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A Large Theropod Metatarsal from the Upper Part of Jurassic Shishugou Formation in Junggar Basin, Xinjiang, China

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じ51ࢣȞじ1᱋ ऐ㘶Ḻߔ➕႒៑ pp. 29-42 2013Ꭰ1ᰴ VERTEBRATA PALASIATICA figs. 1-4 A large theropod metatarsal from the upper part of Jurassic Shishugou Formation in Junggar Basin, Xinjiang, China HE Yi-Ming1, 3ȞJames M. CLARK2ȞXU Xing1 (1 Key Laboratory of Vertebrate Evolution and Human Origin of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences Beijing 100044 [email protected]) (2 Department of Biological Sciences, George Washington University Washington DC 20052, USA) (3 University of Chinese Academy of Sciences Beijing 100049) Abstract The Shishugou Formation in the Junggar Basin was deposited during the late Middle Jurassic to the early Late Jurassic. It is known as a rich source of vertebrate fossils, including specimens of several different kinds of theropod. Here we report an isolated theropod left metatarsal IV (IVPP V 18060) from the upper part of the Shishugou Formation, exposed at the Wucaiwan Locality in the northeastern part of the Junggar Basin. Based on comparisons with the fourth metatarsals of other theropods, we refer this specimen to Allosauroidea. Similarities to the equivalent bone in other allosauroids include: the proximal end has a subtriangular outline with a tongue-like, posteriorly directed posteromedial process; the shaft has a gentle outward curve; a semi-lunate concave of muscle attachment is present on the posterolateral side of the shaft; and the cross-section of the shaft is triangular. Within Allosauroidea, V 18060 is most similar to the fourth metatarsal of Sinraptor dongi, a taxon that occurs in the same IRUPDWLRQLQWKH-LDQJMXQPLDRDUHDRIWKH-XQJJDU%DVLQ9GLVSOD\VVHYHUDOVSHFL¿F resemblances to S. dongi: the outline of the distal end is subtrapezoidal; the depression for muscle attachment on the posterior side of the shaft that borders the shaft’s lateral side lacks a ZHOOGH¿QHGULGJH+RZHYHU9DOVRGLIIHUVLQVRPHUHVSHFWVIURPWKHIRXUWKPHWDWDUVDO of S. dongi: V 18060 is much more robust, and the lateral condyle on the distal end is smaller than the medial one. These morphological differences could be ontogenetic variation or due to sexual dimorphism, taxonomical variations. We prefer the interpretation that V 18060 is a new species that has a close relationship to S. dongi, and a cladistic analysis based on the morphology of metatarsal IV supports this inference. The discovery of V 18060 suggests that the theropods from the Shishugou Fauna are more diversified during the Mid-Late Jurassic than previously thought. The presence of different sinraptorid species in the neighboring areas of Wucaiwan and Jiangjunmiao points to the possibility that these two regions within the Junggar Basin were geographically isolated or ecologically distinct from one another during the Mid-Late Jurassic. Key wordsȞJunggar Basin, Mid-Late Jurassic, Shishugou Formation, theropod, metatarsal ఩ტ㜖♢⻽႒ദ䛽(ឥ۲ण喝40830210)স㒺఩఩ტ⻽႒ദ䛽(㑂ण喝EAR 0922187)䉰ߕȠ ᩢ⽫ᬑ᱋喝2011-10-25 30 ऐȞ㘶ȞḺȞߔȞ➕Ȟ႒Ȟ៑ 51ࢣ ᮄ⭚ޚొᇨⲚഄգ㔫㑾⷇ᷥ≳㒘Ϟ䚼ϔ໻ൟ ݑ㛮㉏䎪偼 ਫ਼1, 3bJames M. CLARK2b༗໲1ྡྷۯ ȞȞȞ(1 ͙఩⻽႒䮎ऐ㘶Ḻߔ➕̺ऐϦㆧⵀ⾢ᝬ喏͙఩⻽႒䮎㘶Ḻߔ➕⑀ࡂ̺Ϧㆧ䊣⎼䛹◥჊侸აȞࡃϘȞ100044) ȞȞȞ(2 㒺఩΀⇧·ࡺⰇ䶫๓႒⩋➕႒㈧ȞࡺⰇ䶫➥ࡦȞ20052) ȞȞȞ(3 ͙఩⻽႒䮎๓႒ȞࡃϘȞ100049) 㙆ㆧᕼک䓳࠱᠘ๆ⻹ܦᨬ㽕喝۲஢ᅀ⯲౜͙һ㒃͂ᮆ᱋ݜᮆһ㒃͂ᬕ᱋↵⼛⮰ⴟᴽ⇋㏰ϓ ⇋⮰๓䛻㘶Ḻߔ➕ࡂⴟȠ᣻䔜β̬Т᫜⮰䛳㜖۲஢ᅀ⯲౜͈ࡃ㑄πᒕ⎪౜ࡦⴟᴽڱ哅౔ 㙆ㆧᕼ哅じఇ䌂俔ᄥکЂڢ㙆ㆧᕼ哅ጒじఇ䌂俔ᴳ᱘(IVPP V 18060)Ƞ䕆䓳̺ک㏰̶䘔⮰ Ђᐮ➥哅䊱⻽じఇ䌂俔⮰ⰤѨᕓ࠱ڢᐮ➥哅䊱⻽(Allosauroidea)ȠႯ̺ڑ喏䄑ᴳ᱘छᑾ℀ 喏ऽโӓ䒧ᓚᑛᰞ⮰俔᎞喏ѹκ俔᎞ܦ⾭⟣㞮䲎̵㻾ᒎᎢᰵ̬ऽऺ᫥ᐢѤ⮰㜸ڟ〚᠘喝䓽 ऺ᫥䲌䓽โӓ䓥⮰ࡶᰴᒎ㖸㖵ܥ䮣喏̵㻾ᒎ俔᎞ὖ᝖䲎Ƞ౔ᐮ➥哅䊱⻽ᑿ͙喏V 18060̺ Ꮕ౜ࡦⴟᴽ⇋㏰͙ࣽ⣜⮰㦏℻͙ࡺⰃ哅(Sinraptor dongi)ᰬͦⰤѨ(䮐βБۇ۲஢ᅀ⯲౜ᄲ 㞮䒚᏿ॴ䓽ᷛᒎ喏俔᎞ऺ䘔㖸ڟ〚ⰤѨᕓ喏V 18060স㦏℻͙ࡺⰃ哅⮰ⰤѨᕓ䔄࠱᠘䔈̶ 㖵ܥ䮣̺ঔఠ䓥⩸ॴࡶᐬᩪ⟢ᔭ)ȠѲ᭛喏V 18060স㦏℻͙ࡺⰃ哅΋ႄ౔ⱬ̬χᬺ᭪⮰ࡦ โ͐俭๓ᄻጚᐮⰤࣹȠ䔅χᒎڱ〚ݗ喝V 18060ᬺ᭪℀㦏℻͙ࡺⰃ哅⮰じఇ䌂俔㇃บ喏䔈 ᔭጚᐮछ㘩᭛⩝κ͖ѿࣽ㗞ᝂ㔱ᕓࣸᒎ䕌᜼⮰喏΋छ㘩Џ㶔ܲㆧ႒ጚᐮȠᄥ℀ⵀ⾢সദκ じఇ䌂俔ᒎᔭԍᖛ䔇㵸⮰ܲᩛ㈧㐋႒ܲ᲼㏿᳈ᰠջऽऺ̬⻹㼏䛶喝V 18060Џ㶔̬͖̹स 㙆ㆧᕼ哅Ƞ᫜ᱼ᫅⮰ࣽ⣜්ߌβ͙ᮆһ㒃ک㈧ᒴ䓽⮰᫜⮰ڟκ㦏℻͙ࡺⰃ哅Ѳ̺ऺ㔱Ϟ㑄 Ꮕ౜ࡦࣽ⣜̹स⮰͙ࡺⰃ哅ㆧᕼ哅ۇ㙆ㆧᕼ哅⮰ܲᐮᏒȠ౔πᒕ⎪সᄲکⴟᴽ⇋ߔ➕㓐͂ ᠳ⹦βᮆһ㒃͂ᬕ᱋۲஢ᅀ⯲౜͙Ⱔ䗧౜ࡦछ㘩ႄ౔ⱬ̬჆⮰⩋ᔭܲᐮᕓᝂ㔱౜⤲䯀⻧Ƞ 㙆ㆧᕼ哅喏䌂俔ک䬂䆡喝۲஢ᅀ⯲౜喏͙ᮆһ㒃͂喏ⴟᴽ⇋㏰喏݇ Ё೒⊩ߚ㉏ো喝Q915.864Ȟ᭛⤂ᷛ䆚ⷕ喝AȞ᭛ゴ㓪ো喝1000-3118(2013)01-0029-14 㜖2000Ꭰ㜟϶喏͙఩⻽႒䮎ऐ㘶Ḻߔ➕̺ऐϦㆧⵀ⾢ᝬ̺㒺఩΀⇧ࡺⰇ䶫๓႒㏰ ᭛ᄥ۲஢ᅀڢ᜼⮰㖀ऴ㔯ᄋ䭋̬Ⱐ౔᫜⪲۲஢ᅀ⯲౜স़৴⯲౜ᐬᆁ⻽႒㔯ᄋ≧ߔ喏ᅐ ⮰䄯ᴑ喏䛳䯲ݜ๓䛻⮰౜䉔႒ڑ⌝䱞⮰ⴟᴽ⇋㏰䔇㵸βܦ∇Ꮕ౜ࡦᎫۇ⯲౜πᒕ⎪সᄲ ⮰៑䕿⋡Ⰲβ哋刂ㆧȟ凰ᒎߔ➕ȟѨਦΟ❘㵸ߔ➕ȟਦΟߔ➕ȟڟসऐ⩋➕႒䉰᫅喏Ⱔ ⵀ⾢Ჭ๓͜ჸβ᜽ڱ⮰აڟ㔨哅সᕼ哅6͖ㆧ㓐⮰15͖᫜ᆊ⻹(㶔1)Ƞ䔅χ䛺โ጑҈সⰤ Иᄥκⴟᴽ⇋㏰⮰䃐䃲(Eberth et al., 2010), ΋ӯ䔇β᜽Иᄥκ㠑᎞䛹㺭ᕼ哅ㆧ㓐ᬕ᱋⑀ ࡂ⮰䃐䃲(Clark and Xu, 2009a; Xu et al., 2010)Ƞ 䱞κ۲஢ᅀ⯲౜͈㑄⮰䭲Ⱔ↵⼛喏᪠ऴκ̶һ㒃㐋㺫ᆝ⾽㏰ܦⴟᴽ⇋㏰᭛̬຃ ͷ̶喏፤㷗़䅣冭㓐ճ᪠ऴᝬ㺲Ƞⴟᴽ⇋㏰ࣆ50~700 m, ͧ㺭ͦ㐫ȟ▜台স㏎㞞⴮䉔 ∑ᇕࣶ∑ᇕᅮ喏̷䘔๥⴮ᇕࣶ⵪ᇕȠ䄑㏰̬㝘㷗̶̷ܾܲͦ͐⃡喝̷䘔㷗䃐ͦЏ㶔 㙆ㆧ䌂俔 31ک1᱋ 䉦̬协ふ: ᫜⪲۲஢ᅀ⯲౜һ㒃㏖ⴟᴽ⇋㏰̶䘔̬๓ಷ ⪲㑂ༀц, 2000; ᫜ڤ͙һ㒃͂⮰̬຃↵⼛喏̶䘔Џ㶔ᮆһ㒃͂⮰̬຃↵⼛(͙఩౜ᅮ ㏰,1981)Ƞ䊡ૈ䔇ふ(1987)ദκ̶̷͐⃡ᇕᕓসࡂⴟ㏰ۅ㐠४ᅀ㜖⇧ࡦࡦഋ౜ᅮ㶔㑂 ,.ऴ⮰ጚᐮ喏ា̷⃡ࢁ⠘঩हͦπᒕ⎪㏰喏Ѳ䔅̬᫥ᵴ⇍ᰵᓃݜᎫ∇ᣑः(Eberth et al Ƞ䓽ᎠᲑ⮰ⵀ⾢㶔ᬺ喏ⴟᴽ⇋㏰⮰౜䉔ᎠЏ㮩♢Ϻ͙һ㒃͂ᐢ㐙ݜᮆһ㒃͂喏(2001 䘔ᅬ䭼κ͙һ㒃͂ᰬᮆ᱋喏̶䘔ᅬ䭼κᮆһ㒃͂ᰬᬕ᱋̷ڢѲᐢ㐙ᬢ䬠Ⱔᄥᒴⴙ喏 (Eberth et al., 2010)Ƞ ⴟᴽ⇋㏰ჸॗ⵱ࡂ᱔ȟᬌ㘶Ḻߔ➕স㘶Ḻߔ➕ࡂⴟ喏Ѳ̶䘔ࡂⴟ䔈℀̷䘔͜ჸȠ ᝖㜟᱘᪳Ⴘ᜼喏ጞ㏻៑䕿⮰㘶Ḻߔ➕ࡂⴟᆊ⻹࠱᠘ѨਦΟ❘㵸ߔ➕ȟਦΟߔ➕ȟ哋刂 27⻹(㶔1)Ƞڝ㙆ㆧکㆧȟ凰ᒎㆧȟ㔨哅ㆧȟ卋㛬ㆧȟ㱑㙆ㆧБࣶ 㸼 1ǂ⷇ᷥ≳㒘Ꮖ᡹䘧ⱘЏ㽕㛞Ợࡼ⠽࣪⷇ Table 1ȞList of the reported vertebrates of Shishugou Formation Taxa Species Reference Horizon Locality Xinjiangchelys junggarensis Yeh, 1986 Upper Jiangjunmiao X. macrocentrale Brinkman et al., 2012 ? Wucaiwan 哋刂ㆧ X. radiplicatoides Brinkman et al., 2012 Upper Wucaiwan Testudines Annemys sp. Brinkman et al., 2012 ? Wucaiwan X. chowi Matgke et al., 2005 ? Wucaiwan Sunosuchus junggarensis Wu et al., 1996 Lower Wucaiwan 凰ᒎㆧ Junggarsuchus sloani Clark et al., 2004 Lower Wucaiwan Crocodylomorpha Nominosuchus matutinus Clark & Xu, 2009b Upper Wucaiwan Yuanotherium minor Hu et al., 2009 Upper Wucaiwan ႀㆧک Bienotherium zigongensis Sun & Cui, 1989 Lower Jiangjunmiao Therapsida B. ultimus Maisch et al., 2004 Upper Jiangjunmiao Gongbusaurus wucaiwanensis Dong, 1989 Upper Wucaiwan 卋㛬ㆧ Jiangjunosaurus junggarensis Jia et al., 2007 Upper Jiangjunmiao Ornithischia Yinlong downsi Xu et al., 2006a Upper Wucaiwan Tienshanosaurus chitaiensis Young, 1937 Upper ? Mamenchisaurus sinocanadorum Russel & Zheng, 1993 Upper Wucaiwan 㱑㙆ㆧ Bellusaurus sui Sauropoda Dong, 1990 Lower Jiangjunmiao Klamelisaurus gobiensis Zhao, 1993 Lower Jiangjunmiao Monolophosaurus jiangi Zhao & Currie, 1993 Lower Jiangjunmiao Limusaurus inextricabilis Xu et al., 2009 Upper Wucaiwan Sinraptor dongi Currie & Zhao, 1993 Upper Jiangjunmiao 㙆ㆧ Haplocheirus sollers Choiniere et al., 2010a Upper Wucaiwanک Theropoda Zuolong salleei Choiniere et al., 2010b Upper Wucaiwan Guanlong wucaii Xu et al., 2006b Upper Wucaiwan 㔨哅ㆧ Sericipterus wucaiwanensis Andres et al., 2010 Upper Wucaiwan Pterosauria ਦΟㆧ Klamelia zhaopengi Chow & Rich, 1984 Upper and lower Laoshangou Mammal Acuodulodon sunae Hu et al., 2007 Upper Wucaiwan ȞȞ ⴟᴽ⇋㏰⮰㘶Ḻߔ➕ࡂⴟ㏰ऴь㐋̶㷗䃐ͦস᜽఩㺫ࢃ౜ࡦ͙ᮆһ㒃͂⮰⇅⏖Ꮕ ߔ➕㓐䲊፤ⰤѨȠ䓽ᎠᲑ⮰᫜ࣽ⣜͜ჸβ᜽Иᄥκⴟᴽ⇋㏰㘶Ḻߔ➕ࡂⴟ㏰ऴ⮰䃐䃲 ➥⠘᫜⪲۲஢ᅀ⯲౜ⴟᴽ⇋㏰ԉႄβ̬͖ܦClark et al., 2006; Clark and Xu, 2009b), ᭪⹦) ⮰͙ᮆһ㒃͂䓳⍍᱋⮰ߔ➕㓐喏౔䔅䛸㷗₏ᐻ঩हͦⴟᴽ⇋ߔ➕㓐ȠϺⴟᴽ⇋㏰ጞ㏻ϓ ࡂⴟ⮰ᗱۡⰷ喏ⴟᴽ⇋ߔ➕㓐̺⇅⏖Ꮕߔ➕㓐ᰵⱬᬺ᭪⮰ࡦݗȠⴟᴽ⇋ߔ➕㓐͙㻾呧ܦ 32 ऐȞ㘶ȞḺȞߔȞ➕Ȟ႒Ȟ៑ 51ࢣ 哅ㆧ(Alvarezsauroidea)Бࣶ㻾ڥ⨒哅ㆧ(Ceratosauria)ȟᯠ哅ㆧ(Tyrannosauroidea)ȟ䭫ᅀ 哅ㆧ(Ceratopsia)ࡂⴟ౔⇅⏖Ꮕߔ➕㓐͙⇍ᰵ䃜ᒁ(Peng et al., 2005)Ƞⴟᴽ⇋㏰͙䔇ₑ⮰ 㮆俔哅ㆧ(Coelurosauria)স㻾哅ࡂⴟ⮰ࣽ⣜̺͙఩͈ࡃ䘔͙ᮆһ㒃͂⮰ࡂⴟ㏰ऴᰠͦⰤ Ѩ(Xu et al., 2010), ᭪⹦ⴟᴽ⇋ߔ➕㓐Ϻ⩋➕౜⤲ܲࡦ̶छ㘩ᰠᣑ䓽͙఩͈ࡃ౜ࡦ喏㶔ᬺ ͙఩һ㒃㏖͙ᮆ᱋ᕼ哅ߔ➕㓐⮰㏰᜼छ㘩ᰠๆ౜ः㏘Ꮢᣓݢ喏౔ࢃࡃ᫥⮰ጚᐮᰠ⩆κ͈ 䲎সڔ㺫᫥⮰ጚᐮ(䔅䰬㺭ऐ⩋➕౜⤲႒᫥䲎ᰠ䔇̬ₑ⮰侸䃭)Ƞᄥκⴟᴽ⇋ߔ➕㓐⮰ᰠ ڔ⮰䃐䃲喏ᄲᰵߕκ᜽Иᰠߌ۲⶚౜ู͙࣋఩͙ᮆһ㒃͂ᕼ哅ߔ➕㓐⮰㏰᜼সᄥκڑ⌝ ⤯ᕼ哅⩋➕౜⤲ࡦ㈧⮰䃐䃲Ƞ 㙆ㆧᕼ哅ᴳ᱘(IVPPک᱘᪳ᄲ៑䕿䛳㜖۲஢ᅀ⯲౜πᒕ⎪౜ࡦⴟᴽ⇋㏰̶䘔⮰̬Т 㙆ㆧᕼ哅喏ఌکV 18060)Ƞᅩネ䄑ᴳ᱘ϱԉႄβじఇ䌂俔喏ѲႯᰵछ㘩Џ㶔β̬⻹᫜⮰ ₐ͜ჸβⴟᴽ⇋ߔ➕㓐⮰㏰᜼Ƞ᱘᪳ᄲᄥ䄑ᴳ᱘䔇㵸ᒎᔭ႒᣻䔜喏ܲ᲼Ⴏ⮰㈧㐋ࣽ㗞ѹ 㒚喏Ꭲ䃔䃦䔅̬ࣽ⣜⮰ᘻ͵Ƞ 1Ȟᴳ᱘᣻䔜 IVPP V 18060ϱԉႄβ̬Ⴘ᪠⮰ጒӓじఇ䌂俔(ప1), 䪫㏒355 mm, 俔᎞͙⃡ὖऽⰠ 㙆ㆧᕼ哅じఇ䌂俔Ⱔ℀喏V 18060䲊፤㇃บȠکЂڢᒰ㏒67 mmȠ̺๓ᄻⰤ䓽⮰ 㞮䲎䓽̵㻾ᒎ喏ᰬ๓ὖऽⰠᒰ㏒135 mm, ᬺ᭪๓κڟ〚V 18060䓽〛ᬺ᭪㛔㗬Ƞ䓽 ᐢ喏Ꭲ౔〛䘔ऽڱ◴㑄ऺࡶӓᑦڱ㞮䲎ڟ〚䮣Ƞ䓽ܥӓ㑄ڱᰬ๓ݹऺऽჩᏒȠ䓽〛㻲喏 ऺӓᐢѤȠ ݹ㻲喏俔᎞ᓚऽโӓᑛᰞ喞ӓ㻲喏俔᎞䒯ͦ᎟ⰠȠ俔᎞ὖ᝖䲎ᕧѿॴ᝭̵㻾ᒎ喞 ӓ䲎౔䲌䓽䓽〛ะᰵ̬̺じ̵䌂俔䓽〛ะⰤ䔊ڱ䊶䲌䓽䓽〛喏ݹऺऽ䪫Ꮢ䊶⾰Ƞ俔᎞ ᣑ⮰᫈ऽ䓽̵㻾ᒎܥ䮣喏ὖऽჩȠ俔᎞ऺӓ䲎᎟ಒ喏Ѳ౔͙䬠ᰵ䒧ᓚᡐࢷᒎ᜼⮰⇋ 〚Ἡ喏⇋ἩϺ䓽〛̬ⰠᐢѤݜᣑ䓽䔈〛ะ喏䪫Ꮢ㏒ࢌ俔᎞䪫Ꮢ⮰3/4Ƞ俔᎞ऺӓ䲎⮰䔈 ܥ䮣喏छ㘩Џ㶔じ̵সじఇ㙆䋪ᅴ㖸(M. lumbricalis)䭰ⱬะȠ䄑ܥ䓽ࡶᰴᒎ̬ڣ1/2ะ 䮣̶䘔䓥㑄ⵠᢋ喏⣜ႄᰬ䪫Ⱐᒰ㏒95.4 mm, ჩ㏒35 mmȠܥ䮣䲌䓽俔᎞โӓ䲎喏̺ঔ ఠ⩸䭼⌱ᮜ喏̺俔᎞โӓ䓥㑄ᣑ㼒ፒᰵ̬㏡ऽᐢѤ⮰㘶喏㘶౔䓽〛ะႄ౔㑦ए喏ҫܥ 䲊፤㇃㈅喏ᰵ๓䛻䭰ⱬ㖸㖵⮰㏲ᄻ㹢⯝̺ڱ䮣ܥ䮣Ⱔᄥ俔᎞ऺ䘔㶔䲎ॴࡶᐬᩪ⟢ᔭȠ ܥ౽Ƞ ڟ㞮๠䔈〛㻲ॴ䓽ᷛᒎ喏ὖऽჩᏒ⼹๓κݹऺऽ倄ᏒȠϱ౔ऺӓᰵ䨜䨪ڟ〚䔈 ӓ俭䓽㘶⟢喏โӓ俭ὖऽჩᏒڱ㞮䲎⮰͙䘔Ƞڟ〚โӓ俭⮰⇋ἩखᐢѤݜ䔈ڱ㞮喏ܲ䯀 ӓ䴓ፒ⿉๓κโӓ䴓ፒ⿉喏Ꭲ℀ऺ㔱⌝Ƞڱӓ俭Ƞӓ䴓ፒ⿉ࣽ㗞喏ڱᬺ᭪๓κ 2Ȟ℀䒯̺䃔䃦 㙆ㆧᕼ哅࠱᠘ࢁ㘶哅Monolophosaurus (Zhaoک۲஢ᅀ⯲౜ⴟᴽ⇋㏰͙ጞࣽ⣜⮰ 哅Guanlong (Xuی ,(and Currie, 1993), ͙ࡺⰃ哅Sinraptor (Currie and Zhao, 1993 et al., 2006b), ∑⒙哅Limusaurus (Xu et al., 2009), ガ᝷哅Haplocheirus (Choiniere 㙆ㆧ䌂俔 33ک1᱋ 䉦̬协ふ: ᫜⪲۲஢ᅀ⯲౜һ㒃㏖ⴟᴽ⇋㏰̶䘔̬๓ಷ ,(ӓ㻲(A)ȟโӓ㻲(B)ȟऺ㻲(C)ȟݹ㻲(D)ȟ䓽〛㻲(E)স䔈〛㻲(Fڱప 1ȞIVPP V 18060ጒӓじఇ䌂俔 ㈧⹦ᘻపڟ㞮䲎Ⱔξڟ〚Бࣶ㦏℻͙ࡺⰃ哅じఇ(G)ȟじ̵(H)সじθ(I)䌂俔䓽 Fig. 1ȞLeft metatarsal IV of IVPP V 18060, in lateral (A), medial (B), posterior (C), anterior (D), proximal (E) and distal (F) views, and left metatarsus of Sinraptor dongi, proximal views of metatarsals IV (G), III (H), and II (I)(from Currie and Zhao, 1993) a. the concave with Met III; b. the depression for muscle (M. lumbricalis) attachment on the posterior side of the shaft; A, B, C, D scale bar=10 cm; E, F scale bar=5 cm 34 ऐȞ㘶ȞḺȞߔȞ➕Ȟ႒Ȟ៑ 51ࢣ et al., 2010a)সጒ哅Zuolong (Choiniere et al., 2010b), Бࣶ͙ࡺⰃ哅ㆧ⮰➅咫ᴳ᱘ IVPP V 15310 (Xu and Clark, 2008)ȠV 18060⼹ᄻκ㦏℻͙ࡺⰃ哅₏ಷᴳ᱘(㶔2), ΋ᄻ 㙆ㆧᕼکκV 15310(ᵥᢚ➅咫ᴳ᱘⮰๓ᄻᄥ℀喏छⴑV 15310ͦⰚݹጞ៑䕿һ㒃㏖ 㙆ㆧᕼ哅ȠکЂڢ哅͙ᅦᄤᰬ๓⮰), Ѳᬺ᭪๓κ۲஢ᅀ⯲౜һ㒃㈧౜ᅮ͙ጞࣽ⣜⮰ 㸼 2ǂIVPP V 18060Ϣ㨷⇣Ёढⲫ啭IVPP V 10060㄀ಯ䎪偼Ⳍ᭄݇᥂ᇍ↨ Table 2ǂComparison of left metatarsal IV of IVPP V 18060 and Sinraptor dongi IVPP V 10060 (mm) Length Middle transverse width Minimum transverse Proximal Distal Minimum Specimen (L) of shaft in anterior view width of shaft in width width circumference C/L C/M (M) anterior view (C) IVPP V 18060 355 67 56 132 79 179 0.50 5.3 Sinraptor dongi 375 47 43 109 61 145 0.39 8.0 IVPP V 10060 㙆ㆧک⮰12⻹ڱβᰠ䔇̬ₑ⶚䃐V 18060⮰㈧㐋ѹ㒚喏᜽Иᄲ࠱᠘V 18060౔ͦ ᕼ哅ᐦ⿷β̬͖ϱദκじఇ䌂俔➥ᒭ⮰➥ᒭ݃㶔স⩔κ㈧㐋ࣽ㗞ܲ᲼⮰ⴕ䭡(࠱᠘12 স14͖➥ᒭ)Ƞ᜽И⩔ܲᩛ㈧㐋႒䒛ТTNT (Goloboff et al., 2008)ᄥ䄑ⴕ䭡ٯㆧࢁ͖ܲ 䔇㵸βܲ᲼Ƞᝬᰵ➥ᒭ౳ͦᬌᎻসふᱯ䛹喏䓼⩔ज़ࣽᐻ᥈㉎喏䛹ู1000⁍喏⃻⁍ԉ 㙆ㆧ㈧㐋ࣽ㗞ⵀ⾢⮰㏿䃦کڟЂᝬᰵ࣮᪜䛳⩔㑦Ⱝ䃪㒚ȠӉᢚݹϦᰵڢႄ10ḡᴽ喏 Weishampel et al., 2004), ᜽И䔵᠕ࣸ㘶哅(Dilophosaurus)҈ͦโㆧ㓐Ƞ㔯㭽ݜ㻾呧哅) ࢁ㈧ᕓ䔇㵸βᑦݢᕓ䃪㒚Ƞܲ᲼㏿᳈᭪⹦喏ᐮ➥哅ȟڢㆧ⮰ࢁ㈧ᕓᓃݜᎫ∇䃐छ喏ᄥ ͙V 18060̺㦏℻͙ࡺڢ㈧ᒴ䓽⮰ᩛ㈧喏ڟस㏰᜼̬͖ᒨₐڝ㦏℻͙ࡺⰃ哅̺V 18060 ㈧ᰬͦჲܳȠܲ᲼⮰㏿᳈ϓ⩋β1͖ᴽ(ᴽ䪫L=31, ̬㜠ᕓᠳ᪜CI=0.548, ԉ⪅ᠳڟⰃ哅 ᪜RI=0.576)(ప2)Ƞ 㙆ㆧᕼ哅ᩛᎻపکప 2Ȟᵥᢚじఇ䌂俔➥ᒭᝬֆ⮰ Fig.
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  • 8. Archosaur Phylogeny and the Relationships of the Crocodylia

    8. Archosaur Phylogeny and the Relationships of the Crocodylia

    8. Archosaur phylogeny and the relationships of the Crocodylia MICHAEL J. BENTON Department of Geology, The Queen's University of Belfast, Belfast, UK JAMES M. CLARK* Department of Anatomy, University of Chicago, Chicago, Illinois, USA Abstract The Archosauria include the living crocodilians and birds, as well as the fossil dinosaurs, pterosaurs, and basal 'thecodontians'. Cladograms of the basal archosaurs and of the crocodylomorphs are given in this paper. There are three primitive archosaur groups, the Proterosuchidae, the Erythrosuchidae, and the Proterochampsidae, which fall outside the crown-group (crocodilian line plus bird line), and these have been defined as plesions to a restricted Archosauria by Gauthier. The Early Triassic Euparkeria may also fall outside this crown-group, or it may lie on the bird line. The crown-group of archosaurs divides into the Ornithosuchia (the 'bird line': Orn- ithosuchidae, Lagosuchidae, Pterosauria, Dinosauria) and the Croco- dylotarsi nov. (the 'crocodilian line': Phytosauridae, Crocodylo- morpha, Stagonolepididae, Rauisuchidae, and Poposauridae). The latter three families may form a clade (Pseudosuchia s.str.), or the Poposauridae may pair off with Crocodylomorpha. The Crocodylomorpha includes all crocodilians, as well as crocodi- lian-like Triassic and Jurassic terrestrial forms. The Crocodyliformes include the traditional 'Protosuchia', 'Mesosuchia', and Eusuchia, and they are defined by a large number of synapomorphies, particularly of the braincase and occipital regions. The 'protosuchians' (mainly Early *Present address: Department of Zoology, Storer Hall, University of California, Davis, Cali- fornia, USA. The Phylogeny and Classification of the Tetrapods, Volume 1: Amphibians, Reptiles, Birds (ed. M.J. Benton), Systematics Association Special Volume 35A . pp. 295-338. Clarendon Press, Oxford, 1988.
  • New Heterodontosaurid Remains from the Cañadón Asfalto Formation: Cursoriality and the Functional Importance of the Pes in Small Heterodontosaurids

    New Heterodontosaurid Remains from the Cañadón Asfalto Formation: Cursoriality and the Functional Importance of the Pes in Small Heterodontosaurids

    Journal of Paleontology, 90(3), 2016, p. 555–577 Copyright © 2016, The Paleontological Society 0022-3360/16/0088-0906 doi: 10.1017/jpa.2016.24 New heterodontosaurid remains from the Cañadón Asfalto Formation: cursoriality and the functional importance of the pes in small heterodontosaurids Marcos G. Becerra,1 Diego Pol,1 Oliver W.M. Rauhut,2 and Ignacio A. Cerda3 1CONICET- Museo Palaeontológico Egidio Feruglio, Fontana 140, Trelew, Chubut 9100, Argentina 〈[email protected]〉; 〈[email protected]〉 2SNSB, Bayerische Staatssammlung für Paläontologie und Geologie and Department of Earth and Environmental Sciences, LMU München, Richard-Wagner-Str. 10, Munich 80333, Germany 〈[email protected]〉 3CONICET- Instituto de Investigación en Paleobiología y Geología, Universidad Nacional de Río Negro, Museo Carlos Ameghino, Belgrano 1700, Paraje Pichi Ruca (predio Marabunta), Cipolletti, Río Negro, Argentina 〈[email protected]〉 Abstract.—New ornithischian remains reported here (MPEF-PV 3826) include two complete metatarsi with associated phalanges and caudal vertebrae, from the late Toarcian levels of the Cañadón Asfalto Formation. We conclude that these fossil remains represent a bipedal heterodontosaurid but lack diagnostic characters to identify them at the species level, although they probably represent remains of Manidens condorensis, known from the same locality. Histological features suggest a subadult ontogenetic stage for the individual. A cluster analysis based on pedal measurements identifies similarities of this specimen with heterodontosaurid taxa and the inclusion of the new material in a phylogenetic analysis with expanded character sampling on pedal remains confirms the described specimen as a heterodontosaurid. Finally, uncommon features of the digits (length proportions among nonungual phalanges of digit III, and claw features) are also quantitatively compared to several ornithischians, theropods, and birds, suggesting that this may represent a bipedal cursorial heterodontosaurid with gracile and grasping feet and long digits.
  • A New Raptorial Dinosaur with Exceptionally Long Feathering Provides Insights Into Dromaeosaurid flight Performance

    A New Raptorial Dinosaur with Exceptionally Long Feathering Provides Insights Into Dromaeosaurid flight Performance

    ARTICLE Received 11 Apr 2014 | Accepted 11 Jun 2014 | Published 15 Jul 2014 DOI: 10.1038/ncomms5382 A new raptorial dinosaur with exceptionally long feathering provides insights into dromaeosaurid flight performance Gang Han1, Luis M. Chiappe2, Shu-An Ji1,3, Michael Habib4, Alan H. Turner5, Anusuya Chinsamy6, Xueling Liu1 & Lizhuo Han1 Microraptorines are a group of predatory dromaeosaurid theropod dinosaurs with aero- dynamic capacity. These close relatives of birds are essential for testing hypotheses explaining the origin and early evolution of avian flight. Here we describe a new ‘four-winged’ microraptorine, Changyuraptor yangi, from the Early Cretaceous Jehol Biota of China. With tail feathers that are nearly 30 cm long, roughly 30% the length of the skeleton, the new fossil possesses the longest known feathers for any non-avian dinosaur. Furthermore, it is the largest theropod with long, pennaceous feathers attached to the lower hind limbs (that is, ‘hindwings’). The lengthy feathered tail of the new fossil provides insight into the flight performance of microraptorines and how they may have maintained aerial competency at larger body sizes. We demonstrate how the low-aspect-ratio tail of the new fossil would have acted as a pitch control structure reducing descent speed and thus playing a key role in landing. 1 Paleontological Center, Bohai University, 19 Keji Road, New Shongshan District, Jinzhou, Liaoning Province 121013, China. 2 Dinosaur Institute, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007, USA. 3 Institute of Geology, Chinese Academy of Geological Sciences, 26 Baiwanzhuang Road, Beijing 100037, China. 4 University of Southern California, Health Sciences Campus, BMT 403, Mail Code 9112, Los Angeles, California 90089, USA.
  • The Princeton Field Guide to Dinosaurs, Second Edition

    The Princeton Field Guide to Dinosaurs, Second Edition

    MASS ESTIMATES - DINOSAURS ETC (largely based on models) taxon k model femur length* model volume ml x specific gravity = model mass g specimen (modeled 1st):kilograms:femur(or other long bone length)usually in decameters kg = femur(or other long bone)length(usually in decameters)3 x k k = model volume in ml x specific gravity(usually for whole model) then divided/model femur(or other long bone)length3 (in most models femur in decameters is 0.5253 = 0.145) In sauropods the neck is assigned a distinct specific gravity; in dinosaurs with large feathers their mass is added separately; in dinosaurs with flight ablity the mass of the fight muscles is calculated separately as a range of possiblities SAUROPODS k femur trunk neck tail total neck x 0.6 rest x0.9 & legs & head super titanosaur femur:~55000-60000:~25:00 Argentinosaurus ~4 PVPH-1:~55000:~24.00 Futalognkosaurus ~3.5-4 MUCPv-323:~25000:19.80 (note:downsize correction since 2nd edition) Dreadnoughtus ~3.8 “ ~520 ~75 50 ~645 0.45+.513=.558 MPM-PV 1156:~26000:19.10 Giraffatitan 3.45 .525 480 75 25 580 .045+.455=.500 HMN MB.R.2181:31500(neck 2800):~20.90 “XV2”:~45000:~23.50 Brachiosaurus ~4.15 " ~590 ~75 ~25 ~700 " +.554=~.600 FMNH P25107:~35000:20.30 Europasaurus ~3.2 “ ~465 ~39 ~23 ~527 .023+.440=~.463 composite:~760:~6.20 Camarasaurus 4.0 " 542 51 55 648 .041+.537=.578 CMNH 11393:14200(neck 1000):15.25 AMNH 5761:~23000:18.00 juv 3.5 " 486 40 55 581 .024+.487=.511 CMNH 11338:640:5.67 Chuanjiesaurus ~4.1 “ ~550 ~105 ~38 ~693 .063+.530=.593 Lfch 1001:~10700:13.75 2 M.
  • Rule Booklet

    Rule Booklet

    Dig for fossils, build skeletons, and attract the most visitors to your museum! TM SCAN FOR VIDEO RULES AND MORE! FOSSILCANYON.COM Dinosaurs of North America edimentary rock formations of western North America are famous for the fossilized remains of dinosaurs The rules are simple enough for young players, but and other animals from the Triassic, Jurassic, and serious players can benefit Cretaceous periods of the Mesozoic Era. Your objective from keeping track of the cards that is to dig up fossils, build complete skeletons, and display have appeared, reasoning about them in your museum to attract as many visitors as possible. probabilities and expected returns, and choosing between aggressive Watch your museum’s popularity grow using jigsaw-puzzle and conservative plays. scoring that turns the competition into a race! GAME CONTENTS TM 200,000300,000 160,000 VISITORS VISITORS PER YEAR 140,000 VISITORS PER YEAR 180,000 VISITORS PER YEAR 400,000 VISITORS PER YEAR Dig for fossils, build skeletons, and 340,000 VISITORS PER YEAR RD COLOR ELETONS CA GENUS PERIODDIET SK FOSSIL VISITORSPARTS 360,000 VISITORS PER YEAR PER YEAR attract the most visitors to your museum! VISITORS PER YEAR PER YEAR Tyrannosaurus K C 1 4 500,000 Brachiosaurus J H 1 3 400,000 ON YOUR TURN: TM SCAN FOR VIDEO Triceratops K H 1 3 380,000 RULES AND MORE! Allosaurus J C 2 Dig3 a first360,000 card. If it is a fossil, keep it hidden. FOSSILCANYON.COM Ankylosaurus K H 2 If it3 is an340,000 action card, perform the action.
  • Implications for Predatory Dinosaur Macroecology and Ontogeny in Later Late Cretaceous Asiamerica

    Implications for Predatory Dinosaur Macroecology and Ontogeny in Later Late Cretaceous Asiamerica

    Canadian Journal of Earth Sciences Theropod Guild Structure and the Tyrannosaurid Niche Assimilation Hypothesis: Implications for Predatory Dinosaur Macroecology and Ontogeny in later Late Cretaceous Asiamerica Journal: Canadian Journal of Earth Sciences Manuscript ID cjes-2020-0174.R1 Manuscript Type: Article Date Submitted by the 04-Jan-2021 Author: Complete List of Authors: Holtz, Thomas; University of Maryland at College Park, Department of Geology; NationalDraft Museum of Natural History, Department of Geology Keyword: Dinosaur, Ontogeny, Theropod, Paleocology, Mesozoic, Tyrannosauridae Is the invited manuscript for consideration in a Special Tribute to Dale Russell Issue? : © The Author(s) or their Institution(s) Page 1 of 91 Canadian Journal of Earth Sciences 1 Theropod Guild Structure and the Tyrannosaurid Niche Assimilation Hypothesis: 2 Implications for Predatory Dinosaur Macroecology and Ontogeny in later Late Cretaceous 3 Asiamerica 4 5 6 Thomas R. Holtz, Jr. 7 8 Department of Geology, University of Maryland, College Park, MD 20742 USA 9 Department of Paleobiology, National Museum of Natural History, Washington, DC 20013 USA 10 Email address: [email protected] 11 ORCID: 0000-0002-2906-4900 Draft 12 13 Thomas R. Holtz, Jr. 14 Department of Geology 15 8000 Regents Drive 16 University of Maryland 17 College Park, MD 20742 18 USA 19 Phone: 1-301-405-4084 20 Fax: 1-301-314-9661 21 Email address: [email protected] 22 23 1 © The Author(s) or their Institution(s) Canadian Journal of Earth Sciences Page 2 of 91 24 ABSTRACT 25 Well-sampled dinosaur communities from the Jurassic through the early Late Cretaceous show 26 greater taxonomic diversity among larger (>50kg) theropod taxa than communities of the 27 Campano-Maastrichtian, particularly to those of eastern/central Asia and Laramidia.