Jehol fossils

CONTR CONTRIBUTORS IBUTORS

of Geology Palaeontology, Chinese Academy of Academy Sciences; No. Nanjing Institute Nanjing Instituteand of Geology and Palaeontology, Chinese of Sciences; No. Cao, Mei-zhen Cao, Mei-zhen

39, East Road, Nanjing 210008, China 39, East Beijing Road,Beijing Nanjing 210008, China and Paleoanthropology, Chinese Academy of Academy of Institute of Vertebrate Institute ofPaleontology Vertebrate Paleontology and Paleoanthropology, Chinese Chang, Mee-mann Chang, Mee-mann

No. 142, Xi-Zhi-Men-Wai Street, Beijing 100044, China Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, Beijing 100044, China Nanjing Instituteand of Geology and Palaeontology, Chinese of Sciences; No. of Geology Palaeontology, Chinese Academy of Academy Sciences; No. Nanjing Institute Chen, Jin-huaChen, Jin-hua 39, East Road, Nanjing 210008, China 39, East Beijing Road,Beijing Nanjing 210008, China Nanjing Instituteand of Geology and Palaeontology, Chinese of Sciences; No. of Geology Palaeontology, Chinese Academy of Academy Sciences; No. Chen, Pei-ji Chen, Pei-jiNanjing Institute 39, East Road, Nanjing 210008, China 39, East Beijing Road,Beijing Nanjing 210008, China of Natural BoxHistory; 50007, SE-104 05 Stockholm, Swedish Museum Swedish MuseumHistory; of Natural Box 50007, SE-104 05 Sweden Stockholm, Sweden Friis, Else Marie Friis, Else Marie and Space Sciences, Peking University; YifuBuilding No.2, Peking School of Earth School of Earth and Space Sciences, PekingBuilding University; Yifu No. 2, Peking Gao, Ke-qin Gao, Ke-qin

University, Beijing 100871, China University, Beijing 100871, China Institute ofPaleontology Vertebrate Paleontology and Paleoanthropology, Chinese and Paleoanthropology, Chinese Academy of Academy of Institute of Vertebrate Hou, Lian-haiHou, Lian-hai Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, Beijing 100044, China No. 142, Xi-Zhi-Men-Wai Street, Beijing 100044, China

Nanjing Instituteand of Geology and Palaeontology, Chinese of Sciences; No. of Geology Palaeontology, Chinese Academy of Academy Sciences; No. Hu, Yan-xia Hu, Yan-xiaNanjing Institute 39, East Beijing Road,Beijing Nanjing 210008, China 39, East Road, Nanjing 210008, China Q

of Natural Central Park Central West at Park 79th West Street,atNew American Museum Hu, Yao-ming Hu, Yao-ming American MuseumHistory; of Natural History; 79th York, Street, New York,

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New York New York 10024, USA 10024, USA Institute ofPaleontology Vertebrate Paleontology and Paleoanthropology, Chinese and Paleoanthropology, Chinese Academy ofAcademy of Institute of Vertebrate No. 142, Xi-Zhi-Men-Wai Street, Beijing 100044, China Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, Beijing 100044, China Jin, Fan

Jin, Fan

Institute ofPaleontology Vertebrate Paleontology and Paleoanthropology, Chinese and Paleoanthropology, Chinese Academy ofAcademy of Institute of Vertebrate No. 142, Xi-Zhi-Men-Wai Street, Beijing 100044, China Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, Beijing 100044, China

Leng, Qin Nanjing Institute Nanjing Instituteand of Geology and Palaeontology, Chinese of Sciences; No. of Geology Palaeontology, Chinese Academy of Academy Sciences; No. 39, East Road, Nanjing 210008, China 39, East Beijing Road,Beijing Nanjing 210008, China Li, Chuan-kui Li, Chuan-kui Institute ofPaleontology Vertebrate Paleontology and Paleoanthropology, Chinese Institute of Vertebrate and Paleoanthropology, Chinese Academy ofAcademy of

Leng, Qin

No. 142, Xi-Zhi-Men-Wai Street, Beijing 100044, China Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, Beijing 100044, China Li, Wen-ben Li, Wen-ben Nanjing Instituteand of Geology and Palaeontology, Chinese of Sciences; No. of Geology Palaeontology, Chinese Academy of Academy Sciences; No. Nanjing Institute

39, East Road, Nanjing 210008, China 39, East Beijing Road,Beijing Nanjing 210008, China Liu, Jun

Liu, Jun

American MuseumHistory; of Natural History; 79th York, Street, New York, of Natural Central Park Central West at Park 79th West Street,atNew American Museum New York New York 10024, USA 10024, USA Institute ofPaleontology Vertebrate Paleontology and Paleoanthropology, Chinese and Paleoanthropology, Chinese Academy of Academy of Institute of Vertebrate No. 142, Xi-Zhi-Men-Wai Street, Beijing 100044, China Sciences; No.Sciences; 142, Xi-Zhi-Men-Wai Street, 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

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Mee-nzann Chang ast decade has witnessed a renewed interest in the Jehol Biota both

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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.

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The discoveries of "feathered" dinosaurs in the Jehol Biota appear to have

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provided the direct evidence in support of the paradigm and therefore aroused

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public's intrigue in the notion of dinosaurs still being with us. Scientifically

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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

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"Jehol" is the "Jehol" two characters engraved on aengraved stone tablet Rehetablet Hot Spring is the two characters on ainstone in Rehe Hot Spring

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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 &

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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

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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 distribution over northernover China, Mongolia, regiori of Siberia, distribution northern China,Transbaikalian Mongolia, Transbaikalian region of Siberia, 12

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present-day Europe. ThisEurope. late Mesozoic oasis provided favorable conditions present-day This late Mesozoic oasis provided favorable conditions

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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

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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.

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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

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plants and animals wereanimals rapidlywere buried and consequently, preserved aspreserved as plants and rapidly buried and consequently,

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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.

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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 as Prolebias then to becyprinodont a Tertiary cyprinodont

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(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 S. reassigned Woodwardthe reassigned theMesozoic fish to the Mesozoic Lycoptera, a B e n u s endemic 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

60 species plants, nearly 90 species vertebrates, Jehol Biota include over include Jehol Biota overof60 species of plants, nearlyof90 species of vertebrates,

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. . 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. western L direct descendants of dinosaurs, noking Howeverl a vocal the gun,.' feathered dinosaurs from birds are 870-1946),

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 i Besides, and came featherlike insistsfeathers that birds from a 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: arestructures were found on dinosaurs that apparently could nor fly. Then what Yuan-lin Sun! PKU) 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,

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" trulyexcmng ..... :e~o~ ~dlesS sensational, but it" is

this kind of work is more tedious and less sensational, bur it is truly exciting

:hinese mala-

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,

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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 •

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realized how significant these discoveries were: the feathers and featherlike gymnospermi: ~ uShun-qing ~ - @ g ~ Wu ( I ~(1999) 9 ) ! a ~andS hShuang-xing ua~-~ Guo gymnosperm,byby 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 and westernand North America. more definite angiosperm Sinocarpusdecussatus 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

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90 Ephialtitid wasp Crephanogaster rara, about 10 mm long.

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Ephialtitid wasp Crephanogaster rara, about 10 mm long.

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89 Sawfly, about 10 mm long.

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89 Sawfly, about 10 mm long.

91 Ichneumon fly Tanychora beipiaoensis, about

~ 971 mmIchneumon fly Tanychora beipiaoensis, about long. 7 mm long.

92 Pelecinid wasp Scorpiopelecinus versatilis, m 9 2about Pelecinid 15 mmwasp long.Scorpiopelecinus versatilis, about 15 mm long.

93 Digger wasp Pompiloperus sp., about

93 15 Digger wasp Pompiloperus sp., about mm long. 15 mm long.

insects gives further proofs that the flowering plants were possibly present, though nOt insects gives further proofs that the flowering plants were possibly present, though not dominant, in the Biota from the Late Jurassic-Early Ctetaceous in the northeastern parr of dominant, theprobably Biota from the Late Jurassic-Early Cretaceous in the northeastern part of China. orne ofinthe pollinating insects of the Jehol Biota are referred to the order

China. Some of the probably pollinating of the Jehol Biota are referred to the order Diptera, e.g., ProtonemestritlSj1trassicllS (Figs. 76,insects 77), Protonemestrim beipiaoensis, Florinemestri1ts Diptera, e.g., Protonemestriusjurassicus 76, characteristic 77), Protonemestrius beipiaoensis, Florinernestrius plllcherrimlfS and flowering bug (Fig.78),(Figs. etc. The feature of these insects is with

pulcherrimus, flowering(proboscis) bug (Fig.78), etc. characteristic feature of these insects is with mouthparts that areThe similar to modern con-familial pollinating special nectaring and nectaring mouthpartsthat (proboscis) that are to modern the probability these insects fed,similar in fact, on juice con-familial of plants or pollinating blood of taxa.special However, However, thatBesides these insects fed, in fact, on taxa, juice of plants orofblood othertaxa. animals could the notprobability be excluded. the aforementioned materials otherof animals not were be excluded. Besides collected. the aforementioned taxa, materials of other Jeholother insects (Figs. could 79-93) also intensively Jehol insects 79"-'93) were alsolow intensively collected. In the Jehol(Figs. Biota, spiders were in diversity and not rich in abundance, with In theinJehol were low in diversity andtonot in Araneida abundance, with representatives some Biota, speciesspiders of the family Araneidae referable therich order within They usually lived amongst wood to branches leaves inwithin the the class Arachnidain(Fig. representatives some94). species of the family Araneidae referable the orderorAraneida forests, orb webs(Fig. that94). wereThey vertical or horizontal or inclined on leaves the spatial thespinning class Arachnida usually lived amongst wooddepending branches or in the of the supporting objects. Usually,orthe spider sat the center of theonweb relationships forests, spinning orb webs that were vertical horizontal or on inclined depending the from spatial to dawn, andofhid plant stemsUsually, or leavesthe near its web during the day, preying on dusk relationships thebetween supporting objects. spider sat on the center of the web from

of small manydusk kinds to dawn, andand hidmedium-sized between plant winged stems orinsects. leaves near its web during the day, preying on (Except where indicated, all fossils in thiswinged chapterinsects. were collected from the Huangbanjigou many kinds of small and medium-sized the lower partindicated, of the Yixian Formation in Beipiao, Liaoning) locality of(Except where all fossils in this chapter were collected from the Huangbanjigou

locality of the lower part of the Yixian Formation in Beipiao, Liaoning) 94

i

Orb-web spider, about 10 mm long.

94 Orb-web spider, about 10 mm long.

]: Jian g- yon g Zhan g, Fan Jin Jiong-yong Zhong, Fan Jill 1 ~

ishes are the most abundant fossils in the Jehol Biota. They

Acipenseriformes (sturgeons), Chondrostei, Actinopterygii. The earliest

acipenseriform fishes were found from the Early Jurassic of Britain and arei he counted tens of thousands. The In scientific Acipenscriformes (curgeons) hondro tei Actinopterygii. The cadic t are the 010 t abundant fos if the Jcholstudy Biota.onThey Germany. But the Late Jurassic-Early Cretaceous ones were only found in fossil fishes of Jehol Biota in western Liaoning started in the second acipenseriform fi hes ere found fr 01 rhe Early Jurassic of Britain and are counted ten of thou and. The cientific study on

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Central ermany. Asia and But the north of East Asia.ie-Early Both fossil and extant acipenseriforms half of the 19th H. Biota E. Sauvage, French anatomist, named the Late Jura retaceou ones werc only found in fo century il fi hes when ofJehol in we atern Liaoning tarced in thethe second appear only in Holarctic and live in fresh waters or in anadromous migration specimens ChinaH.asE. a au newage, species, Prolebias davidi, of the acip nseriform entra! A ia and the norch f East A ia. Boeh fo it and extant half of thefrom 19thnorthern centur when a Fr nch anatOmi t named style since Early Acipenseriforms are predators prey on plankCyprinodontidae. half a hina century an of appear onlyJurassic. in Holarctic and live in fresh water orand in anadromou migration pecimen fromNearly northern as later, a ne\! Amadeus pecies, W. Pro/Grabau, bias tlm'idi, tonic style animals juvenile after acquireare a benthic life.and Adapting American paleontologist,early described and L. joholensis var. an inceinEarly Juraand sic. soon Acipenseriforms predarors prey on plankCyprinodontidae. half aLycoptera centuryjoholensis later, Amadeus '\. rabau themselves to this lifestyle, they develop an elongated jaw and an minor in his book Stratigraphyt,ofdcscribed China in 1928. Subsequently, and ar. tOni animals in juvenile and oon after a quire a benthic inferior life. Adapting American paleontologi L)coptera joho/ellsis Kazuo and L. Saito joho/ensis mouth. Theelve preys shellfishes, and small fishes, them ro are thi primarily life tyle, they developmollusks an elongated jaw and an inferior Fuyuji from Japan, also studied fishes from the area. The first mil70rTakai, in hi book tratigrelpb) olCbina fossil in 192 . ub equently Kazuo aitO and including certain of their own. Acipenser sinensis 95) kis an endemic mouth. Thekinds preys are primarily sheJlfi he ,(Fig. mollu and small fishe , work published Chinese scholars is the monograph Fuyuji Takai, by from Japan, also cudied fo il fi hes Lycopterid from the Fishes area. from The first species of China and is akinds national treasure. is a largesinensis anadromous migration including ertain of their own.ItAcipenser (Fig. 95) i an endemic North China by Hsien-ting Liue scholars and others. In recent years, many new fossil Irolll work published by hine is the monograph L)copteri I FiJbeJ an individual weighing up to 550 kg. TheIt juveniles in the littoral specie of hina and i a national treasure. i a largelive anadromou migration byinHsien-ting andseven other. In recent year,fishes many new fo fish, il with fishesorlb werehilla found the area. ToLiu date, genera of the fossil of Jehol h, with an individual weighing up to 550 juvenile live inThe the littOral of thefi Eastern China Sea and then migrate up kg. theThe rivers to spawn. fi hes found in the area. To date, e en genera of the fo sil fi he ofJehol Biota haveere been published, including Peipiaosteus, Yanosteus, Protopsephurus, of theareas Eastern Chinain ea then and migrate the rivers ro spawn. The are mainly theand Yangtze Pearl up Rivers. Biota ha e been publi hed, including PeipiaoJlem, }'aIlOSleJlS, Prolopsephlfrt/s,spawning Sinamia, Longdeichthys, Lycoptera, and Jinanichthys. spa\ ning areas mainlyarein referred the Yangtze and Pearl Ri fossil ers. family Peipiaosteus and are Yanosteus to a newly erected illall/ia, Longdeichlh)'s, L)coptera, andJinanichlbJls. Peipiaosteus, Yanosteus, and Protopsephurus are fishes belonging to

PeipiaosteltS, YanOSletiS, and Protops 'jJblll'lls are fi he belonging to

Peipiaosf

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and YelnOSfem are referr d to a newly erected fo sil family

95 Acipensersinensis,a large anadromous migration fish, with juveniles living in the littoral of eastern China and adults migrating mainlyto the Yangtze and 95 Acipen er sinensi . a large anadromous migration fi h. with juvenile living in the littoral of eastern China and adults migrating mainly to the Yangtze and Pearl rivers to spawn. (Courtesy: IHB) Pearl rivers to spa n. (Courtesy: IHB)

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96 Peipiaosteus pani, the first fossil acipenseriform fish found in China, usually less than one meter. A very well preserved specimen (Left), with impressionspani. of muscles roes,acipenseriform from Huangbanjigou locality (lower usually part of less Yixian Formation) in Beipiao, Liaoning, and aspecimen close-up view ofwith its 96 Peipiaosteus the firstand fossil fish found in China. than one meter. A very well preserved (Left). digestive tract impression (Right, denoted by a red arrow). (Photo: IVPP) impressions of muscles and roes. from Huangbanjigou locality (lower part of Yixian Formation) in Beipiao. Liaoning. and a close-up view of its digestive tract impression (Right. denoted by a red arrow). (Photo: IVPPl

ml~ 97 Yanosteus longidorsalis, characterized by an extremely long dorsal fin (denoted by a red arrow) nearly up to one third of its total body length. The fish 97 Yanosteus Jongidorsalis. by an (Photo: extremely long dorsal fin (denoted by a red arrow) nearly up to one third of its total body length. The fish may reach one meter characterized in standard length. IVPP) may reach one meter in standard length. (Photo: IVPP)

Peipiaosteidae. Peipiaosteus (Fig. 96), from Yixian and Jiufotang Formations

connections between the drainages of the aforementioned places.

of Beipiao, Liaoning and Fengning, Hebei, is the first fossil acipenseriform

Sinamia is very similar to, but more primitive than, the living bowfin

found in China, with a relatively small body size usually not exceeding one

Amia. Fossil bowfins were usually found in marine deposits in many parts of

meter. Judging from the specimens preserved with roes, the fish probably

the world; in contrast, they have occurred only in freshwater deposits of the

became mature when its body reached 30 cm long. Peipiaosteus differs from

Early Cretaceous of both North and South China and early Tertiary of North

other acipenseriforms in the absence of the caudal fulcra on its caudal fin.

America. The only surviving species of bowfin (Amia calva) lives in fresh

Yanosteus (Fig. 97) was discovered in Lingyuan, Liaoning and Fengning,

waters of eastern part of North America and is called a "living fossil".

Hebei. The remarkable feature of the fish is its extremely long dorsal fin,

Normally, Amia inhabits sluggish, clear, lowland fresh waters preferably rich

nearly up to one third of its total body length. Although approximately

in vegetation. It can withstand high temperature, gulp and expel air at the

fusiform in its body shape, Yanosteus has a relatively straight dorsal margin of

surface, and is even known to aestivate. With its strong and sharp teeth, Amia

the body. The length of the smallest individual is about 20 cm whereas a large

is a voracious predator. The juvenile Amia feeds on smaller animals such as

specimen may reach one meter long. No specific information (such as roes) can

insects, insect larvae, ostracods, other zooplankton and phytoplankton, but

be used to determine the adult size.

after reaching 10 cm long, it begins to feed on other fishes. Adults also eat

Protopsephurus (Fig. 98) is a member of a recent family Polyodontidae (paddlefish). It was collected from Lingyuan, Liaoning, with a length of

crayfish. The teeth of Sinamia are similar to those of Amia. Their food sources might be also similar.

approximately 10 cm in smallest individuals and over one meter in large

Lycoptera (Fig. 100) is the most frequently met fish from the Jehol group.

specimens. Judging from the matured skeletons, Protopsephuruscould be the

It is a small fish referred to a group called teleosts that reach the greatest

smallest in body size among the genera of Polyodontidae. This genus is the

diversity among the recent vertebrates. The fish was found mainly in the

earliest fossil record of the family. An extremely long rostrum, a series of

Yixian Formation in west Liaoning. Endemic to East Asian, Lycoptera was

rostral splints and spine-fringed scales are the most prominent features of the

discovered only in Siberia, Mongolia, Korea and northern China from the late

family. Psephurus is the only extant polyodontid in China who lives in the

Mesozoic. The genus was named by a German anatomist, J. Mtiller, based on

drainage of the Yangtze River and littoral region of the East China Sea

the materials from Transbaikalia of Siberia. The study of Lycoptera in China

whereas Protopsephurusis a stem-polyodontid closely related to Paleopsephurus,

began with the work of Henri E. Sauvage on the fossil teleosts collected from

a Late Cretaceous sturgeon from North America. The living paddlefish is

North China (probably Daxinfangzi, Lingyuan, Liaoning). The fish was

commercially an important freshwater fish. It has naked body surface and

named by Sauvage as Prolebias davidi and was referred to Lycoptera by A. S.

spoon-like rostrum and is thus also called duck-mouthed sturgeon. The

Woodward (a British paleontologist) afterwards. From then on, many

paddlefish eats mainly zooplankton, and is adapted to a wide range of

scientists of both China and foreign countries have worked on the genus and

temperature (2---37°C). A native in the drainage of the Mississippi in the

named about 16 species. Most species of Lycoptera had tiny teeth and ate

United States, the paddlefish has been growing very well in China since it was

plankton, but L. sinensis, L. gansuensis and L. muroii bore relatively large teeth

introduced here.

and were capable of preying on small insects and their larvae. Lycopterais usually

Sinamia (Fig. 99) is supposed to belong to a fossil family of its own, Sinamiidae (Amiiformes). The genus was named by E. A. Stensi6, a Swedish

well-preserved, possibly because they were buried in situ. Entombed in great density, the fish seems to have a habit of swimming in shoals (Fig. 101).

paleontologist, based on the materials from Mengyin, Shandong, China

Lycoptera is also the earliest fossil teleosts discovered in China. The strata

collected by Xi-chou Tan from China and O. A. Zdansky from Austria in

bearing Lycoptera were previously considered the Late Jurassic. The vanishing

1923. Sinamia was found in the Yixian and Jiufotang Formations in Yixian,

of Lyc0ptera was regarded as the indicator of the boundary between the Jurassic

Chaoyang, Liaoning and also in Shaanxi, Gansu, Ningxia, Inner Mongolia,

and Cretaceous. But workers studying ostracods, fossil plant and some other

Anhui and Zhejiang. The wide distribution of Sinamia implies the possible

invertebrates have long regarded the strata as the Early Cretaceous. The

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debate has lasted for decades and drawn much attention of geologists and

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paleontologists. From in-depth studies on L)'coprera in recent years, many paleoichthyologists considered that the fish existed in the Late Jurassic to Early Cretaceous.

Lycoprera became the earliest known osteoglossomorph fish after the British ichthyologist P. H. Greenwood suggested a close relationship between the genus and the superorder Osteoglossomorpha. Fossils of the superorder were found from the Early Cretaceous to Oligocene in nearly all the main continents except Antarctica, but early osteoglossomorphs were mostly recovered from China. Fossil fishes similar to Lycoptera found from the

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Mesozoic terrestrial deposits ofChina were frequently referred to the superorder. Up to now, about 25 genera and 50 species of them were reported from China. The superorder Osteoglossomorpha (bonytongues) is a very early

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branch of teleosts. The number of its fossil genera far exceeds the extant ones; :.;;;",-

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however, the reverse is the case with most other teleost groups. Living

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bonytongues are exclusively fresh-water. The Southeast Asian Sc/eropages (Fig. _100 Lycoptera, the most common fossil vertebrate in western Liaoning and the

_98 Protopsephurus liui, the earliest fossil polyodontid found to date and more than one meter long in large specimens, is a distinct relative of Psephurus living in the drainage of the Yangtze River and littoral region of China. This is a well-preserved specimen from Dawangzhangzi

earliest teleost found in China, is an important member of the Jehol Biota and only distributed in East Asia. This specimen is about 12 cm long. (Photo: IVPP)

locality (middle part ofYixian Formation) in Ungyuan, Liaoning, showing distinct scales and caudal skeleton. (Photo: IVPP)

_99 Sinam;a, found from freshwater deposits of both North and South China, is a distinct relative of living bowfin with strong and sharp teeth, and a voracious predator. This specimen is about 50 em long. (Photo: IVPP) ~

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£7I,j 103 j;nan;cllthys. very similar 10 Ly(optera. was found ,nainly (rom the Jiufotang Formation in

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St',lJ~)sitc bird (sli;41~ l~r-~.r zI~,~ ~ ~:,";,:~',~,::~,, I r ~

like more modern birds, for more adapted flight a long and caudal tail primitive is even more 24"-25body caudal24-25 vertebrae. Its vertebrae. tail is evenItsmore thanprimitive that of than that of like modern birds, adapted flight than aforlong andthan incompact bodyincompact In fact, themore tail bears a lot more to dromaeosaurids Archaeopteryx. InArchaeopteryx. fact, the tail bears a lot resemblance to resemblance dromaeosaurids of A rchaeopteryx.of Archaeopteryx. of the most distinctive features of the fenestra at thethandinosaurs any other birds. For dinosaurs any otherthan known birds. known For instance, the instance, elongated the elongated One of the mostOne distinctive features of Confuciusornis is Confuciusornis the fenestra atisthe end of the 171), humerus (Fig.we171), though we are not its yet sure about its prezygapophysis of the caudal vertebrae are of characteristic of prezygapophysis and chevron ofand the chevron caudal vertebrae are characteristic proximal end ofproximal the humerus (Fig. though are not yet sure about

exact function. exact function.

dromaeosaur dinosaurs, providing for thebirds link between birds dromaeosaur dinosaurs, providing further evidencefurther for theevidence link between

Up to over now,one probably overConfuciusornis one thousandspecimens Confuciusornis have dinosaurs, and theropod dinosaurs,the in dromaeosaurid particular, the dromaeosaurid dinosaurs. and theropod in particular, dinosaurs. Up to now, probably thousand havespecimens many cases, many individuals preserved inThe close been holotype of]eholornis moreasthan 50 seeds holotype ofThe Jeholornis preserved more preserved than 50 seeds imprint in theas imprint in the been discovered. In discovered. many cases, In many individuals were preservedwere in close proximity, indicating (Figs. 172,birds 173).are Other birds 178). It isevidence the first direct evidence for seed-eating adaptation in the belly (Fig.are178).belly It is (Fig. the first direct for seed-eating adaptation in the proximity, indicating mass mortalitymass (Figs.mortality 172, 173). Other relativelythan less abundant than Confuciusornis Confuciusornis.isConfuciusornis is not only characa seedeater. Therelines are of also other lines of evidence Thisa seedeater. bird is clearly Mesozoic. This Mesozoic. bird is clearly There are also other evidence relatively less abundant Confuciusornis. not only characteristic of complete skeletalbut preservation, but it usually contains for this such as the robust jaws, forbeautiful this conclusion, suchconclusion, as the short, deep andshort, robustdeep jaws,and which have onlywhich have only teristic of complete skeletal preservation, it usually contains beautiful feather areas suchneck, as the skull, tail. reduced These teeth veryon reduced teeth onMany the lower jaw. Many seeds also indicate that this the lower jaw. intact seeds alsointact indicate that this feather impressions in impressions areas such asinthe skull, wing, andneck, tail. wing, Theseandvery to those ofand Archaeopteryx and modern birds, with has feathers are birda probably has a well-developed crop. bird probably well-developed crop. feathers are comparable to comparable those of Archaeopteryx modern birds, with

andofbarbs; of them even with Some Confuciusornis rachis of apectoral much derived girdleswith and forelimbs with The The combination ofcombination a much derived girdles pectoral and forelimbs rachis and barbs; some them some even with barbules. Somebarbules. Confuciusornis

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very and primitive tails and hind limbs indicates the mosaic pattern of characters also preserve a pair of long tail feather, they may tails very primitive hind limbs indicates the mosaic pattern of characters specimens also specimens preserve a pair of long tail feather, suggesting thatsuggesting they may that be male individuals is rather often of birds. other words, birds first in early evolution in male early and evolution of birds. In other words,Inearly birds firstearly developed flightdeveloped flight be male individuals (Figs. 171,172,(Figs. 175).171, It is 172, rather175). oftenIt to find male andto find female lyingother nextpreserved to each other preserved on 172). one slab We mayand then capability andmodern then became modern of in the the hind features of and the hind capability became in the features limbs the limbs and the female lying next to each on one slab (Fig. We (Fig. may 172). at least by the time of the Early the sexual further concludefurther that atconclude least by that the time of the Early Cretaceous, theCretaceous, sexual tail.

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of feathers early birds are of much like that modern birdsOathayomis (Fig. Cathayornis Cathayornis 180) isenantiornithine a small-sized enantiornithine Cathayornis (Fig. 180) is a (Fig. small-sized dimorphism of dimorphism feathers in early birds areinmuch like that modern birdsof(Fig. 175).

oroo

bird, than slightly larger than a sparrow in size. Enantiornithine bird, slightly larger a sparrow in size. Enantiornithine birds ("oppositebirds ("opposite

The name birds") are Mesozoic the dominant avian group,by characterized Sapeornis wasthe derived the SAPE, theofabbreviation birds") areofthe dominant avianMesozoic group, characterized its unique by its unique The name of Sapeornis wasofderived from SAPE,from the abbreviation thattoisthat "opposite" to that of articulation between scapulathat andiscoracoid the Society of Avian and Paleontology this bird's was between articulation the scapula andthe coracoid "opposite" of the Society of Avian Paleontology Evolution; and this Evolution; bird's holotype was holotype symposium was held birds. the very first avian specimen discovered SAPE's 5th modern birds. modern Cathayornis is Cathayornis the very firstis avian specimen collected by collected by discovered shortly after theshortly SAPE'safter 5th the symposium meeting, whichmeeting, was heldwhich professional paleontologists in Liaoning, China.inIts1990 discovery inJune 2000. Sapeornis (Fig. 176)from was acollected from a new locality paleontologists in Beijing professional in Liaoning, China. Its discovery has in 1990 has in Beijing in June 2000. Sapeornis (Fig. 176) was collected new locality boosted thebirds studyinofChina early and birdstrigged in China and trigged a seriesofof discoveries of kilometers from the downtown of City. the Chaoyang only a few boostedCity. the study of early a series of discoveries only a few kilometers northwest fromnorthwest the downtown of the Chaoyang feathered dinosaurs birds such asinConfuciusornis is the largest known Early Cretaceous It is not only larger than feathered dinosaurs and early birds and suchearly as Confuciusornis that region. in that region. It is the largestIt known Early Cretaceous bird. It is not bird. only larger than was from the Jiufotang Formation, Archaeopteryx larger than many from dromaeosaurs Cathayornis wasCathayornis from the Jiufotang Formation, which overlies which the overlies the Archaeopteryx but also larger but thanalso many dromaeosaurs the same from regionthe same region Yixian ChapterCathayornis 2). Although Cathayornis is much younger such as Microraptor. Yixian Formation (seeFormation Chapter 2).(see Although is much younger such as Microraptor. thaninArchaeopteryx in age, they share similarities in the such skull as structure such as elongatedrelatively forelimbs,short relatively short hind fused than Archaeopteryx age, they share similarities in the skull structure Its elongatedItsforelimbs, hind limbs, fused limbs, toothed jaws. it has a more expanded braincase than Archaeopteryx. carpometacarpus and short pygostyle indicate toothed On jaws. However, it hasHowever, a more expanded braincase than Archaeopteryx. carpometacarpus and short pygostyle indicate powerful flight powerful capability.flight On capability. It has many advanced the pectoral girdle birdprimitive retains some primitive a short and advanced has many features in thefeatures pectoralingirdle and wing thanand in wing than in the other hand,the thisother bird hand, retainsthis some features such asfeatures a short such and as It and Confuciusornis. to the most primitive Archaeopteryx and theropod The Archaeopteryx coracoid, similar to that ofand Archaeopteryx and Confuciusornis. Compared to Compared the most primitive robust coracoid,robust similar to that of Archaeopteryx theropod dinosaurs. The dinosaurs. (see below), has more of such a large with many small to birds mediumenantiornithine sized birds enantiornithine coexistence Protopteryx (seeProtopteryx below), Cathayornis hasCathayornis more reduced and reduced and coexistence of such a large sized bird with sized manybird small to medium sized digits in the hand. the samethat ageby suggests thatCretaceous by the Early the differentiation shortened digitsshortened in the hand. from the same from age suggests the Early the Cretaceous differentiation Boluochia 181, 182) is another enantiornithine bird early birds greater than previously assumed. Boluochia Boluochia (Figs.Boluochia 181,182)(Figs. is another enantiornithine bird of early birds isofgreater than ispreviously assumed. fromFormation. the Jiufotang Formation. It was theand same horizon and Jeholornis (Figs. ~ primitive 179) is a very bird,theand from Jiufotang It was discovered fromdiscovered the same from horizon Jeholornis Jeholornis (Figs.]eholornis 177 "-" 179) is a177 very bird,primitive and

locality asThis Cathayornis. wasthe named after the locality is the third phylogenetically onlyderived slightlythan more derived thanItArchaeopteryx. as Cathayornis. bird was This namedbird after locality Boluochi, a Boluochi, a phylogenetically only slightly more Archaeopteryx. is the third It locality The most distinctive feature village near City, western have atail. longItsskeletal tail. tail Its long bonyabout tail contains villageabout near Chaoyang City,Chaoyang western Liaoning. The Liaoning. most distinctive feature bird known to bird haveknown a long to skeletal long bony contains

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2_. Aeschnidillm les ·blli 1/111/1 htishankDWtnSe bel b II/kO/l'fllSt 22. 23. ProtonttntJtrius Prot fltm str//ls )111"1 JIm jurassicus

Ga r oj Gastropod 2 . Probaicalia Pl'Obal tli I vitimensis rltl/llt'11 IS 24.

Bi Bivalve 25. Argunitl/a ''/!,1II1It11.7/I11JOIIOIIWJIS 2.5. lingyuannuis

IIII)S(III/·OPItI).\ Prl~ Y. SinoJaUropleryx 9. pri"'aI '(1// itpU/}.\ Jongi 1011'1 10. 1 o. CauJipleryx

St Shrimp n 01 10 (~/f) m aethlls delhll 26. Cricoiaoscelosus

IIIII/'nltbil IIlrllS millenii /r.tI 111// /I. SinornithosaurllJ 11.

12. jinzhoNsaurlls jinzbollJ I//ms yangi )dngi Il. Beipiaosaurus Btipl If) (/11/'11 inexpecllIs III~.\/Jutil 13.

1 PsittacosallrUJ Pitt 1/0J(/"mJ "'tiJe,'ingensis lrtule;lIIgul1Is 14.

Bird 15. Con!lIcillsornis IIlIfr({llIJomi Janetlls J IU til

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