The first theropod tracks from the Middle Jurassic of Gansu,Northwest China:new and rare evidence of quadrupedal progression in theropod dinosaurs
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摘要
A new Middle Jurassic tracksite dominated by non-avian theropod footprints from the Wangjiashan Formation in Pingchuan District,Baojishan Basin,Gansu Province has yielded a unique trackway with four consecutive manus-pes sets.Only three previous examples,all Early Jurassic in age,of theropod trackways are known with convincing examples of manus tracks and in each case,only two tracks were recorded in association with pes tracks with metatarsal impressions and pelvic traces indicating crouching behavior.Thus,this is the first example of manus tracks registered while a theropod trackmaker was walking.This unique configuration is here designated as Grallator pingchuanensis ichnosp.nov.which shows the trackmaker forelimbs registering in a wide straddle gait,much wider than the pes trackway width.G.pingchuanensis confirms previous reports that theropods could occasionally register tridactyl,ectaxonic manus traces.In the case of the Pingchuan trackway,the short step indicates an animal moving at a slow speed,probably due to a soft substrate.
A new Middle Jurassic tracksite dominated by non-avian theropod footprints from the Wangjiashan Formation in Pingchuan District,Baojishan Basin,Gansu Province has yielded a unique trackway with four consecutive manus-pes sets.Only three previous examples,all Early Jurassic in age,of theropod trackways are known with convincing examples of manus tracks and in each case,only two tracks were recorded in association with pes tracks with metatarsal impressions and pelvic traces indicating crouching behavior.Thus,this is the first example of manus tracks registered while a theropod trackmaker was walking.This unique configuration is here designated as Grallator pingchuanensis ichnosp.nov.which shows the trackmaker forelimbs registering in a wide straddle gait,much wider than the pes trackway width.G.pingchuanensis confirms previous reports that theropods could occasionally register tridactyl,ectaxonic manus traces.In the case of the Pingchuan trackway,the short step indicates an animal moving at a slow speed,probably due to a soft substrate.
A new Middle Jurassic tracksite dominated by non-avian theropod footprints from the Wangjiashan Formation in Pingchuan District,Baojishan Basin,Gansu Province has yielded a unique trackway with four consecutive manus-pes sets.Only three previous examples,all Early Jurassic in age,of theropod trackways are known with convincing examples of manus tracks and in each case,only two tracks were recorded in association with pes tracks with metatarsal impressions and pelvic traces indicating crouching behavior.Thus,this is the first example of manus tracks registered while a theropod trackmaker was walking.This unique configuration is here designated as Grallator pingchuanensis ichnosp.nov.which shows the trackmaker forelimbs registering in a wide straddle gait,much wider than the pes trackway width.G.pingchuanensis confirms previous reports that theropods could occasionally register tridactyl,ectaxonic manus traces.In the case of the Pingchuan trackway,the short step indicates an animal moving at a slow speed,probably due to a soft substrate.
引文
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Deng,S.H.,Y.Z.Lu,Y.Zhao,R.Fan,Y.D.Wang,X.J.Yang,X.Li,and B.N.Sun.2017.The Jurassic palaeoclimate regionalization and evolution of China.Earth Science Frontiers 24(1):106-142(in Chinese with English Abstract).
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Gierliński,G.D.1994.Earky Jurassic theroipod tracks with the metatarsal impressions.Prezglad Geologiczny 42:280-284.
Gierliński,G.D.,M.G.Lockley,and G.Nied?wiedzki.2009.A distinctive crouching theropod trace from the lower Jurassic of Poland.Geological Quarterly 53:471-475.
Lallensack,J.N.2018.Automatic generation of objective footprint outlines,Chapter4,105-114.Bonn:Doctoral Thesis.
Li,B.X.,F.X.Xu,Q.H.Ma,H.Z.Pan,S.Q.Wang,and Z.W.Li.1982.Middle Jurassic strata of Wangjiashan Basin,Jingyuan,Gansu.Journal of Stratigraphy 6(1):33-40(in Chinese with English Abstract).
Li,D.Q.,L.D.Xing,M.G.Lockley,L.Pi?uela,J.P.Zhang,H.Dai,J.Y.Kim,W.S.I.V.Persons,and D.L.Kong.2015.A manus dominated pterosaur track assemblage from Gansu,China:Implications for behavior.Science Bulletin 60(2):264-272.
Li,K.,J.Liu,C.Yang,and F.Hu.2011.Dinosaur assemblages from the middle Jurassic Shaximiao formation and Chuanjie formation in the Sichuan-Yunnan Basin,China.Volumina Jurassica 9:21-42.
Lockley,M.G.2009.New perspectives on morphological variation in tridactyl footprints:Clues to widespread convergence in developmental dynamics.Geological Quarterly 53:415-432.
Lockley,M.G.,J.Li,R.Li,M.Matsukawa,J.D.Harris,and L.D.Xing.2013.A review of the tetrapod track record in China,with special reference to type ichnospecies:Implications for ichnotaxonomy and paleobiology.Acta Geologica Sinica 87:1-20.
Lucas,S.G.2007.Tetrapod footprint biostratigraphy and biochronology.Ichnos 14:5-38.
McCrea,R.T.,L.G.Buckley,A.G.Plint,P.J.Currie,J.W.Haggart,C.W.Helm,and S.G.Pemberton.2014.A review of vertebrate track-bearing formations from the Mesozoic and earliest Cenozoic of western Canada with a description of a new theropod ichnospecies and reassignment of an avian ichnogenus.New Mexico Museum of Natural History&Science Bulletin 62:50-93.
Milner,A.R.C.,J.D.Harris,M.G.Lockley,J.L.Kirkland,and N.A.Matthews.2009.Birdlike anatomy,posture,and behavior revealed by an early Jurassic theropod dinosaur resting trace.PLoS One 4(3):e4591.
Olsen,P.E.1980.Fossil great lakes of the Newark Supergroup in New Jersey.In Field Studies of New Jersey Geology and Guide to Field Trips,New York State Geology Association 52ndAnnual Meeting,Rutgers University,ed.W.Manspeizer,352-298.
Olsen,P.E.,J.B.Smith,and N.G.McDonald.1998.The material of the species of the classic theropod footprint genera Eubrontes,Anchisauripus and Grallator(early Jurassic,hart ford and Deerfield basins,Connecticut and Massachusetts,USA).Journal of Vertebrate Paleontology 18(3):586-601.
Ostrom,J.1972.Were some dinosaurs gregarious?Palaeogeography Palaeoclimatology Palaeecology 11:287-301.
Weems,E.1992.Re-evaluation of the taxonomy of Newark Supergroup saurischian dinosaur tracks,using extensive statistical data from a recently exposed tracksite near Culpeper,Virginia.In Proceedings of 26thforum on the geology of industrial minerals.Virginia division of mineral resources publication,Charlottesville,ed.P.C.Sweet,vol.119,113-127.
Xing,L.D.,L.G.Buckley,M.G.Lockley,J.P.Zhang,D.Marty,Y.Wang,J.J.Li,R.T.McCrea,and C.Peng.2016a.A new bird track,Koreanaornis lii ichnosp.nov.,from the lower cretaceous Hekou Group in the Lanzhou-Minhe Basin,Gansu,Northwest China,and implications for early cretaceous avian diversity.Cretaceous Research 66:141-154.
Xing,L.D.,D.Q.Li,P.L.Falkingham,M.G.Lockley,M.J.Benton,H.Klein,J.P.Zhang,H.Ran,W.S.I.V.Persons,and H.Dai.2016b.Digit-only sauropod pes trackways from China-Evidence of swimming or a preservational phenomenon?Scientific Reports 6:21138.
Xing,L.D.,D.Q.Li,J.D.Harris,P.R.Bell,Y.Azuma,M.Fujita,Y.Lee,and P.J.Currie.2013a.A new deinonychosaurian track from the lower cretaceous Hekou group,Gansu Province,China.Acta Palaeontologica Polonica 58(4):723-730.
Xing,L.D.,D.Q.Li,M.G.Lockley,D.Marty,J.P.Zhang,W.S.I.V.Persons,H.L.You,C.Peng,and S.B.Kümmell.2015.Dinosaur natural track casts from the lower cretaceous Hekou Group in the Lanzhou-Minhe Basin,Gansu,Northwest China:Ichnology track formation,and distribution.Cretaceous Research 52:194-205.
Xing,L.D.,D.Q.Li,M.G.Lockley,J.P.Zhang,H.L.You,H.Klein,D.Marty,W.S.I.V.Persons,and C.Peng.2014b.Theropod and sauropod track assemblages from the lower cretaceous Hekou Group at Zhongpu,Gansu Province,China.Acta Palaeontologica Sinica 53(3):381-391(in Chinese with English Abstract).
Xing,L.D.,M.G.Lockley,W.Chen,G.D.Gierliński,J.Li,W.S.I.V.Persons,M.Matsukawa,Y.Ye,M.K.Gingras,and C.W.Wang.2013b.Two theropod track assemblages from the Jurassic of Chongqing,China,and the Jurassic stratigraphy of Sichuan Basin.Vertebrata Palasiatica 51:107-130.
Xing,L.D.,M.G.Lockley,Y.Guo,H.Klein,J.Q.Zhang,J.Zhang,W.S.I.V.Persons,A.Romilio,Y.Tang,and X.Wang.2018.Multiple parallel deinonychosaurian trackways from a diverse dinosaur track assemblage of the lower cretaceous Dasheng Group of Shandong Province,China.Cretaceous Research 90:40-55.
Xing,L.D.,M.G.Lockley,Z.D.Li,H.Klein,J.Zhang,G.Gierliński,Y.Ye,W.S.I.V.Persons,and L.Zhou.2013c.Middle Jurassic theropod trackways from the Panxi region,Southwest China and a consideration of their geologic age.Paleoworld 22:36-41.
Xing,L.D.,D.Marty,H.L.You,D.Q.Li,H.Klein,D.Knaust,J.P.Zhang,R.T.McCrea,and W.S.I.V.Persons.2017.Complex in substrate dinosaur(Sauropoda,Ornithopoda)foot pathways revealed by deep natural track casts from the lower cretaceous Xiagou and Zhonggou formations,Gansu Province,China.Ichnos 24(3):163-178.
Xing,L.D.,G.Nied?wiedzki,M.G.Lockley,J.P.Zhang,X.F.Cai,W.S.I.V.Persons,and Y.Ye.2014a.Asianopodus type footprints from the Hekou Group of Honggu District,Lanzhou City,Gansu,China and the“heel”of large theropod tracks.Palaeoworld 23:304-313.
Young,C.C.1948.Fossil crocodiles in China,with notes on dinosaurian remains associated with the Kansu crocodiles.Bulletin of the Geological Society of China 28(3-4):255-288(in Chinese with English Abstract).
Young,C.C.,and X.J.Zhao.1972.Mamenchisaurus hochuanensis sp.nov.Institute of Vertebrate Paleontology and Paleoanthropology Monographs,Serie A.Vol.8,1-30.
Zhang,H.2009.Sedimentary environments and coal accumulation of the Baojishan-Honghui Basin,eastern Qilian Mountains.Acta Sedimentologica Sinica 27(4):622-631(in Chinese with English Abstract).
Zhang,J.P.,D.Q.Li,M.L.Li,M.G.Lockley,and Z.Bai.2006.Diverse dinosaur-,pterosaur-,and bird-track assemblages from the Hakou formation,lower cretaceous of Gansu Province,Northwest China.Cretaceous Research 27:44-55.
Alexander,R.1976.Estimates of speeds of dinosaurs.Nature 261:129-130.
Avanzini,M.1998.Anatomy of a footprint:Bioturbation as a key to understanding dinosaur walk dynamics.Ichnos 6:129-139.
Bohlin,B.1953.The Sino-Swedish expedition publication 37,VI.Vertebrate Palaeontology 6:Fossil Reptiles from Mongolia and Kansu,113.Stockholm:Statens Etnografiska Museumin.
Deng,S.H.,Y.Z.Lu,Y.Zhao,R.Fan,Y.D.Wang,X.J.Yang,X.Li,and B.N.Sun.2017.The Jurassic palaeoclimate regionalization and evolution of China.Earth Science Frontiers 24(1):106-142(in Chinese with English Abstract).
Du,B.-A.1985.Sporo-pollen assemblages from the middle Jurassic in the Wangjiashan Basin of Jingyuan,Gansu,and their stratigraphic and paleogeographic significance.Geological Review 31(2):131-141(in Chinese with English Abstract).
Gierliński,G.D.1994.Earky Jurassic theroipod tracks with the metatarsal impressions.Prezglad Geologiczny 42:280-284.
Gierliński,G.D.,M.G.Lockley,and G.Nied?wiedzki.2009.A distinctive crouching theropod trace from the lower Jurassic of Poland.Geological Quarterly 53:471-475.
Lallensack,J.N.2018.Automatic generation of objective footprint outlines,Chapter4,105-114.Bonn:Doctoral Thesis.
Li,B.X.,F.X.Xu,Q.H.Ma,H.Z.Pan,S.Q.Wang,and Z.W.Li.1982.Middle Jurassic strata of Wangjiashan Basin,Jingyuan,Gansu.Journal of Stratigraphy 6(1):33-40(in Chinese with English Abstract).
Li,D.Q.,L.D.Xing,M.G.Lockley,L.Pi?uela,J.P.Zhang,H.Dai,J.Y.Kim,W.S.I.V.Persons,and D.L.Kong.2015.A manus dominated pterosaur track assemblage from Gansu,China:Implications for behavior.Science Bulletin 60(2):264-272.
Li,K.,J.Liu,C.Yang,and F.Hu.2011.Dinosaur assemblages from the middle Jurassic Shaximiao formation and Chuanjie formation in the Sichuan-Yunnan Basin,China.Volumina Jurassica 9:21-42.
Lockley,M.G.2009.New perspectives on morphological variation in tridactyl footprints:Clues to widespread convergence in developmental dynamics.Geological Quarterly 53:415-432.
Lockley,M.G.,J.Li,R.Li,M.Matsukawa,J.D.Harris,and L.D.Xing.2013.A review of the tetrapod track record in China,with special reference to type ichnospecies:Implications for ichnotaxonomy and paleobiology.Acta Geologica Sinica 87:1-20.
Lucas,S.G.2007.Tetrapod footprint biostratigraphy and biochronology.Ichnos 14:5-38.
McCrea,R.T.,L.G.Buckley,A.G.Plint,P.J.Currie,J.W.Haggart,C.W.Helm,and S.G.Pemberton.2014.A review of vertebrate track-bearing formations from the Mesozoic and earliest Cenozoic of western Canada with a description of a new theropod ichnospecies and reassignment of an avian ichnogenus.New Mexico Museum of Natural History&Science Bulletin 62:50-93.
Milner,A.R.C.,J.D.Harris,M.G.Lockley,J.L.Kirkland,and N.A.Matthews.2009.Birdlike anatomy,posture,and behavior revealed by an early Jurassic theropod dinosaur resting trace.PLoS One 4(3):e4591.
Olsen,P.E.1980.Fossil great lakes of the Newark Supergroup in New Jersey.In Field Studies of New Jersey Geology and Guide to Field Trips,New York State Geology Association 52ndAnnual Meeting,Rutgers University,ed.W.Manspeizer,352-298.
Olsen,P.E.,J.B.Smith,and N.G.McDonald.1998.The material of the species of the classic theropod footprint genera Eubrontes,Anchisauripus and Grallator(early Jurassic,hart ford and Deerfield basins,Connecticut and Massachusetts,USA).Journal of Vertebrate Paleontology 18(3):586-601.
Ostrom,J.1972.Were some dinosaurs gregarious?Palaeogeography Palaeoclimatology Palaeecology 11:287-301.
Weems,E.1992.Re-evaluation of the taxonomy of Newark Supergroup saurischian dinosaur tracks,using extensive statistical data from a recently exposed tracksite near Culpeper,Virginia.In Proceedings of 26thforum on the geology of industrial minerals.Virginia division of mineral resources publication,Charlottesville,ed.P.C.Sweet,vol.119,113-127.
Xing,L.D.,L.G.Buckley,M.G.Lockley,J.P.Zhang,D.Marty,Y.Wang,J.J.Li,R.T.McCrea,and C.Peng.2016a.A new bird track,Koreanaornis lii ichnosp.nov.,from the lower cretaceous Hekou Group in the Lanzhou-Minhe Basin,Gansu,Northwest China,and implications for early cretaceous avian diversity.Cretaceous Research 66:141-154.
Xing,L.D.,D.Q.Li,P.L.Falkingham,M.G.Lockley,M.J.Benton,H.Klein,J.P.Zhang,H.Ran,W.S.I.V.Persons,and H.Dai.2016b.Digit-only sauropod pes trackways from China-Evidence of swimming or a preservational phenomenon?Scientific Reports 6:21138.
Xing,L.D.,D.Q.Li,J.D.Harris,P.R.Bell,Y.Azuma,M.Fujita,Y.Lee,and P.J.Currie.2013a.A new deinonychosaurian track from the lower cretaceous Hekou group,Gansu Province,China.Acta Palaeontologica Polonica 58(4):723-730.
Xing,L.D.,D.Q.Li,M.G.Lockley,D.Marty,J.P.Zhang,W.S.I.V.Persons,H.L.You,C.Peng,and S.B.Kümmell.2015.Dinosaur natural track casts from the lower cretaceous Hekou Group in the Lanzhou-Minhe Basin,Gansu,Northwest China:Ichnology track formation,and distribution.Cretaceous Research 52:194-205.
Xing,L.D.,D.Q.Li,M.G.Lockley,J.P.Zhang,H.L.You,H.Klein,D.Marty,W.S.I.V.Persons,and C.Peng.2014b.Theropod and sauropod track assemblages from the lower cretaceous Hekou Group at Zhongpu,Gansu Province,China.Acta Palaeontologica Sinica 53(3):381-391(in Chinese with English Abstract).
Xing,L.D.,M.G.Lockley,W.Chen,G.D.Gierliński,J.Li,W.S.I.V.Persons,M.Matsukawa,Y.Ye,M.K.Gingras,and C.W.Wang.2013b.Two theropod track assemblages from the Jurassic of Chongqing,China,and the Jurassic stratigraphy of Sichuan Basin.Vertebrata Palasiatica 51:107-130.
Xing,L.D.,M.G.Lockley,Y.Guo,H.Klein,J.Q.Zhang,J.Zhang,W.S.I.V.Persons,A.Romilio,Y.Tang,and X.Wang.2018.Multiple parallel deinonychosaurian trackways from a diverse dinosaur track assemblage of the lower cretaceous Dasheng Group of Shandong Province,China.Cretaceous Research 90:40-55.
Xing,L.D.,M.G.Lockley,Z.D.Li,H.Klein,J.Zhang,G.Gierliński,Y.Ye,W.S.I.V.Persons,and L.Zhou.2013c.Middle Jurassic theropod trackways from the Panxi region,Southwest China and a consideration of their geologic age.Paleoworld 22:36-41.
Xing,L.D.,D.Marty,H.L.You,D.Q.Li,H.Klein,D.Knaust,J.P.Zhang,R.T.McCrea,and W.S.I.V.Persons.2017.Complex in substrate dinosaur(Sauropoda,Ornithopoda)foot pathways revealed by deep natural track casts from the lower cretaceous Xiagou and Zhonggou formations,Gansu Province,China.Ichnos 24(3):163-178.
Xing,L.D.,G.Nied?wiedzki,M.G.Lockley,J.P.Zhang,X.F.Cai,W.S.I.V.Persons,and Y.Ye.2014a.Asianopodus type footprints from the Hekou Group of Honggu District,Lanzhou City,Gansu,China and the“heel”of large theropod tracks.Palaeoworld 23:304-313.
Young,C.C.1948.Fossil crocodiles in China,with notes on dinosaurian remains associated with the Kansu crocodiles.Bulletin of the Geological Society of China 28(3-4):255-288(in Chinese with English Abstract).
Young,C.C.,and X.J.Zhao.1972.Mamenchisaurus hochuanensis sp.nov.Institute of Vertebrate Paleontology and Paleoanthropology Monographs,Serie A.Vol.8,1-30.
Zhang,H.2009.Sedimentary environments and coal accumulation of the Baojishan-Honghui Basin,eastern Qilian Mountains.Acta Sedimentologica Sinica 27(4):622-631(in Chinese with English Abstract).
Zhang,J.P.,D.Q.Li,M.L.Li,M.G.Lockley,and Z.Bai.2006.Diverse dinosaur-,pterosaur-,and bird-track assemblages from the Hakou formation,lower cretaceous of Gansu Province,Northwest China.Cretaceous Research 27:44-55.