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British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library Library of Congress Cataloging-in-Publication Data A catalog record for this book is available from the Library of Congress ISBN: 978-0-12-374173-8 For information on all Academic Press publications visit our web site at books.elsevier.com Printed and bound in China 08091011 12 1 0 9 8 7 6 5 4 3 2 1 Working together to grow libraries in developing countries www.elsevier.com I www.bookaid.org I www.sabre.org Reconstruction of animal representatives representatives of the Jehol Biota 1. prima Ji Ii et Ji, Ii, 1996 (" (.. feathered" dinosaur) dinosaur) Sinosauropteryx prima Zhangheotherium quinquecuspidens Hu, Wang, Luo et Li, 1997 (mammal) 2. Psittacosaurus meileyingensis Sereno, Zhao, Cheng et Rao, 1988 (dinosaur) (dinosaur) 3. 4. Jeholomis prima prima Zhou et Zhang, 2002 (bird) Jeholornis Caudipteryx dongi Zhou et Wang, 2000 (feathered dinosaur) dinosaur) 5. Callobatrachus sanyanensis Wang et Gao, 1999 (frog) 6. Ii, 1995 (turtle) Manchurochelys liaoxiensis Ji, 7. 8. Jinzhousaurus yangi Wang et Xu, 2001 (iguanodon) Lil, 2001 (pterosaur) (pterosaur) Haopterus gracilis Wang et Lti, 9. (pterosaur) Sinopterus dongi Wang et Zhou, 2002 (pterosaur) 10. (crayfish) Cricoidoscelosus aethus Taylor, Schram et Shen, 1999 (crayfish) 11. Aeschnidium heishankowense (Hong, 1965) (dragonfly) 12. Hyphalosaurus lingyuanensis Gao, Tang et Wang, 1999 (aquatic (aquatic reptile) 13. Lycoptera sp. (fish) 14. 15. Protopteryxfengningensis Zhang et Zhou, 2000 (bird) Protopteryxfengningensis Protopsephurus liui Lu, 1994 (fish) 16. ° 17. Araenidae indet. (spider) 18. Confuciusomis Confuciusornis sanctus Hou, Zhou, Gu et Zhang, 1995 1995 (bird) 19. Ii, Luo, Yuan, Wible, Zhang et Georgi, 2002 (mammal) Eomaia scansoria Ji, Cfour-winged" Microraptor gui Xu, Zhou, Wang, Kuang, Zhang et Du, 2003 (" four-winged" dinosaur) Yanomis Yanornis martini Zhou et Zhang, 2001 2001 (bird) 20. 21. This Page Intentionally Left Blank CONTENTS CONTENTS A Panorama of the Jehol Biota . . . . .4 Contributors . . . . . . . . . . . . . . .8 Choristoderes . . . . . . 11 Mee-mann Chang Mesozoic Pompeii . . . . . . . . . 19 Gastropods . . . . . . . . . . . . . Hua-zhang Pan, Xiang-gen Zhu Bivalves . . . . . . . . . . . . . . . . .37 '41 . . 89 . . 97 . . . . . . 99 Xiao-lin Wang, Zhong-he Zhou . . . . . . . . 109 --l Xing Xu Birds . - Jun Liu Pterosaurs . . . . . . . . . . . . . . Dinosaurs. Xiao-lin Wang, Zhong-he Zhou - Jun Liu, Xiao-lin Wang Squamates. Introduction. . I m c.. m . . . . . . . . . . . . . . . . . . . . . . . . 129 % o ... o Fu-cheng Zhang, Zhong-he Zhou, Lian-hai Hou III III ;:: III Mammals . . . . . . . . . . . . . . . . . . . . . . . . Jin-hua Chen Conchostracans . 44 . Mei-zhen Cao, Yan-xia Hu Shrimps . . . . . . . . . . . . . . . . . . . . . . . . . s3 ' ' J9 . . . . Jun Liu 162 . . . . . . . . . . . . . . . . I79 Qin Leng, Shun-qing Wu, Else Marie Friis . ,186 ' . . ' Wen-ben Li . 77 Yuan Wang, Ke-qin Gao Turtles. Shun-qing Wu Angiosperms . . Spores and Pollen . . . . . . . . . . . . . . Jun-feng Zhang, Hai-chun Zhang Fishes . . . . . . . . . . . . . . . . . . Jiang-yong Zhang, Fan Jin Amphibians . . . . . . . . . . . . . Qi-fei Wang, Hui-nan Lu, Jing-Ein Yang Land Plants . . . . . . . . . . . . . . . . . . Yan-bin Shen Insects and Spiders. Yuan-qing Wang, Yao-ming Hu, Chuan-kui Li Charophytes . . . . . . . . . . . . . . . . . . . Pei-ji Chen Ostracods . . . . . . . . . . . . . IJI 87 Selected References . . List of Taxa . . . . . . . . . . . . . . . . . . . A Composite Picture of the Jehol Biota . . . Abbreviation of Institutions and Organizations . . 188 197 ,207 . 208 CONTR IBUTORS CONTRIBUTORS of Geology Chinese Academy of Sciences; Nanjing Institute Cao, Mei-zhen Nanjing Instituteand of Palaeontology, Geology and Palaeontology, Chinese AcademyNo. of Sciences; No. Cao, Mei-zhen 39, East Beijing NanjingRoad, 210008, China 39, Road, East Beijing Nanjing 210008, China Paleontology and Paleoanthropology, Chinese Academy of Academy of Institute of Vertebrate Chang, Mee-mann Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Chang, Mee-mann Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, BeijingStreet, 100044, China No. 142, Xi-Zhi-Men-Wai Beijing 100044, China of Geology Chinese Academy of Sciences; Nanjing Institute Chen, Jin-huaChen, Jin-hua Nanjing Instituteand of Palaeontology, Geology and Palaeontology, Chinese AcademyNo. of Sciences; No. 39, East Beijing NanjingRoad, 210008, China 39, Road, East Beijing Nanjing 210008, China of Geology Chinese Academy of Sciences; Nanjing Institute Chen, Pei-ji Chen, Pei-ji Nanjing Instituteand of Palaeontology, Geology and Palaeontology, Chinese AcademyNo. of Sciences; No. 39, East Beijing NanjingRoad, 210008, China 39, Road, East Beijing Nanjing 210008, China of Natural History; Box History; 50007, SE-104 05 Stockholm, Friis, Else Marie Swedish Museum of Natural Box 50007, SE-104 05Sweden Stockholm, Sweden Friis, Else Swedish Marie Museum and Space Sciences, Peking University; YifuBuilding No.2, Peking School of Earth Gao, Ke-qin Gao, Ke-qin School of Earth and Space Sciences, PekingBuilding University; Yifu No. 2, Peking University, Beijing 100871, China100871, China University, Beijing Paleontology and Paleoanthropology, Chinese Academy of Academy of Institute of Vertebrate Hou, Lian-haiHou, Lian-hai Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, BeijingStreet, 100044, China No. 142, Xi-Zhi-Men-Wai Beijing 100044, China of Geology Chinese Academy of Sciences; Nanjing Institute Hu, Yan-xia Hu, Yan-xia Nanjing Instituteand of Palaeontology, Geology and Palaeontology, Chinese AcademyNo. of Sciences; No. 39, East Beijing Nanjing 210008, China 39, Road, East Beijing Road, Nanjing 210008, China Q of Natural History; Central Park West at 79th American Museum Hu, Yao-mingHu, Yao-ming American Museum of Natural History; Central Park Street, West atNew 79thYork, Street, New York, u New York 10024, New USA York 10024, USA Paleontology and Paleoanthropology, Chinese Academy of Academy of Institute of Vertebrate Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, BeijingStreet, 100044, China No. 142, Xi-Zhi-Men-Wai Beijing 100044, China Jin, Fan Jin, Fan Leng, Qin of Geology Chinese Academy of Sciences; Leng, Qin Nanjing Institute Nanjing Instituteand of Palaeontology, Geology and Palaeontology, Chinese AcademyNo. of Sciences; No. Paleontology and Paleoanthropology, Chinese Academy of Academy of Institute of Vertebrate Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, BeijingStreet, 100044, China No. 142, Xi-Zhi-Men-Wai Beijing 100044, China 39, East Beijing NanjingRoad, 210008, China 39, Road, East Beijing Nanjing 210008, China Li, Chuan-kuiLi, Chuan-kui Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Academy of Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, BeijingStreet, 100044, China No. 142, Xi-Zhi-Men-Wai Beijing 100044, China Li, Wen-ben Li, Wen-ben of Geology Palaeontology, Chinese Academy of Sciences; Nanjing Institute Nanjing Instituteand of Geology and Palaeontology, Chinese AcademyNo. of Sciences; No. 39, East Beijing Nanjing 210008, China 39, Road, East Beijing Road, Nanjing 210008, China Liu, Jun Liu, Jun American Museum of Natural History; Central Park West at 79th American Museum of Natural History; Central Park Street, West atNew 79thYork, Street, New York, New York 10024, New USA York 10024, USA Paleontology and Paleoanthropology, Chinese Academy of Academy of Institute of Vertebrate Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, BeijingStreet, 100044, China No. 142, Xi-Zhi-Men-Wai Beijing 100044, China CONTRIBUTORS CONTRIBUTORS Lu, Hui-nan Lu, Hui-nan Nanjing Instituteand of Geology and Palaeontology, Chinese Academy of Sciences; Nanjing Institute of Geology Palaeontology, Chinese Academy of Sciences; No. 39, Road, East Beijing Nanjing No. 39, East Beijing NanjingRoad, 210008, China210008, China Pan, Hua-zhang Pan, Hua-zhang of Geology and Palaeontology, Chinese Academy of Sciences; Nanjing Instituteand Nanjing Institute of Geology Palaeontology, Chinese Academy of Sciences; No. 39,Road, East Beijing Nanjing No. 39, East Beijing NanjingRoad, 210008, China210008, China Shen, Yan-binShen, Yan-bin Nanjing Instituteand of Geology and Palaeontology, Chinese Academy of Sciences; Nanjing Institute of Geology Palaeontology, Chinese Academy of Sciences; No. 39,Road, East Beijing Nanjing No. 39, East Beijing NanjingRoad, 210008, China210008, China of Geology and Palaeontology, Chinese Academy of Sciences; Nanjing Instituteand Wang, Qi-fei Wang, Qi-fei Nanjing Institute of Geology Palaeontology, Chinese Academy of Sciences; No. 39,Road, East Beijing Nanjing No. 39, East Beijing NanjingRoad, 210008, China210008, China Wang, Xiao-lin Vertebrate Paleontology and Paleoanthropology, Chinese Institute of Wang, Xiao-lin Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Academy of No. 142, Xi-Zhi-Men-Wai Beijing Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, BeijingStreet, 100044, China100044, China Vertebrate Paleontology and Paleoanthropology, Chinese Institute of Wang, Yuan Wang, Yuan Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Academy of No. 142, Xi-Zhi-Men-Wai Beijing Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, BeijingStreet, 100044, China100044, China Wang, Yuan-qing Vertebrate Paleontology and Paleoanthropology, Chinese Institute of Wang, Yuan-qing Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Academy of No. 142, Xi-Zhi-Men-Wai Beijing Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, BeijingStreet, 100044, China100044, China Wu, Shun-qing of Geology and Palaeontology, Chinese Academy of Sciences; Nanjing Instituteand Wu, Shun-qing Nanjing Institute of Geology Palaeontology, Chinese Academy of Sciences; No. 39,Road, East Beijing Nanjing No. 39, East Beijing NanjingRoad, 210008, China210008, China Xu, Xing Institute of Vertebrate Vertebrate Paleontology and Paleoanthropology, Chinese Institute of Xu, Xing Paleontology and Paleoanthropology, Chinese Academy of Academy of No. 142, Xi-Zhi-Men-Wai Beijing Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, BeijingStreet, 100044, China100044, China of Geology and Palaeontology, Chinese Academy of Sciences; Nanjing Instituteand Yang, Jing-linYang, Jing-lin Nanjing Institute of Geology Palaeontology, Chinese Academy of Sciences; No. 39,Road, East Beijing Nanjing No. 39, East Beijing NanjingRoad, 210008, China210008, China Zhang, Fu-cheng Vertebrate Paleontology and Paleoanthropology, Chinese Institute of Zhang, Fu-cheng Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Academy of No. 142, Xi-Zhi-Men-Wai Beijing Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, BeijingStreet, 100044, China100044, China Nanjing Instituteand of Geology and Palaeontology, Chinese Academy of Sciences; Zhang, Hai-chun Zhang, Hai-chun Nanjing Institute of Geology Palaeontology, Chinese Academy of Sciences; No. 39,Road, East Beijing Nanjing No. 39, East Beijing NanjingRoad, 210008, China210008, China Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Zhang, Jiang-yong Zhang, Jiang-yong Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Academy of No. 142, Xi-Zhi-Men-Wai Beijing Sciences; No. Sciences; 142, Xi-Zhi-Men-Wai Street, BeijingStreet, 100044, China100044, China of Geology and Palaeontology, Chinese Academy of Sciences; Nanjing Instituteand Zhang, Jun-feng Zhang, Jun-feng Nanjing Institute of Geology Palaeontology, Chinese Academy of Sciences; No. 39,Road, East Beijing Nanjing No. 39, East Beijing NanjingRoad, 210008, China210008, China Vertebrate Paleontology and Paleoanthropology, Chinese Institute of Zhou, Zhong-he Zhou, Zhong-he Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Academy of No. 142, Xi-Zhi-Men-Wai Beijing Sciences; No. Sciences; 142, Xi-Zhi-Men-Wai Street, BeijingStreet, 100044, China100044, China of Geology and Palaeontology, Chinese Academy of Sciences; Nanjing Instituteand Zhu, Xiang-gen Zhu, Xiang-gen Nanjing Institute of Geology Palaeontology, Chinese Academy of Sciences; No. 39,Road, East Beijing Nanjing No. 39, East Beijing NanjingRoad, 210008, China210008, China c.. o'",... :z: ." o VI VI VI Mee-nzann Chang L ast decade has witnessed a renewed interest in the Jehol Biota both within the scientific community and among the general public worldwide. The numerous research papers on the Jehol Biota, published in the prestigious journals such as Natllre and Science, have generated heated controversies among scientists and gained a widespread media frenzy. Here is a perfect example of Stephen J. Gould meeting Thomas S. Kuhn: a sudden "scientific revolution" has punctuated a long period of "normal science," with John Ostrom's revival of Thomas H. Huxley's theory of dinosaurian origin of birds as the new (or more precisely, renewed) paradigm. ...., I m The discoveries of "feathered" dinosaurs in the Jehol Biota appear to have "':rm" provided the direct evidence in support of the paradigm and therefore aroused ,..o public's intrigue in the notion of dinosaurs still being with us. Scientifically ""i=en even more important is the Jehol Biota's well-preserved ancient biodiversity, which contains enormous amount of information on the paleoecosystem as well as many evolutionary issues. We Chinese paleontologists have also had our turn of the wheel not only in those fabulous fossil finds but also in our poise to join our country's drive toward excellence in scientific research. To that end, I hope that this book bears some fruits. In the remainder of this chapter, I will present a brief history of the studies on the ]ehol Biota, an outline of its main components, and a highlight of its scientific import. "JehoI" is the transliteration of the two Chinese characters " !~;O[ " in the Wade-Giles romanization system of the Chinese language, which was used until 1979 when the spelling of proper nouns were officially adopted using the Pinyin (Chinese phonetic alphabet) system of romanization in the mainland of China. In the Pinyin system, these two characters are transliterated to "Rehe". However, we must abide by the "International Stratigraphic Guide" (1976) to continue to use the terms of]ehol Group and Jehol Biota (or Fauna). The literal meaning of the two Chinese characters H'~ ;0[" is "Hot River", derived from many hot springs in the area. What are now called western Liaoning, northern Hebei and southeastern Inner Mongolia were municipally included in Jehol Province (Fig. 1) before 1956, when the provincial name was abolished. At present, the only historical reminder of the ." o en II two characters engraved on aengraved stone tablet Rehetablet Hot Spring "Jehol" is the "Jehol" is the two characters on ainstone in Rehe Hot Spring ° ;'9 ) / W of .the summer resort, originally built for the emperors of the Qing of the Qing \ Chengde of the Chengde summer resort, originally built for the emperors .... 7."° to escape Dynasty the to midsummer in the Forbidden (Fig. 2). City (Fig. 2). Dynasty escape the heat midsummer heat in theCity Forbidden • In his paper In "Cretaceous Mollusca from North from China" (923), the (1923), the his paper "Cretaceous Mollusca North China" American geologist Prof. Amadeus W.Amadeus Grabau (Fig. 3) named the3)fossilAmerican geologist Prof. W. Grabau (Fig. named the fossilbearing strata in the strata vicinity County (nowCounty Lingyuan city in bearing in of theLingyuan vicinity of Lingyuan (now Lingyuan city in western Liaoning Province) "Jehol Series". AndSeries". while studying thestudying the western LiaoningasProvince) as "Jehol And while Map of the1 ea tern the China, the Map of China, the eastern in et (taken from Ding, Weng inset {taken from Ding, Weng & ~o Zeng, ed ., 1936. ew Map of New Map of Zeng, eds., 1936. Province ofChina) howing the Mesozoic stratigraphy China in 1928, he first used the nameused "Jehol Mesozoic of stratigraphy of China in 1928, he first the Fauna". name "Jehol Fauna". In 1962, after on various bearing the fossil fishthe Lycoptera Inworking 1962, after workingdeposits on various deposits bearing fossil fish Lycoptera Q° Provincesof China) showing the from different areas of western Zhi-wei Gu Zhi-wei (Fig. 4), Gu a (Fig. 4), a from different areas Liaoning, of westernProf. Liaoning, Prof. comparable area of the "Jehol comparable area of the "Jehol Province" ( haded ar a), after Province" {shaded area), after which the Biota was named. which the Biota was named. malacologist malacologist from the Nanjing of Geology Palaeontology, used from Institute the Nanjing Instituteand of Geology and Palaeontology, used the name "Jehol for the strata for containing conchosrracan EOJeJtheria Eosestheria the Group" name "Jehol Group" the stratathe containing the conchostracan middendorfii middendorfii (previously known as Bairdestheria middendorfii),middendorfi'i), insect larva insect larva (previously known as Bairdestheria Ephemeropsis and fish Lycoptera. Accordingly, he called the he biota "Jehol Ephemeropsis and fish Lycoptera. Accordingly, called the Biota" biota "Jehol Biota" or, in short, the E.-E.-L. (Fig. 5). or, in short, Biota the E.-E.-L. Biota (Fig. 5). The late Mesozoic Jehol Biota and comparable it had a wide The late Mesozoic Jeholthose Biota and those to comparable to it had a wide 12 distribution over northernover China, Mongolia, regiori of Siberia, distribution northern China,Transbaikalian Mongolia, Transbaikalian region of Siberia, II 2 >_ {' .a:~-"9.~: > present-day Europe. ThisEurope. late Mesozoic oasis provided favorable conditions present-day This late Mesozoic oasis provided favorable conditions A i - i, for many ancient animals and animals plants toand thrive. A series of NE/SWfor many ancient plants to thrive. A series oriented of NE/SW- oriented ~t:{ {'>> It was probably because of the frequent volcanic eruptions numerous It was probably because of the frequent volcanicthat eruptions that numerous i - :~] 2, @qg;~ -!i~) ....... }!~,, the Yanshan Orogeny (mountain-building fault basins developed fault basinsduring developed during the Yanshan Orogeny (mountain-building episode), andepisode), were filled and volcanic fluvial-lacustrine deposits. andwith werethick filledvolcanic with thick and fluvial-lacustrine deposits. / A,£7: Korea, and Japan (Fig. The (Fig. size of of the that of the Korea, and6). Japan 6).this Thearea sizealmost of thisapproaches area almostthat approaches o,~4] i: - plants and animals wereanimals rapidlywere buried and consequently, preserved aspreserved as plants and rapidly buried and consequently, exquisite fossils. Theirfossils. catastrophic misery has turned our bestinto luckour best luck exquisite Their catastrophic misery into has turned today: we have collected not collected only complete skeletons burskeletons also soft parts, today: we have not only complete but alsosuch soft parts, such as feathers, and featherlike as preserved impressions, and not onlyand not only as feathers, andstructures featherlikepreserved structures as impressions, gizzard stones bur also stomach contents, especially in especially Chaoyanginand Beipiao and Beipiao gizzard stones but also stomach contents, Chaoyang regions of western regionsLiaoning. of western Liaoning. ,x~.: ~-: ':~ - -i*i:?{!~} '/ ¢:. The earliest studied fossilstudied from the Jehol Biota western is aLiaoning is a The earliest fossil from the in Jehol BiotaLiaoning in western small fish found the found vicinity City. The material wasmaterial collectedwas collected smallinfish in of theLingyuan vicinity of Lingyuan City. The named and in 1880 by in a French ichthyologist, H. E. by L'Abbe David, and David, by L'Abb6 named 1880 by a French ichthyologist, H. E. Sauvage, as Sauvage, Prolebias davidi, thendavidi, thought to thought be a Tertiary then to becyprinodont a Tertiary cyprinodont as Prolebias : (~iib, ...... >,{< : ~¢"}' i : (pupfish). It (pupfish). was not until 1901 renowned ichthyologist A. It was notwhen until the 1901 when theBritish renowned British ichthyologist A. ,. S. Woodward fish to the Lycoptera, a genus endemic Lycoptera, a B e n u s endemic S. reassigned Woodwardthe reassigned theMesozoic fish to the Mesozoic to Siberia, Mongolia, northernand China. To date, the To described fossils of the fossils of the to Siberia,and Mongolia, northern China. date, the described plants, nearly 90 species vertebrates, 60 species Jehol Biota include over include Jehol Biota overof60 species of plants, nearlyof90 species of vertebrates, :;:! iT; c.. :z: '" ..... o o VI VI ~ I] ~2 . . 2 The Rehe Hot Spring in the Chengde slimmer resort. The two red characters on the stone tablet are '1ehol" in traditional Chinese. The Rehe Hot Spring in the Chengde summer resort. The two red characters on the stone tablet are "Jehol" in traditional Chinese. almost aspecies thousand species of invertebrates. WithBiota the Jehol onbeing some keyonissues evolution as origin andasearly of major of major and almost and a thousand of invertebrates. With the Jehol beingBiota origindiversification and early diversification someinkey issues insuch evolution such hotbed of paleontological present, that number is increasing groups likegroups birds (Hou et a1.,1995), and angiosperms (flowering (flowering a hotbed ofa paleontological research at research present, at that number is increasing like birds (Houetmammals, al.,1995), mammals, and angiosperms rapidly. rapidly. plants), origin of avian flight, tempo and mode evolution, paleobiogeography, plants), origin ofavian flight, tempoofand mode ofevolution, paleobiogeography, paleoecology, and paleoenvironments. The Jehol Biotathe possesses the dualthat qualities entice an unbridled paleoecology, and paleoenvironments. The Jehol Biota possesses dual qualities entice that an unbridled The most The notable theamong recent the fossil findings western fossils are extremely from professionals and lay persons alike. are Theextremely mostamong notable recent fossilfrom findings from western enthusiasm enthusiasm from professionals and lay persons alike. The fossils Liaoning are of course "feathered" dinosaurs. Feathers never been Liaoning arethe of course the "feathered" dinosaurs.had Feathers had never been well beautiful, preserved, and beautiful, and They abundant. have implications important implications well preserved, abundant. have They important view had not was not until 1973 when Prof. John Ostrom , the small theropod Deino view had not received general acceptance. Ie was not until 1973 when Prof. and found of Archaeopteryx, John Ostrom at the Yale University studied the small theropod Deinoll)'chm, theand earlie: |s might be the found that its skeleton was astonishingly similar to that of Archaeopteryx, the earliest bird. And he suggested that small theropods might be the with birds of bitds. Despite the many skeletal e more convincing ancestors characters that than reiareactuallydinosaurs seeing are convinced that withthe birds, for most people, nothing canTo bethose more who convincing than actually ;, the feathered dinosaurs from To those who are convinced that seeing the dinosaurs covered with feathers. 870-1946), western L direct descendants of dinosaurs, noking Howeverl a vocal the gun,.' feathered dinosaurs from birds are minority Ii insists that birds came from a gun". However, a vocal western Liaoning are nothing bur the "smoking 3 Amadeus W. Grabau (1870-1946), minority insistsfeathers that birds from a i Besides, and came featherlike primitive it of leading paleornithologists still (Courtesy: all American geologist who proposed primitive group of reptiles called thecodonts. Besides, feather/ike }arently couldfeathers not fly.and Then what the Jehol Fauna in J928. (Courtesy: Yuan-lin Sun! PKU) arestructures were found on dinosaurs that apparently could nor fly. Then what for courtship, are those feathers for? Were they for insulation,tgmate: ' for " camouflage, ~ Did the proto-birds or for defense? And how did the avian flight originate? the proto-birds aing on theDid ground and then on thefrom ground attaintheir theirwings flight capability by walking and running raising ~rby gliding the and tree then to raising wings learn howtheir to fly ( to fly ("ground-up" theory), or by gliding from the tree to learn how to fly ("tree-down" theory)? We cannot expect that all these questions, togeth questions, tOgether with the origin of birds, be answered JUSt with the discoveries of new discoveries of new materials. More careful and synthetic work has to be done before more persua ersuasive h otheses emerge on more solid ground. before more persuasive hypotheses emerge on more solid ground. Admittedly, this :hinese mala- mor " trulyexcmng ..... :e~o~ ~dlesS sensational, but it" is this kind of work is more tedious and less sensational, bur it is truly exciting and intellectually stimu ins. ~on LtrariWise, clever argumentation and reli .... and intellectually stimulating. Contrariwise, clever argumentation and reli4 Zhi-wei Gu (b.1918), a Chinese malahO proposed gious war are bound tO d~trii~ental to the progress in science. cologist of the NIGP who proposed gious war are bound to be detrimental to the progress in science. 962. The fossil ~ , ~ats'e~, .~ciallyangiosperms, . . . . . . . from western Lhoning are atso~. ......... the E.-E.-L. Biota in 1962. The fossil plants, especially angiosperms, from western Liaoning are also _ very interesting. T ~ ~ordsofangiosperms can ~ traced back to the works ~- ' very interesting. The records of angiosperms can be traced back to the works discovered previously in animals other than birds. Yet, feathers or featherlike f, ofH. ~ ofH. Yabe and S. Endo in 1930s. They described Potamogeton jeholensis. Due to the structures have been found abundantly associared with a number of dinosaurs to the poor preservation of the material, little attention was paid to their work. ~dipteryx, Later, S. Miki (19~4) questioned the identification of Potamogeton, and ~: ~i from Liaoning such as Sinosauropteryx (Chen er al.,1998), Caudipteryx, Later, S. Miki (1964) questioned the identification of Potamogeton, and the plant a:i ~nun~ln-~ instea& ~ore recently, Zheng~yao ~aol Beipiaosallms, Protarchaeopteryx, Sinomithosallrlls, and Microraptor. If we only considered considered the plant as Rammctl/fIS instead. More recently, Zheng-yao Cao say without others (I997) and Shu-ying Duan (:1997) reported monocots and look at the skeletons of these creatures, dinosaur workers would say without and and others (1997) and Shu-ying Duan (1997) reporred monocots and ad hesitation that they are dinosaurs. Yet all of them bear feathers or feather/ike fructificat (Cyperaceae)a, and fructification with carpels. Cao et al.'s Liaoxia chenii(Cyperaceae) oglsts soon , ~chan .,~gu":..((.)ramineae) .(Gramineae) We~ structures. Though somewhat bewildered at first, many paleontologists soon Eragrosite Eragrosites changii werelater laterdes~nated designatedas._gnetales, as gnetales,a group a groupofof • 7 realized how significant these discoveries were: the feathers and featherlike gymnospermi: ~ - @ g ~ Wu ( I ~(1999) 9 ) ! a ~andS hShuang-xing ua~-~ Guo gymnosperm,byby~ uShun-qing Guoan~ and~a~-~i Xiang-:I i~i~ structures found in those dinosaurs betrayed the dinosaurs' affinity to birds! wu Wu (2000) while Duan's Chaoyangia /iangii is probably also a gnetalean, In fact, about 130 years ago, Thomas H. Huxley thought the dinosaur not an angiosperm. Similar forms have been found in the Lower Cretaceous was the direct ancestor of birds. And a few daring scientists even predicted strata of Mongolia and were described by V. A. Krassilov (1982) under other that feathers would sooner or later be found on dinosaurs. But that point of names: Cypel'acites sp., Potamogeton-like spike, and Gurvamlla dictyptera. is regarded the oldest record of the genus. Thegenus. genusThe is atgenus present Archaefructus liaoningensisliaoningensis was described Ge Sun,byD.Ge L. Sun, Dilcher andDilcher others anditothers it is as Archae/ructus was by described D. 1. regarded as the oldest record of the is at present (1998) as an angiosperm although this is not yet accepted byaccepted surviving in California. For lackingFor cone and the leafand cuticular (1998) surviving only in California. lacking cone the leafstructure, cuticular structure, as an angiosperm although thisunquestionably is not yet unquestionably by only most paleobotanists. Its age, however, not bemay the not Latebe Jurassic, these as it these must be it treated caution at present. More recently, Zhou and Zhou and most paleobotanists. Its age,may however, the LateasJurassic, must with be treated with caution at present. MoreZhi-yan recently, Zhi-yan authors suggested, but is morebut likely to belikely the same, Early Shao-lin Zheng (2003) reported the ovulate organs of Ginkgo the from the is more to bei.e., the the same, i.e.,Cretaceous, the Early Cretaceous, authors suggested, Zheng (2003)that reported that the ovulate organs from of Ginkgo Shao-lin as that of the earlyofangiosperms previously previously discovered discovered from Europe, Mongolia Yixian Formation show striking to those oftothe extant species as that the early angiosperms from Europe, Mongolia Yixian Formation showsimilarities striking similarities those of the extant species Sinocarpusdecussatus and westernand North America. more definite angiosperm Ginkgo biloba, indicating a morphological stasis in Gin~ego's western NorthAAmerica. A more definite angiosperm Sinocarpus decussatus Ginkgo biloba, indicating a morphological stasis inreproductive Ginkgo's reproductive was described recently by Qin Leng and Leng E. M.and FriisE. (2003). structure for over 100formillion years. Withyears. ever increased better-preserved was described recently by Qin M. FriisAnother (2003). Another structure over 100 million With everand increased and better-preserved significant significant plant from plant the Jehol is a gymnosperm Sequoiajeholensis. specimens, we anticipate thorough research ofresearch the Jeholof the Jehol fromBiota the Jehol Biota is a gymnosperm SequoiaAnd jeholensis. And specimens, we more anticipate morepaleobotanical thorough paleobotanical ..... I m <til ..o :z: o " C/I C/I ;:: C/I 15 .5 Three representatives of the Jehol Biota in the early studies, the conchostracan Eosestheria (Upper left), 5 Three representatives of the Jehol Biota in the early studies, the conchostracan Eosestheria(Upper left), insect larva Ephemeropsis (Upper Right) and fish Lycoptera (Lower). (Photo: IVPPj insect larva Ephemeropsis(Upper Right) and fish Lycoptera(Lower). (Photo: IVPP) Biota in Biota the near future well. as well. in the nearasfuture others. Although thislist faunal is far from complete, glimpse at others. Although this faunal is far list from complete, it catchesit acatches glimpsea at the vast panorama of the Jeholwhich Biota,awaits whichfurther awaitsexploration. further exploration. The diversity, abundance, and exquisite preservation of the Jehol Biota the vast panorama of the Jehol Biota, The diversity, abundance, and exquisite preservation of the Jehol Biota The Jeholalso Biota has significant on reconstructing the make it make one ofita one handful of extraordinary "Fossil-Lagerstatten" (strata (strata The Jehol Biota has also significant bearingsbearings on reconstructing the of a handful of extraordinary "Fossil-Lagerstiitten" relationships among thelandmasses, ancient landmasses, i.e., paleobiogeography. containing abundant and welland preserved fossils) infossils) the world. relationships among the ancient i.e., paleobiogeography. During During containing abundant well preserved in the Recently world. Recently theof lifetime of the Jehol thepart eastern part ofwas Eurasia was anarea. isolated area. describeddescribed vertebrate fossils include fishes, salamanders, the lifetime the Jehol Biota, theBiota, eastern of Eurasia an isolated vertebrate fossils bony include bonyarchaic fishes,frogs, archaic frogs, salamanders, It was separated the part western part of by Eurasia by theStrait, Turgaiand Strait, and aquatic reptiles, early mammals, etc. Thereetc. areThere also areItalso was separated from thefrom western of Eurasia the Turgai aquatic lizards, reptiles,turtles, lizards,dinosaurs, turtles, dinosaurs, early mammals, fromAmerica North America by the comparatively wideStrait Bering Strait (Fig. 7). paleoabundantabundant invertebrates: mollusks,mollusks, ostracods, conchostracans, insects and by the comparatively wide Bering (Fig. 7). Paleoinvertebrates: ostracods, conchostracans, insectsfrom and North Qinling and Dabie Mountains on the southern of this must have Qinling and Dabie Mountains on the southern border ofborder this area mustarea have some kind ofthat barrier that prevented the exchange of organisms acted as acted some as kind of barrier prevented the exchange of organisms and south.toOwing to this isolation, Jehol Biota consists between between the norththe andnorth south. Owing this isolation, the Jeholthe Biota consists many endemic forms, seen on other continents, and not of many of endemic forms, not seen not on other continents, and not even seeneven in seen in is a fish never the southern part of China. For example, part of China. For example, Lycoptera Lycoptera is a fish never found in found any in any the southern otherout places of the distribution area of the JeholFishes Biota.such Fishes other places of theoutdistribution area of the Jehol Biota. as such as polyodontid Protopsephurus and Yanosteus well as amiiformes Sinamia were polyodontid Protopsephurus and Yanosteus as well asasamiiformes Sinamia were neverinfound in their contemporaneous else. In Late never found their contemporaneous deposits deposits anywhereanywhere else. In Late Cretaceous, the Asian-Alaskan Land came Bridge came into when thewhen Asian-Alaskan Land Bridge into Cretaceous, however,however, many formstorelated to the aforementioned fishestostarted existence,existence, many forms related the aforementioned fishes started appearto appear and in develop North America of their close relatives still and develop NorthinAmerica (Fig. 8). (Fig. Some8). of Some their close relatives still Hiodon (mooneye) North as America as relics even For instance, survive insurvive North in America relics even today. Fortoday. instance, Hiodon (mooneye) .6 Amia (bowfin) to Sinamia, (paddlefish) to to Lycoptera, is relatedistorelated Lycoptera, Amia (bowfin) to Sinamia, Po/yodon Polyodon (paddlefish) to Distribution oftheJehol Biota (green area) at its peak period. m6 Distribution of the Jehol Biota (green area) at its peak period. Protopsephurus and Yanosteus; a few other fishes seem to Protopsephurus and Yanosteus; and a fewand other fishes seem to adopt theadopt Norththe North as theirday present day refugium. Youthem can find them AmericanAmerican freshwaterfreshwater system assystem their present refugium. You can find else nowadays, and hardly you canfind hardly their relatives ancient relatives in and you can theirfind ancient in nowhere nowhere else nowadays, Biota. other parts of theeither. world,Yet either. they occurred in the Jehol other partS of the world, they Yet occurred in the Jehol Biota. This is This is another aspect showing Jeholscientific Biota's scientific importance. another aspect showing the Jeholthe Biota's importance. Asage for of thethe age of the Jeholthe Biota, thehas debate for several J ehol Biota, debate lastedhasforlasted several As for the aretwo mainly two different age" the Late There areThere mainly different opinions opinions about theabout age: the Late decades. decades. Jurassic (ca. 145 years millions years before or present, or versus 145 Ma) 145 Ma) theversus Early the Early Jurassic (ca. 145 millions before present, Cretaceous (ca. 125 Ma). Recently, C. C.III Swisher III et al.2002) (1999, C. C. Swisher et al. 0999, and2002) and Cretaceous (ca. 125 Ma). Recently, Ching-hua Lo et al.dated (1999) theFormation Yixian Formation respectively. The Ching-hua Lo et al. (999) thedated Yixian respectively. The former suggested age of125 around 125 Ma (Early Cretaceous) using single former suggested an age ofan around Ma (Early Cretaceous) using single 140 crystal . sanidine dating. And the used for the 4°Ar/39Ar sam'd'me crysta Ar/39 Ar 4°Ar/39Ar datmg. And the latter usedlatter biotite forbiotite the 4°Arj39Ar laser single-grain fusionand method camean upage with age of147 around of an around Ma 147 Ma laser single-grain fusion method cameand up with (Late Jurassic). Many from workers frominstitutions various institutions are interested in the (Late Jurassic). Many workers various are interested in the . . 7 Paleogeographic map of the world in the Early Cretaceous. (Modified frolll dating, and the work is still underway. The debate would probably is still underway. The debate would probably go on for go on for dating, and the work Fenton, Rich & Rich, 1989) some But expect we canbetter expectresults betterwith results with more dating refined dating Buttime. we can more refined some time. 7 Paleogeographic map of the world in the Early Cretaceous. (Modified from andsampling careful sampling and laboratory work in futute. the near future. techniquestechniques and careful and laboratory work in the near Fenton, Rich & Rich, 1989) This book book provides provides the the Biota's Biota's temporal and and spatial spatial relationships, relationships, This Asian-Alaskan Land Bridge showcases some some of of its most most wonderful fossils, fossils, and and summarizes summarizes our our preliminary showcases understandings of of them. In In aa book that involves many authors authors and and covers covers understandings groups, it isis almost inevitable inevitable that inconsistencies in age many taxonomic groups, and stratigraphic stratigraphic correlations correlations exist. In most most chapters, chapters, the Jehol assignments and few Group includes the Yixian and JJiufotang iufotang Formations only, whereas in a few other chapters the scope of the Group varies to include certain strata either overlying or underlying those formations. We cannot overstate scientific impacts of the JJehol ehol Biota on recent progress in paleontology. We hope that our readers will find the following pages visually pleasing, scientifically interesting, and intellectually rewarding. It is worth noting that the Chinese names have, over the years, confused our overseas colleagues and bibliographers bibliographers alike. To ease the situation, situation, we decide to list the Chinese names according to the English convention, i.e., the custom with surname surname first. To first name first -~ a a reversal of the Chinese custom Ancient Landmass P Subduction lone ..(' further ease the pronunciation pronunciation and potential potential confusion, we hyphenate hyphenate the first name with with two characters, characters, e.g., Shu-an Shu-an Ji. Without Without the hyphen, "Shuan" "Shuan" name with only one could be misconstrued misconstrued as a single syllable, i.e., a first name 8 Paleogeographic map of the world in the Late Cretaceous. (From: http://www.scotese.com) 8 Paleogeographic map of the world in the Late Cretaceous. (From: http://www.scotese.com) character. ACKNOWLEDGMENTS: Needless Needless to say, in a project project as vast as this, there there are numerous numerous unsung unsung heroes behind behind the the contributors contributors to this volume. I would like to name name just just a few whose assistances assistances are particularly particularly appreciated: appreciated: E. M. Friis for her help help with with Chapter Chapter 19 and her willingness willingness in joining joining in to add add Chapter Chapter 20, Zhi-yan Zhi-yan Zhou Zhou for his generosity generosity in providing providing much-needed much-needed consultation consultation on on fossil plants, plants, Er-mi Er-mi Zhao, Zhao, Zhe-xi Zhe-xi Luo, and and Ke-qin Ke-qin Gao Gao for helping helping with with figures for amphibian, amphibian, mammal, mammal, and and choristodere choristodere chapter, chapter, respectively, respectively, Dong Dong Ren for supplying supplying information information and and providing providing figures on on insects, lizards. We insects, and and Shu-an Shu-an Ji Ji for offering offering figures figures for some some of of the the Jehol Jehollizards. We express express our our heartfelt heartfelt appreciation appreciation to to Xiao-lian Xiao-lian Zeng Zeng and and Anderson Anderson Yang Yang for their their elegant elegant artwork. artwork. We We would would also also like like to to express express our our gratitude gratitude to to the the Chaoyang Chaoyang City City Government Government and and Land Land and and Resources Resources Bureau Bureau of of Liaoning Liaoning Province Province for their their support support in in facilitating facilitating our our fieldwork, fieldwork, and and to to the the Museum Museum of of Natural Natural Science Science (Taiwan) (Taiwan) for supporting supporting and and participating participating in in our our research research work work (Fig. (Fig. 9). 9). This This project project isis financed financed by by the the grants grants from from the the Chinese Chinese Academy Academy of ofSciences, Sciences, the the National National Natural Natural Science Science Foundation Foundation of of China, China, the the Ministry Ministry of of Science Science and and Technology Technology of of China, China, and and the the National National Geographic Geographic Society Society of of the the US. US. 9 The IVPP field team joined by Prof. Yen-nian Cheng (back row, 2nd from the right) and 9 The IVPPfield joined by Yen-nian (back 2nd fromatJianshangou, the right) and his crew from team the Museulll ofProf. Natural HistoryCheng (Taiwan) in row, an excavation his crew from the Museum of Natural History Jianshangou, western Liaoning in May, 1999, when the vice (Taiwan) presidentin ofan theexcavation CAS, Prof. at Yi-yu Chen (back row, 6thLiaoning from the and the deputy mayor of the Chaoyang City, Xiao-kun Chen western inright) May, 1999, when the vice president of the CAS, Prof.Mr. Yi-yu Chen (back 7th theand right) fossil site.Chaoyang City, Mr. Xiao-kun Chen (back6th row, row, from thefrom right) the visited deputy the mayor of the (back row, 7th from the right) visited the fossil site. ZhOll Xioo-/in Wang. Zhont:-he . l .. .. . . T P mpeii but d the historical city 0 , 1748 , the ' 79 AD not only destroye ' unt Vesuvius In , was rediscovered In he eruptIon of Mo I When the lost cIty . . ns Just as the death of many peop e. 'shes of the volcamc eruptlo . also caused '11 f, d lying in the thICk a bers of the Jehol d ' als were Stl oun AD so were the mem , 'ed b y the volcanic eruptions of79,' window through which we people an PompeII..,s resl'de nts were VICtlmlZ Th' "Mesozoic Pompell.... has provIded us a am~ IS 'II' years ago. C ceous reta . h I' d over one hundred ml JOn 'and quickly radiated , relatively short time , onderfullife t at Ive can observe t e IIW h J hoi Biota developed In a f trial vertebrates dunng aking tee d' tions 0 terres e of the largest ra la d fossils, and thus Geologica y s p e , onproduced nu merous beautifully preserve in a large area in East Asia. JIt hrepresents I Biota has Biota in the Early h eriod The e 0 I /'f, . I te y of the terrestria I e. f Liaoning Province In a d de several revealed many evolutlona.r evendt: thered dinosaurs in the western part 0 the Cretaceous P , ' ~ . fb rds an lea I . worldwide. In the last eca, h The discoverIes 0 normous attention of paleonto oglsts , ion by workers mainly from t e I calities have been found in thIS reg d he N anjing Institute of 1980s and 1990s attracte .e b' d and dinosaur 0 I dozens of major Ir I o y (IVPP) an t h nd Paleoanthropo og , hb .ng areas of nort ern e Paleonto ogy a ' I n the nelg OrI Institute of Verte rat NIGP) Chinese Academy ofSCIences. mous Region). there were b te;n Inner Mongolia (Nei Mongol Auhtonfio crews of the IVPP have , d he sout eas , 1997 t e Ie Hebei Province an t . 'I those from Liaoning. Since , Fld' in Liaoning, Fengmng d' veries sImI ar to " Chaoyang, UXIn also important ISCO I scale excavations in Belplao, f' 'fkant vertebrate specimens launched about half a dozen arge-M golia and collected hundreds 0 birds and mammals. . 'heng in Inner o n , 'd rosaurs, dinosaurs, , . d in Hebel and N Ingc tic reptiles, IIzar s, pte . d aleontologlsts an hibians, turtles, aqua . . h ar worldWIde an p d h dreds of Visitors eac ye including fishes, amp has since hoste un I£ Geology and Palaeontology ( h sl~m o Western Liaomng Ok with ample rainfal or . T g laypersons alI e. . when the climate In t he area was warm, t cor animals and plants ed at a time , I env ironmen (I hehJehol Biota emer 'ded an Idea bl Ode and h I 'matic background proVI iS f the water bodies were proba y WI at ons Suc C h of t e seas most . widespread. orne 0 tions were frequent · and differentiate. Lakes were . s of the lakes. Volcanic erup to t rIve . I the margin . I k depOSIts. ., A flourishing a ong deep. The plants were ed in rather deep a e , d olcanic actiVIties. t b tes were preserv d the Increase v that time. Most verte ra the Jehol Biota witnesse . and were responsible h E rly Cretaceous, h Yixian Formation , Throughout t e a d ' the deposition of t e f h Yixian Formation ' tions occurred UrIng The volcanic rocks 0 t e d least three major erup 'lcanic eruption cycles. tion were mainly forme ' 'f! t lake depoSlt-vo , f h Yixian Forma 'd d ' s The lake depOSIts 0 t e l l _ s i z e d intermediate-aCl for the four slgm ICan mainly consist of basalts and an. eS.lte. ediate-basic eruptions although sma at the Interva Is between t he nnaJor Internn ° --i I m c.. rn :z: o ~ ." o en en ;: en I~
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