliao Biota, preceding the Early Jehol Biota in NE China has yielded a rich collection of plant, and vertebrate . But contrary to the Jehol Biota which is rich in freshwater vertebrates, in the Yanliao Biota the aquatic are absent, and 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 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 , in the Yanliao deposits in northeastern China (Western Biota the freshwater reptiles are absent; , northern Hebei and southeastern and turtles and crocodiles have hitherto ) have yielded a rich collection not been reported (Xu et al., 2016, 2019 ; of assemblages, termed Yanliao Biota. Zhou & Wang, 2017). In this paper we Like the famous Jehol report on the first turtle remains from Biota that covers the same area, remains the Yanliao Biota. The material consists of plants, 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, , Province, China. , 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 tianyiensis (Sullivan et al., 2014), the 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 qi (Xu et al., in the northern part of Hebei Province 2015) and the 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 . It is divided in two phases: and Human Origin of Chinese Academy of the - Daohugou phase Sciences, IVPP, Beijing. (early Yanliao Biota, Jiulongshan/ Haifengshan formations, about 168-164 million years ago) and the 4. Systematic palaeontology Linglongta phase (late Yanliao Biota, Tiaojishan/Lanqi formations, 164-159 Testudines Linnaeus (1758) million years ago). The fossil-bearing beds 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 suture whether or coveredexternus byis mostly the lowerhidden jaw,by the the lower hyoid jaw. fused.they are united by suture or fused. apparatus,Anterior toa cervical the basisphenoid vertebra and and other between bone fragments,the pterygoids, so therethat ismany a short structures slit extending and Hyoid boneHyoid The bone right cornusThe right branchiale cornus I is suturesanteroposteriorly are not clearly along thevisible. midline, The branchialecomplete whereas I is complete the left onewhereas is damaged. the left It trituratingwhich we surface, interpret as partlyas a remnantexposed onof the oneis rod is damaged.-like and Itcurved, is rod-like with and a curved,narrow Volume 7, Number 1, January-June 2021 First turtle remains from the Middle-Late Jurassic 5 leftinterpterygoid side, appears vacuity to be narrow (Figure with 2C-D a sharp and withposterior a narrow end. posterior end. labial ridge. The pterygoids are partly Yanliao Biota, NE China exposed;Figure 3B). the The foramenprocessus posterius pterygoideus canalis externuspterygoidcarotici isinterni, withmostly a transverse locatedhidden by at suture thethe lower posterioron which jaw. Anterioraend pair of of the foraminato pterygoid,the basisphenoid posterius is visible canalis and on between caroticithe left thepalatinumside. pterygoids, The basisphenoid is therelocated. is a shortThe is slightly basisphenoidslit extending shorter anteroposteriorlyextendsthan the laterally main body alongto form of the thethe midline, basipterygoidbasioccipital. which weprocesses,Anteriorly, interpret but the as asbasisphenoid thea lateralremnant portion contacts of of thethe interpterygoid vacuity (Fig. 2C-D and Fig. basisphenoidpterygoid with is a transversenot completely suture onexposed, which 3B).a pair The of foraminaforamen posteriusposterius canaliscanalis carotici interni,the shape located and atsize the ofposterior the processes end of theare notpalatinum determinable. is located. A pair The of basisphenoid grooves, the pterygoid,extends laterally is visible to formon the the leftbasipterygoid side. The basisphenoidsulcus caroticus, is slightly extends shorter anteromedially than the mainwithinprocesses, body the basisphenoid. ofbut the as thebasioccipital. lateral A pair portion of Anteriorly, swellings, of the thedelimitedbasisphenoid basisphenoid by is acontacts notfurrow completely the posteriorly, pterygoid exposed, with is apresentthe transverse shape posterior and suture size to onofthe the whichsulcus processes acaroticus. pair areof foraminaThenot determinable.basisphenoid posterius contacts A paircanalis theof grooves,basioccipital carotici the palatinumposteriorlysulcus caroticus, is withlocated. extendsa straight The anteromedially basisphenoidsuture. The Figure Figure 3. Micro-computed3. Micro-computed scanning scanning slice slice of of extendsbasioccipitalwithin thelaterally basisphenoid. is robust,to form the A the pair main basipterygoid of body swellings, of its AnnemysFigAnnemysure 3. sp. Micro sp. (IVPP (IVPP- computedV16730) V16730) from scanning from the the Upper Upperslice of processes,delimited but by as a thefurrow lateral posteriorly, portion of the is JurassicAnnemysJurassic Tiaojishan Tiaojishansp. (I VPPFormation Formation V16730) of of Bawanggou. Bawanggou. from the ventral exposure is roughly as long as wide, Scale bar = 2 cm. Abbreviations: see Figure 2. basisphenoidpresent posterior is not tocompletely the sulcus exposed, caroticus. the UpperScale barJurassic = 2 cm. Abbreviations:Tiaojishan Formation see Figure 2.of shapewith theand lateral size bordersof the slightlyprocesses convergent are not Bawanggou. Scale bar = 2 cm. posteriorly.The basisphenoid The tuberculacontacts the basioccipitale basioccipital determinable.posteriorly withA pair a ofstraight grooves, suture. the sulcus The Abbreviations:Shell Thesee Fig. carapace, 2. presented caroticus,are reduced extends and set anteromedially close one to withinanother. the A basioccipital is robust, the main body of its in ventral view, has a smooth and likely basisphenoid.strong occipital A pair condyle of swellings, is preserved. delimited The upturned anterolateral margin. The cervical bysupraoccipital a furrow posteriorly, bears a relatively is present long posterior crista notchShell Theis relatively carapace, deeppresented and delimitedin ventral to the sulcus caroticus. The basisphenoid view, has a smooth and likely upturned supraoccipitalis. The lower jaw is somewhat by the peripherals 1 (Figure 2A-B and contactsdisarticulated, the basioccipital allowing theposteriorly right condylus with a anterolateral margin. The cervical notch is straight suture. The basioccipital is robust, Fig.3A).relatively Thedee nuchalp and isdelimited trapezoidal, by withthe mandibularis to be exposed. The articular the anteriorly convergent lateral margins. thesurface main is body concave of itsand ventral wider thanexposure long. is peripherals 1 (Fig. 2A-B and Fig.3A). The roughly as long as wide, with the lateral Thenuchal neural is trapezoidal, 1 appears towith be six-sidedthe anteriorly with borders slightlyLower jawconvergen The lowert posteriorly. jaw is robust, The shortconvergent posterolateral lateral margins. sides (Figure The 3A).neural The 1 tuberculawith a rough basioccipitale outer surface are reduced similar and to thatset peripheralappears to 1 is benarrow; six- sidedthe sutural with contact short closeof the one shell. to Theanother. left ramusA strong is separatedoccipital withposterolateral the costal sides1 is short. (Fig. The 3A).peripheral The condylefrom the is right preserved. one by Thecrushing, supraoccipital it is not 2peripheral is roughly 1 isas narrow;long as thewide sutural and slightlycontact bears a relatively long crista with the costal 1 is short. The peripheral 2 is clear whether they are united by suture or narrowed posteriorly. The peripherals 3-4 supraoccipitalis. The lower jaw is somewhat areroughly wider as than long long. as Thewide costal and 1 bearsslightly a disarticulated,fused. allowing the right condylus narrowed posteriorly. The peripherals 3-4 sharp ridge on its ventral surface. Hyoid bone The right cornus The preserved part of the plastron branchiale I is complete whereas the left one is damaged. It is rod-like and curved, consists of a complete anterior lobe, but with a narrow posterior end. the epiplastra are slightly disarticulated. The anterior lobe, of 47 mm. long and 67 mm. wide at the base, is relatively long and narrow, with a rounded and slightly scalloped anterior border. The lateral margins are gently curved and convergent anteriorly. The length/width ratio of the anterior lobe is about 70%, comparable to that of Annemys 6 Lu Li, Jialiang Zhang, Xiaolin Wang, Science Technology and Engineering Journal (STEJ) Yuan Wang & Haiyan Tong

Shell The carapace, presented epiplastron. The humeropectoal sulcus is in ventral view, has a smooth and likely not discernible. Some scute sulci on the upturned anterolateral margin. The cervical detached fragment of the plastron likely notch is relatively deep and delimited indicate the presence of inframarginals by the peripherals 1 (Figure 2A and (Figure 2). Figure 3). The nuchal is trapezoidal, with Axial and appendicular elements the anteriorly convergent lateral margins. At least three cervical vertebrae are The neural 1 appears to be six-sided with preserved, all disarticulated and presented short posterolateral sides (Figure 3). The in dorsal view. They are relatively peripheral 1 is narrow; the sutural contact short and wide, with divergent pre- and with the costal 1 is short. The peripheral postzygapophyses that form an X in 2 is roughly as long as wide and slightly dorsal view. narrowed posteriorly. The peripherals 3-4 are wider than long. The costal 1 bears Both coracoids are preserved, but a sharp ridge on its ventral surface. none of them are completely exposed. The scapular process and acromion process The preserved part of the plastron form a wide angle of about 100°. consists of a complete anterior lobe, but the epiplastra are slightly disarticulated. Both humeri are presented in ventral The anterior lobe, of 47 mm. long and 67 view. The humerus, of 39 mm. long, has mm. wide at the base, is relatively long an enlarged proximal end. The medial and narrow, with a rounded and slightly process and the lateral process have roughly scalloped anterior border. The lateral margins the same length. The shaft is straight. The are gently curved and convergent anteriorly. distal end is enlarged, but the trochlea and The length/width ratio of the anterior lobe is capitellum are not distinct. about 70%, comparable to that of Annemys The right forearm is preserved. wusu (Rabi et al., 2013). The plastron is The radius and ulna have roughly the same loosely attached to the carapace, a series length. The metacarpals and phalanges are of pegs with striations on the surface is somewhat mixed. The claws are long and visible on the lateral border of the axillary slender. buttresses. The axillary buttresses are long, extending anteromedially to reach the peripheral 2. The epiplastron has a similar 6. Comparisons and discussion: size as the entoplastron. The left and right epiplastra meet along the midline with a Based on the skull and shell morphology, IVPP long suture, preventing the entoplastron V16730 is assigned to Xinjiangchelyidae. to be exposed on the front margin of the In IVPP V16730, the internal carotid canal plastron. The entoplastron is an oval plate is incompletely floored and a remnant of and much longer than wide as in other the interpterygoid vacuity is present. This xinjiangchelyids. configuration is comparable to that of The intergular reaches the anterior xinjiangchelyids such as Annemys spp. from end of the entoplastron. The gularohumeral the Late Jurassic of Shar Teg, Mongolia, sulcus is located at about midlength of the A. sp. from the Shishugou Formation of the and Qigu Formation Volume 7, Number 1, January-June 2021 First turtle remains from the Middle-Late Jurassic 7 Yanliao Biota, NE China

of the Turpan Basin, and Kalasinemys (Protoxinjiangchelys, Chengyuchelys, prasarttongosothi from the Phu Kradung Tienfuchelys and Yanduchelys) in having Formation of Thailand (Brinkman et al., a loose plastron/carapace attachment 2013 ; Rabi et al., 2014 ; Tong et al., 2019 ; (Hu et al., 2020 ; Tong et al., 2012b, 2012a). Wings et al., 2012). The morphology of In this respect, it is closer to Xinjiangchelys the plastron of IVPP V16730 matches and Annemys (Matzke et al., 2005, 2004 ; well that of xinjiangchelyids in having a Nessov, 1995 ; Rabi et al., 2014, 2013 ; well-developed anterior lobe with a rounded Tong et al., 2015a ; Ye, 1986). front margin, mesiolaterally elongated The skull morphology of IVPP epiplastra, and an oval, longer than wide V16730 is similar to that of Annemys. entoplastron (Tong et al., 2012b). The The skull of IVPP V16730 is relatively lack of obvious ornamentation on the shell broad with roughly parallel lateral borders surface and a longer than wide entoplastron as in Annemys, different from a more distinguishes IVPP V16730 from the basal triangular skull of Xinjiangchelys. In IVPP trionychoid Sineaspideretes wimani V16730, the interpterygoid vacuity is not from the Late Jurassic Shangshaximiao completely closed, but remains open as a Formation of the Sichuan Basin (Danilov slit. This primitive feature is also present & Parham, 2006 ; Tong et al., 2014 ; Young in Annemys (A. levensis and A. sp. from & Chow, 1953). the Junggar and Turpan Basins) and K. Xinjiangchelyidae is the dominant prasarttongosothi (Brinkman et al., 2013 ; turtle group in the freshwater ecosystems Rabi et al., 2014 ; Tong et al., 2019; Wings during the Middle-Late Jurassic in Asia. et al., 2012). In A. wusu (Rabi et al., 2013) According to the recent reviews, the from the Qigu Formation, Turpan Basin, family Xinjiangchelyidae consists of the interpterygoid vacuity is an oval-shaped eight genera in mainland Asia, namely opening. The skull of Xinjiangchelys is more Protoxinjiangchelys from the Middle derived, e.g. in X. radiplicatoides from Jurassic Xiashaximiao Formation and the Shishugou Formation, Junggar Basin, Xintiangou Formation, Chengyuchelys, the interpterygoid vacuity is completely Tienfuchelys and Yanduchelys from the closed. In addition, the main body of the Late Jurassic Shangshaximiao Formation basioccipital in IVPP V16730 is roughly of the Sichuan Basin ; Xinjiangchelys from square, with the lateral borders slightly the Middle-Late Jurassic of Lufeng, Turpan convergent posteriorly, with reduced and and Junggar bassins, Annemys from the close-set tubercula basioccipitale as in Middle-Late Jurassic of China, Mongolia, Annemys whereas in Xinjiangchelys, the Kirgizstan and Siberia ; and Phunoichelys basioccipital is wider than long, with nearly and Kalasinemys from the Late Jurassic parallel lateral borders and more developed Phu Kradung Formation of NE Thailand tubercula basioccipitale which are more (Averianov et al., 2016 ; Rabi et al., 2014, distant from one another. The crista 2013, 2010 ; Sukhanov, 2000 ; Tong, 2017 ; supraoccipitalis in IVPP V16730 is relatively Tong et al., 2019, 2015a, 2015b). long as in Annemys, in contrast to a short crista in Xinjaingchelys. Kalasinemys differs IVPP V16730 is distinguished from from IVPP V16730 in having a very long xinjiangchelyids from the Sichuan Basin basisphenoid and a reduced basioccipital 8 Lu Li, Jialiang Zhang, Xiaolin Wang, Science Technology and Engineering Journal (STEJ) Yuan Wang & Haiyan Tong

(Tong et al., 2019). On the basis of these 7. Conclusions skull characters, IVPP V16730 is assigned to Annemys. The partial skeleton (IVPP V16730) from During the Middle-Late Jurassic, the Upper Jurassic Tiaojishan Formation of the Annemys was widespread in Qinglong, Hebei Province, China, described northern China and Central Asia. The in the present paper is the first turtle genus contains originally two : specimen reported from the Yanliao Biota. It is A. levensis and A. latiens, both from the assigned to Annemys sp. (Xinjiangchelyidae). Late Jurassic of Shar Teg, Transaltai Gobi, This discovery adds a new element to Mongolia (Rabi et al., 2014 ; Sukhanov, the poorly known freshwater vertebrate 2000 ; Sukhanov and Narmandakh, 2006). fauna of the Yanliao Biota and extends the Originally assigned to Xinjiangchelys, geographical distribution of Annemys to A. wusu from the late Qigu northeastern China. Further discoveries Formation of the Turpan Basin, China of more complete turtle material from has been referred later to Annemys the Yanliao Biota are expected for (Averianov et al., 2016 ; Rabi et al., 2013), a better understanding of the Yanliao the assignment being supported by the fauna and the evolutionary history of remnant of interpterygoid vacuity, the xinjiangchelyid turtles. narrow vertebral scutes 2-4 and the vertebral 1 wider than the vertebrals 2-4. A. sp. from 8. Acknowledgments the Bathonian Itat Formation of Berezovsk mine in western Siberia represents This research is supported by the Strategic the oldest record of the genus (Averianov Priority Reasearch Project (B) of the Chinese et al., 2016). The Qigu Formation of the Academy of Sciences (XDB18000000). We Turpan Basin has yielded a large number of thank two anonynous reviewers for their Annemys sp. skeletons (Wings et al., 2012). valuable comments. Addition material of A. sp. has been reported from the Middle-Late Jurassic Shishugou Formation (Callovian or Oxfordian) of the 9. References Junggar Basin and the Balabansai Formation, Fergana, Kirgizstan. (Brinkman et al., 2013 ; Averianov, A.O., Martin, T., Skutschas, Nessov & Kaznyshkin, 1985). Although P.P., Danilov, I.G., Schultz, J.A., Annemys contains the best material among Schellhorn, R., Obraztsova, E., Xijiangchelyidae, the study of many of the Lopatin, A., Sytchevskaya, E., specimens is still outstanding. Because of Kuzmin, I., Krasnolutskii, S. the preservation of our specimen, we assign & Ivantsov, S. (2016). Middle IVPP V16730 to Annemys sp. Annemys sp. Jurassic vertebrate assemblage of from Qinglong is the first xingjiangchelyid Berezovsk coal mine in Western turtle reported from NE China and also Siberia (Russia). Global Geology, represents the easternmost distribution of 19, 187-204. the genus Annemys. Volume 7, Number 1, January-June 2021 First turtle remains from the Middle-Late Jurassic 9 Yanliao Biota, NE China

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