First definitive stegosaur from the Middle Jurassic of Kyrgyzstan

0 Paläontologische Zeitschrift 00, Vol. /, p., --00 First definitive stegosaur from the Middle Jurassic of Kyrgyzstan ALEXANDER O. AVERIANOV, St. Petersburg; AIZEK A. BAKIROV, Bishkek & THOMAS MARTIN, Bonn with figures and table Introduction AVERIANOV, A.O.; BAKIROV, A.A. & MARTIN, T. 00. First definitive stegosaur from the Middle Jurassic of Kyrgyzstan. Paläontologische Zeitschrift ():, figs., tab., Stuttgart,.. 00. Abstract: Four articulated posterior dorsal vertebrae from the Middle Jurassic (Callovian) Balabansai Svita at Sarykamyshsai locality in the northern Fergana Valley represent the first record of Stegosauria in Kyrgyzstan. The vertebrae cannot be attributed to any known stegosaurian genus. This is one of the oldest records for the Stegosauria. Keywords: Stegosauria Middle Jurassic Callovian Balabansai Svita Fergana Valley Kyrgyzstan Tashkumyr Zusammenfassung: Vier artikulierte posteriore Dorsalwirbel aus der mitteljurassischen (Callov) Balabansai Svita von der Lokalität Sarykamyshsai im nördlichen Fergana-Tal stellen den ersten Nachweis von Stegosauria in Kirgistan dar. Die Wirbel lassen sich keiner bekannten Stegosaurier-Gattung zuordnen. Es handelt sich um einen der ältesten Nachweise der Stegosauria. Schlüsselwörter: Stegosauria Mittlerer Jura Callov Balabansai Svita Fergana-Tal Kirgistan Tashkumyr The Middle Jurassic Balabansai Svita in the Fergana Depression, Kyrgyzstan is a source of a diverse and interesting vertebrate fauna, albeit known mostly from fragmentary specimens (see review in AVERIANOV et al. 00, 00; MARTIN & AVERIANOV 00, 00). The dinosaur record is particularly incomplete, with the majority of discoveries represented by isolated teeth and bone fragments; a partial skeleton of the sauropod Ferganasaurus verzilini ALIFANOV & AVERIANOV, 00 collected in at the Balabansai locality near Tashkumyr is a notable exception. However, since 000, the situation has considerably improved: a joint field project and subsequent excavations led by one of us (AB) in 00 and 00 at two localities close to Tashkumyr (Sarykamyshsai and Urusay) have produced more complete and partially associated dinosaur remains. Here we describe the first of these findings, a series of associated posterior dorsal vertebrae of Stegosauria indet. from the Sarykamyshsai locality, representing the first definitive record of stegosaurs in the Balabansai Svita, as well as in Kyrgyzstan. Institutional abbreviation: IGB, Institute of Geology, National Academy of Sciences of Kyrgyz Republic, Bishkek, Kyrgyzstan. Systematic paleontology Order Ornithischia SEELEY, Suborder Thyreophora NOPCSA, Infraorder Stegosauria MARSH, Stegosauria indet. Figs. Material: IGB 00, four associated posterior dorsal vertebrae. Locality and horizon: Site FTA-, nearby site FTA- (see AVERIANOV et al. 00 for details), Sarykamyshsai locality, about four km SE of Tashkumyr, northern Fergana Depression, Kyrgyzstan. Upper (red-colored) part of Balabansai Svita, Middle Jurassic (Callovian). Description: The vertebrae are similar in morphology, except that the neural spine becomes progressively more posteriorly inclined through vertebrae one to four. The centrum is longer than high and wide (ACW/CL ratio Addresses of the authors: Alexander O. Averianov, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., St. Petersburg, Russia; e-mail <lepus@zin.ru>. Aizek A. Bakirov, M.M. Adyshev s Institute of Geology, National Academy of Sciences of Kyrgyz Republic, Erkindik, Bishkek 0, Kyrgyzstan; e-mail <aizek_bakirov@mail.ru>. Thomas Martin, Institut für Paläontologie, Universität Bonn, Nussallee, Bonn, Germany; e-mail <tmartin@uni-bonn.de> 00 00/0/00 $. 00 E. Schweizerbart sche Verlagsbuchhandlung, D 0 Stuttgart

First definitive stegosaur from the Middle Jurassic of Kyrgyzstan Fig.. IGB 00, four associated posterior dorsals of Stegosauria indet. in lateral view. Site FTA-, Sarykamyshsai, northern Fergana Depression, Kyrgyzstan; upper part of the Balabansai Svita, Middle Jurassic (Callovian). Numbers denote vertebrae. Scale bar = cm. %; see Tab. for measurements), with a deeply concave ventral margin in lateral view (Fig. ). Anterior and posterior intercentral articulation surfaces are slightly concave and circular to oval in outline, with their height equal to or slightly exceeding their width (Figs. c, e). The dorsal margin of the intercentral articulation surface is incised by the neural canal (Figs. c, e). Laterally, there is a pronounced longitudinal ridge on the centra of vertebrae three and four close to the neurocentral suture, flanking dorsally a poorly defined lateral depression (Figs., d). In ventral view the centrum is spool-shaped, without a ridge or a groove (Fig. a; a poorly developed ridge is present on the centrum of vertebra one). The neural arch pedicel is high,. (vertebra four) to. (vertebra three) times greater than the height of the centrum. The neural canal is pear-shaped (Figs. c, e) and higher on vertebra four than it is on vertebrae two and three ( % and % of the centrum height respectively). In lateral view the anterior and posterior margins of the neural arch pedicel converge towards the base of the diapophysis and zygapophyses (Figs., d). Two weakly defined laminae support the transverse process ventrally and converge towards the parapophysis. The prezygapophyses are united forming a U- shaped articulation surface (Figs. b, c). There is a longitudinal median ridge between the posterior end of the prezygapophyseal articular surface and the base of the neural spine (Figs. b, c). The postzygapophyses are also united, and are triangular in cross section (Fig. e). Below the postzygapophyses there is a prominent median ridge extending ventrally towards the dorsal margin of the neural canal (Fig. e). The parapophysis is large, oval shaped, and slightly convex, with almost no pedicle. It is located at the base of the transverse process (Figs., d). Its dorsal margin is below the dorsal margin of the prezygapophysis and approximately at the level of the ventral margin of the postzygapophysis. The transverse process is directed posterodorsally at º (vertebra four) and º (vertebra three) to the horizontal. The transverse process is triangular in cross section, with flat posterodorsal side and salient reinforcement rib along the anteroventral margin, connecting ventrally with the parapophysis. As in other stegosaurs (GALTON ; GALTON & UPCHURCH 00), the transverse processes unite anteromedially to form a vertical surface immediately posteriorly to the prezygapophyses (Fig. c). The diapophysis (complete only in vertebra two) is triangular and convex. The neural spine is plate-like and not noticeably higher than the diapophyses (Figs., c e). It is placed posteriorly on the neural arch, between the diapophyses and the posterior margin of the postzygapophyses. The anterior margin of the neural spine is slanting and slightly concave while the posterior margin is straight and almost vertical. Comparison: IGB 00 is referable to Stegosauria because in vertebra three the neural arch is. times the height of the centrum. According to SERENO & DONG 0

ALEXANDER O. AVERIANOV et al. 0 Fig.. IGB 00, posterior dorsal (vertebra ) of Stegosauria indet. in ventral (a), dorsal (b), anterior (c), lateral (d), and posterior (e) views. Site FTA-, Sarykamyshsai, northern Fergana Depression, Kyrgyzstan; upper part of the Balabansai Svita, Middle Jurassic (Callovian). Abbreviations: di, diapophysis; nc, neural canal; ns, neural spine; pa, parapophysis, prz, prezygapophysis; psz, postzygapophysis. Scale bar = cm () and GALTON & UPCHURCH (00), stegosaurs are characterized by neural arches that are at least. times the height of the centrum. As in other stegosaurs, except Dacentrurus LUCAS,, the centrum length is greater than the transverse centrum width (GALTON & UPCHURCH 00). The reverse condition is observed in a posterior dorsal vertebra of a stegosaur from the Early Cretaceous of Thailand (BUFFETAUT et al. 00: fig. ), although this has been accentuated by anteroposterior crushing of the specimen. BUFFETAUT et al. (00: ) considered this condition (centrum shorter than wide) to be normal for dorsal vertebrae of Stegosauria; this is obviously incorrect and apparently followed a misprint in GALTON (: ; compare with GALTON & UP- CHURCH 00: ). IGB 00 can be coded for three characters from the recent phylogenetic analysis of MAIDMENT et al. (00): (), transverse processes of dorsal vertebrae at high angle to horizontal (all stegosaurs); (), neural arches of posterior dorsal vertebrae elongated a great deal above neural canal (all stegosaurs except Huayangosaurus DONG, TANG & ZHOU, and Hesperosaurus CARPENTER, MILES & CLOWARD, 00); and ( or ), prezygapophyses fused on midline in middle and posterior dorsals (), or in all dorsals () [state : Huayangosaurus and Kentrosaurus, state : Hesperosaurus, Stegosaurus and ankylosaurs]. The Balabansai stegosaur is similar to other stegosaurs except Hesperosaurus because it has a relatively low neural spine, projecting not far above the level of the diapophyses (CARPENTER et al. 00). Pedicel height of IGB 00 falls between the low and medium neural arch categories of CARPENTER et al. (00: fig..). The Balabansai

First definitive stegosaur from the Middle Jurassic of Kyrgyzstan Tab.. Measurements (in mm) of IGB 00, posterior dorsal vertebrae of Stegosauria indet. Site FTA-, Sarykamyshsai, Fergana Depression, Kyrgyzstan; upper part of the Balabansai Svita, Middle Jurassic (Callovian). Abbreviations: ACH, anterior height of centrum (without hypapophysis); ACW, anterior width of centrum; ANW, anterior width of neural arch (between lateral margins of prezygapophyses); CL, centrum length (ventral, without odontoid process for axis); NAL, neural arch length (between anterior and posterior margins of ventral floor of neural canal); NSL, neural spine length (maximum); PCH, posterior height of centrum; PCW, posterior centrum width; PNW, posterior width of neural arch (between lateral margins of postzygapophyses). Vertebra stegosaur is more derived than Huayangosaurus, Lexovisaurus HOFFSTETTER,, and the Phu Kradung stegosaur (BUFFETAUT et al. 00) because it lacks the ventral notch between the postzygapophyses, according to GALTON () and GALTON & UPCHURCH (00). IGB 00 bears no characters or character combinations that allow referral to any known stegosaurian genus. The specimen bears no autapomorphies or unique character combinations and is therefore regarded as Stegosauria indet. The vertebrae IGB 00 belong to a rather large stegosaur, some % larger than Stegosaurus stenops MARSH, from the Late Jurassic of North America, reaching approximately m in length (GILMORE ; GALTON & UPCHURCH 00). Discussion Asiatic stegosaurs are known predominantly from China (a possible Indian Late Cretaceous stegosaur Dravidosaurus YADAGIRI & AYYASAMI, is not considered here because at that time India was not part of the Asiatic continent). A number of stegosaurian genera were described from the Jurassic and Early Cretaceous of China but several have not survived a thorough alpha-level taxonomic revision (MAIDMENT 00; MAIDMENT & WEI 00). Discoveries of stegosaurs in other parts of Asia were made as early as (Tuva), 0 (Yakutia), or (Mongolia), but were recognized and/or published only recently (see the list below). Recent discoveries of stegosaurs in the Jurassic and Early Cretaceous of Siberia, together with the new finding in Kyrgyzstan show that stegosaurs were a constant albeit rare element of dinosaurian faunas of these times. Below we summarize the record of Asiatic Stegosauria (see also Fig. ). () Dashanpu Quarry, Sichuan Province, China; Lower Shaximiao Formation, Middle Jurassic (?Bajocian-Bathonian). Complete and fragmentary skeletons of Huayangosaurus taibaii DONG, TANG & Measurements ACH ACW ANW CL NAL NSL PCH PCW PNW # 0... ~. #...0. ~...0. #..0...0.... #.0. 0.... ZHOU, (DONG et al. ; DONG ; SE- RENO & DONG, ; MAIDMENT et al. 00). () Berezovsk Quarry, Krasnoyarsk Territory, Russia. Itat Formation, Middle Jurassic (Bathonian). Disarticulated bones of Stegosauria indet. (ALIFANOV et al. 00; AVERIANOV et al. 00b). () Sarykamyshsai, Kyrgyzstan; Balabansai Svita, Middle Jurassic (Callovian). Articulated series of posterior dorsals (this report). () Taipingstai, Yunghsing, Sichuan Province, China; Upper Shaximiao Formation, Middle Jurassic (?Callovian). Fragmentary skull and skeletons of Chialingosaurus kuani YOUNG, (YOUNG ; DONG ). This taxon is invalid according to MAIDMENT & WEI (00). The age of the Upper Shaximiao Formation is often and without any arguments considered as Upper Jurassic, Oxfordian (e.g., WEISHAMPEL et al. 00). The vertebrate fauna from this formation is more likely to be Middle Jurassic in age (see LUCAS and AVERIANOV et al. 00 for review). Invertebrates from this stratigraphic unit also suggest a Middle Jurassic age (CHEN ). () Wujiabai, Zigong City, Sichuan Province, China; Upper Shaximiao Formation, Middle Jurassic (?Callovian). Partial skull and skeleton of Tuojiangosaurus multispinus DONG, LI, ZHOU & CHANG, (DONG et al. ; DONG ; MAIDMENT & WEI 00). () Mia-esni, Chongqing Municipality, China; Upper Shaximiao Formation, Middle Jurassic (?Callovian). Incomplete skull and skeleton of Chungkingosaurus jiangbeiensis DONG, ZHOU & CHANG, and partial skeletons of Stegosauria indet. (DONG et al. ; DONG ; MAIDMENT & WEI 00). () Pengtang, Zigong City, Sichuan Province, China; Upper Shaximiao Formation, Middle Jurassic (?Callovian). Partial skeleton of Gigantspinosaurus sichuanensis OUYANG, (OUYANG ; MAIDMENT & WEI 00). 0

ALEXANDER O. AVERIANOV et al. 0 Fig.. Map of Asia showing records of Stegosauria. Symbols: asterisk, Middle Jurassic localities; triangle, Late Jurassic localities; quadrate, Early Cretaceous localities. Localities:, Dashanpu Quarry;, Berezovsk Quarry;, Sarykamyshsai;, Taipingstai;, Wujiabai;, Mia-esni;, Pengtang;, Yingshan;, unspecified locality for Changtusaurus laminaplacodus [nomen nudum], not shown on the map;, Jiangjunmiao;, Ban Khok Sanam;, Kalbak Kyry;, Lawulashan;, Bol shoi Kemchug;, Urho;, Yangpo;, Shestakovo ;, Khamryn Us;, Kempendyai. () Yingshan, Sichuan Province, China; Upper Shaximiao Formation, Middle Jurassic (?Callovian). Complete skull and skeleton Yingshanosaurus jichuanensis [nomen nudum] (ZHAO ; ZHOU & TANIMOTO ; MAIDMENT & WEI 00). () Unspecified locality, China; Dabuka Group, Middle Jurassic. Changtusaurus laminaplacodus [nomen nudum] (ZHAO ; MAIDMENT & WEI 00). () Jiangjunmiao, Xinjiang-Uygur Autonomous Region, China. Shishugou Formation, transitional

First definitive stegosaur from the Middle Jurassic of Kyrgyzstan Middle Upper Jurassic (Callovian Oxfordian). Partial skull and skeleton of Stegosauria indet. (CLARK et al. 00). An isolated stegosaur dorsal vertebra from an unspecified locality in the southern part of the Junggar Basin within the Qigu Formation (Oxfordian) is described by WINGS et al. (00). () Ban Khok Sanam, Thailand. Phu Kradung Formation, Upper Jurassic. Isolated vertebra of Stegosauria indet. (BUFFETAUT et al. 00). () Kalbak Kyry, Tuva, Russia. Saldam Formation, Upper Jurassic. Partial braincase and isolated postcranial elements of Stegosauria indet. (ALIFANOV et al. 00; AVERIANOV et al. 00). () Lawulashan, Tibet, China. Loe-ein Formation, Upper Jurassic or Lower Cretaceous. Fragmentary skeleton of Monkonosaurus lawulacus ZHAO in DONG (ZHAO ; DONG ). This taxon is invalid according to MAIDMENT & WEI (00). () Bol shoi Kemchug, Krasnoyarsk Territory, Russia. Ilek Formation, Lower Cretaceous (Hauterivian? Barremian?). Rare teeth of Stegosauria indet. (AVERIANOV et al. 00, 00a; LESHCHINSKIY et al. 00). () Urho, Xinjiang-Uygur Autonomous Region, China. Hutubihe Formation, Tugulu Group, Lower Cretaceous (Barremian). Fragmentary skeleton of Wuerhosaurus homheni DONG, (DONG, ). () Yangpo, Inner Mongolia, China; Ejinhoro Formation, Lower Cretaceous (Barremian). Incomplete skeletons of Wuerhosaurus ordosensis DONG, (DONG ). This taxon is considered undiagnostic [nomen dubium] by MAIDMENT (00). () Shestakovo, Kemerovo Province, Russia. Ilek Formation, Lower Cretaceous (Albian Aptian). Rare teeth of Stegosauria indet. (AVERIANOV et al. 00; LESHCHINSKIY et al. 00). () Khamryn Us, Mongolia; Khukhtyk or Dzunbain Formation, Lower Cretaceous (Aptian Albian). Fragmentary skeleton of Stegosauria indet. (BARS- BOLD et al. ; ALIFANOV et al. 00). () Kempendyai, Yakutia, Russia. Sangar Series, Lower Cretaceous. Isolated teeth and postcranial elements of Stegosauria indet., initially assigned to ankylosaurs (ROZHDESTVENSKY ; MAR- YASKA ; NESSOV ; KURZANOV et al. 000, 00). The earliest skeletal records for Stegosauria are from the Bajocian Bathonian of Sichuan Province, People s Republic of China, while tracks attributable to these dinosaurs are known from Early Jurassic rocks of Europe and North Africa (GALTON & UPCHURCH 00). Abundance and diversity of stegosaurs in the Middle Jurassic of Asia may suggest an Asiatic origin for the group. Currently, the Balabansai stegosaur is the most western record for the Stegosauria in Asia (Fig. ). Acknowledgments We are grateful to reviewers Susannah Maidment (University of Cambridge) and Kenneth Carpenter (Denver Museum of Nature and Science) for constructive criticism and helpful suggestions. 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