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,
u
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,
this
mor
" 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,
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: ~ 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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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 the southern part of China. For example, is a fish never 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 North as America as relics even For instance, Hiodon (mooneye) survive insurvive North in America relics even today. Fortoday. instance, Hiodon (mooneye)
.6
to Lycoptera, Amia (bowfin) to Sinamia, (paddlefish) to 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, other parts of theeither. world,Yet either. they occurred in the Jehol Biota. 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 western inright) May, 1999, when the vice president of the CAS, Prof.Mr. Yi-yu Chen (back from the and the deputy mayor of the Chaoyang City, Xiao-kun Chen 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
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.
P mpeii but
d the historical city 0 ' 79 AD not only destroye , 1748 , the ' 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 people an , 'ed b y the volcanic eruptions of79,' window through which we PompeII..,s resl'de nts were VICtlmlZ Th' "Mesozoic Pompell.... has provIded us a
T
am~
ceous IS 'II' years ago. reta . h I' d over one hundred ml JOn 'and quickly radiated onderfullife t at Ive , relatively short time , can observe t e IIW h J hoi Biota developed In a f trial vertebrates dunng aking tee d' tions 0 terres Geologica y s p e , e of the largest ra la d fossils, and thus onproduced nu merous beautifully preserve in a large area in East Asia. JIt hrepresents I Biota has Biota in the Early h
C
I /'f, . I te eriod The e 0 y of the terrestria I e. f Liaoning Province In a revealed many evolutlona.r evendt: thered dinosaurs in the western part 0 d de several
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 1980s and 1990s attracte .e I calities have been found in thIS reg d he N anjing Institute of
b' d and dinosaur 0 I dozens of major Ir
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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 including fishes, amp d h dreds of Visitors eac ye has since hoste un I£ Geology and Palaeontology ( h
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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 ons Suc C iS f the water bodies were proba y WI at 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 .
flourishing a ong I k depOSIts. ., A ed in rather deep a e , d olcanic actiVIties. t deep. The plants were 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 for the four slgm ICan d ' s The lake depOSIts 0 t e l l _ s i z e d intermediate-aCl mainly consist of basalts and an. eS.lte. ediate-basic eruptions although sma
at the Interva Is between t he nnaJor Internn °
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locality, several highly fossiliferous with exquisitely fossils volcanic eruptions occurred occasionally. The volcanic less were volcanic eruptions occurred occasionally. The activities volcanic were activities less several locality, highly fossiliferous layers withlayers exquisitely preserved preserved fossils indicated some mass mortality eventsand of feathered birds and dinosaurs. feathered dinosaurs. frequent and relatively weak at the timeat of Jiufotang Formation.Formation. indicated some frequent and relatively weak thethetime of the Jiufotang major massmajor mortality events of birds The vertebrate mainly preserved grayish black lacustrine shale The volcanic long-lasting impact on impact the development and Theand Theeruptions volcanic had eruptions had long-lasting on the development vertebrate fossils are fossils mainlyare preserved in grayishinblack lacustrine shale They are frequently a layer ofashes volcanic evolution evolution of the Jehol With the With intermediate-acid eruptions,eruptions, a lot of aand They are frequently covered bycovered of Biota. the Jehol Biota. the intermediate-acid lot mudstone. of and mudstone. a layer by of volcanic (tuff)ashes (tuff) 10). all fossils are preserved in articulation; and dinosaurs poisonouspoisonous gases were spread the air,to which could have caused gases weretospread the air, which could havethe caused the10). (Fig. (Fig. Nearly all Nearly fossils are preserved in articulation; birds and birds dinosaurs with skins feathers, tissues in of theimpression form of impression or deterioration of the whole ecological system. For instance, the Sihetun deterioration of the whole ecological system. For at instance, at the Sihetun or often withoften feathers, and skins other and soft other tissuessoft in the form
~0
10 Stratigraphic section at the Sihetun locality. showing lacustrine deposits (gray layers) with intercalated tuffs (yellow layers).
m 10 Stratigraphic section at the Sihetun locality, showing lacustrine deposits (gray layers) with intercalated tufts (yellow layers).
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11 Distribution of major fossil vertebrate localities in western Liaoning and the 11 Distribution of major fossil vertebrate western basins; Liaoning localities neighboring areas. in 1, Tertiary 2, and the neighboring areas. I. Tertiary basins; 2. Cretaceous basins; 3, Jurassic-Cretaceous 3.jurassic-Cretaceous basins; 4,Cretaceous faults (F);5, basins; Provincial boundary; basins; 4. faults (F); 5. Provincial 6, Vertebrate sites. A,Jingangshan, Yixian;boundary; 6. Vertebrate A.jingangshan. B, Sihetun, Beipiao; C,sites. Dawujiazi, Fuxin; Yixian; B. Sihetun. Beipiao; C. Dawujiazi, D, Boluochi, Chaoyang; E, Fanzhangzi, Fuxin; D,F,Boluochi. Chaoyang; E, Fanzhangzi. Lingyuan; Daohugou, Ningcheng; (;, Lingyuan; F, Daohugou. ingcheng; G, Shangheshou, Chaoyang; Q, Liaohe Chaoyang; (D, Liaohe Basin; @,Shangheshou. Fuxin-YixianBasin; @,JinlingsiBasin; ~, Fuxin-Yixian Basin; Q).jinlingsiYangshan Basin; @, Beipiao-Chaoyang Yangshan Basin; ,Beipiao-Chaoyang Basin. @, Jianchang-Kazuo Basin. @, Basin; @, jianchang-Kazuo Basin; ®. Lingyuan-Sanshijiazi Basin. ®, Lingyuan-Sanshijiazi Basin; (f), Pingzhuang-Ningcheng Basin; @, Pingzhuang-Ningcheng Basin; @. Chifeng-Yuanbaoshan Basin. (geologic in4 Chifeng-Yuanbaoshan Basin. (Geologic information of basins partly from Liaohe formation of basins partly from Liaohe 6 Petroleum Administration Bureau) Bureau) Petroleum Administration
In sum,Inafter a century's study of the Jehol especially sum,over afterhalf over half a century's study of the Biota Jehol and Biota and especially
the unusual discoveries of the past years, knowledge of the Jehol eggs, gastroliths and theand stomach contents with remains identifiable as plant eggs, gastroliths the stomach contents with remains identifiable as plant of theten past ten our years, our knowledge of the Biota Jehol Biota the unusual discoveries has increased greatlygreatly compared with what was known in the inpast seeds, seeds, lizard lizard and mammal skeletons. It is most environmental and mammal skeletons. It is likely most that likelythe that the environmental has increased compared with what was known the as past as by thebyEosestheria-Ephemeropsis-Lycoptera assemblage (E.-E.-L. changes resulted from the volcanic eruptions causedcaused the mass of represented changes resulted from the volcanic eruptions the mortality mass mortality of represented the Eosestheria-Ephemeropsis-Lycoptera assemblage (E.-E.-1. fauna fauna as traditionally defined). The recent finds of the of vertebrate fossils fossils in birds and other After aAfter briefafloating transportation on the on surface birds and vertebrates. other vertebrates. brief floating transportation the surface as traditionally defined). The recent finds the vertebrate in western Liaoning have shed the on study of the of origin of birdsofand of the of lakes, the dead quicklyquickly descended into, and in, thein, the the lakes, thebodies dead bodies descended into,were andburied were buried western Liaoning havenew shedlight newon light the study the origin birds and their flight, the origin of feathers, early radiation of birds, mammals and and deep water. The abundant volcanic ashes speeded up theup deposition process; deep water. The abundant volcanic ashes speeded the deposition process; their flight, the origin of feathers, early radiation of birds, mammals angiosperms, and also our understanding of the of Early therefore the dead were preserved rapidlyrapidly and completely. In a sense, therefore thebodies dead bodies were preserved and completely. In a sense, angiosperms, andfurthered also furthered our understanding the Cretaceous Early Cretaceous this is this no different from the great of Pompeii. is no different from the burial great burial of Pompeii.
continental ecosystem. continental ecosystem.
Among the famous vertebrate fossil localities of the of Jehol Biota Biota are: are: Among the famous vertebrate fossil localities the Jehol
Like the meticulous grid system at archaeological excavation sites, the the meticulous grid system at archaeological excavation sites, the Like
rock layers have tohave be systematically studiedstudied in order determine where where the the Sihetun and Lujiatun localities in Beipiao; Fanzhangzi and Shanzui Sihetun and Lujiatun localities in Beipiao; Fanzhangzi and Shanzui rock layers to be systematically in to order to determine fossils fossils are andaredecipher the geological age ofage the of fossils embedded in them. (Dawangzhangzi) localities in Lingyuan; Wanfotang, Hejiaxin, Wujiatun (Dawangzhangzi) localities in Lingyuan; Wanfotang, Hejiaxin, Wujiatun and decipher the geological the fossils embedded in them. Long before the recent discoveries of birdsofand dinosaurs, the rocks Long before the recent discoveries birdsfeathered and feathered dinosaurs, the rocks and Xierhuqiao localities in Yixian, Jinzhou; Shangheshou, Dapingfang, and Xierhuqiao localities in Yixian, Jinzhou; Shangheshou, Dapingfang, producing them have by several generations of paleontologists producing LianheLianhe and Dongdadao localities in Chaoyang, western Liaoning Province; thembeen have studied been studied by several generations of paleontologists and Dongdadao localities in Chaoyang, western Liaoning Province; and geologists. It wasItthe geologist Amadeus W. Grabau who who Sichakou and Senjitu localities in Fengning, northern Hebei Hebei Province; and and and geologists. wasAmerican the American geologist Amadeus W. Grabau Sichakou and Senjitu localities in Fengning, northern Province; first proposed the "Jehol Series"Series" and "Jehol Fauna"Fauna" in 1920s. In 1962, Prof. Prof. Daohugou and Xitaizi localities in Ningcheng, southeastern Inner Inner Mongolia first proposed the "Jehol and "Jehol in 1920s. In 1962, Daohugou and Xitaizi localities in Ningcheng, southeastern Mongolia (Fig. 11). (Fig. 11).
Zhi-wei Gu introduced the Jehol into the literature. The Jehol is Zhi-wei Gu introduced the Group Jehol Group into the literature. The Group Jehol Group is
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Ca/hayorttis-Chaoyangia Avifauna a set of late Mesozoic rocks consisting of intercalated Sinorn;s santensis Psi/lacosaurus Fauna Cathayornis-Chaoyangia Avifauna Klsh a setlacustrine of late Mesozoic rocks of intercalated Boluochia zhengi Uaoxi/riton zhongjiani volcanic and deposits richconsisting in tuffaceous Psittacosaurus F a u n a Cmhayornis yandica Sinornis santensis Sinop/erus dong; Ma Liaoxitriton Boluochia zhengi Chaoyangop/eros zhangi zhong]ianiCO/hayornis calida/lis }inanich/hy.f110 Ichthyofauna andcomprises lacustrinethe deposits in tuffaceous materials. volcanic The group Yixian rich FormaSinopterus dongi yandica Uaoningop/eros gui Eoeo/hayornis Cathayornis }inanich/hys longieephalus \IIalkeri hyofauna Chaoyangopterus zhangi mongoliensi.f LongchengarnisCathayornis sanyanensiscaudatus Longdeieh/hys Jinanichthys luojioxioensisk h tPs;ttocoSaUnLf materials. The group comprises tion and the overlying Jiufotang Formation.the Yixian FormaJinanichthys longicephalus Eocathayornis walkeri Ps;uaco.murIL'i Liaoningopterus meileyingensis gui Cuspirostrisornis houi Peipioos/eus sp. V luojiaxiaensis Psittacosaurusmongolier~is sanyanensis Largiros/rornisLongchengornis sexden/oris Pr%psephurosLongdeichthys sp. Sauropoda inde!. tion and the isoverlying Jiufotang in Formation. The Jehol Group mainly distributed northPsittacosaurus meileyingensis 800-1200 m Cu~wirostrisornis houi Chaoyongio beishanefLfis Yanos/eos sp. Peipiaosteus sp. MicroraplOr zhaoianus V o "~ Sauropoda indet. Largirostrornis sexdentoris Songlingornis linghensis Microrap/or glli Sinomia sp. Protopsephurus sp. It. Ilfl 8 0 0 - 1 2 0 0 m ern Hebei Province, The Jehol Group is mainly distributed western Liaoning Province and in northYanosteus sp. Microraptor zhaoianusLongipteryx chao)'angefLfis Chaoyangia beishanen.~is Sinamia sp. Microraptor gui Songlingornis linghensis Yanornis marlini e~ o Inner Mongolia in ortheast China. ern Hebei Province, western Liaoning Province and Yixianornis groballi Longipteryx chaoyangensis southeastern Conjilci'lSornisYanornis sp. martini The deposits were formed a series ofinnortheast southeastern Innerin Mongolia Northeast China. Sapeomis chooyangensis Nxianornis grabaui leholornis prima Confuciusornis sp. -'~ _..__ faulting basins in the late Mesozoic Manchurochelys manehou~1lOensis The deposits were formedofinnortheastern a series of northeast Sapeornis chaoyangensis MonjurosuchllS splendellS Jeholornis prima Asia. Among Yabeinosaurus Manchurochelys lenui.\' the major Fuxin-Yixian Lycop/era Ichthyofauna manchoukuoensis faulting basinsbasins in theare latethe Mesozoic of northeastern 200-300 m Pterodactyloidea ~ Lyeop/era muroii Monjurosuchus splendens "D ID m
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Lycoptera I c h t h y o f a u n a Yabeinosaurtt~ tenuis Manchllroehelys sp. Lycoptera muroii Pterodactyloidea
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Basin, Jianchang-Kazuo Basin, Lingyuan-Sanshijiazi Basin, Jinlingsi-Yangshan Basin, Beipiao-Chaoyang Basin, Pingzhua,ngingcheng Basin, andLingyuan-Sanshijiazi ChifengBasin, Jianchang-Kazuo Basin,
Hyphalosaurus lingyuanensis
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Yuanbaoshan Basin. The vigorous collisions of plates
Yabeinosaurus Uaoningosallrus paradoXlis tenuis Pterodactyloidea linzhollSallros yangi Lycop/era davidi ConJuciusornis Avifauna Psittacosaurus sp. Sinosallropteryx Lyeop/era /okunagai sp. Liaoxiornis delieal1ls Liaoningosaurus paradoxus 2'.0 0 ..... -400 m LioDxiornis sp. Peipiaos/eusJengninge,.,is SinornilhosallnlS sp. d a v Yixianosaurus idi Jinzhousaurus yangi libeinio luanhera Confuciusornis Avifauna ::::.:::.:.:::.:.:::: ~ Pr%psephurosLycoptera lilli longimanlls L ycoptera tokunagai Sinosauropteryx sp. Liaoxiornis delicatus ....... "':"7'" Yanos/eas longidorsalis Sinobaa/ar IingyJJanefLfis ProlOpteryxJengningellSis Peipiaosteusfengningensis Sinornithosaurussp. ConjiJcillSornisLiaoxiornis sp. sp. Eomaio seal/soria
oill o ~ it_ ~ 122.5 Protopsephurus liui m Yixianosaurus longimanus 122.2 Ma, Ma ~ 100-1S0 m Yanosteus longidorsalis LiaobatrachllS Sinobaatar graballi lingyuanensis N Iiioii"~ C
Jibeinia luankera Protopteryxfengningensis Confuciusornis sp.
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in the western rim of the Pacific have resulted in the
the Eurasian continent at that time. Consequently,
intensive tectonic activities in the eastern margin of
tectOnic activities and frequent volcanic eruptions
the Eurasian continent at that time. Consequently,
1
17'1
Jiufotang Shahai of Formation characteristics and theFormation, fossil assemblages the Shahaiand Fuxin studies show from that the lithographic and Fuxin Formation. FormationsRecent are much different those
Sinornithosaurus millenii Zhangheotherium quinquecuspidens Psi/lacosallrosJeholodens sp. jenkinsi
139.4Ma
3
Eoenantiornis buhleri
Zhongheo/herium quinqueelispitlefLf Caudipteryx dongi 125.0 Ma jenki,.,i leholodefLf Beipiaosaurus inexpectus
139.4 Ma
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SinornilhosQunJ."i millenii zoui Caudipteryx
128.4 Ma
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Yuanbaoshan Basin. The vigorous collisions of plates
intensive tectOnic activities in the eastern margin of
complicated the depositional history in different Calloba/raehllsEomaia sanyonefLfis scansoria tectonic activities and frequent volcanic eruptions Mesophryne dt~ 122.2 Ma, 122.5 Ma beipiaoefLfi ~ 100-150 m Liaobatrachus basins; hence the correlation of the deposits in these Manchllroehelys liaoxiensis grabaui N O'I splendefLf us sanyanensis complicated the depositional history in different .~ 122.9 Ma MonjllrosuchllSCallobatrach e" Yabeinosallros Mesophryne /enllis m beipiaoensis basins often becomes difficult. ConJllciusornis sane/liS Lycop/era sine,.,is Eosipleros yangi Manchurochelys liaoxien.~is m ~ basins; hence the correlation of the deposits in these Lycop/era jUxinefLfis ConJucillSorni suniae Haop/enlS gracilis Monjurosuchus splendens The traditional Jehol Group, chronologically Dendrorhynchoides curvitientolllS Peipiaos/ellS pani Yabeinosaurus tenuis ConfucilLwrni ehuonzhous 150-300 m basins often becomes difficult. ConjileillSornisConfuciusornis dlli Psi/locosollrosEosipterus Yanas/eos longidorsalis sp. Lycoptera sine~is yangi from old to new, comprises the Yixian Formation, sanctus Q Changchengornis hengdaoziellsis Sinosallrop/eryx prima IIl '.. '. .'. ~ ProtopsephunlSLycoptera liui fuxinensis Haopter~ gracilis Confuciusorni suniae The traditional Jehol Group, chronologically Liaoningornis longidi/ris Pro/archaeopteryx rob,wa : : :.: .:.: .: .:.: .: .:.:. "7'" Sinamia sp. Peipiaosteus pani Dendrorhynchoides c~rvidentatus Confuciusorni chuonzhous Jiufotang Formation, Shahai Formation and Fuxin Eoenantiornis buhleri Coudip/eryx zoui O Yanosteus longidorsalis Psittacosauru~ sp. Confuciusornis dui ol oldstudies to new,show comprises Yixian Formation, Caudiplery;< tlongi Protopsephurus liui c "'~I=;- 124.6 Ma. 125.0 Sinosauropteryx prima Ma Formation.from Recent that thethe lithographic Changchengornis hengdaoziensis inexpeclllS Sinamia sp. Beipiaosauros Protarchaeopteryx robusta e. Liaoningornis longiditris
==
Cl
Basin, Pingzhuang-Ningcheng Basin, and Chifeng-
in the western rim of the Pacific have resulted in the
Anura inde!.
characteristics and Formations. the fossil assemblages of the Shahai of the Yixian and Jiufotang The Yixian
leholosaunlS shal/gyJJanefLfis Liaocera/ops yanzigoliellSis Anura indet. Incisivosaunts gaurhieri Psittacosaurus sp. Sinovenator changii Jeholosaurus shangyuanensis Repenomamas robIiS/IL' Liaoceratops yanzigouensis Gobiconodon zofiae
Fuxincomprises Formations are much different from those Formationand mainly basalts and andesites,
4
of the Yixian and Jiufotang Formations. with interbedding lacustrine sediments (tuffaceousThe Yixian Formation comprises and andesites, sandstones, gray andmainly gray-black shales,basahs mudstone
Incisivosaurus gauthieri Sinovenator ehangii Repenomamus robustus 5 Gobiconodon zofiae
r777l • L-J Gill 2 3 4
sediments and tuffs). with Four interbedding fossil-bearing lacustrine beds can now be recog-(tuffaceous
6 r-:=:==:l ~ 5
7 6
sandstones, and gray-black shales, mudstone nized from the Yixiangray Formation. The Jiufotang
7
Formationand mainly lacustrine sediments tufts). comprises Four fossil-bearing beds can now be recog-
(grayish, gray-yellow, andYixian gray-black sandstones, nized from the Formation. The Jiufotang 1, basalt and andesite with volcanic breccia (Lava); 2, conglomerate with volcanic breccia; 3, sandstone and conglomerate; 4, tuffaceous sandstone and tuff; Formation mainly comprises lacustrine sediments 5, shale and tuff; 6, silt and silty mudstone; 7 subvolcanic rock and gray-black sandstones, 12 Biostratigraphic (grayish, sequencesgray-yellow, of the Jehol Group. 1, basalt and andesite with volcanic breccia (lava); 2, conglomerate with volcanic breccia; 3, sandstone and conglomerate; 4, tuffaceous sandstone and tuff; 5, shale and tuff; 6, silt and silty mudstone; 7, subvolcanic rock
! !!i;1 lii?:i:!i:is:ii:!:!:!i:ii:i ii
12 Biostratigraphicsequences of the Jehol Group.
siltstones, shales and mudstones, with intercalated tufts). The Shahai and
Jiufotang Formations. Their appearance in the Yixian Formation or slightly
Fuxin Formations are mainly composed of coal deposits and clastics, seldom a major event Formation in the Early Jiufotangrepresents Formations. Theirbiological appearanceradiation in the Yixian or slightly siltstones, shales and mudstones, with intercalated tuffs). The Shahaiearlier and deposits with volcanic contents. The typical Jehol elements such as the Eosestheria- Cretaceous, and most of them existed until the Jiufotang time. Therefore, the Fuxin Formations are mainly composed of coal deposits and clastics, seldom earlier deposits represents a major biological radiation event in the Early Ephemeropsis-Lycoptera assemblage as well as the feathered dinosaurs, early Yixian and Jiufotang Formations record a complete history of the Jehol Biota. Cretaceous, and most of them existed until the Jiufotang time. Therefore, the with volcanic contents. The typical Jehol elements such as the Eosestheriabirds and some other distinctive vertebrates are only found in the Yixian and Currently, the Jehol Group is generally accepted only to comprise these Yixian and Jiufotang Formations record as a complete history of the two Jehol Biota. Ephemeropsis-Lycoptera assemblage as well as the feathered dinosaurs, early birds and some other distinctive vertebrates are only found in the Yixian and
Currently, the Jehol Group is generally accepted as only to comprise these two
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13 Lujiatun localit (basal pan of Yixian Formation) in hangyuan. Beipiao. I.iaoning. showing tuffaceous sand tones. mudstones and overl ing basalts (Lava. 128.4Ma).
1:3 Lujiatun locality (basal part of Yixian Formation)in Shangyuan, Beipiao, Liaoning, showing tuffaceous sandstones, mudstones and overlying basalts (Lava, 128.4Ma).
14 Daohugou locality (basal part ofYixian Formation. but also arguably as Middle Jurassic Jiulongshan Formation by some other researchers) in Shantou.
ingcheng.
Inner Mongolia. showing IVPP excavation sites of the year 2003.
m 14 Daohugou locality (basal part of Yixian Formation, but also arguably as Middle JurassicJiulongshan Formation by some other researchers) in Shantou, Ningcheng, Inner Mongolia, showing IVPP excavation sites of the year 2003.
formations.
locality, yet the age of this deposit is still controversial, ranging from the
Middle Jurassic to the Early Cretaceous according to different workers. We locality, yet the age of this deposit is still controversial, ranging from the now regard the bed at Daohugou comparable to the Dabeigou Formation in different age, with distinctive vertebrate assemblages (Fig. 12).beds They Recently, we have recognized five fossil-bearing (or are, "members") of Middle Jurassic to the Early Cretaceous according to different workers. We northern and it might be comparable to or slightly lower than the from the bottom top:with Lujiatun Bed of the lowest Yixian Formation differenttoage, distinctive vertebrate assemblages (Fig. 12). They are, Hebei, now regard the bed at Daohugou comparable to the Dabeigou Formation in Lujiatun Bed in western Liaoning. At the neighboring Xitaizi locality (Fig. (Jeholosaurus-Repenomamus assemblage), Jianshangou Bed of the lower Yixian from the bottom to top: Lujiatun Bed of the lowest Yixian Formation northern Hebei, and it might be comparable to or slightly lower than the with its statra overlying the Daohugou fossil horizon, abundant FormationfJeholosaurus-Repenomamus (Lycoptera sinensis-Confuciusornis fauna assemblage), assemblage),avian Jianshangou Bed of the lower 15) Yixian Lujiatun Bed in western Liaoning. At the neighboring Xitaizi locality (Fig. acipenseriform fishes Protopsephurus and Yanosteus, bird Confuciusornis and Dawangzhangzi Bed of the middle Yixian Formation (Lycoptera davidiFormation (Lycoptera sinensis-Con/uciusornis avian fauna assemblage), 15) with its statra overlying the Daohugou fossil horizon, abundant dinosaur Psittacosaurus have been collected, and this horizon is clearly compaHyphalosaurus assemblage), Jingangshan Bed of the upper Yixian Formation Dawangzhangzi Bed of the middle Yixian Formation (Lyeoptera davidiacipenseriform fishes Protopsephurus and Yanosteus, bird Con/uciusornis and rable to the lower part of the Yixian Formation. Hyphalosaurus assemblage), Jingangshan Bed of the Yixian Formation (Lycopteramuroii-Manchurochelys manchoukuoensis assemblage), andupper the Boluochi dinosaur Psittaeosaurus have been collected, and this horizon is clearly compaJianshangou Bed (Member) of the lower Yixian Formation The Bed of the(Lyeoptera Jiufotang Formation (linanichthys-Cathayornis avian and fauna muroii-Manehuroehelys manehoukuoensis assemblage), the Boluochi rable to the lower part of the Yixian Formation. Jianshangou Bed is mainly distributed in Sihetun and the neighboring areas, assemblage). Most of the fossil localities of the Jehol Group can be referred avian to Jianshangou Bed (Member) of the lower Yixian Formation The Bed of the Jiufotang Formation (Jinaniehthys-Cathayornis fauna corresponding to Member 3 of the Sihetun composite section (Fig. 16). It is Most of the fossil localities of the J ehol Group can be referred to Jianshangou Bed is mainly distributed in Sihetun and the neighboring areas, one of the assemblage). above five beds. best represented in the Sihetun and Jianshangou localities. Around 20 corresponding to Member 3 of the Sihetun composite section (Fig. 16). It is Lujiatun Bed the lowest Yixian Formation This bed is one of the(Member) above fiveofbeds. localities in Sihetun and the neighboring areas are mainly distributed in an mainly distributed in theBed Sihetun and neighboring areas of the Jinlingsi-This bed is best represented in the Sihetun and Jianshangou localities. Around 20 Lujiatun (Member) of the lowest Yixian Formation area about 12 N 14 km from north to south and 4 N 5 km from east to west. Yangshan mainly Basin, representing newly recognized fossil-bearingareas horizon of Jinlingsi- localities in Sihetun and the neighboring areas are mainly distributed in an distributed ina the Sihetun and neighboring of the the YixianYangshan Formation. It corresponds to athe lowest bed (Member 1) in the horizon of area about 12-14 km from north to south and 4-5 km from east to west. Basin, representing newly recognized fossil-bearing ¸:i1411; ~ composite the stratigraphic section at It thecorresponds Sihetun area. where 1) !iiii!ili Yixian Formation. toThe the main lowestlocalities bed (Member in the Recently, we have recognized five fossil-bearing beds (or "members") of formations.
the Lujiatun Bed is stratigraphic exposed include Lujiatun, Xiaobeigou, Shuiquan and at the Sihetun area. The main localities where composite section Sihetun ofthe Beipiao City Bed and Liutai of Yixian County, Jinzhou City. It is Shuiquan best Lujiatun is exposed include Lujiatun, Xiaobeigou, and represented in theof Lujiatun Sihetun Beipiaolocality. City and Liutai ofYixian County, Jinzhou City. It is best The represented Lujiatun alluvial deposits locality. were formed in the margin of early in the Lujiatun developing stages of the basins. alluvial They mainly comprise The Lujiatun deposits weretuffaceous formed conglomerate, in the margin of early sandstonesdeveloping and silty mudstones 20---40 meters thick (Fig.tuffaceous 13). Thisconglomerate, bed stages of theabout basins. They mainly comprise representssandstones a nearly and simultaneous massabout mortality that wiped 20-40event meters thick (Fig. out 13). This bed silty mudstones numerousrepresents adult and juvenile of vertebrates, including the that small-wiped out simultaneous mass mortality event a nearlyindividuals sized ornithischians such and asJeholosaurus, Psittacosaurus and Liaoceratops, thethe smallnumerous adult juvenile individuals of vertebrates, including small-sized theropods like Sinovenator and Incisivosaurus, the primitive mamsized ornithischians such asJeholosaurus, Psittacosaurus and Liaoceratops, the mals Repenomamus andtheropods Gobiconodon, frogs. Noand invertebrate fossil been mamSinovenator Incisivosaurus, thehasprimitive small-sized likeand recognized. few plant fragments and spore-pollen samples have been malsARepenomamus and Gobiconodon, and frogs. No invertebrate fossil has been collected.recognized. Most of the ALujiatun fossils were collected in 2000 andsamples 2001. have been few plant fragments and spore-pollen Since the fall Most of 1998, theLujiatun Daohugou locality 14) inin Ningcheng of the fossils were (Fig. collected 2000 and 2001. collected. County, Chifeng City,theInner Mongolia, adjacent and north to Lingyuan City, Since fall of 1998, the Daohugou locality (Fig. 14) in Ningcheng Liaoning,County, has produced salamanders asffeholotriton Chifengabundant City, Inner Mongolia, such adjacent and northparadoxus to Lingyuan City, and Chunerpeton tianyiensis, the haired pterosaur ffeholopterus ningchengensis andparadoxus Liaoning, has produced abundant salamanders such asJeholotriton featheredand theropods such as the arboreal Epidendrosaurus Chunerpeton tianyiensis, the hairedmaniraptoran pterosaurJeholopterus ningehengensis and
ningchengensis and other vertebrates. noteworthy that thousands of maniraptoran Epidendrosaurus feathered theropods such as Ittheis arboreal 15 Xitaizi of Yixian Formation) in Ningcheng, Chifeng, Chifeng, 15locality Xitaizi (lower localitypart (lower part of Yixian Formation) in Ningcheng. Inner Mongolia. beautifully preserved insects and plants have also collected that fromthousands this ningchengensis and other vertebrates. It isbeen noteworthy of Inner Mongolia.
beautifully preserved insects and plants have also been collected from this
[,f"
mainly gray, gray-black sandstones, shales and mudstones with rich The mostThe notable localities are Sihetun, most among notablethese among these localities are Jianshangou, Sihetun, Jianshangou, gray, gray-black sandstones, shales tuffaceousmainly components. This bed also represents the and mostmudstones important with rich Zhangjiagou, Huangbanjigou, Hengdaozi, Libalanggou, Heitizigou, etc. Zhangjiagou, Huangbanjigou, Hengdaozi, Libalanggou, Heitizigou, etc. tuffaceous components. This bed also represents the most important horizon preserving birds and feathered dinosaurs. Fossil-bearing beds comparable to the Jianshangou Bed are also in found Fossil-bearing beds comparable to the Jianshangou Bedfound are also in horizon preserving birds and feathered dinosaurs. Nearly 40 genera and species of vertebrates have been discovered from other basins of western Liaoning, northern Hebei and southeastern Inner other basins of western Liaoning, northern Hebei and southeastern Inner Nearly 40 genera and species of vertebrates have been discovered from the Jianshangou Bed, including birds such as Con/uciusornis and Liaoningornis, Mongolia,Mongolia, including including Shangyuan Bed in Beipiao, Zhuanchengzi Beds in Beds Shangyuan Bed inand Beipiao, and Zhuanchengzi in the Jianshangou Bed, including birds such as Confuciusornis and Liaoningornis, feathered dinosaurs Sinosauropteryx, Beipiaosaurus, Sinornithosaurus, Caudipteryx Yixian County. Yixian County. feathered dinosaurs Sinosauropteryx, Beipiaosaurus, Sinornithosaurus, Caudipteryx The Jianshangou Bed represents 2-3 sedimentary cycles of coastal and Protarchaeopteryx; ornithischian dinosaur Psittacosaurus, pterodactyloids The Jianshangou Bed represents 2 " 3 sedimentary cycles of coastal and Protarchaeopteryx; ornithischian dinosaur Psittacosaurus, pterodactyloids Haopterus and Eosipterus, rhamphorhynchoid pterosaur Dendrorhynchoides; lake, shallow semi-deep lake to deep Thelake. sediments comprise comprise lake, lake, shallow lake, semi-deep lakelake. to deep The sediments Haopterus and Eosipterus, rhamphorhynchoid pterosaur Dendrorhynchoides; amphibians Callobatrachus and Mesophryne; primitive mammals Zhangheotherium Manchurochelys Iiaoxiensis liaoxiensis 125.0 Ma 125.0 Ma Manchurochelys
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amphibians Callobatrachusand Mesophryne;primitive mammals Zhangheotherium and]eholodens, and fishes Lycoptera sinensis, L.fuxinensis, Peipiaosteus pani. andJeholodens, and fishes Lycopterasinensis, L. fuxinensis, Peipiaosteuspani. Among them, Lycoptera sinensis, L. /uxinensis, Peipiaosteuspani, Con/uciusornis Among them, Lycoptera sinensis,and L. fuxinensis, Confuciusornis and feathered dinosaurs are most abundant notable.Peipiaosteuspani, The Con/uciusornis and feathered dinosaurs are most abundant and notable. The Confuciusornis avian fauna in the Jianshangou Bed represents the earliest known avian fauna avian fauna in the Jianshangou Bed represents the earliest known avian fauna in the Jehol Biota. Invertebrates are also abundant in this bed including in the Jehol Biota. Invertebrates are also abundant in this bed including gastropods, bivalves, conchosrracans, ostracods, insects as well as numerous gastropods, bivalves, conchostracans, ostracods, insects as well as numerous plants (e.g., stoneworts, spores and pollen). plants (e.g., stoneworts, spores and pollen). Due to the fragmentary material of Sinornis and Cathayornis (both Due tothethemost fragmentary material of Sinornis and described in 1992), remarkable vertebrate discovery in Cathayornis the Jehol (both
Lycoptera sinensis sinensis Peipiaosteus Lycoptera pani Sinamia sp. Peipiaosteus pani Sinamia Liaobatraclllls grabauisp. Liaobatrachus 124.6 Ma Ca//obatraclllls sanyanensisgrabaui described in 1992), the most remarkable vertebrate discovery in the Jehol Ma Callobatrachussanyanensis Biota of Liaoning beipiaoensis 125.2 Ma 124.6Mesophryne had not been made until 1993 when the earliest beaked bird 125.2 Ma Mesophrynebeipiaoensis Manchurochelys Iiaoxiensis 125.0 Ma Biota of Liaoning had not been made until 1993 when the earliest beaked bird .'2 125.0Monjl/rosuchus Ma Manchurochelys liaoxiensis Con/uciusornis splendens sanctus was discovered at the Jianshangou locality in Beipiao Monjurosuchus splendens Yabeinosal/rus tenuis Confuciusornis sanctus was discovered at the Jianshangou locality in Beipiao Yabeinosaurus Da/inghosaurus longidigitl/stenuis City (Fig. 17). The first important mammal fossil from this region, Dalinghosaurus longidigitus Dendrorhynchoides cl/rvidentatl/s City (Fig. 17). The first important mammal fossil from this region, Haoptenls gracilis Dendrorhynchoides curvidentatusZhangheotherium quinquecuspidens, a symmetrodon, was also collected at Sinosauropleryx prima gracilis Haopterus Zhangheotherium quinquecuspidens, a symmetrodon, also collected at approximately the same time. And the putative angiospermwas Archae/ructus Pro/archaeopteryx Sinosarobusta uropteryx prima Caudip/eryx Protarchaeopteryx zoui robusta approximately same time. AndHuangbanjigou the putative angiosperm liaoningensis was found the in the neighboring locality. InArchaefructus the Caudip/eryx Caudipteryx dongi zoui liaoningensis was found in the neighboring Huangbanjigou locality. In the Beipiaosaurus inexpec/us dongi 128.4 Ma Caudipteryx following years, about 20 new localities such as Sihetun, Zhangjiagou, millen;; inexpectus 128.4Sinornithosaunls Ma Beipiaosaurus following years, about 20 new localities such as Sihetun, Zhangjiagou, Psillacosaurus sp. •"~~ Sinornithosaurus millenii Heitizigou have been found in the nearby areas, preserving vertebrateConfl/ciusornis sanclUS Psittacosaurus sp. have beentofound in the nearby preserving vertebrateCanjilciusornis sllniae Confuciusornis sanctus Bedareas, of the lower Yixian bearing Heitizigou deposits belonging the Jianshangou Confuciusornis dui Confuciusornis suniae bearing deposits belonging to the Jianshangou Bed of the lower Yixian Changchengomis hengdaoziensis Formation. ConJuciusornis dui Liaoningornis longiditri Formation. :'.~ Changchengornis hengdaoziensis Eoenantiornis bl/hleri Starting from 1995, the farmers of the Sihetun Village, Beipiao City Liaoningornis longiditris Zhangheotherium ql/inql/ecuspidens Eoenantiornis buhleri Starting from 1995, the farmers of the Sihetun Village, Beipiao City have also been involved in fossil digging. They have come actoss some leholodens jenkinsi Zhangheotherium quinquecuspidens
139.4 Ma
\
Anura indet. 139.4 Ma
Jeholodens jenkinsi
Psillocosourus sp. Anura indet. leholosounlS shangyuonensis Psittacosaurus sp. Lioocerolops yOllzigouensis Jeholosaurus shangyuanensis Incisivosaums gourhieri Liaoceratops yanzigouensis SinovenalOr chongii Incisivosaurus gauthieri Repenomomus robusllls changii GobiconodonSinovenator zofioe
Repenomamusrobustus Gobiconodonzofiae
16 Stratigraphic sequences of the lower part of the Yixian Formation in
Sihetun the neighboring areas of Legends see part Fig. of 12.the Yixian Formation in 16and Stratigraphic sequences the lower Sihetun and the neighboring areas Legends see Fig. 12.
have also been involved in fossil digging. They have come across some primitive birds near their village. Most of the fossil birds were later recognized primitive birds near their village. Most of the fossil birds were later recognized as Con/uciusornis. The field crew of the IVPP began large-scale excavations at as Confuciusornis. The field crew of the IVPP began large-scale excavations at this locality in 1997, and in the four consecutive seasons, we collected thisoflocality in fossils 1997, as and in asthenumerous four consecutive seasons, we collected vertebrate well invertebrates and plants. hundreds hundredshave of vertebrate fossils as attentions, well as numerous and plants. Many findings drawn worldwide and theinvertebrates Sihetun locality Many findings have drawn worldwide attentions, and the Sihetun locality immediately became the spotlight in paleontological community (Fig. 18). immediately became the spotlight in paleontological community (Fig. 18). Among the many interesting vertebrates from this locality are primitive birds Among the many interesting vertebrates from this locality are primitive birds
suchasasConfudusornis Con/uciusornisand and£iaoningornis Liaoningornislongiditris, longiditris, such feathereddinosaurs dinosaurssuch suchasasSinosauropteryx Sinosauropteryxprima, prima, feathered
Beipiaosaurttsinexpectus, inexpectus,Sinornithosaurus Sinornithosaurusmillenii milleniiand and Beipiaosaurus thecommon commonornithischian omithischiandinosaur dinosaurPsittacosaurus, Psittacosaurus, the pterosaur such such asas Haopterus Haopterusgracilis, gracilis, amphibian amphibian pterosaur CallobatrachltSsanjanensis sanyanensisand andthe theprimitive primitivemammal mammal Callobatrachus ]eholodensjenkinsi. jenkinsi. Amazingly, Amazingly, all allthese thesefossils fossils are are Jeholodens ptimarily preserved preserved inin the the shale shale of ofabout about seven seven primarily meter's thick thick(Fig. (Fig. 19). 19). meter's The colleagues colleagues of ofthe the Museum Museum of ofNatural Natural The Science(Taiwan) (Taiwan)joined joinedthe theIVPP IVPPfield field crew crewfor for the the Science 1999 long long field field season season inin the the Sihetun Sihetun area, area, and and 1999 collectedlots lots of offishes, fishes, dinosaurs, dinosaurs, birds, birds, insects insects and and collected plants. In In order order toto better betterunderstand understand the the biostratibiostratiplants. graphic framework, framework, we we also also collected collected about about 150 150 graphic meters of ofthe the core coresamples samples from from the the drillings drillings done done atat meters ofthe the Sihetun, Hengdaozi Hengdaozi and and Zhangjiagou. Zhangjiagou. Some Some of Sihetun, samples are are currently currently under under study study atat the the Museum. Museum. samples The Zhangjiagou Zhangjiagou locality locality (Fig. (Fig. 20) 20) isis about about The three kilometers kilometers north north of of the the Sihetun Sihetun locality, locality, and and three has produced produced important important feathered featheted dinosaurs dinosaurs such such as as has
Caudipteryx zoui, zoui, C. C. dongi dongi and and Protarchaeopteux Protarchaeopteryx robusta. robusta. Caudipteryx We carried carried out out two two large-scale large-scale excavations excavations in in 1998 1998 We and 2001, 2001, and and collected, collected, in in addition addition to to the aforemenand the aforemen-
Confuciusornis, Psittacosaurus, Psittacosaurus, turtles turtles and and tioned taxa, taxa, Confudusornis, tioned rhamphorhynchoid pterosaur pterosaur Dendrorhynchoides Dendrorhynchoides rhamphorhynchoid curvidentatus. curvidentatus. Dawangzhangzi Bed Bed (Member) (Member) of of the the middle middle Dawangzhangzi
The Dawangzhangzi Dawangzhangzi Bed Bed is is best best Yixian Formation Formation The Yixian represented by by the the Fanzhangzi Fanzhangzi and and Shanzui Shanzui localities localities represented in Dawangzhangzi, Dawangzhangzi, about about 20 20 km km southwest southwest of of the the in town of ofLingyuan City (Fig. (Fig. 21). 21). town Lingyuan City The Dawangzhangzi Dawangzhangzi Bed Bed is is approximately approximately The A. W. W. equivalent to to the the "Jehol "Jehol Series" proposed by by A. equivalent Series" proposed
17 Jianshangou Jianshangou locality (lower (lower part part of of Yixian Yixian 17 Formation) Formation) in in Beipiao, Beipiao, Liaoning. Liaoning.
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(lower part of Yixian 18 Site 1 of Sihetun locality (lower partFormation) of Yixian in Beipiao,
Formation)Liaoning. in Beipiao, Liaoning.
19 Close-up view of Site 1, 19 Close-up view of Site I, excavation showing IVPP showing IVPP excavation of the year 2000, where a of the year 2000, where a complete Confuciusornis complete COIl!lIC;lISorn;s specimen was discovered specimen was discovered during the excursion of during the excursion of t h e 5 th International meetthe 5th International meeting of SAPE and the syming of SAPE and the symposium on the Jehol Biota. posiulll on the Jehol Biota.
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IVPP excavation the year U 2 0 _20 IVPPexcavation site of site the of year at Zhangjiagou locality 200 I 2001 at Zhangjiagou locality of Yixian Formation) (lower (lower part ofpart Yixian Formation) in Beipiao. Liaoning. in Beipiao, Liaoning.
.
.
.
.
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21 Shanzui site (vicinity) and Shanzui site (vicinity) and Fanzhangzi site (beyond) in Fanzhangzi site (beyond) in Dawangzhangzi (middle part of Dawangzhangzi (middle part of Yixian Formation) of Lingyuan, YixianChaoyang, Formation) of Lingyuan, Liaoning. Chaoyang, Liaoning.
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29
Grabau (1923). The sediments comprise mainly gray, gray-black sandstones, shales and mudstones with rich tuffaceous components. Fossil-bearing beds comparable to the Dawangzhangzi Bed are the Daxinfangzi Bed in Lingyuan, Dakangpu and Hejiaxin beds in Yixian, Liaoning; and Sichakou and Senjitu beds in Fengning, Hebei. Among the known vertebrate fossils fossils are Lycoptera Lycoptera
davidi and L. tokunagai, ascipenseriform ascipenseriform Peipiaosteus Peipiaosteus fengningensis and Protopsephurus Monjurosuchus and Protopsephurus liui, turtle Manchurochelys, Manchurochelys, choristoderes Monjurosuchus Hyphalosaurus, dinosaurs Sinosauropteryx, Hyphalosaurus, Sinosauropteryx,Sinornithosaurus, Sinornithosaurus, andjinzhousaurus, andJinzhousaurus, birds Liaoxiornis, Liaoxiornis, Protopteryx Protopteryxand a few Confuciusornis, Confuciusornis,mammals Sinobaatar Sinobaatarand Eomaia, putative angiosperm Archaefructus Archaefructus sinensis. sinensis. Lycoptera Lycoptera Eomaia, as well as the putative davidi and Hyphalosaurus Hyphalosaurus are most abundant abundant in this bed. Since 1998, important important vertebrate and plant fossils fossils have been discovered from the Dawangzhangzi Dawangzhangzi Bed of the middle middle Yixian Formation Formation at the Fanzhangzi and Shanzui localities of Lingyuan City. The main fossils fossils from this bed include the juvenile juvenile enantiornithine Liaoxiornis delicates, enantiornithine Liaoxiornis delicates, the choristodere
Hyphalosaurus lingyuanensis, multituberculate mammal Sinobaatar Sinobaatar reptile Hyphalosaurus lingyuanensis, the multituberculate 22 Hejiaxin locality (middle part ofYixian Formation) in Toutai, Yixian, Liaoning.
m 2 2 Hejiaxin locality (middle part of Yixian Formation) in Toutai, Yixian, Liaoning.
lingyuanensis, the earliest known eutherian mammal Eomaiascansoria, Eomaia'scansoria, the lingyuanensis, eutherian mammal Sinosauropteryx and putative angiosperm Archaefructus Archaefruetus primitive compsognathid Sinosauropteryx
Dongtuyaosite, site,Senjitu Senjitu(Left) (Left)and andJiecaigou Jiecaigollsite, site,Sichakou Sichakou(Right)(middle (Right) (middlepart partof ofYixian Formation) inFengning, Fengning,Hebei. Hebei. ~ 2 3 23 Dongtuyao Yixian Formation)in
sinensis as well as some unpublished materials of feathered theropods, pterodactyloids, Confuciusornis, insects and plants.
and feathered dinosaur Yixianosaurus longimanus.
The primitive birds discovered from the Yixian Formation in the sinensis as well as some unpublished materials of feathered theropods, and feathered dinosaur Yixianosaurus longimanus. Various fossil vertebrates have been also known from the Dawangzhangzi Senjitu-Sichakou Basin, Fengning County, northern Hebei Province (Fig. 23) pterodactyloids, Confuciusornis, insects and plants. The primitive birds discovered from the Yixian Formation in the Bed of Yixian Formation at the Hejiaxin, Wangjiagou, and Wanfotang includeJibeinia luanhera and the most primitive enantiornithine Protopteryx Various fossil vertebrates have been also known from the Dawangzhangzi
Senjitu-Sichakou Basin, Fengning County, northern Hebei Province (Fig. 23)
localities, Yixian County, Jinzhou, Liaoning (Fig. 22). They include hundreds
jengningensis. Some very primitive acipenseriform fishes, Protopsephurus liui and
of Hyphalosaurus, the iguanodontid]inzhousaurus yangi, birds, pterodactyloids
Yanosteus longidorsalis, were also found from these localities.
fengningensis. Some very primitive acipenseriform fishes, Protopsephurusliui and Bed of Yixian Formation at the Hejiaxin, Wangjiagou, and Wanfotang include]ibeinia luanhera and the most primitive enantiornithine Protopteryx of Hyphalosaurus, the iguanodontidJinzhousaurus yangi, birds, pterodactyloids Yanosteus longidorsalis, were also found from these localities.
localities, Yixian County, Jinzhou, Liaoning (Fig. 22). They include hundreds
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25 Field excursion to the Boluochi locality
a25
Field excursion to the Boluochi locality Uiufotang Formation) in Chaoyang, (Jiufotang Formation)in Chaoyang, Liaoning, the 5th Inrernationalmeeting Liaoning, the the 5thsymposium International of SAPE and on meeting the Jehol 2000). ofBiota SAPE(May, and the symposium on the Jehol Biota (May, 2000).
j.
26 lVPP excavation sites of the year 2000
~26 .o
(Below) and 200 Isites (Right) IVPP excavation of at theShangheshou year 2000 locality (Jiufotang Formation) (Below) and 2001 (Right) at Shangheshouin Qidaoquanzi, Chaoyang, Liaoning. locality (Jiufotang Formation) in Qidaoquanzi, Chaoyang, Liaoning.
Jingangshan Bed (Member) of the upper Yixian Formation The
tenztis, some undescribed pterodactyloid pterosaurs, and birds. Lycoptera muroii
]ingangshan fossil Bed consists of the lake deposits of the upper Yixian Jingangshan Bed (Member) of the upper Yixian Formation The Formation at the ]ingangshan locality in Yixian County (Fig. 24). The Jingangshan fossil Bed consists of the lake deposits of the upper Yixian sediments are mainly gray, gray-black sandstones, shales and mudstones with Formation at the Jingangshan locality in Yixian County (Fig. 24). The rich tuffaceous components. The vertebrate assemblage here includes only a sediments are mainly gray, gray-black sandstones, shales and mudstones with few taxa such as Lycoptera muroii, Manchurochelys manchoukuoensis, Yabeinosaurus rich tuffaceous components. The vertebrate assemblage here includes only a
istenuis, especially abundant in pterodactyloid this bed. some undescribed pterosaurs, and birds. Lycopteramuroii Boluochi Bed (Member) of the Jiufotang Formation The Boluochi is especially abundant in this bed. fossil Boluochi bed is represented by theofdeposits at theFormation Boluochi, Dapingfang, Bed (Member) the Jiufotang The Boluochi Dongdadao, and Shangheshou localities in Chaoyang (Fig. 25). fossil bed isLianhe represented by the deposits at the Boluochi,City Dapingfang, is about 50 km west to Chaoyang. The Shangheshou The Boluochi Lianhe localityand Dongdadao, Shangheshou localities in Chaoyang City (Fig. 25).
few taxa such as Lycoptera muroii, Manchurochelysmanchoukuoensis, Yabeinosaurus
The Boluochi locality is about 50 km west to Chaoyang. The Shangheshou
suburb of the Chaoyang City. Other Other localities with locality lies in the west suburb comparable fossil-bearing deposits include Meileyingzi and Shengli, Chaoyang; Wujiatun Wujiatun and Xierhuqiao, Xierhuqiao, Yixian. The vertebrate assemblage comprises more than 20 genera and species including Jinanicbtbys, Sinopterus dongi, includingJinanichthys, Sinopterus dongi,
Chaoyangopterus zhangi, Liaoningopterus Chaoyangopterus Liaoningopterus gui, Microraptor Microraptor zhaoianus, zhaoianus, M. gui, Cathayornis, Chaoyangia, Chaoyangia, Longipteryx, Yanornis, Yixianornis, Sapeornis Sapeornis and Cathayornis, Longipteryx, Yanornis, Jeholornis. Cathayornis avian fauna from this bed represents the second Jeholornis. The Cathayornis avian fauna of the Jehol ]ehol Biota. In 1987, a farmer in Shengli, Chaoyang, Liaoning Province discovered the first fossil fossil bird in the Province, later studied by Paul Sereno and Chenggang Rao and named as Sinornis Sinornis santensis. santensis. In 1990, Zhong-he Zhong-he Zhou, then a paleoichthyologist working at the IVPP found several bird specimens from the Lower Cretaceous Cretaceous Jiufotang ]iufotang Formation Formation in the locality of Boluochi, Chaoyang while collecting fishes. Among them was a rather complete bird and was described by Zhou and his colleagues in 1992 as Cathayornis Cathayornisyandica. yandica. This was the first bird from the Jehol ]ehol Biota collected by a professional
paleontologist in this region. More than three dozens of bird specimens were Dapingfang locality Uiufotang Formation) in the vicinity of Chaoyang, Liaoning. ~_ 2 727 Dapingfang locality (Jiufotang Formation) in the vicinity of Chaoyang, Liaoning.
unearthed from Boluochi in the following four years by the IVPP field crew. In 2000, the IVPP team excavated abundant birds, fishes fishes and insects in the Jiufotang Shangheshou locality, Chaoyang (Fig. 26). ]iufotang Formation at the Shangheshou The following year, another even larger-scale excavation there resulted in the
discoveries of over a dozen birds, feathered theropods, pterosaurs and turtles as well as abundant fishes. fishes. The Shangheshou locality has since become one of productive localities of the ]iufotang the most productive Jiufotang Formation, much like the Sihetun locality of the Yixian Formation. At the same time, various important
vertebrate fossils fossils were also collected in the neighboring areas. Consequently,
Longipteryx several new vertebrate taxa have been recognized such as the birds Longipteryx
chaoyangensis and Yanornis Yanornis martini. It chaoyangensis It is worth worth noting that that the smallest dinosaur Microraptor "Archaeoraptor" Microraptor zhaoianus zhaoianus as well as the infamous specimen "Archaeoraptor"
that was composed of the tail of the dinosaur Microraptor Microraptor zhaoianus zhaoianus and the body of the bird Yanornis Yanornis martini were also collected from the the]Jiufotang iufotang Formation in these areas. During (2001---2003), hundreds During the past past three years (2001-2003), hundreds of beautifully preserved vertebrate fossils fossils have been collected from the Dapingfang and the neighboring Dongdadao Dongdadao and Lianhe localities, Chaoyang (Fig. 27). These fossils andJeholornis,pterosaurs Sinopterus, fossils include birds Sapeornis Sapeornis andJeholornis, Sinopterus, Chaoyangopterus Chaoyangopterus 28 Wujiatun locality Uiufotang Formation) in Yixian, Jinzhou. Liaoning.
m 2 8 Wujiatun locality (]iufotang Formation) in Yixian,Jinzhou, Liaoning.
and Liaoningopterus Liaoningopterus and dromaeosaur Microraptor Microraptor gui.
usingmethod, 40 Arj39Ar method, with total fusion and incremental-heating analysis At approximately the same many important fossils4°Ar/39Ar At approximately the same time, manytime, important vertebrate vertebrate fossils using with total fusion and incremental-heating analysis Wujiatun and Xierhuqiao localities (Fig.of28),ofalso of werefrom collected from the and were collected the Wujiatun Xierhuqiao localities (Fig. 28), also of sanidine and biotites from the tuff samples from the Sihetun sanidine and biotites from the tuff samples from the Sihetun locality andlocality and Yixian County. They include fishes Protopsephurus, Yanosteus and Sinamia, the Hengdaozi locality Lo et collected al. (1999)their collected biotite samples. the Yixian the County. They include fishes Protopsephurus, Yanosteusand Sinamia, the Hengdaozi locality where Lo etwhere al. (1999) biotitetheir samples. for the Jianshangou resultconfirmed not only confirmed the ornithurine bird Yixianornis grabaui, the turtle the A4anchurochelys, the the ornithurine bird Yixianornis grabaui, turtle Manchurochelys, The the result The not only the 125 Mathe age125 forMa the age Jianshangou Bed, but Bed, but feathered caudipterid as well as pterodactyloids and some alsothe indicated theofpresence of trapped Ar in the biotites, which helped explain feathered caudipterid as well as pterodactyloids and some other birds.other birds. also indicated presence trapped Ar in the biotites, which helped explain thepaleontological extensive paleontological the Jehol in the why the was not geologically Further, they also dated Despite theDespite extensive research onresearch the JeholonBiota in theBiotawhy the biotite age biotite was notage geologically meaningful.meaningful. Further, they also dated age ofremains sanidine from tuff the Biota to be anproblem enduring problem samples from the Tuchengzi which underlies last decade,last thedecade, precise the ageprecise of the Biota to remains be an enduring sanidine from tuff samples from the Tuchengzi Formation,Formation, which underlies the Yixian Formation unconformably. as inherited well as anone. inherited one. Currently, theredifferent are threeopinions differentasopinions as 139.4 for part the upper part as well as an Currently, there are three the Yixian Formation unconformably. The 139.4 The Ma age for Ma the age upper of the Tuchengzi lends furtherforcredence forCretaceous the Early Cretaceous regard age ofBiota, the Jehol i.e., the LateLate Jurassic, Late regard to the age to of the Jehol i.e., Biota, the Late Jurassic, Jurassic to Jurassic of thetoTuchengzi Formation Formation lends further credence the Early age ofBiota the Jehol Early Cretaceous, and Early Cretaceous. Paleontologists and base geologists Early Cretaceous, and Early Cretaceous. Paleontologists and geologists agebase of the Jehol (Fig. Biota 29). (Fig. 29). their age determination mainly on paleontological and Stratigraphic Now, the agesLujiatun, for all the Lujiatun,Jianshangou, Dawangzhangzi, their age determination mainly on paleontological and stratigraphic compari- compariNow, the isotope agesisotope for all the Jianshangou, Dawangzhangzi, as well dating. as isotope dating. using Recently, using the biostratigraphic in Jingangshan Beds and the Formation Jiufotang Formation are The available. sons as wellsons as isotope Recently, the biostratigraphic work in work Jingangshan Beds and the Jiufotang are available. YixianThe Yixian ranges approximately from Valanginian to Barremian, of the tuff in the sediments, most workers concert with the dating concert with the dating of the tuff samples in samples the sediments, most workers Formation Formation ranges approximately from Valanginian to Barremian, and the and the Jiufotang corresponds Formation corresponds to the Aptian. Recent discoveries vertebrate discoveries that Jehol the whole Biota the Early Cretaceous. have agreedhave thatagreed the whole BiotaJehol belongs to belongs the Earlyto Cretaceous. Jiufotang Formation to the Aptian. Recent vertebrate havefurther provided furthersupporting evidence supporting the Early Cretaceous However, still some workers argue For otherwise. ForLoinstance, Lo et al.have (1999) However, still some workers argue otherwise. instance, et al. (1999) provided evidence the Early Cretaceous age of the age of the Biota.most Although most workers now have the Early Cretaceous proposed the Late forpart the lower of the Yixian Formation, proposed the Late Jurassic ageJurassic for the age lower of the part Yixian Formation, Jehol Biota.Jehol Although workers now have accepted theaccepted Early Cretaceous
based on the 4°Ar/39Ar of the biotite Ma). (147 Ma). 40 Arj39Ar dating of (147 the biotite based on thedating
age for the Jehol more stratigraphic work remains to beit done age for the Jehol Biota, moreBiota, stratigraphic work remains to be done and will and it will be aissue debatable issueyears for many years to come. a joint project by the scientists from bothand theCanada IVPP and probably Canada beprobably In 1995, In 1995, a joint project by the scientists from both the IVPP a debatable for many to come. 40Arj39 Ar thetofirst to datehorizons the major horizons used4°Ar/39 the laser used the laser Ar method for method the first for time datetime the major the Jehol Group. They successfully obtained the volcanic age (from volcanic 20i of the Jeholof Group. They successfully obtained the age (from 10i deposits) ofdeposits) the Dawangzhangzi and Jingangshan beds of the Yixian of the Dawangzhangzi and Jingangshan beds ofFormathe Yixian Forma-
20
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.""
10
0i
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0
-
tion as 122.2 Ma, and 121.4+ 1.1--'121.6+0.4 Ma, tion___0.2--122.5+0.3 as 122.2±0.2-122.5±0.3 Ma, and 121.4± 1.1-121.6±0.4 Ma, respectively.respectively. It is noteworthy that in 1993,that theinage of the It is noteworthy 1993, thebasalts age ofoverlying the basaltsthe overlying the 0.04 0.04 0.03 Jiufotang Formation Inner Mongolia dated as 110_ 0.52 Ma. 0.52 Ma. 0.03 Jiufotang in Formation in Innerwas Mongolia was dated as 110± 0.02 0.02 0.01 fromcooperated the IVPP with cooperated with C. C. Swisher In 1999, paleontologists In 1999, paleontologists from the IVPP C. C. Swisher 0.01 III from theIIIBerkeley Center on the dating Yixian from theGeochronology Berkeley Geochronology Center on of thethe dating of the Yixian " 401 n = 24 2 4 ~n 40 Formation. crystal sanidine from the sediments, tuffs of the sediments, Formation. They dated They singledated crystalsingle sanidine from the tufts of the
4°ArP,¢*
11
that the J ianshangou Bed Confudusornis, that bears Con/uciusornis, Zhangheotherium and that the Jianshangou Bed that bears Zhangheotherium and
,-
105 100
!1oo
95
••
CalK
80
80
,,= 24
Analyses
Analyses
4~n =19
t :0
40
n = 19
...,..99L-HDZI
(12501 ±O I'I(ISD)
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~004
30
(SEll 1. 99L-TI 1m =( 139 44 ± 019 (ISD)± 005(SE))M.
20
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= (125.03 ± 0.18 (1 SO) ± 0.04 (SE» Ma
10
V-Pb method, resulted an age_+of ± 0.9 works Ma. These works using U-Pb using method, resulted in an age ofin125.2 0.9125.2 Ma. These
furtherthe confirmed the Early age Cretaceous age for the Jehol Biota, and indicated further confirmed Early Cretaceous for the Jehol Biota, and indicated 22
) 10
95
~
part of the Yixian 124.6 _+ is0.2---124.6 + 0.3 Ma. In 2001, part of Formation the Yixian is Formation 124.6 ± 0.2-124.6 ± 0.3 Ma. In 2001, 20. al. (2001) dated crystal with zircon from thesame tuffs locality of the same locality Wang et al. Wang (2001) etdated with zircon from crystal the tufts of the
110 : 105
"'Ar!"Io"
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thus aobtaining of the sediments the in first thus obtaining direct agea direct of the age sediments for the firstfortime thetime in the 30 Bed ofFormation. the Yixian According Formation.toAccording to their study, the lower JianshangouJianshangou Bed of the Yixian their study, the lower
~
0 122
124
126
124
128
126
130
128
132 Age/Ma
130
132
134AgeIMa136
134
138
136
140
138
140
142
142
. . 29 Graphics showing results of a recent isotopic dating on the lower feathered dinosaurs about 125isMa, about Maabout younger than the oldest featheredis dinosaurs about 125 20 Ma, 20 Ma younger than the oldest 29 Graphics showing of a recent isotopic dating on the part ofresults the Yixian Formation (125.0 Ma) and the lower underlying upper part of the Yixian Formation (125.0 Ma) and the underlying upper Tuchengzi Formation (139.4 Ma) in Beipiao, liaoning. part ofthe bird A rchaeopteryx. bird Archaeopteryx. part of the Tuchengzi Formation (139.4 Ma)in Beipiao, Liaoning. (See Swisher III et aI., 2002 for details) In 2001, and his Chinese collaborators In 2001, Swisher andSwisher his Chinese collaborators re-dated newre-dated samplesnew samples (See Swisher Ill et al., 2002 for details)
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Hua-zhan g Pan, Xian g- gen Zh.u
Hua-zhang Pan, Xiang-gen Zhu
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bottoms, while others swim or float at the surface, often forming a conspicuastropods (snails and slugs) are the most abundant, most diverse, (snails slugs)ofareextant the most abundant, while others swim or float at the surface, oftenare forming a conspicuand astropods perhaps the bestand known mollusks. Theymost havediverse, ous partbottoms, of the plankton, but the overwhelming majority of snails crawling and perhaps37,500 the best known of extant mollusks. They part ofInthe but the overwhelming majority of snailsand are crawling approximately living species, and represent some 80 have on the ous bottoms. theplankton, inland freshwaters, gastropods are also common,
G
approximately species, and represent 80 on the bottOms. In the inland fresh waters, gastropods are also common, and percent of the living mollusks, 37,500 only less living than insects in number among some all they are represented by some forms of the Prosobranchia as well as the percent ofMore the living mollusks, in number all they are represented by some forms of the Prosobranchia as well as the invertebrates. than half of these only snailsless and than slugs inseCts are marine animals, among and Basommatophora from the Pulmonata, e.g., viviparids, bithynids, planorbids, More than half of these snails and slugs are marine animals, and BasommatOphora from the Pulmonata, e.g., viviparids, bithynids, planorbids, theyinvertebrates. are the only molluscan class to have spread both into freshwater and on lymnaeids, etc., commonly found in lakes or rivers. Their fossils often they are the only molluscan class to have spread both into freshwater and on etc., commonly found in lakes or rivers. Their fossils often lymnaeids, land. Gastropods include three subclasses: Prosobranchia, Opisthobranchia, occurred in non-marine deposits from the Mesozoic to Cenozoic. Some forms land. Gastropods include three subclasses: Prosobranchia, Opisthobranchia, occurred in non-marine deposits from the Mesozoic to Cenozoic. Some forms and Pulmonata. Both the prosobranchs and the opisthobranchs are mostly of Stylommatophora may be seen in terrestrial habitats from deserts to and Pulmonata. Both the prosobranchs and the opisthobranchs are mostly of StylommatOphora may be seen in terrestrial habitats from deserts to marine inhabitants with high diversity and wide distribution. Generally, tropical rainforests and montane altitudes of nearly 5,000 m above the sea marine inhabitants with high diversity and wide distribution. Generally, tropical rainforests and montane altitudes of nearly 5,000 m above tbe sea many marine prosobranchs have well-developed and colorful shells with level. The variations and special adaptations of these snails reflect a wide range many marine prosobranchs have well-developed and colorful shells with level. The variations and special adaptations of these snails reflect a wide range beautiful sculptures. A great number of them are found on rocky, sandy, and of habitats. The majority of gastropods are carnivores, herbivores or occasionally, beautiful sculptures. A great number of them are found on rocky, sandy, and ofhabitats. The majority ofgastropods are carnivores, herbivores or occasionally, muddy bottoms in the shallow sea areas. The Pulmonata are the only scavengers. Identification of fossil gastropod taxa mainly depends on protoconch muddy bottoms in the shallow sea areas. The Pulmonata are the only scavengers. Identification offossil gastropod taxa mainly depends on protoconch molluscan subclass that lead a successful life on land, with some returning to characters, shape of shell, aperture and ornamentation. molluscan subclass that lead a successful life on land, with some returning ro characters, shape of shell, aperture and ornamentation. a secondarily freshwater dwelling. Some gastropods burrow into the soft Fossil gastropods are very common in the Jehol Biota, and widely a secondarily freshwater dwelling. Some gastropods burrow into the soft Fossil gastropods are very common in the Jehol Biota, and widely
30 Shell of Probaicalia gerassimovi (3.34 mm long, 30 Shell of ProbaicaJia gerassimovi (3.34 mm long, 1.3 mm wide), a micromelaniidsnail in apertural 1.3 mm wide), a micromelaniid snail in apertural and ventral views, from Jianshangou locality and ventral views, from Jianshangou locality (Yixian Formation)in Beipiao, Liaoning. (Left) (Yixian Formation) in Beipiao, Liaoning. (Left)
31 Shell31of Pseudarinia yushugouensis (1.81 mm long, Shell of PseLidarinia yLlshLigoLiensis (1.81 mm long, 0.80 mm0.80 wide), a cyclophorid snail in apertural mm wide), a cyclophorid snail in apertural view, from Pijiagou locality (]iufotang view, from Pijiagou locality Uiufotang Formation) in Yixian, Liaoning. (Right) Formation) in Yixian, Liaoning. (Right)
31
38
~8
32 Shells ~of (2.58 philippii mm long. 1.00mm mmlong, wide). an mm ellobiid snail apertural. 3 2Ptychostylus Shells ofphilippii Ptychostylus (2.58 1.00 wide), an in ellobiid snail ventral. in apertural, ventral, apertural and apertural apical views. (Yixianlocality Formation) in Formation) Beipiao. Liaoning. and from apicalSihetun views, locality from Sihetun (Yixian in Beipiao, Liaoning.
33 Shell of Gyraulus sp. (0.75 mm long. 2.25 mm wide). a planorbid snail in apertural. apical and basal
~33
Shell of Gyraulus sp. (0.75 mm long, 2.25 mm wide), a planorbid snail in apertural, apical and basal views, from Sihetun locality (¥ixian Formation) in Beipiao, Liaoning.
views. from Sihetun locality (Yixian Formation) in Beipiao. Liaoning.
eastern Asia (including Transbaikalian area of Russia; ItsouthIt is noteworthy these gastropod faunas include mostly very tiny forms, is noteworthy that these that gastropod faunas include mostly very tiny forms, distributeddistributed in eastern in Asia (including Transbaikalian area of Russia; southeastern Mongolia; Anhui provinces, southern Hebei, general 5 mm long. general in less than 5less mmthan long. eastern Mongolia; Zhejiang, Zhejiang, Anhui provinces, southern China andChina Hebei,and in other provinces northern China). Based on its stratigraphic distribution fossil characteristics, the Based on its stratigraphic distribution and fossil and characteristics, the Shandong,Shandong, Henan andHenan some and othersome provinces of northernofChina). They are They are mainly found in the Dabeigou, and Formations Jiufotang Formations fauna of Biota the Jehol Biotarelated is closely those of the Middle gastropod gastropod fauna of the Jehol is closely to related those oftothe Middle mainly found in the Dabeigou, Yixian andYixian Jiufotang of northernof northern andLiaoning, western Liaoning, and equivalent of other The Purbeck Hebei andHebei western and equivalent horizons ofhorizons other areas. Theareas. fauna represented is largely represented by Probaicalia (Fig. 30), Pseudarinia 31), western fauna is largely by Probaicalia (Fig. 30), Pseudarinia (Fig. 31), (Fig.
Purbeck Bed of Dorset,England, southern the England, theFormation Serpulit Formation Bed of Dorset, southern Serpulit of north- of northwestern Germany, Lower Cretaceous of southeastern and Germany, the Lowerthe Cretaceous of southeastern Mongolia Mongolia and
of Reesidella, of the Prosobranchia, 32) Transbaikalia. Russian Transbaikalia. This that indicates that the gastropod opercula ofopercula Reesidella, viviparids viviparids of the Prosobranchia, PtychostylusPtychostylus (Fig. 32) (Fig. Russian This indicates the gastropod fauna bearsfauna obvi-bears obviand Zaptychius of Ellobiidae, Cyraulus (Fig. 33) of Planorbidae Calbacharacteristics ously characteristics those of Cretaceous. the Early Cretaceous. of those ofofthe Early and Zaptychius of Ellobiidae, Gyraulus (Fig. 33) of Planorbidae and Galbaand ously of Lymnaeidae (Fig. 34). Sometimes are low relatively low inGastropodsGastropods range ofbut habitats, butsensitive they aretosensitive have wide have rangewide of habitats, they are slight to slight sphaira of sphaira Lymnaeidae (Fig. 34). Sometimes the faunas the are faunas relatively in or even absenthorizons. in some Once horizons. were inchanges in environment, such asofstability of thetype bottom, type of sediments, environment, such as stability the bottom, of sediments, diversity ordiversity even absent in some theyOnce were they foundwere theyfound were theychanges of individuals. The gastropod fauna, scattered always in a large salinity, food water supplies, water depth, temperature, oxygen and content, and salinity, food supplies, depth, temperature, oxygen content, always found in a found large number ofnumber individuals. The gastropod fauna, scattered in the tuffaceous of Bed 9 below bird-bearing ofpart the lower part turbidity. Thus, are gastropod different gastropod assemblages in different turbidity. Thus, there are there different assemblages in different in the tuffaceous siltstone ofsiltstone Bed 9 below bird-bearing rocks of therocks lower of the Yixian Formation nearvillage, Sihetun village,Liaoning, western Liaoning, a low Many modern settings. Many scholars have the examined the environments scholars have examined environments of modern of snails, so snails, so of the Yixian Formation near Sihetun western has a low hassettings. diversity a highe.g., density, e.g., one (227 specimen (227 cm 2 in area) with theofhabitats of fossil gastropods can be extrapolated from the comparisons the117 habitats fossil gastropods can be extrapolated from the comparisons diversity and a highand density, one specimen c m 2 in area) with 117 of fossil gastropods are living counterparts. Different assemblages fauna Amplovalvata contains Amplovalvata sp., Probaicalia individuals. with their with livingtheir counterparts. Different assemblages of fossil gastropods are individuals. The faunaThe contains sp., Probaicalia vitimensis, vitimensis, found deposits in severalthat deposits that are interpreted to different representkinds different harpaeformis and Gyraulus found in several are interpreted to represent of kinds of ProbaicaliaProbaicaliagerassim(JVi, gerassimovi,PtychostylusPtychostylusphilippii, philippii, PtychostylusPtychostylus harpaeformis and Gyraulus habitat areas. but it is dominated overwhelmingly by Ptychostylus, constituting about areas. habitat sp., but it sp., is dominated overwhelmingly by Ptychostylus, constituting about to theircounterparts, modern counterparts, of the fossil 66.5% the totalofnumber of specimens in the gastropod fauna. PseudariniaAnalogousAnalogous to their modern populationpopulation of the fossil 66.5 % of the totalofnumber specimens in the gastropod fauna. Pseudarinia increased individual number but body decreased yushugouensis, Cyraulus loryi and from the Jiufotang of the gastropods PtychostylusPtychostylus increased individual in numberinbut decreased in body in yushugouensis, Gyraulus loryi and Galba fromCalba the Jiufotang FormationFormation of the gastropods usuallyaindicates brackish water living environment. Pijiagou section western in Yixian, westernare Liaoning are persevered in purple siltstone. size usuallysize indicates brackish awater living environment. PtychostylusPtychostylus mainly mainly Pijiagou section in Yixian, Liaoning persevered in purple siltstone. in theFormation Serpulit Formation of North-West Germany associated with occurred inoccurred the Serpulit of North-West Germany associated with and neomiodontids the brackish water Hydrobia Hydrobia and neomiodontids (bivalve), (bivalve), showing showing the brackish water environment; from Inter-marine Beds Building or Upper Stone Building Stone environment; PtychostylusPtychostylus from Inter-marine Beds or Upper nearCinder marine Cinder andBeds Corbula Beds of Middle Beds Purbeck near marine Beds, andBeds, Corbula of Middle Purbeck in Beds in Durlston Bay,isEngland is associated with fish and bivalve Neomiodon Durlston Bay, England associated with turtles, fishturtles, and bivalve Neomiodon brackish water There affinities. are a fewbeds gypsum in part the lower of brackishofwater affinities. are aThere few gypsum in thebeds lower of part of Yixian Formation near Sihetun village,Liaoning. western All Liaoning. All the factors the Yixianthe Formation near Sihetun village, western the factors indicate that Ptychostylus-dominated from the indicate that Ptychostylus-dominated assemblageassemblage from the lower partlower of thepart of the Yixian Formation nearVillage, Sihetunw.Village, w. might Liaoning be of brackish Yixian Formation near Sihetun Liaoning be might of brackish water setting, butloryiGyraulus loryi- and Calba-dominated water setting, but Gyraulus and Galba-dominated assemblageassemblage from the from the may represent drifting assemblage JiufotangJiufotang FormationFormation may represent a drifting a assemblage frequentlyfrequendy occurred thea shore of lake. a river or lake. occurred along the along shore of river or (All photographs in this chapter taken byYuan/NIGP) Liu-Ping Yuan/ NIGP) (All photographs in this chapter were takenwere by Liu-Ping
~34
Shell sphaira of Galbo(1.20 sphaira Shell 34 of Galba mm(1.20 long, mm 0.74long, mm 0.74 wide),mm a wide), a Iymnaeid snail in apertural, ventral apical views, from lymnaeid snail in apertural, ventral and apicaland views, from Pijiagou locality Uiufotang Formation) in Yixian, Liaoning. Pijiagou locality (Jiufotang Formation) in Yixian, Liaoning.
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B
ivalvia are a class of bivalved mollusks, also known as Pelecypod a and LameUibranchiara in rhe literature. The bivalves are aquatic animals with a shell consisting of two matching lateral valves united by a dorsal horny ligament. The
valves can be drawn togethet by a pair of adductor muscles (reduced to one in some) against the opening counter-fotce of the ligament. Today the bivalves are among the most common benthic invertebrates, especially on marine shelves, although there are many species in fresh-water or brackish-water environments and even in abyssal habitats. The laterally compressed shape characteristic of the great majority of bivalves renders them well adapted for burrowing in sandy or muddy substrates. Some bivalves are borers in solid
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worms"). Among the benthic epifauna, bivalves form an important element. Some rely on
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the weight or the shape of their shell to maintain their position on the seafloor; others attach
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themselves to stationary objects by means of their byssus or by cementation of one of the valves. Some highly mobile bivalves, for example the pectinids, can "hop" near the sea bottom for short distances. No living bivalves are pelagic, except for larval stages and very few species with strongly reduced shells. However, some extinCt thin-shelled bivalves are considered to have possibly lived pelagically. The first occurrences of Bivalvia are known from the Lower Cambrian at a few localities in the United States, southern Australia, and China. During the Early Paleozoic periods, bivalves were not abundant. Since the Devonian, bivalves became very frequent and diversified. The class reached the acme of its development in the Mesozoic and Cenozoic. In most instances, Mesozoic bivalves are useful for stratigraphical correlation. The Jehol Fauna arose after an important biotic extinction event, which tOok place between the latest Middle Jurassic and early Late Jurassic in China, accompanied by the strong tectOnic movements during the Oxfordian and early Kimmeridgian (Late Jurassic). Owing to this extinction event, the Eolamprotttla-Psiittnio fauna and the Psettdocatdinia fauna, flourished in the Middle Jurassic of the Chinese continent, are missing from the Jehol Fauna. The bivalves of the Jehol Fauna manifested themselves with a new feature
35 Multiple species of Arguniella (15- 25 111m long of the individuals). a sibireconchid bivalve in aggregation. from the roadside of Sihetun to Libalanggou (lower part ofYixian Formation) in Beipiao. Liaoning. (Photo: Da-jian Li/ CAS)
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Internal moulds of Mengyinaia. a unionid bivalve. from Ningjiagou locality (Mengyin Formation) in Mengyin. Shandong. a. Mengyinaia F~
mengyinensis (65 mm long): b. Mengyinaia lugrigensis (90 mm long).
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Internal moulds of Sp!Jaerium. a mall·sized pisidiid bivalve. from Xiwa locality (Xiwa Formation) in :i'~' ~,!~:
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greatly differing from the Middle Jurassic ones. Among them, three fresh-
locality of this primitive fossil bird (Fig. 35). Although it is very abundant in
water assemblages are recognizable on the basis of bivalve samples respec-
number of individuals, the assemblage possesses only four species in two
genera,locality Arguniella Sphaerium jeholense andabundant S. tively from the formations the indices the Jehol Fauna. The three three freshofthilingyuanensis, primitive foA. ilyanshanensis, bird (Fig. 35). Although it i very in greatly differing from yielding the Middle Juras icofone. Among them, anderssoni, indicating emerging the but assemblage still low diversity with species some in two number of individuals, po ecompared es only four water a semblages are recognizable on the basis of bivalve samples re pecassemblages are as follows. other groups (e.g., ostracods and insects) phaerimfl and somejeholeme verte- and . genera,ofAinvertebrates rgll1li lIa ling)'lIanemiJ, A. )'fmJhanemiJ tively from the formation the indiceincludes of th Jehol Fauna. of The 1. Arguniella assemblage.yielding The assemblage two species a three as emblage as follows. single genus, i.e.,are Arguniella lingyuanensis and A. yanshanensis, found from the
brates cmdemolli, (e.g., birds)indicating in the sameemerging phase. Therefore, radiation in thewith orne but still the lowbivalve diversity compared
Jehol was delayed, probably owing slow and evolutionary ratessome of verteArglllliellain aeastern emblage. a emblage twO species of a Fauna other groups of invertebrates (e.g.,to0 the tracods insects) and Dabeigout.Formation HebeiThe province, showinginclude a very low diversity genu, Arglmiella or ling)'lIanensiJ and of A. the )'a/lShanemiJ, found from the same phase. Therefore, theshells bivalve single brate (e.g., birds) in thewere these bivalve mollusks, which protected by their solid andradiation most in the of the bivalves at i.e., the beginning the early phase faunal development Dabeigou Formation in eastern Heb i province, with howing diversity Jehol \ a todelayed, probably owing toat the probably hadFauna adapted the adverse environments that low time.evolutionary rates of of the Jehol Fauna. The assemblage is comparable that ainvery the low Upper of theMayakskaya bivalve at the beginning earlyreaches ph e of e bivalve mollusk, which were protected by theirIt is olid hells 3.theMengyinaia-Nakamuranaia-Sphaerium assemblage. from theand mo t Jurassic Formation in or thethe upper of the the faunal Argundevelopment River, probably had adapted of the Jehol Russia. Fauna. The assemblage i comparable with that in the pper to the adverse enJehol ironments that time. fossiliferous horizons of the late phase of the Fauna,atthe Jiufotang eastern Siberia, 3. western Mengyinaiaakamllrcmaiaphaerillm a emblage.is It i from the Juras ic Mayakskaya Formation in the of thebelongs ArguntoRiver, Formation in Liaoning and its coevals. The assemblage more 2. Arguniella-Sphaerium assemblage. Thisupper bivalvereaches assemblage fos and iliferou horizon of the pha e two, of the Jehol Fauna, eastern iberia, Rus ia. abundant diverse in bivalves thanlate the above consisting of morethe than]iufotang the main phase of the Jehol Fauna, found from the lower Yixian Formation 2. Arguniella- phaerillln as emblage. Thi bivalve as emblage belong to Formation in west rn Liaoning and its coeval. The as embtage is more in western Liaoning and northern Hebei provinces. Particularly rich bivalves 30 species designated to five genera including five species to Mengyinaia (Fig. abundant and diverse in bivalves than the above twO con i ting of more than the main pha e of the Jehol Fauna, found from the lower Yixian Formation 36), three to Nakamuranaia (Fig. 37), 12---13 to Sphaerium (Fig. 38), six to of the assemblage occur just below the bird (Confuciusornis)-bed at the type in we tern Liaoning and northern Hebei province. Particularly rich bivalve 30 pecies de ignated to five genera including five pe ie to Alengyiflaia (Fig. Arguniella and five to Weichangella (Fig. 39), 36), three to akamlmmaia (Fig. 37), 12-13 to phaerilllll (Fig. 38), six to of the assemblage occur JUSt below the bird (Coll!lIciIlJomiJ)-bed at the type representing a distinct faunal radiation. This Arglmiella and five to Weichallgella (Fig. 39) assemblage was succeeded by the Nippononaia repre enting a distinct faunal radiation. Thi sinensis-N, cf. tetoriensis-Tetoria cf. yokoyamai assemblage was succeeded by the ippollonaia assemblage found from the Shahai Formation JinemiJ- . cf. letoriemiJ-Tetoria cf. yoko)'amai in Liaoning, which overlays the Jiufotang assemblage found from the hahai Formation Formation and can be correlated with the in Liaoning, which overlay the Jiufotang beds of the upper Itoshiro Subgroup of the Formation and can be correlated with the Tetori Group in Japan. The latter beds of beds of the upper lroshiro ubgroup of the Japan have been recently dated, based on Tetori roup in Japan. The latter bed of ammonites and marine bivalves, as Japan have been recently dated, based on Hauterivian (Early Cretaceous). Therefore, ammonites and marine bivalve, a the present assemblage is regarded to be Hauterivian (Early Cretaceoll ). Therefore, around Valanginian in age. the pre ent assemblage i regarded to be around Valanginian in age.
Shells of Weichangella qingquanen; (29-34 mm long), a plicatounionid bivalve, from halingou locality Qiufotang Formation) in Weichang, Hebei. (Photo: Pu Wang! GPH)
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onchostracans, or clam shrimps, are small, bivalved branchiopod onchostracans, or clam are laterally small, bivalved branchiopod crustaceans (Fig. 40). Theyshrimps, have a short compressed body
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crustaceans (Fig.two 40).lateral They valves have a that shortconstitute laterally compressed enclosed between the chitinousbody
enclosed between twO lateral valves that constitute the chitinous carapace with an interior membranous lining. The head is located anteriorly carapace with an inrerior membranous lining. The head is located anreriorly
and bears sessile, paired compound eyes with well-developed ocelli. Posteriorly,
and bears sessLle, paired compound eyes with well-developed ocelli. Posteriorly,
the telson is distinguished by two flattened, upwardly curved processes or
the telson is distinguished by two flattened, upwardly curved processes or
claws. The antennules are small and simple, but the antennae are modified
claws. The anrennules are small and simple, but the anrennae are modified
into powerful biramous swimming organs. During mating, males use them
inro powerful biramous swimming organs. During mating, males use them
to aid in seizing the female carapace. Trunk legs (swimming legs) vary from to
aid in seizing the female carapace. Trunk legs (swimming legs) vary from
10 to 32 pairs.
10 to 32 pairs.
These animals occur in non-marine facies sporadically, but often
These animals occur in non-marine facies sporadically, but often
abundantly from the Devonian to the present day. Many paleontologists still
abundantly from the Devonian to the presenc day. Many paleonrologists still
refer to them informally using the pre-occupied generic name "Estheria", refer to them informally u ing the pre-occupied generic name" Estheria", originally originallyused usedforfora fly. a fly. The flourished twice in the Late Paleozoic of Europe and and Theclam clamshrimps shrimps flourished twice in the Late Paleozoic of Europe ininthe Asia with 200 genera in 27 families, but but rapidly theMesozoic Mesozoicofof Asia withabout about 200 genera in 27 families, rapidly declined with only 1414 genera in in 5 families stillstill living today. theCenozoic Cenozoic with only genera 5 families living tOday. declinedininthe The reside in in small andand temporary inland ponds, Theliving livingconchostracans conchosrracans reside small temporary inland ponds, flood-plain ditches, andand almost anyany shallow flood-plainpools, pools,rice ricefields, fields,roadside roadside ditches, almost shallow depressions water. They have also been reported from spring water, depressionsfilled filledwith with water. They have also been reported from spring water, along coastal saltsalt flats. They appear in the alongmargins marginsofoflarge largelake lakeand andonon coastal flats. They appear in the Yangtze spring or or in in North China, Northwest YangtzeRiver Riverdrainage drainagearea areaduring during spring orth China, orrhwest China,Northeast orrheastChina, China,and andInner Inner Mongolia during summer early China, Mongolia during summer andand early autumn.The Themost mostfavorable favorabletemperature temperature majority of living forms autumn. forfor thethe majority of living forms is is
13toto34°C. 34°C. 13 Thereare aretwo twOkinds kindsofof conchosrracan eggs: thin-shell eggs hatch There conchostracan eggs: thethe thin-shell eggs cancan hatch afterseveral severaldays daysofof ovulationbyby females; thick-shell dormanr eggs could after ovulation females; thethe thick-shell dormant eggs could withstandlong longdesiccation desiccationororfrozen frozenof of pools, sub eguently withstand thethe pools, so so as as subsequently to to becomedispersed disper edbybywind, wind,water water birds hatch even a number of years become oror birds andand hatch even a number of years later.This Thismay mayexplain explainthe theworldwide worldwide distribution fossil clam shrimps, later. distribution of of fossil clam shrimps, whichhas hasgreat greatpotential potencialininbiostratigraphical biostratigraphical research, especially which research, especially for for the the non-marine) urassic and Cretaceous in eastern Asia. non-marine Jurassic and Cretaceous in eastern Asia.
The taxonomy of living conchostracans is based on the structures of soft body, which is seldom preserved as fossils. Thus, we have to pay more attention to Thethe taxonomy variationofofliving carapace concho andtracans the minute is basedornamentation on the structures of of oft body, which is eldom as fossils. Thus we to pay more growth bands of valve in fossilpre clamerved shrimps. For example, the have euestheriids attention to the bear variation of carapace ornamentation of and loxomegaglyptids reticulate sculptureand and the the minute orthestheriids radial of valve in have fossil the clam shrimps. For example the euestherijds lirae, growth whereasbands the eosestheriids mixed sculpture of reticulation and loxomegaglyptids bear reticulate culprure and the orrhestheriids radialand lirae. From the Jehol Biota, three conchostracan assemblages can be radial lirae, whereas the eosestheriids have rhe mixed sculpture of reticulation and clearly identified chronologically as follows. radial lirae. From the ]ehol Biota, three conchosrracan emblages Nestoria-Keratestheria assemblage: Besides its namesakeas forms, the can be clearly includes identifiedSentestheria, chronologically as follows. assemblage Abrestheria, Ambonella, andJibeilimnadia. estot'ia-Keratestheria a semblage: Be ides ir namesake forms, the Generally, they have a large carapace with several brawny growth lines assemblage Abresthel'ia, and Jibeilimnaclia. sometimes bearingincludes spines or mtesthet'ia, nodes. Growth bands areAmbomlla, broad and flatted with Generally, they have a large carapace withfound several growth line big reticulate sculpture (Fig. 41). These fossils were frombrawny the Dabeigou
~40
Eocyzicusmongolianus, a recent conchostracan
sometimes bearingHebei spinesProvince, or nodes. the Growth bands are or broad and flarred with Formation of northern Baoshi, Murei, Baiyingaolao
arthropod from Zhenglanqi, Inner Mongolia. (Photo: Ji-liang Mao/NIGP) a recent conchosrracan 40 EocyzicLls mongolianLls,
big reticulare sculpture (Fig. 41). These fossils wereand found from Formathe Dabeigou Formations of Inner Mongolia of China and the Argun Ust'kal Formarion of norrhern Hebei Province, the Baoshi, Murei or Baiyingaolao
m
arthropod from Zhenglanqi, Inner Mongolia. (Photo: Ji-liang Mao/ NIGP)
Formarions of Inner Mongolia of Chjna and the Argun and U t'kal Forma-
:z: or-
tions of eastern Transbaikalia.
tions of eastern Tran baikaJja.
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45
m 41 Carapace valves of Nestoriapissovi (14 mm long), a loxomegaglyptid 41conchostracan Carapace valves Nestoria pis ovi with of several brawny (14 mm long), a loxomegaglyptid growth lines ornamented by large concho tracan severalfrom brawny reticulations in with between, growth lines ornamented by large Dadianzi locality (Dabeigou in between, from reticulations Formation) in Luanping, Hebei. Dadianzi locality (Dabeigou (Photo: Da-jian Li/CAS) Formation) in Luanping, Hebei. (Phoro: Da-jian LiI CAS)
Eosestheria-Diestheria assemblage: These two genera were absolutely predominant in quantity among this assemblage when they lived in the
Eosestheria-Diesfheria assemblage: twO genera were Paleo-Amur River valley including InnerThese Mongolia, Northeast andabsolurely North predominant in and quantity among Their this assemblage when from they the lived in the China, Mongolia Transbaikalia. fossils are collected Yixian Paleo-Amur River valley including Inner North Formation of western Liaoning Province, NE Mongolia, China and theortheast relevant and horizons China, and well-known Transbaikalia. Their fossils are collected from the Yixian in otherMongolia regions. The forms of this assemblage are Eosestheria aff. Formation Liaoning Province (Fig. NE China the relevant horizons middendorfiiof western (Fig. 42), E. lingyuanensis 43), and E. ovata (Fig. 44), E. other regions. Diestheria The well-known forms thisand as emblage are In Eosestheria in jingangshanensis, yixianensis (Fig.of45) D. jeholensis. addition,aff.
middenc/o/Iii (Fig. 42), E. lingyllanensis (Fig. 43), E. ovata (Fig. 44), E. some other species of Eosestheriopsis has been recognized recently from the jingangshanensis, Diestheria yixianensis (Fig. 45) and D.jeholensis. In addition, Jehol Fauna, but in the past were found only from the Upper Jurassic deposits some other specie of Eosesfheriopsis has been recognized recently from the such as the Penglaizhen and Tuodian Formations of southwestern China. Jehol Fauna, but in the past were found only from the Upper Jurassic deposits
Eosestheria has a relative large carapace with moderate growth lines but
uch as the Penglaizhen and Tuodian Fotmations of sourhwestern China.
without spines or nodes. Growth bands are ornamented with irregular Eosestheria has a relative large carapace with moderate growth lines but without spines or nodes. Growth bands ate ornamented with irregular
42 Carapacevalves of Eosestheria aft. middendorfii (13--14 mm long), an eosestheriid conchostracan with mixed sculpture of reticulation and radial lirae on growth 42 Carapace valves ofEosestheria aff.middendorjii (13-14mmlong), an eosestheriid bands (enlarged detail), from Zaocishan locality (lingangshan Bed of the upper conchostracan with mixed sculpture of reticulation and radial lirae on growth Yixian Formation) in Yixian, Liaoning. (Photo: Da-jian Li/CAS, Mao-fang Pang/ bands (enlarged detail), from Zaocishan locality Uingangshan Bed of the upper NIGP) Yixian Formation) in Yixian, Liaoning. (Photo: Da-jian Lit CAS. Mao-fang Pang! NIGP)
m 4 3 Carapace valves of Eosestheria lingyuanensis (11 mm long), an eosestheriid conchostracan, from Sihetun locality (Jianshangou Bed of the lower Yixian _43 Carapace valves of EosestlJeria lingyuanensis (J J mm long). an eosestheriid Formation) in Beipiao, Liaoning. (Photo: Da-jian Li/CAS) conchostracan, from Sihetun locality Uianshangou Bed of the lower Yixian Formation) in Beipiao. Liaoning. (Photo: Da-jian Lil CAS)
medium medium reticulations reticulation transferring tran ferring gradually lirae near gradually into intothe theradial radiallirae nearvenventral and posterior po teriormargins marginsofofthe thevalve. alve. tral and The The ornamentation ornamentationofofDiestheria Diestheriais issimisimilar to to that that of ofEosestheria, Eosestheria,with withadditional additional overlapped big big reticulations reticulationsnear nearthe the overlapped posteroventral margin marginofofthe thevalve. valve.The Th posteroventral impression of ofthese theseoverlapped overlappedreticureticuimpressions andflattened flattenedtubercles tubercles lations are are large large and lations in the the mould mould fossils. fossils. in EEosestheriao s e s t h e r i a - Y Yanjiestheria anjiestheria assemblage: includesAllestheria, Allestheria assemblage: ItIt includes YUlTlenestheria, Diestheria,ininaddition addition to Yumenestheria, Diestheria, to its namesake forms. During this period, its namesake forms. During this period, Eosesthet'ia wa bearing numerou Eosestheria was bearing numerous growth lines and the corre ponding growth lines and the corresponding gtowth bands changed inro narrow growth bands changed into narrow ornamentation with relatively mall ornamentation with relatively small reticulate sculpture only near the dorsal reticulate sculpture only near the dorsal side but fine and crowded radial Ii rae in side but fine and crowded radial lirae in the rest parts of the valve. The genu 44 Carapace valve of Eosestheria ovata (19 mm long). an eosestheriid conchostracan with mixed sculpture of reticulation and ~ 4 4 Carapace valve Eosestheria (19bands mm long), an eosestheriidsmall conchostracan withplatforms mixed sculpture of grooves reticulation andexternal the rest parts of the valve. The genus radiallirae onof the chitinousovata growth and corresponding and polygonal and radial in the Yanjiestheria was evolved from radial lirae on the chitinous growth bands and corresponding small and polygonal platforms and radial grooves in the external Yanjiestheria was evolved from mould. from Sihetun locality Oianshangou Bed of the lower Yixian Formation) in Beipiao. Liaoning. (Photo: Da-jian Li/ CAS) Eosestheria. It has smaller reticulate mould, from Sihetun locality (]ianshangou Bed of the lower Yixian Formation) in Beipiao, Liaoning. (Photo: Da-jian Li/CAS) Eosestheria. It rhe has dorsal smallerside reticulate culpcure near and finer
sculpture the dorsal side lirae and finer and morenear crowded radial near and more crowded radial lirae near ventral and posterior parts of the ventralThe and posterior parts the valve. conchostracan fossil ofofthi valve. The conchostracan fossils of this assem blage were collected from the assemblage were collected from facies the Jiufotang Formation in lacustrine Jiufotang faciesof of westernFormation Liaoning in orlacustrine imilar rocks of western Liaoning or similar rocks of other regions of China, Mongolia, other regions of China, Mongolia, Transbaikalia Korea peninsula and Transbaikalia, Korea peninsula, and outhwestern Japan. southwestern Japan. 45 Carapace valve of Diestheria yixianensis (21 mm long). a diestheriid Bed of the conchostracan. Zhoujiatun locality (Dakangpu 45 Carapace valve offrom Diestheria yixianensis (21 mm long), a diestheriid middle Yixian Formation) in Yixian,locality Liaoning. (photo:Da-jian conchostracan, from Zhoujiatun (Dakangpu Bed ofLi/CAS) the
middle Yixian Formation) in Yixian, Liaoning. (Photo:Da-jian Li/CAS)
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stracods are tiny of theofphylum Arthropoda. stracods are crustaceans tiny crustaceans the phylum Arthropoda. TheyThey are important microfossils and useful in geological are important microfossils and useful in geological prospecting. The most conspicuous featurefeature of these is the is the prospecting. The most conspicuous ofanimals these animals bearing of a bivalved carapace in which the soft body enclosed (Fig. 46). The bearing of a bivalved carapace in which the softisbody is enclosed (Fig. 46). The two valves are not and are constituted by calcareous layer and two valves aresymmetrical, not symmetrical, and are constituted by calcareous layer and chitinous coating. The left is normally somewhat larger larger and overlapping chitinous coating. Thevalve left valve is normally somewhar and overlapping the right one. one. The exterior surface of valves may be ornamented or smooth. the right The exterior surface of valves may be ornamented or smooth. The The ostracod carapace is generally between 0.5 and mm length, with with ostracod carapace is generally berween 0.51.5 and 1.5inmm in length, less than 0.5 or mm or more thanmm, 1.5 and mm,very andfew very few reaching few either a fewa either less than 0.5 mm more than 1.5 reaching 70 mm as Paramoelleritia fromDevonian the Devonian carbonate rocks of Guangxi 70 mm such such as Paramodleritia from the carbonate rocks of Guangxi Province, China. Province, China. a worldwide distribution. They first Ostracods histOry Ostracods have have a longa long history and aand worldwide distribution. They first appeared in the Cambrian of 500 million years ago and are still floutishing appeared in the Cambrian of 500 million years ago and are still flourishing a high salinity and temperature tOlerance and can tOday. today. TheyThey have have a high salinity and temperature tolerance and can live in live in but they are especially common in seawater, freshwater and brackish water, seawater, freshwater, and brackish water, but they are especially common in shallow-sea and lakes. or sulfur springs. Most ostracods shallow-sea and lakes. TheyThey even even live inlive hotinorhot sulfur springs. Most ostracods are crawling benchos, but some are planktOn, and still others swim or burrow are crawling benthos, but some are plankton, and still others swim or burrow inco mud. Ostracods are gonochoristic and can reproduce all year round. In into mud. Ostracods are gonochoristic and can reproduce all year round. In suitable environmencal conditions, they reproduce bisexually, whereas in the suitable environmental conditions, they reproduce bisexually, whereas in the unsuitable environmencs they are parthenogenetic. Ostracods are oviparous unsuitable environments they are parthenogenetic. Ostracods are oviparous animals. Their eggs, round or oval in shape, can withstand desiccation and animals. Their eggs, round or oval in shape, can withstand desiccation and cold, spreading for long distance and hatching in a suitable condition. The cold, spreading for long distance and hatching in a suitable condition. The ontogeny of ostracods is discontinuous. As is in other arthropods, after the ontogeny of ostracods is discontinuous. As is in other arthropods, after the larva hatched Out from the egg, with tbe growth of the soft body tbe bard larva hatched out from the egg, with the growth of the soft body the hard carapace shed when the new carapaces are formed. There are approximately carapace shed when the new carapaces are formed. There are approximately eight molts during their ontOgeny. And the shape and ornament of the eight molts during their ontogeny. And the shape and ornament of the carapace change with the growth. The chitin layers of the carapace are very carapace change with the growth. Thesoft chitin layers of thetocarapace are very only thin and rarely preserved and the body is prone decay. Therefore, thinthe andcalcareous rarely preserved soft body is prone to decay. Therefore, only as fossils. layers and are the usually preserved the calcareous layers are usually preserved as fossils. The fossil ostracods of the]ehol Biota from the northern Hebei and The fossil ostracods the Jehol Biotaand fromalltheofnorthern Hebei and Fossil western Liaoning areofvery abundant freshwater rypes. w e s tostracods ern Liaoning are very abundant and all of freshwater types. Fossil usually occur in great abundance on tbe rock surface of rhe ostracods usually occur great abundance on like the arock surface of the Dabeigou, Yixian, andinJillfotang Formations, "sesame bread" (Fig. 47). Dabeigou, Yixian, andthan Jiufotang Formations, like a to "sesame bread" Up to now, more 130 species belonging 25 genera of (Fig. fossil 47). ostracods Up have to now, more than in130 25 genera of fossil been found thespecies Jebolbelonging Biota, oftowhich 5 species in ostracods 5 genera were have been found Jehol Biota, of which species inof5 11 genera were species genera from the collected from in thethe Dabeigou Formation, over580 collected the Dabeigou Formation, overpecies 80 species 11 genera the than 80 of 19 of genera fromfrom the Jillfotang Yixianfrom Formation, and more Yixian Formation, and more than 80 species of 19 genera from the Jiufotang Formation. Formation.
UO
The fossil ostracods from the Dabeigou Formation are so far only known from Dabeigou, Jingshang and Zhangjiegou areas, Luanping County, Hebei Province. They are dominated by the large-sized (about 2 mm in length) and The fo il
rracod from the Dabeigou Formation are
0
far only known
smooth-surfaced Luanpingella (Fig. 48) and Torinina (Fig. 49), associated from Dabeigou,) ingshang and Zhangjiegou areas, luanping ounty, Heb i with small-sized (less than I ram) Rhinocypris with small spines and tubercles Province. They are dominated by the large- ized (about 2 mm in length) and on the surface (Fig. 50) as well as the smooth-surfaced Darwinula. Torinina mooth- urfaced Llltlnpingell1 (Fig. ) and Toril1/fld (Fig. 9), a ociated 46 Cyprinotu p.. a recent ostracod first appeared in theized Late(les Jurassic the Argun with mallthan Turga I mm)Formation Rhi"ol)prisinwith mall River pine valley and tubercle arthropod in lateral vi w. from of EastonTransbaikalia. In the past, some ostracod specialists identified the urface (Fig. 50) as well as the mooth- urfaced OClI'll'i,,"/c{ . Torinina f Wuhan. Hubei. Torinina the Dabeigou Formation as Eoparacypris. Because of River the fir from t appeared in the Late) uras ic Turga Formation in th both Argun alley Fossil ostracods, on the rock surface like a "sesame bread" ( × 4), from Senjitu Dabeigou and Turga Formations share the Torinina, they may be closely of East Tran baikalia. In the pa t, orne ostracod peciali tS identified locality (Yixian Formation) in Fengning, Hebei. (Photo: Hai-chun Zhang/NIGP) Torini,,({ fr m the Dabeigou Formation a Eop u-oLJpris. Becau e both of the m 46 Cyprinotussp., a recent ostracod arthropod in lateral view, from Wuhan, Hubei.
~47
47 Fos il ostracod, on the rock surface like a "sesame bread" (x 4), from Senjitu locality (Yi ian Formation) in Fengning, Hebei. (Photo: Hai-chun Zhang! IGP)
Dabeigou and Turga Formation
hare the Tori"ilJ(1, they may be clo ely
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i ~ 4 8 Left valve of Luanpingella postacuta ( x 27), external lateral Jingshang postacuta locality (Dabeigou Formation) in .48 Leftview, valve from of Luanpingella (x 27), external lateral Luanping, Hebei. view, from )ingshang locality (Dabeigou Formation) in
Right internal mould of Torinina tersa ( × 30), external from Jingshang locality .49lateral Right view, internal mould of Torinina tersa (Dabeigou (x 30), external Formation) in Luanping, Hebei. lateral view, from )ingshang locality (Dabeigou
~49
Luanping, Hebei.
Formation) in Luanping, Hebei.
~ 5 0 Carapace of Rhinocyprisjurassica ( x 80), external lateral "'50 Carapace Rhinocyprisjurassica 80), external view, of from Sihetun locality(xtlower part of lateral Yixian
51 Carapace of Cypridea (Cypridea) sihetunensis "'51( x Carapace of Cypridea (Cypridea) sihetunensis 50), external lateral view, from Sihetun (x 50), external lateral view, from locality (lower part of Yixian Formation)Sihetun in locality (lower part of Yixian Formation) in Beipiao, Liaoning.
view, from Sihetun locality (lower part of Yixian Formation) in Beipiao, Liaoning. Formation) in Beipiao, Liaoning.
Beipiao, Liaoning.
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~I~52 Carapace of Cypridea (Cypridea) dabeigouensis 52 Carapace of Cypridea (Cypridea) (x 50), external lateral view,dabeigouensis from Dabeigou (x 50). external lateral fromFormation/in Dabeigou locality (lower part view, of Yixian locality (lowerttebei. part of Yixian Formation) in Luanping, .......... Luanping, Hebei.
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~ 5 3 Carapace ofCypridea (Ulwellia) beipiaoensis .53 Carapace Cypridealateral (Ulwellia) beipiaoensis ( x 50), of external view, from (x 50), external lateral view, from Libalanggou locality (lower part of Yixian Libalanggou locality (lower part of Yixian Formation) in Beipiao, Liaoning. Formation) in Beipiao, Liaoning.
~ 5 4 Carapace of Timiriasevia jianshangouensis ( x 80), .54 Carapace of view, Timiriasevia jianshangouensis (x 80), : external tateral from Jianshangou locality external lateralFormation)in view, from Beipiao, )ianshangou locality (lower pa~ of Yixian Liaoning. (lower part ofYixian Formation) in Beipiao, Liaoning.
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56 Carapaceof Cypridea(Cypfidea)jingangshanensis( x 45), m~ljn57Carapace ofgypridea (Cypridea)zaocishanensis(x 45), lateraiofview, from:(Cypridea) Zaocishanzaocishanensis iocali~ (upper . . external 57 Carapace Cypridea (x 45), pa~ external of¥ixian lateral Formation) ¥ixian, Liaoning,locality (upper view,infrom Zaocishan
Carapace of Darwinulaleguminella(× 70)i external
. . 55: lateral Carapace offrom Darwinufa feguminella x 70). pa~ external view, Sihetun locali~ ((lower of . ... .. 56 external locality:(upper Carapace ofCypridea (Cypridea)jingangsflOnensis (x 45), lateral view, from Sihetun locality (lower part of Yixian FOrmation)inBeipiaoi L i a o n i n g l ' .... part of Yixian Formation)in Yixian, Liaoning external lateral view, from Zaocishan locality (upper Yixian Formation) in Beipiao, Liaoning. part of Yixian Formation) in Yixian, Liaoning.
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58 Carapaceof Yumeniacasta x
. . . . . . . . . . . . . . . . . . . . . . ... 58 Carapace of .Yumenia casta (x 40), external lateral view, from Pijiagou locali~lupper pa~ofjiufotang view, from Pijiagou locality (upper part ofJiufotang Formation) Yixian,Liaoning, Liaoning. Formation)ininYixian;
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59 Cara aceofYumeniajianchangensis × 40), external
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( 4 0 ) , external
................ .............................. . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 ............................... Carapace: of.........Yixianella marginuJata (x 40), external lateral view, from Pijiagou t0cality (Upper pa~ of lateral view, from Pijiagou locality (upper part of Jiufotang Formation)in ¥ixian,:giaoning~ : Jiufotang Formation) in Yixian, Liaoning.
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60....
P ............. . . . . . . . . . 59 Carapace of Yumenia jianchangensis. (x 40), external lateral view, from Pijiagou locality(upper part of lateral view, from Pijiagou locality (upper part of JiUfotang ... Formation} in YiXian, giaoning, ... Liaoning. . . . . . Jiufotang Formation) in Yixian,
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part of Yixian Formation) in Yixian, Liaoning.
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correlated: .... mentioned facts, the the lower partpart 0f the Formation should be late Theirage ageisisLate LateJurassic, Jurassic. mentioned facts, lower of Yixian the Yixian Formation should be late correlated. Their OstracMs from the Y~an Formation are ve~ rich in quantity and more : Tithenian or:late Tithonian to Berriasian in age, Oscracods from the Yixian Formation are very rich in quantity and more Tithonian or lare Tithonian to Berriasian in age. diverse isthe predominant Ostrac0ds fromfrom the the Jiufotang Formation are extremely abundant and and thanthose tho efrom fromthe theDabeig0u Dabeigou Formation. Cypridea is the predominant Ostracods divetsethan form. The lower diverse, They maymay be grouped also also into into two twO assemblages: The lower form.These These osi:racods osttacodsmay maybebegrouped groupedinto intotwo tWOassemblages, assemblages. The lower highly highly diverse. They be grouped assemblage. The lower assemblage; abundant large2 is comparatively to that fromfrom the ~the e r upper part part of theofYixian assemblage assemblage, in inaddition additiontoCypridea(Figsl to C),pridea (Fig.5151 -53), 53),includes includes abundant large-~sembiage is comparati similar ely similar to that the Yixian sized predominant withwith forms of Cypridea (Cypridea) The. The fossilfossil ostra:ostraand smooth'surfaced smooth-surfacedforms formssuch suchasasYansbanina, Yanshanilla,Djungaricaandsmall-Formation Djllngal'ica and small- Formation predominant sized and sized forms Timiriasevia (Figl upper upper part are abundantly sized forms Timiri(/s~via (Fig. 5 ) Darwimtla (Fig. 55), etc.; The upper cods from the upper part of the Jiufotang Formation are abundantly assemblage in Which thethe represented:With rapidly evolved and and widely distributed formsforms such such as as assemblage consists consistschiefly chieflyofofspecies speciesofofgenus genusCypridea CyP"idea in which represented with rapidly evolved widely di tributed Y",nmia (Figs. 58, 59), Lilllnocypfidea and a lot of Cheilocypridea, Yixianella subgenus Cypridea (Cypridea) (Figs. 56, 57) with left valve larger than the :
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right, occupied a dominant position. The ostracods from the lower part of the (Fig. 60) and CYPfidea (Ulwellia) with the right valve larger than the left one. Yixian Formation Formationare aresimilar similartothetwo to the twoostracodassemblages ostracod assemblages the lowerT h iThis assemblage is characteristically identical to Early the Early Cretaceous Yixian in in thelower s assemblage is characteristically identical tothe Cretaceous part 0fthe of the PUrbeCk Purbeck Gr0up GroupofEngiand of Englandand andmay maybebecorrelated correlated Valanginian-Barremian ostracodsand in and outside China. part withith thethe Valanginian-Barremian ostraCodSin oUtSide China. oscracod assemblage as emblagefrom fromthe theBulum BuJumMember Memberofof Kuri Formation (Except where indicated, all photo in this chapter ostracod thetheKuti Formation in in thethe (Except where indicatedl all photos in this chapter werewere takentaken by by Argun River valley of the eastern Transbaikalia. Judging from the aboveYong-qiang Mao/ IGP) Argun ~ver valley of the eastern Transbaikaiia. Judging from the above- Yong-qiang Mao/NIGP) : . . . .
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Yan-i,i, Shen
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Yan-bin Shen superfamilies Astacoidea and Parastacoidea. The astacoids live in the Northern
highly sporadic fossil record of freshwater shrimps is
•Tl • h e
i m p r o v e d by the discovery of a b u n d a n t specimens of
Hemisphere. They are divided into the family Astacidae (Europe, western
superfamiJies Ascacoidea and Parascacoidea. The asca oid live in che orchern poradic fo sil record of fresh\ ater shrimps North is America) and the family Cambaridae (North America and East Asia). Hemi phere. They are divided inco che family Ascacid e (Europe, \ escern improved by the di covery of abundant pecimens The of parastacoids are restricted to the Southern Hemisphere. he highly
astacids and spelaeogriphids from the Jehol Group. The former are
a large well-known crayfish (Decapoda, Astacidea) and the latter a new
ta ids and p laeogriphids from the Jehol Group. The former are
member of the family Spelaeogriphidae (Hemicaridea), Liaoningogripbus.
alar e well-known crayfish (Dec pod a, Astacidea) and rhe larrer a new
Crayfish Astacids are commonly referred to as "crayfish", and are also
orch America) and che family
ambaridae ( orch America and Easc A ia).
The Astacidea is characterized by subcylindrical cephalothorax, wellThe parascacoid are rescricc d to che ouchern Hemisphere.
developed rostrum and abdomen, simple cervical groove, chelate first three The Ascacidea is characcerized by ubcylindrical cephalochorax, \ eJl-
member of the family p laeogriphidae (Hemicaridea), Liaollillgogriphm.
pereiopods with the first of them the largest, well-developed abdominal pleura, known by many other common names in the different parts of the world: developed r crum and abdomen, imple cervical groove, chelace fir r chree Crayfish A ta id ar common! referred to, .. rayfi h", and are al 0 telson with transverse suture, and uropods with diaeresis. crawfish, paper-shell crabs, ecrevisse, yabbies, mud-bugs, flusskrebs, raks, ditch pereiopods wich che firsc of chern che largesc, well-developed abdominal pleura kno\ n by many ocher common names in che differenc parc of che world: Males of the superfamily Astacoidea are typically with the first and bugscrawfi and koonacs. With over 500 known species, they are almost distributed cel on wich cran ver e urure and uropod wich dia re i . h, paper-shell rabs, ecre i e, yabbi , mud-bug, flu skreb rak, dicch second pleopods modified into a rather elongated, styliform blade to assist in worldwide. bug and koonac . Wich over 500 kno\ n specie, rhey are a1mosc di cribuced
Traditionally, two large groups of freshwater crayfish are recognized, the worldwide.
Male
f che uperfamily Ascacoide are cypically wirh che fir c and
the transfer of the spermatophore. Females of the family Cambaridae bear an
Tr dirionally, rwo large groups offreshwacer crayfi h are recognized, che
econd pleopods modified inco a racher elongaced
cyLiform blade co assisc in
che tran fer f che spermatOphore. Female of che family
ambaridae bear an
c..
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~61
Recent crayfish Procambarus clarkii. Dorsal view of a male (Left).Ventral view of a male, showing the tubular form of pleopods 1 and 2 (Middle).ventral view of female,
61 Recent crayfish Procambarus c1arkii. Dorsal view of a male (Left). Ventral view of a male. howing the tubular fonn of pleopod I and 2 (Middle). ventral view offemale. showing thethe annulus ventralis and and pleopods l N51-5 (Right). (Photo" Shi-wei Zhao/NIGP) howing annulu ventralis pleopods (Right). (Photo: Shi-wei Zhao/ IGP)
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A male Cricoidoscefosus aethus in ventral view. ~62 87 mm long from rostrum to distal telson. from 87 Dawangzhangzi locality (Yixian Formation) in Linyuan. Liao ing. (Photo: Da-jian LiI CAS)
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cheliped to the last pleomere. from Dawangzhangzi locality (Yixian Formation) ~.~63 A female Cricoidoscelosus ae~hus in lateral view, 99 m m long from top of in Linyuan. Liaoning. (Photo: Da-jian LiI CAS)
cheliped to the last pleomere, from Dawangzhangzi locality (Yixian Formation) in Linyuan, Liaoning. (Photo: Da-jian Li/(;AS)
64 Tail fan in ventral view (27 mm wide) of a female Cricoido5celo5u5 aethu5. from Dawangzhangzi locality (Yixian Formation) in Linyuan. Liaoning.
i64
Dong-xing Deng/ Tail (Photo: fan in ventral view (27 mmNIGP) wide) of a female Cricoidoscelosus aethus,
from Dawangzhangzi locality (Yixian Formation) in Linyuan, Liaoning. (Photo. Dong-xing Deng/NIGP) 65 A molted crayfish exoskeleton (115 mm long) of a female
i65
Cricoidosce/osus aet/1!Is. from Dawangzhangzi locality (Yixian Formation) in Linyuan, Liaoning. (Photo: Da-jian LiI CAS) A molted crayfish exoskeleton (115 mm long) of a female
Cricoidoscelosus aethus, from Dawangzhangzi locality (Yixian Formation) in Linyuan, Liaoning. (Photo: Da-jian Li/CAS)
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A female Cricoido5celo5u5 aethu5 in lateral view. 99 mm long from top of
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ventralis, a specific of spermatheca, in thebetween areas between the crayfish areinfound in the temperate of the Northern Hemisphere, and annulusannulus ventralis, a specific form ofform spermatheca, in the areas the the Northern Hemisphere, and crayfish are found the temperate zones ofzones fourth fifth pereiopods Theofmales of the Cambaridae sometimes the temperate and tropical of the Southern Hemisphere. fourth and fifthand pereiopods (Fig. 61).(Fig. The61). males the temperate the Cambaridae sometimes and tropical zones ofzones the Southern Hemisphere. is polygraphist and on a of variety living dead animal have aset robust set of directed processes, ischialon hooks, on the through second through CrayfishCrayfish of directed processes, or ischialorhooks, the second is polygraphist and feeds onfeeds a variety livingofand deadand animal have a robust andmaterials. plant materials. A few of species of theAstacidae family Astacidae also adapted to fourth pereiopods. However, the members of the Astacidae lackhooks ischial hooks of the Astacidae lack ischial fourth pereiopods. However, the members and plant A few species the family are also are adapted to in theand males and annulus structure in the females. in the males annulus ventralisventralis structure in the females.
the cold-water Typically, is restricted to the warmer the cold-water habitats.habitats. Typically, moltingmolting is restricted to the warmer summersummer
The crayfish specimens of the Jeholwere Biota were obtained the Yixian The crayfish specimens when feeding takeInplace. In contrast, the members of the family of the Jehol Biota obtained from thefrom Yixian months,months, when feeding can takecan place. contrast, the members of the family Formation in Dawangzhangzi, Daxinfangzi, Songzhangzi, and Niuyingzi Formation in Dawangzhangzi, Daxinfangzi, Songzhangzi, Cambaridae show variations greater variations andtheir reflect theirtoability to a wider and Niuyingzi Cambaridae show greater and reflect ability adapttotoadapt a wider areas, Lingyuan and Houshuangshanzi village ofCounty, Yixian County, areas, Lingyuan City, andCity, Houshuangshanzi village ofYixian JinzhouJinzhou of habitats the Astacidae. A number of of species of the Cambaridae range ofrange habitats than thethan Astacidae. A number of species the Cambaridae City, Liaoning Province. The excellently preserved specimens City, Liaoning Province. The excellently preserved specimens include include molted molted the toability make burrows or burrow which them enabletothem to have thehave ability maketoburrows or burrow systems,systems, which enable exoskeleton, adults,and males and females, well asventral dorsal, and ventral inhabit and water exoskeleton, juveniles,juveniles, adults, males females, are that periodically dried out. inhabitbodies waterthat bodies are periodically dried out.species Other occur species as well asasdorsal, Other in occur both in both lateral remains of the They thorax.were They were as named as Cricoidoscelosus aethus (Figs. of the thorax. named Cricoidoscelosus aethus (Figs. lateral remains andhabitats lentic habitats underinrocks, in vegetation, among of riparian lotic andlotic lentic under rocks, vegetation, among the rootsthe of roots riparian and Palaeocambarus licenti by Taylor, and(Family Shen (Family 62-65)62"-'65) and Palaeocambarus licenti by Taylor, Schram Schram and Shen in leaf litter also burrow thebeds gravel beds of streams. trees, intrees, leaf litter and theyand alsothey burrow into the into gravel of streams. Cricoidoscelosidae) (Figs. The isformer is distinguished the latter Liaoningogriphus 67).66, The67). former distinguished from thefrom latter Cricoidoscelosidae) (Figs. 66, Liaoningogriphus The Spelaeogriphidae family Spelaeogriphidae possesses a typically The family possesses a typically in its flagellate or crenulate pleopods, latter bearspleopods. paddle pleopods. in its flagellate or crenulate pleopods, whereaswhereas the latterthe bears paddle peracaridan brood with pouch five of oostegites, thus placed ofpairs oostegites, and wasand thuswas placed peracaridan brood pouch fivewith pairs very to similar recent cambarideans their modified first the Both ofBoth themofarethem veryare similar recenttocambarideans in their in modified first within Superorder Peracarida (Order Hemicaridea). The living within the Superorder Peracarida (Order Hemicaridea). TheSpelaeogriphus living Spelaeogriphus pleopodpleopod in the males annulus ventralisventralis in the females. in theand males and annulus in the females.
firstfrom found from underground caveinpools in theMountain, Table Mountain, was firstwas found underground cave pools the Table South South
Living crayfish mostly inhabits freshwater Some can speciesAfrica. can Acadicaris Living crayfish mostly inhabits freshwater habitats.habitats. Some species Africa. Acadicaris novoscotica, only fossil known fossil species, was reported novoscotica, the onlythe known species, was reported from from
56
IG
tolerate tolerate high salinity, e.g., in the high salinity, e.g.,Caspian in the Caspian andSea, Black they were Sea and Sea Black butSea, theybut were the Carboniferous Lower Carboniferous blackinshale in the Maritime of Canada. the Lower black shale the Maritime ProvinceProvince of Canada. never found real marine More than 500than species neverinfound in real environments. marine environments. More 500 of species of freshwater freshwater Recently, some other Recently, somefossil otherspelaeogriphaceans fossil spelaeogriphaceans the Cretaceous Lower Cretaceous from thefrom Lower
66 A female Iicenti in licenti lateralinview, 47.5 mm long from of chelate 66 APalaeocambarus female Palaeocambarus lateral view, 47.5 mm longtop from top of chelate
67~ 6A7male Palaeocambarus licenti, showing (by a(by reda arrow) the the firstfirst A male Palaeocambarus licenti, showing red arrow)
to the last from Dawangzhangzi locality locality (Yixian Formation) in Linyuan, topleomere, the last pleomere, from Dawangzhangzi (YixianFormation) in Linyuan,
pleopods as styliform bladeblade ( x 2),( × from Dawangzhangzi locality (Yixian pleopods as styliform 2), from Dawangzhangzi locality (Yixian
CAS) Li/CAS) Liaoning.Liaoning. (Photo: Da-jian (Photo: LiJ Da-jian
Formation) in Linyuan, Liaoning. (Photo: Dong-xing Deng/ NIGP) Formation) in Linyuan, Liaoning. (Photo: Dong-xing Deng/NIGP)
69 A nearly complete individual (16 mm long) of Liaoningogriphus quadripartitlls in lateral view, from Sihetun locality (lower part ofYixian Formation) in ~ i 69 A nearly complete individual (16 mm long) of Liaoningogriphusquadripartitus Beipiao, Liaoning. (Photo: Da-jian Li/ CAS) in lateral view, from Sihetun locality (lower part of Yixian Formation) in Beipiao, Liaoning. (Photo: Da-jian Li/CAS) 68 A nearly complete individual (ventral view. 16 mm long) of Liaoningogrip/ws quadripartitus from Sihetun locality (lower part ofYixian Formation) in Beipiao. ~aG8 A nearly complete individual (ventral view, 16 mm long) of Liaoningogriphus Liaoning. showing thorax. abdomen and tail. (Photo: Hai-chun Zhang! NIGP) quadripartitusfrom Sihetun locality (lower part of Yixian Formation) in Beipiao, i i
...t!" .. ' .......
Liaoning, showing thorax, abdomen and tail. (Photo: Hai-chun Zhang/NIGP) deposits of Las Hoyas, Spain are under description.
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A new spelaeogriphacean, LictOningogriphltS q1fadripartitltS. was recently deposits of Las Hoyas, Spain are under description. described by Shen, Taylor and Schram (Figs. 68-70) from the Yixian Formation A new spelaeogriphacean, Liaoningogriphus quadripartitus, was recently in Sihetun, Jianshangou, and Huangbanjigou areas of Beipiao City, and near described by Shen, Taylor and Schram (Figs. 68-"70) from the Yixian Formation Dakangpu village ofYixian County, Liaoning Province. Irs body is elongated and in Sihetun, Jianshangou, and Huangbanjigou areas of Beipiao City, and near cylindrical in outline with a maximum observed length about 20 mm. Thinly Dakangpu village of Yixian County, Liaoning Province. Its body is elongated and sclerotized carapace covers the head and most of the first two thoracomeres. cylindrical in outline with a maximum observed length about 20 mm. Thinly LiaJningogriphllS possesses a very shorr, broadly rounded rostrum. The antennules sclerotized carapace covers the head and most of the first two thoracomeres. are biramous and the antennae uniramous. Thoracopods 2-8 are well developed Liamingogriphus possesses a very short, broadly rounded rostrum. The antennules with endopods. The last six thoracomeres are freely exposed. Very welJ developed, . . 70 Reconstruction of Liaoningogrip/ws quadripartitlls in dorsal (a) and lateral (b) are biramous and the antennae uniramous. Thoracopods 2---8 are well developed elongate, biramous pleopocls are equally present on the pleomeres 1- 5. The distal aspects. (Courtesy: F. R. Schram/ ZMUA) with endopods. The last six thoracomeres are freely exposed. Very well developed, proropod margin is sigmoidally curved, with two-segmented exopods and anm 70 Reconstruction ofLiaoningogriphus quadripartitus in dorsal (a) and lateral (b) elongate, biramous pleopods are equally present on the pleomeres 1---5. The distal aspects. (Courtesy: F. R. Schram/ZMUA) endopod. The telson possesses two pairs of short, robust median terminal spines protopod margin is sigmoidally curved, with two-segmented exopods and an organisms, such as conchostracans, ostracods, bivalves, gastropods, insects and a fine transversal line in its lower middle part. The mopods are more than twice endopod. The telson possesses two pairs of short, robust median terminal spines and plants. Three living spelaeogriphacean forms, SpelaeogriphllS lepidops, as long as the telson. The exopocl has twO ovoid segments. and a fineThe transversal line in its lower middle The uropods more than twice organisms, such as conchostracans, ostracods, gastropods, Potiicom brasilensis and Mangk1ftl/ mityrtlcl, bivalves, are swiftly swimminginsects animals with living spelaeogriphaceans atepart. testticted to theareouthern Hemisphere.
as Conversely, long as the telson. The exopod has two ovoid segments. and plants. living body. spelaeogriphacean forms, Spelaeogriphus lepidops, undulating They are found in caverned freshwater pools and rapidly Three all known fossil taxa are found from the Northern Hemisphere. The living spelaeogriphaceans are restricted to the Southern Hemisphere. Potiicora brasilensis and Mangkutu mityula, are swiftly swimming animals with underground aquifer. However, based on associated conchostracan habits There is a trend of size reduction: the recent taxa appear to be considerably Conversely, all known fossil taxathan are the found from the Northern smaller (about 7 mm long) fossil forms (9.4-20 Hemisphere. mm long).
c.. 1ft
rapidly undulating body. They arebeds foundwith in caverned freshwater pools materials, and and the shrimp-bearing abundant tuffaceous
There isLiaoningogt'iphm a trend of size reduction: the abundant recent taxabut appear to be considerably underground aquifer. must However, associated conchostracan habits is extremely completely lack of diversiry. Liaoningogriphl/S have based lived inonground-water lakes and pools instead of and cave the shrimp-bearing bedswater. with Itabundant materials,climatic smaller 7 mmpreserved long) than the fossil forms tuffaceous (9.4"20 mm long). They (about are usually in yellowish-gray shales and iltstones pools or underground was warm tuffaceous temperate-subtropical
Liaoningogriphus must have lived in ground-water lakes and pools instead of Liaoningogriphus extremelyand abundant but completely lack of diversity. with fine laminatedisstructure often associated wirh terrestrial freshwarer environments during the late Mesozoic in eastern Asia. They are usually preserved in yellowish-gray tuffaceous shales and siltstones
cave pools or underground water. It was warm temperate-subtropical climatic
with fine laminated structure and often associated with terrestrial freshwater
environments during the late Mesozoic in eastern Asia.
Ul
51
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f]3 Jun-f en g Zhan g, Hai-chun Zhan g
lUl1:feng Zhang, Ha;-chllll Zhang s they had been and c o n t i n u e d to be, insects were
A
Hexagenitidae of the order Ephemeroptera, is very important in the correla-
of the order Ephemeroptera, is very important in the correlation ofHexagenitidae non-marine strata in East Asia. Its nymphs, with abundant individuals, undoubtedly s they hadthe been mostand successful continued animals to be, in the insects Jehol were tion non-marine strata East Asia. Its nymphs, lived in theofwarm, shallow and in stagnant waters near the with shore,abundant crawling individuals, on undoubtedly most successful animals the Jehol Biota. This has beenthe confirmed by the extremely high in diversity
warm, shallow and stagnant waters near aquatic the shore, crawling on lived inor the the bottom swimming in the waters and preying on other insects. Biota. This has been confirmed by the extremely high diversity of at least 500 species referable to over 100 families within 17 orders based orland swimming in short-lived the waters and and poor preying onThis othermayfly aquatic the bottom at least 500specimens species referable overthe 100 families within 17northern orders based The adults lived on and were fliers. is insects. onofover 10,000 collectedtofrom non-marine rocks in The adults lived oncomponents land and were short-lived and fliers. This on over collected from non-marine rocks in northern distinguished from other of Hexagenitidae bypoor its large size, andmayfly is Hebei and 10,000 westernspecimens Liaoning provinces. This the extremely high diversity and distinguished fromevolution other components of Hexagenitidae by its large size, and Hebeipreservation and westernofLiaoning extremely high diversity is a cecal branch in the of this family in the late Mesozoic. superb imprintsprovinces. are quite This rare in the geological history and is a cecal branch in the evolution family in dragonfly the late Mesozoic. superb preservation of imprints are have quite rare ina the history Aeschnidium heishankowense (Fig. 7 3) of is athis kind of large referred throughout the world. These fossil insects provided vividgeological picture of the Aeschnidiuffl heishcmkowense 73) is a kind throughout world.some These provided vivid picture to Aeschnidiidae of Odonata. Its larvae(Fig. are numerous, andoflarge lived indragonfly a similar referred late Mesozoic, the in which offossil them insects flew inhave the sky, some acrawled on theof the to Aeschnidiidae of Odonata. Its larvae are numerous, and lived in a similar late Mesozoic, which some of them the sky, some crawled on the environment to that of Ephemeropsis trisetalis. They preyed on other aquatic ground, and some in swam in the water; some flew fed oninplant leaves, seeds or fruits, environment to that of Ephemeropsis tl~isetalis. They preyed on other aquatic ground, and some swam in the water; some fed on plant leaves, seeds or fruits insects dominated by wigglers of mosquitoes and flies or small, young aquatic some were saprophagous, and some preyed on other insects or sucked blood insects dominated by wigglers of mosquitoes and flies or smalJ, young aquatic some were saprophagous, and some preyed on other insects or sucked blood of birds or beasts. Among these fossil insects, some are familiar to us, such as insects and fishes. The adults lived upon land as long-lived and excellent fliers. insects and fishes. The adults lived upon land as long-lived and excellent fliers. of birds or beasts. Among these fossil insects, some are familiar to us, such as wasps (Order Hymenoptera), mosquitoes and flies (Order Dipera), beetles wasps (Order Hymenoptera), mosquitoes and flies (Order Dipera), beetles (Order Coleoptera), cockroaches (Order Blattaria), dragonflies (Order Odonata) (Order Coleoptera), cockroaches (Order Blattaria), dragonflies (Order Odonata) and grasshoppers (Order Orthoptera). These insects open a window through and grasshoppers (Ordet Orrhopteta). These insects open a window through which we have caught a glimpse of the late Mesozoic entomofauna in East which we have caught a glimpse of the late Mesozoic entomofauna in East Asia. Comparing them with modern insect faunas, we also know more about Asia. Compating them with modern insect faunas, we also know more about insect phylogeny and evolution. Moreover, many components had special insect phylogeny and evolution. Moreover, many components had special forms and ecological characteristics, which enable us to reconstruct the forms and ecological characteristics, which enable us to reconstruct the paleogeography, paleoclimate and paleoecology. One of the most interesting paleogeography, paleoclimate and paleoecology. One of the most interesting aspects is many kinds of flower-visiting insects found from this entomofauna, aspects is many kinds of flower-visiting insects found from this entomofauna, such as flower bugs, tumbling flower beetles, brachyceran flies, digger wasps such as flower bugs, tumbling flower beetles, brachyceran flies, digger wasps and cockroaches. From the same locality and the same rocks, the flowering and cockroaches. From the same locality and the same rocks, the flowering plants have also been recovered. These wonderful discoveries have provided plants have also been recovered. These wonderful discoveries have provided important data in the discussion of the origin of flower-visiting insects and important data in the discussion of the origin of flower-visiting insects and
flowering plants as as well as their coevolutionary relationships. flowering plants well as their coevolutionary relationships. InInthethe Jehol Biota, certain geographically widespread and and dominant Jehol Biota, certain geographically widespread dominant insect have been found in almost every terrestrial basins in northern insectspecies species have been found in almost every terrestrial basins in northern Hebei Liaoning provinces. They usually had had abundant indi-indiHebeiand andwestern western Liaoning provinces. They usualJy abundant viduals areare easy to ro collect, with some important and and familiar representavidualsand and easy collect, with some important familiar representativesasasfollows. follows. tives
Ephemeropsis trisetalis (Figs. a giant mayfly assigned to family the family Epbemeropsis trisetalis (Figs. 71,71, 72),72), a giant mayfly assigned to the
72 Adult mayflymayfly Ephemeropsis trisetalis, about 60 mm long. 72 Adult EpIJemeropsis trisetaJis. about 60 mm long.
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73 Adult, female dragonfly Aeschnidiumheishankowense,about 130 mm wide when wings stretched. Red arrow denoting the long ovipositor.
73 Adult. female dragonny At.' chnidium hel hankowen e. about 130 mm wide when wing
tretched. Red arrow denoting the long
0
ipo itor.
Mesolygaeus laiyangensis (Fig. 74), a medium or small-sized aquatic bug This large species influences, directly or indirectly, on rhe quantity variation This large species influences, directly or indirectly, on the quantity variation Mesolygaeus laiyangensis (Fig. 74), a medium or small-sized aquatic bug belonging to an extinct family Mesolygaeidae referable to Heteroptera within of almost all other insects, or even on their rise and fall, for both larvae and of almost all other insects, or even on their rise and fall, for both larvae and belonging to an extinct family Mesolygaeidae referable to Heteroptera within Hemiptera, plays an important role in the correlation of non-marine strata in adults ofthis species are the largest carnivorous insects ofthe J ehol entomofauna. adults of this species are the largest carnivorous insects oftheJehol entomofauna. Hemiptera, plays an important role in the correlation of non-marine strata in East Asia. Both juveniles and adults, with abundant individuals, lived in the Its con-generic relative has been found from the lower Tithonian in Solnhofen, Its con-generic relative has been found from the lower Tithonian in Solnhofen, East Asia. Both juveniles and adults, with abundant individuals, lived in the Germany, and thus it plays an important role in ascertaining the age of the warm, shallow and stagnant waters near the shore, crawling and swimming Germany, and thus it plays an important role in ascertaining the age of the warm, shallow and stagnant waters near the shore, crawling and swimming in the waters, bur sometimes they also disembarked. The adults could fly off Jehol insect fauna. The family Aeschnidiidae was an interesting group in the Jehol insect fauna. The family Aeschnidiidae was an interesting group in the in the waters, but sometimes they also disembarked. The adults could fly off Late]urassic-Early Cretaceous, and as the typically early-staged representathe waters, preying on other small aquatic insects, such as the wigglers of Late Jurassic-Early Cretaceous, and as the typically early-staged representathe waters, preying on other small aquatic insects, such as the wigglers of tive of this group. This Chinese species shows some primitive characteristics mosquitoes and flies. This species only lived in the late Mesozoic bur was tive of this group. This Chinese species shows some primitive characteristics mosquitoes and flies. This species only lived in the late Mesozoic but was such as the dense veins and much small cells on fore- and hind-wings. closely telated to extant shore bugs (the family Saldidae), and both of them such as the dense veins and much small cells on fore- and hind-wings. closely related to extant shore bugs (the family Saldidae), and both of them
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~:~ili"~?¸ ..... ~iii~ bug Mesolygaeus/aiyangel1sis, ......~ i:~i : from Nanligezhuang _74 Saldoid about 7 mm long, _75 Chaoborid mosquito CiJirollomaptera gregaria. about 6 mm long. from ~ 7 5 Chaoborid mosquito locality Chironomaptera gregaria, about 6 mmShandong. long, trom locality (Laiyang Formation) in Laiyang, Shandong. Nanligezhuang (Laiyang Formation) in Laiyang. 74 Saldoid bug Mesolygaeus laiyangensis, about 7 mm long, from Nanligezhuang Nanligezhuang locality (Laiyang Formation) in Laiyang, Shandong. locality (Laiyang Formation) in Laiyang, Shandong. 76 Brachyceran fly Protonemestrius ~ 7 6 Brachyceran fly about Protonemestrius J 3 mm long. red jurossicus. jurassicus, about 13 mm long, red arrow denoting the proboscis.
arrow (Courtesy: denoting Dong the proboscis. Ren/ CNUl (Courtesy: Dong Ren/CNU)
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\ 77 Sketch drawing of Protonemestrius
jurassicus. (Courtesy: Dong Ren/ 77 Sketch drawing of Protonernestrius CNUl jurassicus. (Courtesy: Dong Ren/
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~W78 Flower bug, about 10 mm long.
~179
Echinosomatid earwig, about 20 mm long.
~180 Katydid, about 25 mm long.
~81 Mesoblattinid cockroach. about 25 mm long.
~182 Froghopper, about 15 mm long.
~83 Elater (elaterid beetle). about 1S mm long.
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""84 Cupedid beetle Notocupes laetus, about 15 mm long. 7':::'~': ~
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i84 Cupedid beetle Notocupes laetus, about 15 mm long. 86 Snakefly Alloraphidia longistigmosa, about 20 mm long. 86 Snakefly Alloraphidia longistigmosa, about 20 mm long.
~7 ~
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..., 85 Dung beetle, about 20 mm long.
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85 Dung beetle, about 20 mm long.
Chaoborid mosquitos Chironol11aptera
vesca, abollt 10 mm long. 87 Chaoborid mosquitos Chironomaptera
vesca, about 10 mm long.
belong to the same superfamily Saldoidea. As for lifestyle, however, the belongs to the Chironomapterinae, a Mesozoic extinct subfamily, the belongs to the Chironomapterinae, a Mesozoic extinct subfamily, the belong to the same superfamily Saldoidea. As for lifestyle, however, the former could live in water and is regarded more primitive than the latter that phylogenic position of which in the family Chaoboridae remains unclear. As former could live in water and is regarded more primitive than the latter that phylogenie position of which in the family Chaoboridae remains unclear. As are often seen along shores of ponds or lakes. the food of other medium-sized and small predatory insects, these wrigglers the food of other medium-sized and small predatory insects, these wrigglers are often seen along shores of ponds or lakes. Chironomapteragregaria (Fig. 75), belonging to phantom midges (the were important in the ecological chains of the lacustrine systems in the period Chironomaptera gregaria (Fig. 75), belonging to phantom midges (the were important in the ecological chains of the lacustrine systems in the period extant family Chaoboridae), is a small mosquito. It is also an important they lived. extant family Chaoboridae), is a small mosquito. It is also an important they lived. component in the correlation of non-marine strata in East Asia. Their Though not large in quantity, the Jehol probably flower-associated component in the correlation of non-marine strata in East Asia. Their Though not large in quantity, the ]ehol probably flower-associated wrigglers, with abundant individuals, lived in the warm, shallow and insects are of great importance to the Biota, giving an example of co-evolution wrigglers, with abundant individuals, lived in the warm, shallow and insects are of great importance to the Biota, giving an example of co-evolution stagnant waters near the shore, wriggling in the waters. The adults were between these ancient probable pollinators and earliest representatives of stagnant waters near the shore, wriggling in the waters. The adults were between these ancient probable pollinators and earliest representatives of terrestrial, flying over the marshes where plants were prosperous. The species angiosperms (flowering plants). The identification of the flower-related terrestrial, flying over the marshes where plants were prosperous. The species angiosperms (flowering plants). The identification of the flower-related
65
B8 Brachyceran fly Protonemestrius sp., about 1S mm long.15 mm long. .88 Brachyceran fly Protonemestrills sp., about
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90 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
F
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
lIS
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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ClC/l C/l
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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; :.;;;",-
-
however, the reverse is the case with most other teleost groups. Living
-
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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102) is the most prized and expensive osteoglossomorphs. Because of the two
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Sr/eropog4!S JiJnH()Sll~ r·foftune fish
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a living
usteoglossoid fish. is thla nlost prized and
expensive ostl:aoglossomorphs.
t1
group iliso
includes l.y("optera and Jinonicht llys as prilni-
£7I,j 103 j;nan;cllthys. very similar 10 Ly(optera. was found ,nainly (rom the Jiufotang Formation in
live melnbers, fFrofll: http:,>\\ww.af()\,vana.
\\'t.astlarn
Liaoning. This
spt!(ilnl11l
is abollt 9 em long. (Photo: IVPI')
(OOl,t\\')
barhels. ~]orious large
St',lJ<:S
anJ old history of tht: ti~h . it \vas <:alled .. dragon
fish" in <:hina ilnd \\'as b(:liC:Vl'd to ha\'e th<: rx)\\'c:r
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2JV' 101 Lycoptera is usually preserved in great density. The specimen was found in Daxinfangzi locality (middle part ofYixian Formation) in Lingyuan, liaoning. (Photo: IVPP)
dt:strihcd by
th(: carlit'st kno\\'n oSft,:'oglossonlorphs. Thcrc,t)rc. sonlC: s u ~ ~~ (; S ( edt h a{ Eas t As i a
Illtk ilnd t()rtllnc:.
and
lycoptl'ra. Jinunichthys and ochc:r fossils of the supt:rordtr t()und in Chin,l ~lTt.'
1O.~) \vas 1l1ainly t~)lInd frolll thl' J;uforang Fornlarion
F(,:'n~-zh<:n
!vIa and.J ia-ru Sun b.lsed on the fossils
()~tt·oJ.doSS()nl0rphs.
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p~lleoiththyol()~isrs
i I-: h t h t" the ~1 nee s r rill () r i ~ i n ( l" n r e r () f
An Afri(an origin \vas also sUt:gesrcd. In rc:cent YC:~lrs ..
(roIl)
SOnll' rest:arch{·rs argu<.:d rhat tht: rransotcanic distribution \\'as r<:sultl"d (ronl
southern JiJin Provin<:<.:. The aurhors h<..'Ii<;vl:d th~lf the sp<.'cinlcns froIll Jilin
the splir of ,1n lU1Ct·scral Llnd block according to "icariarl<:t: priot..-iplla. Thil( is
\vt·rt· sinlilar to 1.'.l(fJPIt~I'1 /fJ",~i,-tl}lhll".l but diftl·rt."nr fronl otll<:r spc...·it:s of tht:
to S~lY.
ge n lIS. 'rhercforc ~ rhey g ~lV(: a ne\\' lctOnl:ric nan1 L' ••Jill" ,))(hl /..1\ .. to borh
till· last hrc:ak of tht· supt:rconrinc:nt P41n~<:a.
\\\tS
/-Jt'o/JIt r~, /flllgi(t!,lhllliJ ,lnd t h(.' sp<:cinl<: ns frolll Jilin.
Extant ostco,glossonlorph, surviv(.. d in rht· tfopii...al or
the t'arly l'\'olution of osteoglossonlorphs had alrcildy tinishcd bt·t'')re
J.J,IIg(./t·j(lJ/!;.l".\
~lIbtr()pical frt·sh
\vas another teleost found in \vcstern Liaoning. It \vas
coJlt"cted fron) Jiufotanj..: Formation of 't'ixiilll and Hcishan. Irs rotal body
reach 2"~
The original narnc: \\'~lS ~i\'en by Zhi-thc..'n~ Liu,
\""art..'rs of North AnlCric4.1. ~outh '\lllcrica, Ausrralia. ~ollth{'~lst ,\si.l. Indi'l.
length
and A trilLl. ~() cOllsensus
hast:d on tht· sp<:cinlt"tlS froln LOJlgde. Ningxial I.lnd \·ik(.'zhaoln<.."n~. Inn<..'r
11.lS
b(;('n rtac ht·d hy il ht hyulo~ists up
to
no,,' ~tbour
the intl'rpretiltions of rht" transoct:~uli( dist rihuridn of rl1o\(' frt"sh,\· .Her tJsht·s.
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Yuan Wang, Ke-qin, GaoGao Yuan Wang, Ke-qin ~
mphibians ("double life" life" in Greek) are a are classa of primitive tetrapods mphibians ("double in Greek) class of primitive tetrapods sions ofsions soft parts as such eyes,as gills, andgills, skins. They alsoThey include earliestthe earliest of softsuch parts eyes, and skins. alsothe include
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(vertebrates withwith four four limbs) that spend at least their (vertebrates limbs) that spend at part leastofpart oflives their lives knownknown record record of several taxonomic groups. groups. These incredible finds havefinds not have not of sevetal raxonomic These incredible in water and and part part on land, although some some are entirely aquaticaquatic or only in water on land, although are entirely or rejuvenated only rejuvenated amphibian paleontology alsoa drawn a worldwide amphibian paleontology but also but drawn worldwide
terrestrial. Amphibians provide a biological link between fishes fishes and the true terrestrial. Amphibians provide a biological link between and the true attention the scientific community. attention beyondbeyond the scientific community. as they arefirst the group first group of tetrapods that invaded Frogs Frogs land-living vertebrates, land-living vertebrates, as they are the of tetrapods that invaded Frogs are tailless amphibians in the order Anura Frogs are tailless amphibians classifiedclassified in the order Anura some million Living amphibians are classified the the landland some 370 370 million yearsyears ago. ago. Living amphibians are classified in the in the (belonging to the superorder Salientia, which also includes the primitive (belonging to the superorder Salientia, which also includes the primitive Subclass Lissamphibia closely related ancesrors in forms). fossil forms). proanurans). Theyunusual have unusual body structures are specialized for Subclass Lissamphibia (with(with theirtheir closely related ancestors in fossil proanurans). They have body structures that are that specialized for not only the familiar forms such as the frogs and toads (Order They include jumping. Such structures rodlike urostyle formed of tail They include not only the familiar forms such as the frogs and toads (Order jumping. Such structures include include a rodlikea urostyle formed by fusion by of fusion tail Anura), salamanders and newts (Order Urodela), but also the less familiar vertebrae and the greatly elongated hind limbs with highly modified tarsal Anura), salamanders and newts (Order Urodela), but also the less familiar vertebrae and the greatly elongated hind limbs with highly modified tarsal forms such as the limbless caecilians (Order Gymnophiona). Extinct archaic elements. The earliest fossil representarives of Salientia were from the Early forms such as the limbless caecilians (Order Gymnophiona). Extinct archaic elements. The earliest fossil representatives of Salientia were from the Early amphibians include labyrinthodonts and lepospondyls that flourished in the Triassic of Madagascar and Poland. Mter a long time evolLltion, now the order amphibians include labyrinthodonts and lepospondyls that flourished in the Triassic of Madagascar and Poland. After a long time evolution, now the order Late Paleozoic, from either of which the ancestral stocks of the living includes some 4,800 living species with a global distribLltion except for Late Paleozoic, from either of which the ancestral stocks of the living Anura Anura includes some 4,800 living species with a global distribution except for amphibians may have evolved. extreme northern latitudes, Antarctica, and most oceanic islands. amphibians may have evolved. extreme northern latitudes, Antarctica, and most oceanic islands. Compared with other vertebrate groups, amphibians, especially the The fossil frogs known from the Liaoning beds include several archaic Compared with other vertebrate groups, amphibians, especially the The fossil frogs known from the Liaoning beds include several archaic lissamphibians (having thin and fragile bones unfitted for fossilization), are forms that documented an important diversification of anurans during the lissamphibians (having thin and fragile bones unfitted for fossilization), are forms that documented an important diversification of anurans during the generally less well documented in the fossil record. This is especially, and Early Cretaceous time in the area. One of the frogs known from Liaoning is generally less well documented in the fossil record. This is especially, and Early Cretaceous time in the area. One of the frogs known from Liaoning is painfully, true of the Mesozoic Era, during which impOrtant events of origin Ca/tobat'rachlts sanyanensis (Fig. 104, 105), named and described on the basis painfully, true of the Mesozoic Era, during which important events of origin Callobatrachus sanyanensis (Fig. 104, 105), named and described on the basis and early diversification of modern amphibians took place. Therefore, of a nearly complete skeleton from the Sihetun site. The fossil beds yielded a and early diversification of modern amphibians took place. Therefore, of a nearly complete skeleton from the Sihetun site. The fossil beds yielded a discoveries of any lissamphibian fossils of this time are usually newsworthy, radiometric date of 125 million year BP indicating the Early Cretaceous age discoveries of any lissamphibian fossils of this time are usually newsworthy, radiometric date of 125 million years B P, indicating the Early Cretaceous age if not outright sensational. In the past few years, many well-preserved of the fossil. Taxonomic studies have revealed that Ca/tobatrachm sanyanensis if not outright sensational. In the past few years, many well-preserved fossil. Taxonomic studies have revealed that Callobatracbussanyanensis specimens of lissamphibians were recovered from rhe Mesozoic beds of inthe is a primitive member of the Discoglossidae, a basal family group of the living specimens of lissamphibians from theInner Mesozoic bedsininChina. is a primitive member of the Discoglossidae, a basal family group of the opisthocoelous living Mongolia western Liaoning, northern were Hebeirecovered and southeastern Callobatl"achfls is basal (or primitive) in having nine anurans. western northern and southeastern Innerof Mongolia in China. anurans. Callobatraclous is basal (or primitive) in having ninediscoglossids. opisthocoelousIt further MostLiaoning, of the fossils areHebei important components the Jehol Biota that presacral vertebrae, differing from eight in other Most of the fossils important components of the Jehol presacral vertebrae, differing fromofeight in other It further flourished in Eastare Asia about 130-110 miLLion years ago.Biota The that recovered differs from other members the family in discoglossids. the combination of the following flourished in East Asia about 130---110 million years ago. The recovered differscharacters: from other members the family in the combination following fossils constitute a diverse lissamphibian fauna in the late Mesozoic of Asia, lacking a of dorsal protuberance but havingofathe weak dorsal crest on fossils a diverse lissamphibian fauna in the late of Asia,evolucharacters: dorsal protuberance but having a weak dorsal crest on sculpand constitute provide crucial information on understanding theMesozoic biogeographic ilium;lacking havinga bicondylar sacro-urostylar articulation; lacking dermal andtion provide crucial information understanding biogeographic evolu- ilium; tures having articulation; lackingsacral dermal sculpof early salamanders andonfrogs of modernthe amphibian affinities. onbicondylar skull roof,sacro-urostylar and having anteriorly expanded diapophysis. tion of early salamanders and frogs fossil of modern amphibian affinities. tures on skull roof, discoglossids and having anteriorly expanded diapophysis. amphibians from China were rarher Before these discoveries includesacral five species in a single genus Living in China Before these discoveries, fossil amphibians from China were rather discoglossids in China include five known species as in the a single genus limited in both quantity and taxonomic diversity. Prior to 1998, all the Living Bombil/a, with Bombina oriel/talis popularly "oriental fire-bellied limited both quantity and taxonomic diversity. Prior totomiddle 1998, PleistOcene), all the Bombina, with Bombina popularly known as the "oriental knownin fossils were the Cenozoic in age (early Miocene 106). orientalis Before the discovery of Ca/tobatl'ach/ls, nofire-bellied discoglossid fossils toad" (Fig. known fossils the Cenozoic in agefrom (early Miocene to middleinPleistocene), tOraling tenwere species of five genera five major localities northern China. toad" (Fig. 106). Before discovery of the Callobatrachus, no in discoglossid were known in thethe same range of extant group East Asia. fossils Consequently, totaling species of five genera five majorChina localities northern China. as many Since ten then, new findings from from northeastern haveinadded nearly were known in the same range of the group in Eastftog Asia. Consequently, Ca/lobatl'ach/lS represents rhe extant first discoglossid known from China, and species the findings rare Mesozoic lissamphibians tohave the Chinese record. The fossils Since then,ofnew from northeastern China added nearly as many Callobatrachus represents the first frog Asia. known from China, and the earliest fossil record of discoglossid the group from hundred lissamphibians well-preservedtoskeleton, some with clear impresinclude several species of the rare Mesozoic the Chinese record. The fossils (Fig.from 107)Asia. is another Mesozoic frog reported from the earliest Mesopl:J1J!l7e fossil recordbeipiaoemis of the group
include several hundred well-preserved skeletons, some with clear impres-
Mesophryne beipiaoensis(Fig. 107) is another Mesozoic frog reported from
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77
thebeds. Liaoning beds. It is represented a nearly skeleton completesplit skeleton split on part the Liaoning It is represented by a nearlyby complete on part and counterpart a shale slab. It has obviously different osteological and counterpart of a shaleofslab. It has obviously different osteological from Callabatrachus, including theofpresence of procoelous including the presence procoelous presacrals presacrals structures structures from Callobatrachus, andshortened greatly shortened column. It noting is worthhere noting and greatly vertebral vertebral column. It is worth that here that alsopresacrals, has nine presacrals, albeit having a short column. vertebral column. MesophryneMesophryne also has nine albeit having a short vertebral Theofnumber of vertebrae presacral vertebrae the few significant crucially significant The number presacral is among is theamong few crucially in the evolution of the frogs. In theknown earliest known frog, anatomicalanatomical features infeatures the evolution of frogs. In earliest frog,
Triadobatrachus (a proanuran) of the EarlyofTriassic of Madagascar, this Triadobatrachus (a proanuran) of the Early Triassic Madagascar, this many as 14;an Vieraella, an Early Jurassic frog from Argentina, has number is number as many isasas14; Vieraella, Early Jurassic frog from Argentina, has ten presacrals. Nine presacrals can in be the found in theto Middle to Late Jurassic ten presacrals. Nine presacrals can be found Middle Late Jurassic frog Notobatrachus, as in the aforementioned as well as as in well the aforementioned two Early two Early ArgentineArgentine frog Notobatrachus, Chinese Thisisnumber eightinorall fewer in frogs, all living frogs, number eight orisfewer living CretaceousCretaceous Chinese taxa. Thistaxa. 1 cm
forprimitive two very forms, primitive forms,(North Ascaphus (North American tailed frogs) Ascaphus American tailed frogs) except for except two very and (New Leiopelma (Newfrogs), Zealand frogs), both having nine presacrals. Zealand both having nine presacrals. Thus, the Thus, the and Leiopelma
. . 105 Skeletal reconstruction of CallobatrachLls sanyanensis. I, sacrum; 2, presacral
presacralhas number has been recognized as antrend obvious trend in the decrease indecrease presacralin number been recognized as an obvious in the 105 3,Skeletal reconstruction Callobatrachus sanyanensis. 1, sacrum; 2, presacral vertebrae; urostyle; 4, ilium. (Art:ofYuan WanglIVPP) vertebrae; 3, urostyle; 4, ilium. (Art: Yuan Wang/IVPP)
frog evolution. frog evolurion. its large number, presacralMesophryne number, Mesophryne also has primitive its largetopresacral also has primitive In additionIntoaddition
18
features as theofpresence ribs onthree anterior three presacrals, features such as thesuch presence free ribsofonfree anterior presacrals, and the and the of an intermedium bone in the carpalThus, region. retaining retaining of an intermedium bone in the carpal region. it isThus, not it is not that a recent phylogenetic study has recognized it asclade a distinct clade surprising surprising that a recent phylogenetic study has recognized it as a distinct of basal(Fig. anurans of basal anurans 108).(Fig. 108).
Liaobatrachus the first Mesozoic frog described Liaobatrachus grabaui is grabaui the firstis Mesozoic frog described from the from the Liaoning beds; its however, its systematic is still questionable Liaoning beds; however, systematic position is position still questionable at present.at present.
Liaobatrachus briefly in described in early Liaobatrachus grabaui wasgrabaui namedwas andnamed brieflyand described early 1998 based1998 on based on an incomplete skeleton with a disarticulated poorly skull. preserved an incomplete skeleton with a disarticulated and poorlyand preserved It hasskull. It has a snout-pelvic length75 of mm, aboutintermediate 75 mm, intermediate between Callobatrachus between Callobatrachus a snout-pelvic length of about (94 Mesophryne mm) and Mesophryne (71.3 The taxonomic sanyanensissanyanensis (94 mm) and beipiaoensisbeipiaoensis (71.3 mm). Themm). taxonomic status of this animaltoremains to be investigated, as theofreferral status of this animal remains be investigated, as the referral it to theof it to the family Pelobatidae (by theresearchers) original researchers) based on several characfamily Pelobatidae (by the original was based was on several characters in such question, as theof presence of procoelous andofthe lack of ters in question, as thesuch presence procoelous presacrals presacrals and the lack free ribs, which be confirmed duepreservation. to poor preservation. free ribs, which cannot be cannot confirmed due to poor
Salamanders Salamanders tailed amphibians classified in the the Salamanders Salamanders are tailedare amphibians classified in order(belonging Urodela (belonging to the superorder Caudata, also includes order U rodela to the superorder Caudata, which alsowhich includes some primitive non-urodeles). In few the years, past few years,hundred several of hundred of In the past several some primitive non-urodeles). Bombina orienta/is, a living discoglossid frog mainly distributed in East Asia, and an extant relative of CallobatracllUs. (Courtesy: Er-Illi Zhao/ C1B)
salamander from beds the fossil beds in China. northern China. salamander fossils havefossils been have foundbeen fromfound rhe fossil in northern
Some were recovered the Fengshan bed in northern Hebei Some recovered from the from Fengshan fossil bedfossil in northern Hebei 106 Bombinaorientalis, a living discoglossid frog mainly distributed in East Asia,were Province, and the Jiufotang western These Liaoning. These the Jiufotang FormationFormation in westerninLiaoning. strata can strata can and an extant relative of Callobatrachus.(Courtesy: Er-mi Zhao/CIB)Province, and
107 Holotype of Me ophryne beipiaoensis (Slab A. dorsal iew. snout-pelvis length about 71 mm). a primitive frog representing a distinct basal anuran clade. from Heitizigou
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(lower part ofYixian Formation) Liaoning. (Photo: IVPP)about 71 ram), a primitive frog representing a distinct basal anuran clade, from Heitizigou 107locality Holotype of Mesophryne beipiaoensis (Slabin A,Beipiao. dorsal view, snout-pelvis length locality (lower part of Yixian Formation) in Beipiao, Liaoning. (Photo: IVPP)
i!iiiii!i~¸¸¸/ i~!i!!;i!i!i!i~ii~!~i/ii~iiii!i!!( ~il ....!~ ~i~i~i~: .... 5¸¸¸¸¸¸¸¸¸¸¸¸
Caudata (M Jur-Ree) Triadobalraehus (E Tri) Czalkobalrachus (E Trl) Prosalirus (E Jur) Nolobalraehus (M-L Jur) Vieraella (E Jur) Gobiales (L ere) Mesophyme (E ere) Ascaphus (Ree) Leiope/ma (P/g-Ree) Callobalrocllus (E Cre)
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DiscogloS5US (Mio-Ree) Barbourula (Ree) Bombina (Mio-Ree) Pelobales (Mio-Ree) Eopelobales (Eoc-PIi) Megophrys (Ree) Pelodytes (Mia-Ree) Pipa (Qua-Ree) Xenopus (Pal-Ree) Rhinophrynus (Oli-Ree)
Pa/aeobalraclws (L Cre-PIi)
_108 C1adogram showing showing the the hypothesized hypothesized phylogenetic phylogenetic relationships relationships of of m 108 Cladogram major lineages lineages of ofarchaic archaic anurans anurans and and the the relationships relationships within within the the major Discoglossidae, with two Jehol frogs included (in red). Discoglossidae, with two Jehol flogs included (in red).
109 Aspecimen of LaCCOlriton subsolanus (dorsal 109 A specimen ofLaccotriton (dorsala view. snout-pelvis lengthsubsolanus about 40 mm).
view, length about 40 mm), a small. snout-pelvis primitive metamorphic salamander. small, primitivelocality metamorphic salamander, from Fengshan in Fengning, Hebei. from Fengshan localityAM in Fengning, Hebei. (Photo: Mick Ellison/ H) (Photo: Mick Ellison/AMNH)
110 Holotype Holotypeof ofSinerpetonfengsl1anen (dorsalview, view, snout-pelvis snout-pelvis length length 47 47 mm), mm), 110 Sinerpetonfengshanensisis (dorsal a primitive salamander with neotenic features indicated by the ossified a primitive salamander with neotenic features indicated by the ossified ceratobranchial (denoted by a red arrow) and ossified carpals (denoted by ceratobranchial (denoted by a red arrow) and ossified carpals (denoted by yellow arrows). from Fengshan locality in Fengning, Hebei. (Photo: Mick yellow arrows), from Fengshan locality in Fengning, Hebei. (Photo: Mick Ellison! AM H) Ellison/AMNH)
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be be included included in in the the Jehol ]ehol Group Group (sensu (sensu lato) lato) that that has has yielded yielded the the Jehol ]ehol Biota. Biota.
except for for one one taxon, taxon, Chunerpeton, Chunerpeton, which which isis referred referred to to the the Cryptobranchidae. Cryptobranchidae. except
Some fossils were were recovered recovered from from the the Daohugou Daohugou locality locality of ofsoutheastern southeastern Some other other fossils
Laccotritonsubsolanus mbsolanus (Fig. (Fig. 109) 109) isis the the first first Mesozoic Mesozoic salamander salamander reported reported Laccotriton
Inner Inner Mongolia, Mongolia, the the strata strata older older in in age age (refer (refer to to Chapter Chapter 22 for for stratigraphic stratigraphic
from China. China. ItIt isis aa small-sized small-sized metamorphosed metamorphosed salamander, salamander, represented represented by by aa from
information). ofMesozoic Mesozoic salamanders salamanders from from China China has has special special information). The The discovery discovery of
large number number of ofarticulated articulated skeletons skeletons from from aa small small quarry quarry in in Fengshan Fengshan Basin Basin large
implications, implications, as as they they are are the the earliest earliest known known representatives representatives of of modern modern
ofnorthern northern Hebei Hebei Province. Province. Laccotriton Laccotriton subsolanus subsolanus isis characterized characterized by by having having of
salamander salamander groups, groups, and and thus, thus, provide provide important important insights insights into into the the early early
16 presacrals presacrals and and unicapitate unicapitate ribs ribs with with aa broadened broadened base base (most (most living living 16
evolution evolution of ofsome some anatomical anatomical structures strucrures and and the the biogeographic biogeographic history history of of
salamanders have have bicapitate bicapitate ribs, ribs, except except for for hynobiids hynobiids and and cryptobranchids). cryptobranchids). salamanders
these these tailed tailed amphibians. amphibians. Living Living salamanders salamanders form form China Chinahave have been beengrouped grouped inin three three families: families: the the
primitively retains retains lacrimal lacrimal and andprefrontal prefrontal bones bones on on the the skull, skull, and and has has five five ItItprimitively separate bones bones inin the the lower lower jaw. jaw. The The phalangeal phalangeal formula formula (number (number of of separate
Hynobiidae, Hynobiidae, the the Cryptobranchidae, Cryptobranchidae, and and the theSalamandridae. Salamandridae. None Noneof ofthe thefossil fossil
phalangesininhand handand andfoot) foot)isis2-2-3-2 2-2-3-2ininthe thehands, hands,and and2-2-3-4-2 2-2-3-4-2ininthe thefeet. feet. phalanges
salamanders salamandersreported reportedfrom fromnorthern northernChina Chinacan canbe beassigned assignedtotoany anyof ofthese thesefamilies families
Sinerpeton fengshanensis(Fig. (Fig. 110) 110)isisanother anothersalamander salamanderfrom fromthe theFengshan Fengshan Sinerpetonfengshanensis
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113 Holotype of Chunerpeton tianyiensis 113 and Holotyp of CllUnerpeton tianyiensi (part counterpart of a slab of shale, (part and counterpart ofaa basal lab f hale, body length about 180 mm), body length ab ut I 0 mm), cryptobranchid salamander and the a ba al cryptobran hid alamand rand th only Mesozoic member of the group of the only e ozoic m mber found to date, from Daohugou locality r up from Da (Photo: hugou locality found toInner date,Mongolia. in Ningcheng, in ingchen. Inner Mongolia. (Photo: Mick Ellison/AMNH) Mick Elli
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site, and the fossils were collected from the same quarry as that of Laccotriton. andthe themetamorphosed fossils were collected from the quarry as of Laccotriton. Differentsite, from Laccotriton, thissame salamander hasthat ossified
Different(asfrom metamorphosed Laccotriton, salamander has ossified ceratobranchials bonythe support for external gill filamentsthis in life) and ossified ceratobranchials (as bony support for external gill filaments in life) and ossified carpals and tarsals (ossification of these bones only seen in adulthood). The carpals and tarsals (ossification of these bones only seen in adulthood). The combination of these features suggests a mature individual with larval external combination these features suggests a mature individual with larval gills, a condition calledofneoteny in modern biology. Sinerpeton also differs from external gills, condition neotenyformula in modern biology. in Sinerpeton alsoand differs from Laccotriton in ahaving the called phalangeal of 1-2-3-2 the hands Laccotriton in having the phalangeal formula of 1-2-3-2 in the hands and 1-2-3-4-2 in the feet. 1-2-3-4-2 in the feet. ]eholotriton paradoxus (Figs. 111, 112) is reported from the Daohugou .Jeholotriton paradoxus 111, This 112) apparently is reported isfrom the Daohugou locality, Ningcheng County, Inner (Figs. Mongolia. an aquatic
locality, Ningcheng Innerlaterally Mongolia. This apparently is an aquatic salamander as indicated by its County, external gills, compressed tail, presence salamander as indicated by its external gills, laterally compressed tail, presence of well-developed haemal arches on caudal vertebrae, and the lack of ossified of well-developed haemal arches on caudal vertebrae, and the lack of ossified carpal and tarsal elements.]eholotriton is a special Mesozoic salamander showing carpal and tarsaland elements. is aindicate special Mesozoic a combination of larval adult Jeholotriton features that neoteny.salamander The larval showing a combination of larval and adult that indicate neoteny. The larval features include the presence of external gills,features a tooth-bearing coronoid bone on include presence of external a tooth-bearing coronoid the lowerfeatures jaw, the larval the shaped pterygoids and gills, a short maxillary arcade with bone on the lower jaw, the larval shaped a short maxillary arcade with underdeveloped maxilla in the cranial part.pterygoids The adult and features include extensive underdeveloped in the part. adult features include medial contact of the twomaxilla nasals and thecranial presence of The a posteriorly directed toothextensive medial contact of the two nasals and the presence of a posteriorly directed tooth row in the palate. ]eholotriton is characterized by having 17 presacrals, the the palate.Jeholotriton characterized having 17 presacrals, the vertebraerow withinshort transverse processes,isand a prominentby dorsal process on the
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vertebrae with transverse processes, and a prominent dorsal process premaxillae. Its ribs areshort like those of Fengshan salamanders as unicapitate and on the premaxillae. Its ribs are like those of Fengshan salamanders as unicapitate and proximally expanded. The phalangeal formula is 2-2-3-2 for the hands and proximally expanded. The phalangeal formula is 2-2-3-2 for the hands and 2-2-3-3-2 for the feet. 2-2-3-3-2 tianyiensis for the feet.(Fig. 113) is another salamander from the Chunerpeton Chunerpeton tianyiensis represents (Fig. 113) isa basal anothermember salamander Daohugou site. This salamander of thefrom the Daohugou the site.family This including salamander represents Asian a basalgiant member Cryptobranchidae, the endangered sala- of the familyand including the American endangeredhellbender Asian giant salamander Cryptobranchidae, Andrias (Fig. 114, the Upper) the North mander Andrias (Fig. 114, Upper) and the shares North American hellbender Cryptobranchus. Morphologically, Chunerpeton with extant Cryptobranchus. Morphologically, Chunerpeton with extant cryptobranchids several derived characters, such as the nasalsshares being much severalwidth; derived characters, such as theabsent; nasals being narrowercryptobranchids than the interorbital nasal-prefrontal contact and much narrowerprocess than the interorbital width;along nasal-prefrontal contact and the anterolateral of parietal extending the lateral border of absent; the theprimarily anterolateral of parietal extending along the lateral border of the differsprocess from extant cryptobranchids, however, in lacking frontal. It frontal. It primarily fromofextant cryptobranchids, however, in lacking the frontal-maxillary contact;differs retention a palatal fenestra between vomers; 114 Andrias davidianus (Upperl and Batracllllperus pinchonii (Lower), a living representative of the Cryprobranchidae and Hynobiidae. respectively; the two families are widely accepted asand the Batrachuperus most basal groups of (Lower), living 114 Andrias davidianus (Upper) pinchonii a
contact; retention of a palatal between presencethe offrontal-maxillary a distinct medial process of pterygoid; andfenestra ossification of vomers; presence distinct medial process Because of pterygoid; and ossification of basibranchial II asofa atrident-shaped structure. no pre-Paleocene
basibranchial II asthe a trident-shaped structure. Because no pre-Paleocene living known for family, the Chunerpeton fossils from the fossils were Hynobiidae, respectively; the fossils were known for the family, the Chunerpeton fossils from the Daohugou site document the first Mesozoic and the earliest known record of two families are widely accepted as the most basal groups of living Daohugou site document the first Mesozoic and the earliest known record of salamanders. (Courtesy: Er-mi Zhao/CIB)
salamanders. (Courtesy: Er-mi C18) representative of theZhao/ Cryptobranchidae and
the Cryptobranchidae. The fossils also provide evidence supporting the
type unicapitate ribs with an expanded proximal end. The phalangeal formula
hypothesis that the divergence of the Cryptobranchidae from the Hynobiidae
is 2-2-3-2 in the hands and 1-2-3-4-3 in the feet. This salamander is similar
had taken place during the Jurassic in Asia.
to some living hynobiids (Fig. 114, Lower) in several osteological features. It
Liaoxitriton zhongjiani (Fig. 115) is the only salamander taxon of the
is noted that the species is represented by a series of fossils showing different
Jehol Biota that is known from the Jiufotang Formation. Fossils of this
developmental stages, which allows a possible ontogenetic study of the
the CryptObranchidae. The fossils also provide evidence supporting the hypothesis thar the divergence of the CryptObranchidae from the Hynobiidae
salamander were recovered from a site near Huludao City of western had raken place during the Juras ic in Asia.
type unicapitate ribs with an expanded proximal end. The phalangeal formula
is 2-2-3-2 in the hands and 1-2-3-4-3 in the feet. Thi salamander is similar
Liaoning. As the zhongjiani formation is(F;g. dated115) as about million years BP,taxon Liaoxitriton is the110 only salamander of the Liaoxitriton
animal in the near future. to some living hynobiids (Fig. 114, Lower) in several osteological features. It In general, amphibian fossils (especially of salamanders) from is noted that the the species is represented by a seriesthose of fossils showing different
documents so faristhe youngest occurrence of salamander fossils J ehol Biota that known fromstratigraphic the J iufotang Formation. Fossils of this
the Mesozoic bedsstages in northern China are important forontogenetic their superb preservation, developmental which allows a possible tlIdy of the
in the Jeholwere Group. The species by dozens of of articulated salamander recovered from isa represented site near Huludao City western
large quantity, and considerable taxonomic diversity. The discoveries of these animal in the near future.
skeletonsAsinthevarious preservation and was theBP, first Chinese Liaoning. formation i dated as conditions, about 110 million years Liaoxitriton
wonderful fossils provide solid paleontological evidence answer some major In general, the amphibian fos ils (especially thosetoof salamanders) from
Mesozoic so lissamphibian to be reported with well-preserved soft tissue documents far the youngest stratigraphic occurrence of salamander fossils
questions on the of amphibians; and the studies these preservation, fossils have the Mesozoic bedsevolution in northern China are important for theirofsuperb
(e.g., theThe skinspecies & eye impressions). Theby diagnostic features of this inimpressions the Jehol Group. is represented dozen of articulated
opened a new window to view thetaxonomic evolutionary history,The including the origins, and considerable diversity. di coveries of these large quantity
animal include the presence of 16 presacral vertebrae skeletons in various preservation conditions, and and wa transverse the first processes Chinese
taxonomic diversification and geographic radiation, of modern amphibians. wonderful fossils provide solid paleontological evidence to answet some major
Mesozoic lissamphibian to be reported with well-preserved soft tissue of vertebra about half-length of the centrum. It also has the cryptobranchoid
questions on rhe evolution of amphibians· and the studies of these fossils have
impressions (e.g., the skin & eye impressions). The diagnostic features of this
opened a new window to view the evolutionary histOry, including the origins,
animal include the presence of 16 ptesacral vertebrae and transverse processes
taxonomic diversification and geographic radiation, of modern amphibians.
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116A Adorsal-ventrally dorsal-ventrallycompressed compressedskeleton skeleton ofof .116 Manchurochelys liaoxiensis(about (about3030cm cmlong long MancllUrochelys liaoxiensis fromhead headtototail), tail),a asinemydid sinemydidturtle, turtle,from from from Jianshangoulocality locality(lower (lowerpart partofofYixian Yixian jianshangou Formation) Formation)ininBeipiao, Beipiao,Liaoning. Liaoning.(Photo: (Photo:IVPP) IVPP)
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ecause they are still living, turtles are commonplace objects to
lUll Lill had once been regarded as the ancestor group of the turtles, but none of these
us; were they entirely extinct, their shells ~ the most remar-
animals has convincing derived characters shared with turtles. Some paleon-
D "Because
kable defensive armor everliving, assumed by a are tetrapod~would tologists think thebeen turtles may beasclosest to the marine had once regarded the ancestor group reptile of the sauropterygian, turtles, bur none of these they are still turtles commonplace be objects to a cause for wonder."--A. S. Romer, 1945. but current evidence is not strongderived enoughcharacters to supportshared this view. origin of paleonus; were they entirely extinct, their shells - the most remaranimals has convincing withThe turtles. Some Turtles are unique among living animals. Their trunk is embedded in the is an think unsolved be still kable defensive armor ever assumed by a tetrapod-wouldturtles tOlogists the puzzle. turtles may be closest ro the marine reptile sauroptetygian, bony ashell. The can be -divided dorsal1945. and ventral parts (carapace and cause forshell wonder." A. S.into Romer,
Many well-preserved haverobeen found western but current evidenceturtle is not specimens strong enough suppOrt thisinview. The origin of
Turtles are includes unique among living animals. trunkone is embedded the turtles is an can unsolved puzzle.to Manchurochelys belonging to the plastron), and usually inner and outer layer:Their the outer is horny, inLiaoning. Moststill of them be referred bony shell. The shell can be divided into dorsal and ventral parts (carapace and Many well-preserved turtle specimens made up by many scutes; the inner one is bony, made up by many plates. Not family Sinemydidae. Three species have been describedhave so farbeen of thefound genus:in western layer: the outer plastron), and usually includes inner and Liaoning. Most of themM. can be referred Manchm'ochelys belonging to the only do the vertebrae grow together with shell, butouter also the carapace bony one platesis horny, Manchurochelys manchoukuoensis, donghai and M.toliaoxiensis (Fig. 116). M. up ribs. by many the inner oneterrestrial is bony, made up by by many Not family Sinemydidae. have beenShikama described far of made are fused with Theyscutes; differ from all other vertebrates the plates. ribs manchoukuoensis was named byThree Riuji species Endo and Tokio in so 1942, thethe genus: only the vertebrae grow together with shell,them bur also the carapace bony plates mancho/lktloensis, liaoxiensis (Fig. 116). M. outside thedo shoulder and pelvic bones, which enables to retract their heads holotypeManch/lyochelys of which was lost during WWII.M.M.donghai donghaiand wasM. from the coalmine are fused They differand from all other terrestrial vertebtates by the ribs mancholtk/loensis was named bythan Riujithe Endo and Tokio Heilongjiang Shikama in 1942, the and limbs into with shellsribs. for protection self-defense. in Jixi (possibly from a higher horizon Jehol Group), holotype of which was lost during WWII. M. donghai was from the coalmine outside the shoulder and pelvic bones, which enables them to retract their heads The turtles are one kind of specialized reptiles; they also are one branch Province, named by Shao-liang Ma in 1986. M. liaoxiensis was established on and limbs into shells for protection and self-defense. in Jixi (possibly fram a higher horizon than the Jehol Group), Heilongjiang of ancient reptiles. They used to be classified as primitive anapsids (i.e. reptiles a specimen from Jianshangou village, Beipiao, Liaoning Province by Shu-an The turtles are one kind of specialized reptiles; they also are one branch Province, named by Shao-liang Ma in 1986. M. liaoxiemis was established on with no temporal opening on the skull), but also regarded as diapsids (i.e. Ji in 1995. Some smaller-sized turtles were also collected from the Yixian and a specimen from Jianshangou village, Beipiao, Liaoning Province by Shu-an of ancient reptiles. They used to be classified as primitive anapsids (i.e. reptiles reptiles with two temporal openings) by other people. The turtles primarily Jiufotang Formations of the Jehol Group, whereas their systematic position with no temporal opening on the skull), but also regarded as diapsids (i.e. Ji in 1995. Some smaller-sized turtles were also collected from the Yixian and can be divided into two groups: the Pleurodira and the Cryptodira. These remains to be studied (Fig. 117). reptiles with two temporal openings) by other people. The turtles primarily Jiufotang Formations of the Jehol Group, whereas their systematic position Characteristic features of Manchurochelys include: skull very low, nasal names refer to the manner in which the living members of these groups retract can be divided into two groups: the Pleurodira and the Cryptodira. These remains to be studied (Fig. 117). small, prefrontal in contact with vomer, a paired pit present on ventral surface their names necks. refer The pleurodires do so by lateral flexure of the cervical vertebrae to the manner in which the living members of these groups retract Characteristic features of Manchttrochelys include: skull very low, nasal of basisphenoid, the supramarginal scales absent, shellpit very flat, plastron and the cryptodires by vertical flexure. All the living pleurodires inhabit fresh their necks. The pleurodires do so by lateral flexure of the cervical vertebrae small, prefrontal in contact with vomer, a paired present on ventral surface
cruciform, mesoplastra absent. Based on these characters, Manchurochelys waterand of the continents (Africa, Australian South America), but fresh thesouthern cryptodires by vertical flexure. All the and living pleurodires inhabit of basisphenoid, the supramarginal scales absent, shell very flat, plastron could be classified into Sinemydidae, Cryptodira. And it is the closest to extinct ones may have lived in a marine environment. This group has a water of the sourhern continents (Africa, Australian and South America), but cruciform, mesoplastra absent. Based on these characters, Manch/lrochelys Dracochelys, of turtlesinto from the Early worldwide in thelived Cretaceous and Paleogene. Cryptodires are has extinctdistribution ones may have in a marine environment. This group a coulda group be classified Sinemydidae, Cryptodira. And it is rhe closest to Cretaceous of Xinjiang, China. muchworldwide more diverse in the modern than areand the Paleogene. pleurodires.Cryptodires Modern distribution in thefauna Cretaceous are Dracochelys, a group of turtles from the Early are cold-blooded likeChina. most other cryptodires may bediverse classified intomodern three groups: Testudinoidea, includingModern Turtles much more in the fauna the than are the pleurodires. Cretaceous of Xinjiang, reptiles; their modern groups are mainly the tortoise and most freshwater thegrau] Chelonioidea, the sea turtles, cryptodires may be classified turtles; into three s: the Testudinoidea, including Turtles are cold-blooded like most other distributed in the temporal, zones. and most freshwater theshell Chelonioidea, reptiles; their moderntorrid groups are mainly with the the rortoise limbs specialized as flippersturtles; and the reduced; the and sea theturtles, live in terrestrial environment, with the limbs specialized as flippers and the shell reduced; and Most the turtles distributed in the temporal, torrid zones. Trionychoidea, the soft-shelled turtles. Trionychoidea, soft-shelled turtles. Most turtles in terrestrial normally in river, lakelive or swamp. Only environment, a few Where did the the turtles come from? How the shell and the strange Where did The the most tuales come turtle from?known How by thenow, shell and the strange in river, lake or swamp. Only a few groups normally are completely terraneous. There were structure were formed? primitive Proganochelys, structure were in formed? The most primitive turtle known now, Proganochelys, groups completely terraneous. There were many lakes in are western Liaoning during the which was found the Upper Triassic of Germany, had by a completely which was found in the Upper Triassic of Germany, had a completely many lakes in western Liaoning during the time of Manchurochelys, and this kind of turtle "normal" shell. The Jurassic turtles almost had the structure of modern "normal" shell.from The the Jurassic turtles almost hadwas theonce structure ofto modern time of Manchllrochelys, andPerhaps this kind may have lived in the lake areas. its of turtle turtles. Eunotosaurus Permian of South Africa thought turtles. Eltnotosaltrus from the Permian of South Africa was once thoughtlifestyle to may have lived in the lake areas. Perhaps its is similar to that of modern freshbe the ancestor of the turtles, but in fact it is not related to turtles at all. be the ancestor of the turtles, but in fact it is not related ro turtles at all. lifestyle is similar to that of modern freshwater turtles. Among reptiles preceded the turtles, pareisaurs, procolophonoids, captorhinids Among reptiles preceded the turtles, pareisaurs, procolophonoids, caprorhinids water turtles. m 117 A small turtle (about 7 cm long) from Shangheshou locality _117 A small turtle (about 7 cm long) from Shangheshou locality 0iufotang Formation)in Chaoyang, Liaoning. (Photo: IVPP) Oiufotang Formation) in Chaoyang, Liaoning. (Photo: IVPP)
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Jun Liu, Xiao-lin Wang ' ~ he Choristodera is a clade of distinctive but poorly known
lUll
no plantar tubercles. Monjurosuchus can be distinguished from all other Lill, Xiao-/in Wang
aquatic reptiles. For more than one century, knowledge of
choristoderes by the combination of the small supratemporal fossa and closed
hethis Chorisrodera a cladetooftwo di rincri bur poorly genera: known group was ilimited highly e specialized
no plantar rubercles. Monjll1'os/{chllS can be disringui hed from all orher subtemporal fossa.
aquaric For more rhan America one cenrury, knowledge of Champsosaurus andreprile Simoedosaurus from North and Europe. Over the
choristOderes by rhe combinarion of rhe small fossa and closed Many specimens of Monjurosuchus have supraremporal been discovered in western
rhi group was limired ro have rwobeen highly specialized genera: last two decades, additional nine genera referred to the Choristodera.
subremporal fossa. years, some of which bear exquisite integumentary Liaoning in recent Many specimens ofimpressions Monj/{rosffchllS discovered in wesrern (Fig.have 120).been Gao and others suggested
T
Champsoscm1"l/s andhave imoedosa/{rt/s from orrh America and Europe. Over rhe Thus, we now a better understanding of lasrthese rwo decades, generarange have been animals. addirional They had anine temporal span-referred ro rhe ChoristOdera. Thus, a berreryears under randing of ning we at now least have 190 million from the Late rheTriassic e animals. had a remporal spanto theThey late Oligocene and a range geographical ning ar leasr 190 million years from rhe Lare distribution from western North America to Triassic to rhe lare Oligocene and a geographical Japan via Eurasia. disrriburion from we rern orrh America co The Jehol Biota is famous for its fossil Japan ia Eurasia. vertebrates found in recent years, especially birds The Jehol Biota is famou for ir fossil and feathered dinosaurs. The earliest reported verrebrare found in recent year, e pecially bird tetrapod of the Jehol Biota was a small reptile and fearhered dino aur . The earlie t reporred Monjurosuchus splendens (Fig. 118). It was named retrapod of rhe Jehol Biota was a small reprilein 1940 by R. Endo, and was classified as a Monjll1'osllch/{s splendens (Fig. L18). It ~ a named primitive archosaur (Thecodontia). The holotype in 1940 by R. Endo, and wa classified as a of the taxon was reportedly lost during WWII. primitive archosaur (Thecodontia). Thethe holotype
Shikama10named another taxon, of R. rheEndo raxonand was T. reporredly t during rhe WWI1. 1942 based on raxon, a speciR. Rhynchosaurus Endo and T. orientalis, hikamainnamed anorher men from the same locality horizon R!Jynchosallrt/s orientalis, in 1942and based on a where pecithefrom holotype of Monjurosuchus splendens waswhere found. men the same localiry and horizon taxon of was classified intosplerldens Rhynchocephalia, rheThis holorype Monj/{ros/{ch/{s was found. a group related to lizards intO and Rhynchocephalia, represented today by This raxon was classified a
Liaoning in recent years, some of which bear exquisire inregumenrary that the overall appearance of the integument may impre ion (Fig. L20). Gao and orher suggesred be like that of the living Chinese crocodile lizard, thar rhe overall appearance of rhe integument may Shinisaurus crocodilurus (Fig. 121), a semi-aquatic be like thar of rhe living hine e crocodile lizard, lizard that feeds on small fish, amphibians and hinisa/{rllS crocodilffrtls (Fig. 121), a semi-aquatic invertebrates, and Monjurosuchus may have had a lizard that feeds on small fish, amphibians and similar lifestyle. inverrebrates, and Monj/{ros/{chllS may have had a There also is one kind of "long-necked" imilar life tyle. choristoderes, the hyphalosaurs, in the Jehol Biota. There also is one kind of "long-necked" The specimens of Hyphalosaurus were excavated choristOdere the hyphalosaurs, in theJehol Biota. from Bed of the Yixian FormaThe Dawangzhangzi pecimens of H)phalosallrtls were excavated
tion 123 Ma). It is referred to choristoderes from(ca. Dawangzhangzi Bed of the Yixian Formabased on the following characteristics: vertebral tion (ca. L23 Ma). Ie is referred to choristoderes centrum threecharacteri sacral vertebrae; dorsal based onplatycoelus; the following rics: vertebral Line drawing of the losr holorype of MonjllroslIchus splendens (skull length 92 mm), recently identified as a choristodere. from Danangou (Tanankou) locality (middle pan ofYixian) in Lingyuan, Liaoning. (Art: Mick Ellison! AMNH; Courtesy: Ke-qin Gaol PKU)
ribs pachyostotic; epipodial (radius, ulna, centrum platycoelus; rhreesegments sacral verrebrae; dorsal tibia, fibula) greatlyepipodial shorter than the humerus ribs pachyostOtic; segments (radius,and ulna, femur. But itsgrearly neck is greatly elongated. tibia, fibula) horter rhan rhe humerus and There mainly two fossil localities for femur. Bur are ir neck is grearly elongated. There are rwo localiries Hyphalosaurus, onemainly of them liesfossil in the north offor
Hypha/osallrlls, one of City, rhem Liaoning lies in the norrh of Fanzhangzi of Lingyuan Province.
group relared New to lizard and Tuatara, represented tOday These by the isolated Zealand Sphenodon. two kinds of reptiles were
Fanzhangzi of Lingyuan Ciry 1998. Liaoning A nearly complete fossil skeleton was found here in August, ThenProvince. it was
thecompared i olared by ewF.Zealand Tuarara, phenodoll. e twOthat kinds of repriles were F. von Huene (1942), who The believed they are the same
A nearlybycomplete skeletOn wasfrom found in Augusr was studied Gao andfossil his collaborators thehere IVPP, CAS, and199 was. Then namediras
compared by should F. F. von 09into 2), Rhynchocephalia. who believed rhat Monjurosuchus rhey are the same taxon and be Huene classified was
rudied by Gao and his collaboratOr and was named Hyphalosaurus lingyuanensis (Figs. 122, from 123).rhe TheIVPP, paperCA was published in the as Hypha/osa/{ms ling)'/{anensis L2_, L23). paper was published was in the journal Vertebrata PalAsiatica(Fig. of January, 1999.The However, its counterpart journal as Vertebrclla PalAsiatica ofJanuary,by 1999. However, counterpart named "Sinohydrosaurus lingyuanensis" Jian-jun Li andirsothers from thewas
taxfinally n andreferred shouldtobe clas ified into Rhynchocephalia. MOllj/{ros/{chm Choristodera by Ke-qin Gao and others in 2000 basedwa on finally referredspecimen co Chorisrodera Ke-qin orhersThey in 2000 based the on the neotype (Fig. 119)bythat they Gao newlyand selected. diagnosed rhetaxon neorype specimen (Fig.combination 1 L9) thar rhey newly elected. They diagno ed rhe by the following of characters: dorsoventrally flattened taxon follo\ ing posterior combination character: dor ventrally flarrened skull;bya rhe deeply incised skull of margin; parietal foramen absent; conical skull; a deeply incised po rerior kull margin; parieral foramen absent; conical subthecodont teeth, with striae; three sacral vertebrae; fibula with wide distal ubthecodont reerh, wirh rriae; rhree sacral vertebrae; fibula wirh wide disral but narrow proximal head; fifth metatarsal with expanded proximal end but bur narrow proximal head; fifrh meratarsal wirh expanded proximal end bur
named aNatural " inohydrostlllms ling)'l'anensis" by J ian-jun Li Harris and orher from arhe Beijing History Museum. J. B. Smith and J. D. published Beijingin Journal aeural ofHistory MuPakontology eum. J. B.to discuss mirh and D. Harris publi hed a paper Vertebrate theJ.validity of two names paper injo/{mal o/Vertebrate Paleolltology ro di cu s rhe validiry of rwo names in 2001; he pointed out that the two names are actually a synonymy, and he in 2001; he pointed our rhar the rwo names are acrually a synonymy, and he chose Hyphalosaurus lingyuanensis as the valid name. cho e Hypha/osallms lingY',anemis a rhe valid name.
119 Neotype ofMonjurosuchus splendens(skull length 58 mm) from Niuyingzi locality (middle part of Yixian 119 Neotype of Monjllrosllcllll 58 mm) from Niuyingzi (middle parr ofYixian Formation) in Lingyuan,splendens Liaoning. (skulliengrh (Photo: Mick Ellison/AMNH; Courtesy:locality Ke-qin Gao/PKU) Formarion) in Lingyuan, Liaoning. (Photo: Mick Ellisonl AM H; Courtesy: Ke-qin Gaol PKUj
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120 Skin impressions on a specimen of Monjurosuchus splendens. (Photo: IVPP)
120 Skin impressions on a specimen ofMonjurosuchus splendens. (Photo: IVPP)
the fieldwork The of holotype of Hyphalosaurus lingyuanensis is along 116specimen em long specimen of China-Canada Project. Sigogneau-Russell The holotype Hyphalosaurus lingyuanensis is a 116 cm during theduring fieldwork of China-Canada Dinosaur Dinosaur Project. Sigogneau-Russell had in theofpresence a broad closely packed It has a small skull, several needle-like in ventral view. exposed inexposed ventral in view. It has a small skull, several needle-like teeth lyingteeth in lying had noted thatnoted in thethat presence a broad of snout and snout closelyand packed teeth withteeth with thepart anterior of There the skull. leastof13 ribs and than mosttosimilar to the Paleocene genus rectangular bases, lkechosaurus the anterior of thepart skull. are There at leastare 13atrows ribsrows andofmore thanmorerectangular bases, Ikechosaurus was most was similar the Paleocene genus gastralia ("abdomen The column vertebralconsists columnofconsists of 19 Simoedosaurus, alsoseveral held by several other researchers. still rows of("abdomen 20 rows of20gastralia ribs"). Theribs"). vertebral 19 Simoedosaurus, a view alsoa view held by other researchers. However, However, still thought lkechosaurus is more closely to Champsosaurus othersIkechosaurus cervicals, 16-17three dorsals, three than 55Therefore, caudals. Therefore, cervicals, 16--17 dorsals, sacrals, andsacrals, more and thanmore 55 caudals. others thought is more closely related to related Champsosaurus than to than to the mostfeature striking on this is specimen is theofpresence of greatly elongated Simoedosaurus. A new species, lkechosaurus gaoi (Fig. 125) was erected the most striking on feature this specimen the presence greatly elongated Simoedosaurus. A new species, Ikechosaurus gaol (Fig. 125) was erected based on based on neck 20 and em long) neck (about 20 (about cm long) a longand tail.a long tail.
a fragmentary skeleton from the Jiufotang of Inner Chifeng, Inner a fragmentary skeleton from the Jiufotang FormationFormation of Chifeng,
The proportionally small head,snout, pointed snout, needle-like and Mongolia by Jun-chang Lil and the third choristodere The proportionally small head, pointed needle-like teeth and teeth Mongolia by Jun-chang Lti and others. Thisothers. is the This thirdischoristodere reported reported greatly elongated neck of Hyphalosaurus from Biota. the Jehol Biota. stronglyitsindicate its fish-eating dietthe Jehol greatly elongated neck of Hyphalosaurus strongly indicate fish-eating diet from Many well-preserved skeletons foundand in Yixian and Chaoyang, life. The ribs are pachyostotic and thickened distally. Functionally,Many well-preserved in life. Theindorsal ribsdorsal are pachyostotic and thickened distally. Functionally, skeletons have been have foundbeen in Yixian Chaoyang, Liaoning recently, Province which recently, which could betoreferred to Ikechosaurus. Their this thickening increase specific of the body, enabling this thickening appears toappears increasetothe specificthe gravity of gravity the body, enabling Liaoning Province could be referred Ikechosaurus. Their is flat, having elongated and largeopenings, temporal with openings, with a skull animal submerged to remain submerged with a minimum effort. It also showsskull other the animalthe to remain with a minimum of effort. Itofalso shows other is flat,skull having elongated snout and snout large temporal a skull em. length The total lengthreptiles of these to the 2 m, and the length that aquatic including lifestyle, including platy- of 30 morphological features morphological features that reflect an reflect aquaticanlifestyle, platylength cm. of The30total of these canreptiles be up can to 2 be m,up and is more than the total length. length the tail coelous vertebrae, poor ossification of the ends of bones, the limb length bones, of the coelous vertebrae, poor ossification of the distal endsdistal of the limb tail of is more than half of thehalf totaloflength. reduced ossification of the short epipodials. reduced ossification of the carpals andcarpals tarsalsand andtarsals short and epipodials. It is interesting to at note that leastfish six of fossil of the genus Lycoptera It is interesting to note that least sixatfossil the fish genus Lycoptera were preserved on slab the same slabholotype with theof holotype of Hyphalosaurus. were preserved on the same with the Hyphalosaurus. One of One of to the mouth of Hyphalosaurus 123) as ifnot it could not them is sothem close istosotheclose mouth of Hyphalosaurus (Fig. 123) (Fig. as if it could escape being on when Hyphalosaurus escape from beingfrom preyed on preyed when Hyphalosaurus opened itsopened mouth. its mouth. of Hyphalosaurus found in Wangjiagou, Many skeletons complete of skeletons Many complete Hyphalosaurus were foundwere in Wangjiagou, This area andofHejiaxin ofYixian in fall, WanfotangWanfotang and Hejiaxin Yixian County in County fall, 1999. This1999. area became thebecame the
second fossil locality of locality Hyphalosaurus. second fossil of Hyphalosaurus. known specimens of Hyphalosaurus were excavated from tuffaceous All knownAll specimens of Hyphalosaurus were excavated from tuffaceous shales in deposited in environment. lacustrine environment. rocksthe recorded the frequent shales deposited lacustrine These rocksThese recorded frequent volcanic eruptions. volcanic eruptions. One can imagine scenery that time:that the time: the One can the imagine theduring scenery during
volcanoes volcanoes threw masses of masses ashes into the atmosphere, and as the and ashes threw of ashes into the atmosphere, as fell the ashes fell of greenhouse gases out and buried everything. down, theydown, spreadthey outspread and buried everything. Volumes ofVolumes greenhouse gases and toxic with volcanic eruptions; was devastated, and toxic gases camegases alongcame with along volcanic eruptions; the habitat the washabitat devastated, and hundreds of thousands animals died Some together. Some individuals of and hundreds of thousands of animalsofdied together. individuals of
Hyphalosaurus were buried as together, as seenspecimens in many specimens Hyphalosaurus were buried together, seen in many (Fig. 124).(Fig. 124). ~ ii!!ill ~! ~.... ~.....
The first specimen as choristodere in China was a snout The first specimen mentionedmentioned as choristodere in China was a snout fragment from Qi the District Otog QiinDistrict in the Ordos Basin, Inner Mongolia fragment from the Otog the Ordos Basin, Inner Mongolia
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It was by D. Sigogneau-Russell in 1981, and by E. Buffetaur. studied by studied E. Buffetaut. It was revised by revised D. Sigogneau-Russell in 1981, and 121 Shinisaurus crocodi/urus. Chinese crocodile lizard. a living semi-aquatic lizard. to
a new taxon, lkechosaurus sunailinae named based on that A specimen. a new taxon, Ikechosaurus sunailinae was namedwas based on that specimen. few which Monjurosuchus may have a similar overallto appearance. mA few 121 Shinisauruscrocodilurus, Chinese crocodile lizard, a livingintegumental semi-aquatic lizard, years later, many well-preserved, articulated specimens of lkechosaurus were years later, many well-preserved, articulated specimens of Ikechosaurus were which Monjurosuchus may haveChen) a similar integumental overall appearance. (Photo: Chull-xuan in Luohandong Zhidan Group (Early Cretaceous) (Photo: Chun-xuan Chen) discovereddiscovered in Luohandong Formation,Formation, Zhidan Group (Early Cretaceous)
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uanensis (tota 122 Holotype of Hyphalosaurus lingy
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istodere, from length 116 em), a long-necked chor t of Yixian par Fanzhangzi loca lity (mid dle Province. (Photo: Formation) in lingyuan, liaoning IVPP)
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skull of the holotype of . . 123 A close-up view of the a fossil haJosaurus Jingyuanensis, showing
Hyp ) fish near its mouth. (Photo: IVPP
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124 Four individuals of Hyphalosaurus on one slab, with two adults and two young, as if one family died in a single accident. (Photo: IVPP)
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A crushed skull of the holotype of Ikecho ourus goo; (skull about 19 cm long), a gavial·like chori todere, from a locality ofJiufotang Formation in Chifeng, Inner Mongolia. (Photo: IVPPI
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keleton of Yabeinosaurus from Gezidong locality • Uiulong • • han ••
261: A~ ilnCompiete skeleton of Yabeinosaurusi f r ~ Gezidong local i~ Oiul~ngsha~ . ......,......-~,ormat~on), Lmgyu~n, Liaoning (Photoi I V P P ) :
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An unde cribed fo il keleton of Yabeinosaurus from Dawangzhangzi locality (middle part ofYixian Formation) in Lingyuan, Liaoning, red arrow denoting a po ible condition of autotomy in the tail. (Photo: IVPP)
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Juu Liu he Squamata (lizards and snakes) is the most successful
lUll group of modern reptiles. Squamates usually are slender animals with scutella covering their bodies. This group can be
T
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quamata (lizards and snakes) i the mo t
uc essful
identified by skull characteristics" a high degree of skull mobility and an
with the emergence of the first snake.
Liu Yabeinosaurus tenuis is one of the earliest found tetrapods in the Jehol
Biota and the earliest studied fossil lizard in China. This species was named with the emergence of the fir t snake.
in 1942 based on the material from Zaocishan, Yixian County of Liaoning Yc,beinosclIIrtls tenllis i one of the earliest found tetrapod in the Jehol group of modern reprile. quamare u uaJly are slender incomplete lower temporal bar. The Order Squamata is divided into six Province by R. Endo and T. Shikama, who erected a new family for this genus, animals with scurella covering rheir bodie . Thi group can be
infraorders" Iguania, Gekkota, Amphisbaenia, Scincomorpha, Anguimorph
identified by kull charaCteristics: a high degree of kull mobility and an
Biota and the earlie r studied fo il lizard in
hina. Thi specie wa named
but it was later classified into the family Ardeosauridae by R. Hoffstetter in in 1942 based on the material from Zaocishan, Yixian
ounty of Liaoning
and Serpentes. Serpentes are snakes, and the other five groups are generally
1964. Another specimen from Lingyuan, Liaoning Province was referred to Province by R. Endo and T. hikama, who erected a new family for thi genu, incomplere lower temporal bar. The Order quamata is divided into six this species by C.-c. Young in 1958 (Fig. 126). The specimen of Lingyuan was called lizards.
infraorders: Iguania, Gekkora, Amphisbaenia, cincomorpha, Anguimorph
but it was later classified into the family Ardeo auridae by R. Hoff: tetter in
regarded as from the Late Jurassic originally, but it more probably is from The undoubted oldest-known lizards wereother foundfive in the Middle 1964. Another specimen from Lingyuan, Liaoning Province was referred to and erpentes. erpentes are snakes, and rhe groups areJurassic. generally Jiulongshan Formation of the Middle Jurassic, and perhaps represents a They are represented by several genera of about three infraorders. Thus, thi species by .-c. Young in 1958 (Fig. 126). The pecimen of Lingyuan was called lizards. species. The the holotype of Yabeinosaurus tenuis wasit lost in probably World War lizards must have originated earlierlizards than this the Middle earliest Juras lizard ic. different regarded as from Late Jurassi originally, but more is from The undoubred olde r-known weretime, foundand in the Ji andFormation his collaborators proposed a newperhaps specimen from shouldarehave appeared on at least from Triassic, view of the. date iuJongshan of the recently Middle Jura sic, and represents a They represented by earth everal genera ofthe abour threeininfraorder Thu , II.J Shu-an YixianThe County as theofYabeinosallrlls neotype of this lenllis species, and some newly War of the origin of theoriginared sphenodontians. found Permiandifferent specie. holotype was lost in World mu t have earlier Santaisaurus, than thi time, andnear thethe earlie r lizard Jingangshan, lizards materials maycollaborators also be referred to the proposed genus (Fig. 127). pecimen from Triassic boundary in Xinjiang, was referred to lizards 11. IlU-an J i and his recently a new hould have appeared on earthnorthwestern at least fromChina, the Triassic, in view of rhe by dare discovered Two other lizard species were the alsoneotype added toofthe Biotaand by Shu-an han Yixian ountyas thiJehol species, orne newly of the paleontologists. origin of the sphenodontians. ScwtaisclIIYlfs, found rhe Jurassic, Permian- Jingang some Squamates increased rapidly from thenear Middle and others materials in recent may years:also Dalinghosaurus longidigitus be referred to the genus (Fig. l27). Trias boundary in Xinjiang, northwesrern hina to lizards then ic they had another evolutionary radiation in the was Earlyreferred Cretaceous alongby Ji discovered (Fig.lizard 128), specie andJeholacertaforrnosa withtowellTwotail other were also added the Jehol Biota by hu-an some paleontologists. quamates increased rapidly from rhe Middle Jurassic, with a long J i and others recent DalinghosallYIIs IOllgidigillls then they had another evolutionary radiation in the Early Cretaceous along preserved scales in (Fig. 129).years: The diversity of lizards may
128 Holotype of Dalinghosaurus longidigitus, a long-tailed lizard, from Sihetun locality (lower part Yixian Formation) in Beipiao,a 128 Holotype ofofDaJingho auru Jongidigitus. Liaoning. (Courtesy: Shu-an Ji/PKU)localiry long-tailed lizard. from Sihetun (lower parr ofYixian Formation) in Beipiao, Liaoning. (Courtesy: Shu-an ji/ PKU)
with a that longthe tailSquamata (Fig. 12 was ), andJeholacerta formosa highly with wellindicate one kind of animals pre erved (Fig. 129).ofThe of lizards may adaptive to thecales environments that diversity time. indicate that the quamata was one kind of animals highly
Holotype ofof Jeholacerta formosa, a adaptive to athe129 environments that time.
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Jehol lizard with well-preserved skin impressions, a localityformosa. (Yixian a 129 Holorype from of jehoJacerta Formation) in Pingquan, Hebei. jehollizard with well-pre erved skin (Courtesy: Shu-anfrom Ji/PKU) impressions, a localiry (Yi ian Formarion) in Pingquan, Hebei. (Courtesy: Shu-an jil PKU)
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Xiao-lin Wang, Zhong-he Zhou terosaurs are flying reptiles and the first successful flying
Many pterosaurs have been discovered from the Mesozoic deposits and
vertebrate in earth history. They first appeared in the Late
most of them were preserved in the marine deposits. The Solnhofen is among
Triassic together with dinosaurs, approximately 230 million years
the most notable pterosaur localities in the world, and the Solnhofen
(Myr) before present, and became extinct by the end of the Late Cretaceous.
limestone has produced the fossils of both Rhamphorhynchoidea and
Pterosaurs were the dominant creatures in the Mesozoic sky until birds joined
Pterodactyloidea. These pterosaurs lived in the Late Jurassic of approximately
in toward the late Mesozoic.
145""" 150 Myr ago (Tithonian). The Santana Formation in the northeastern
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All pterosaurs are grouped in the order Pterosauria, which comprises
Brazil has produced only pterodactyloid pterosaurs, which lived in the Early
two suborders: Rhamphorhynchoidea and Pterodactyloidea. The former
Cretaceous (Aptian-Albian, about 100""" 110 Myr before present). The Late
mainly occurred from the Late Triassic to the Late Jurassic (some
Cretaceous Niobrara Formation (Santonian, around 85 Myr ago) in western
extended into Early Cretaceous), and are relatively primitive with
Kansas, USA is famous for producing thousands of large-sized pterodactyloids.
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short neck, long tail (except the short tailed Anurognathidae),
Recently, several dozens of pterosaurs have been discovered from the lacus-
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short metacarpals, and long pedal digit V; the latter
trine deposits of the Lower Cretaceous Jehol Group in western Liaoning
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ranged from the Late Jurassic to the Late Cretaceous,
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and represents a more advanced group, with
Many pterodactyloid pterosaurs have been reported from the Lower
long neck, short tail, long metacarpal and
Cretaceous terrestrial deposits in other parts of northern China comparable to the Jehol Group. Among them are Dsungaripterus weii and Noripterlls
short pedal digit V.
complicidens from the Tugulu Group of the Dsungar Ounggar) Basin of Xinjiang, and Huanhepterlls quingyangensis from the Zhidan Group of the Ordos Basin, Gansu Province. Lots of pterosaur skeletons have been discovered from both the Yixian and Jiufotang Formations of the Jehol Group. These fossils comprise mostly
j ,.
pterodactyloids and a few rhamphorhynchoids. Among the known pterodactyloids are Eosipteru.s yangi and Haopterus gracilis from the J ianshangou
.
Bed of the Yixian Formation at the Hengdaozi locality and Sihetun locality
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ofBeipiao, Liaoning respectively, and Sinopterus dongi, Chaoyangoptents zhangi and Liaoningopterlls gui from the Jiufotang Formation at the Dongdadao and
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Dapingfang localities in Chaoyang, Liaoning. Rhamphorhynchoids comprise
_130 Holotype of Haopterus gracilis (subadult, wingspan about 1.35 m), a pterodaetylid pterosaur, from Sihetun locality (lower part ofYixian Formation) in Beipiao, Liaoning. The wing digit appears to be in the mouth (denoted by a red arrow), possibly reflecting the struggle of the animal shortly before being killed by the poisonous gases from volcanic eruption. (Photo: IVPP)
fit
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fit
99
reducedreduced foot suggests that it probably has strong and the and the Dendrorhynchoides curvidentatus from thefrom Zhangjiagou locality locality in Beipiao, foot suggests that it probably has flight strongcapability flight capability Dendrorhynchoides curvidentatus the Zhangjiagou in Beipiao, suspended by the hind limb in limb resting Liaoning andJeholopterus ningchengensis from the Daohugou locality locality in body body was suspended by the hind in position. resting position. Liaoning and Jeholopterus ningchengensis from the Daohugou in was Ningcheng, Inner Mongolia. Ningcheng, Inner Mongolia.
Jeholopterus (Figs. 131 ~ 133) was named 2002,inand represents a Jeholopterus (Figs. 131N 133) was in named 2002, and represents a
nearly completely articulated pterosaur with excellently preserved fibers infibers in Shu-anJi and his (1997,1998) reported two pterosaurs nearly completely articulated pterosaur with excellently preserved Shu-an Ji colleague and his colleague (1997,first 1998) first reported two pterosaurs the wingthe membrane and "hairs" the body. newThe species be further from thefrom Yixian EosipterusEosipterus yangi and Dendrorhynchoides curvidentatus. wing membrane and in "hairs" in theThe body. new can species can be further theFormation: Yixian Formation: yangi and Dendrorhynchoides curvidentatus. referred to the to "strange" short-tailed rhamphorhynchoid family family Eosipterus is represented only byonly fragmentary postcranial materials, and the "strange" short-tailed rhamphorhynchoid Eosipterus is represented by fragmentary postcranial materials, and referred Anurognathidae. It is the It most complete and largest of the of the Dendrorhynchoides, an incomplete skeleton.skeleton. The former a smallis to medium Anurognathidae. is the most complete andknown largest individual known individual Dendrorhynchoides, an incomplete The isformer a small to medium sized pterodactyloid with a wingspan about 1.2 m long, to the tofamily, a wingspan of aboutof90about cm. Its include:include: a skull a skull sized pterodactyloid with a wingspan about 1.2 mreferable long, referable the with family, with a wingspan 90 characteristics ca. Its characteristics Pterodactylidae. It has a It relatively robust forelimb, with thewith ulnathe andulna radius wider than long, resembling that of athat frog;ofaashort metacarpal short and Pterodactylidae. has a relatively robust forelimb, and radius wider than long, resembling frog;neck; a short neck; metacarpal short and approximately 1.3 times1.3astimes long as wing metacarpal. The femur slightly less thanless 1/4than of the the radius; long pedal V (about 1/4length of theof length of the extremely radius; extremely longdigit pedal digit V (about approximately long as wing metacarpal. Theisfemur is slightly smaller smaller than 2/3than of the the tibia. ulna, first the wing digital 1.5 times1.5 thetimes length the third pedal comprising two longtwo phalanges; 2/3length of the of length of theThe tibia. Thetheulna, first wing digital theoflength of the thirddigit) pedal digit) comprising long phalanges; phalanxphalanx and the and tibiathe aretibia approximately of equalof length. is andisa short and tail. a short tail. are approximately equal Dendrorhynchoides length. Dendrorhynchoides represented by a small individual, with a wingspan of aboutof40about cm. 40 ca. The wing and "hair"-like structures represent one of the represented by juvenile a small juvenile individual, with a wingspan Themembrane wing membrane and "hair"-like structures represent one of the This pterosaur was initially referred referred to the Rhamphorhynchoidae, and is now featuresfeatures of the of newthepterosaur. The propatagium, This pterosaur was initially to the Rhamphorhynchoidae, and is most now distinctive most distinctive new pterosaur. The propatagium, member of the Anurognathidae ("frog-jawed" generally agreed to be a to cheiropatagium and the and uropatagium of the wing can be well generally agreed be a member of the Anurognathidae ("frog-jawed" cheiropatagium the uropatagium of themembrane wing membrane can be well pterosaurs). Its characteristics include robust vertebrae, short metacrecognized. The cheiropatagium clearly attaches to both to sides of sides the legs as legs as pterosaurs). Its characteristics includecervical robust cervical vertebrae, short metacrecognized. The cheiropatagium clearly attaches both of the arpals only about of the the radius, shorter humerus, far as thefarankle. uropatagium is between the two the legstwo andlegs composed of arpals only1/4 about 1/4length of theof length of the tibia radius, tibia than shorter than humerus, as theThe ankle. The uropatagium is between and composed of Ou
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metatarsals I ~ IV ofIapproximately equal length, pedal digit V comprisfibers that are that shorter those ofthose the cheiropatagium. Short fibers were alsowere also metatarsals - I V of approximately equal and length, and pedal digit V comprisfibers are than shorter than of the cheiropatagium. Short fibers ing two ing longtwo phalanges. long phalanges.
preserved associated with thewith pedalthe digits, the fifth the digit, indicating preserved associated pedalincluding digits, including fifth digit, indicating
the IVPP, have excavated abundant Since 1998, fieldthe crews that thethat footthe wasfoot webbed and thatand thethat medially curved robust digit Sincethe 1998, fieldofcrews of theCAS IVPP, CAS have excavated abundant was webbed the medially curved fifth robust fifth digit pterosaurs specimens. We have five specimens (including four provided both attachment and control the uropatagium. pterosaurs specimens. Wedescribed have described five specimens (including four provided both attachment and for control for the uropatagium. nearly complete skeletonsskeletons and one and skull), representing a new genus nearly complete oneeach skull), each representing a newand genus and The purported webbedwebbed foot injeholopterus may indicate that thisthat this The purported foot in Jeholopterus may indicate pterosaur probablyprobably lived near thenear water, could swim. pterosaur species: species: Haopterus gracilis,jeholopterus ningchengensis, Sinopterus chaoyangensis, pterosaur lived theand water, andeven could evenThis swim. This pterosaur Haopterus gracilis,Jeholopterus ningchengensis, Sinopterus chaoyangensis,
Chaoyangopterus zhangi, and Liaoningopterus gui. Chaoyangopterus zhangi, and Liaoningopterus gui.
may eatmay insects otheroranimals such as such fishes.asIts extremely long wings eat or insects other animals fishes. Its extremely long wings
HaopterusHaopterus (Fig. 130) was130) named 2001 in after theafter late Prof. Yi-chun Hao suggest strong flying (Fig. wasinnamed 2001 the late Prof. Yi-chun Hao suggest strongcapability. flying capability. in memory of her contribution to the study the Jehol is a nearly in memory of her contribution to theofstudy of theBiota. JeholItBiota. It is a nearly It is a controversial issue as to whetheli pterosaurs are warm-blooded and It is a controversial issue as to whetheg pterosaurs are warm-blooded and complete skeleton,skeleton, and probably represents a subadult with a wingspan of "haired" vertebrates. Injeholopterus, "hairs" are short, and curved; they complete and probably represents a subadult with a wingspan of "haired" vertebrates. In Jeholopterus, "hairs" arethick short, thick and curved; they about 1.35 m. 1.35 The m. skull is long low,and lacking any sagittal crest, and the andalso about The skull and is long low, lacking any sagittal crest, thetaperalso from thefrom base the to the theyand are they associated with thewith whole taper basetip; to and the tip; are associated thebody whole body rostrumrostrum is relatively pointed.pointed. The upper lower arejaws eachare equipped is relatively Theand upper andjaws lower each equipped from thefrom neckthe to neck the tail ~ 133). Functionally, the "hairs" to region the tail(Figs. region131 (Figs. 131--133). Functionally, the "hairs" with 12 with posteriorly curved and sharp in the anterior 12 posteriorly curved andteeth, sharpmainly teeth, distributed mainly distributed in the anterior of pterosaurs could becould used be forused thermoregulation, flight orflight reducing noises noises of pterosaurs for thermoregulation, or reducing part of part the jaws. forelimb is extremely robust with of theThe jaws. The forelimb is extremely robusta long with wing a long wing during flight. "hairs" may alsomay indicate that some duringThe flight. The injeholopterus "hairs" inJeholopterus also indicate thatpterosome pterometacarpal. The metatarsals are slender veryand small. I ~ III areI---IIIsaurs metacarpal. The metatarsals are and slender veryMetatarsals small. Metatarsals are might warm-blooded. The "hairs" bear some saursbemight be warm-blooded. The ofjeholopterus "hairs" ofJeholopterus bearresemsome resemless thanless 1/5than of the the wing metacarpal. The sternum is large and 1/5length of theof length of the wing metacarpal. The sternum is largeblance and to the hairlike integumental structures of the feathered dinosaurdinosaur blance to the hairlike integumental structures of the feathered fan-shaped with well-developed keel, andkeel, its length width about fan-shaped with well-developed and itsand length andare width arethe aboutSinosauropteryx the Sinosauropteryx and Beipiaosaurus, which might that thethat "hairs" and Beipiaosaurus, whichsuggest might suggest the of "hairs" of same.
same.
pterosaurs and the and fiber-like protofeathers in Sinosauropteryx are homologous pterosaurs the fiber-like protofeathers in Sinosauropteryx are homologous HaopterusHaopterus has a large and a pointed rostrum.rostrum. The front teeth has skull a large skull and a pointed The frontareteethstructures. are structures.
sharp and slender, suggesting a piscivorous feeding feeding habit. Its extremely (Figs. 134, 135) is a135) recently described pterodactyloid from from sharp and slender, suggesting a piscivorous habit. Its extremely Sinopterus Sinopterus (Figs. 134, is a recently described pterodactyloid
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131 Holotype ofJeholopterus ningchengensis (Slab A, adult or subadult with wingspan about 90 cm), a short tailed pterosaur belonging to the Anurognathidae ("flog-jawed" pterosaurs), Daohugou locality (lowest part of,adult YixianorFormation) in Ningcheng, Inner 1 3 1 p from fjt/lo/Opt ru nmgchengtn i (lab ubadult with \ ing pan ab ut Mongolia. 90 m), a (Photo: h rt tailIVPP) d pter aur b Ion III to the Anuro nathidae rfro -Ja\\ d" pt ro aur I, fr m D ohu
u I ali
(I \ e t part of Yi ian rormauon)
III
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hen, Inner
on olia. (ph to: IVPPI
132 Wing membrane and the "hair "on
jellOloplerus ningchengen is. repre132 Wing membrane and the "hairs" on senting the mo t complete wing Jeholopterus ningchengensis, repremembrane and "hair" found to date
senting the most complete wing
in ptero aur . The wing membrane
membrane and "hairs" found to date
is relatively long and straight fibers.
in pterosaurs. The wing membrane
"hair" are generally short. wav
is relatively long and straight fibers.
and curved and present all over the
"hairs" are generally short, wavy
body. The "hair" of this pterosaur
and curved and present all over the
resembles the hairlike integument
body. The "hairs" of this pterosaur of the dino aur Sinosauropleryx. ugresembles the hairlike integuments gesting a po sible homolog
of the dinosaur Sinosauropteryx, sugthese structures. (Photo: IVPP)
gesting a possible homology of these structures. (Photo: IVPP)
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133 Reconstruction of Jeholopterus ningchengensis. (Artwork by. 133 Reconstruction of je/l0/oplerus Rong-shan Li/IVPP) ningchellgen i . (Art\ ork by: Rong- han Li/IVPP)
of theofPterodactylidae were were previously known only in EuroMembers the]ehol Biota.Biota. It is referred to thetofamily Tapejaridae, representing its first the Jehol It is referred the family Tapejaridae, representing its first Members the Pterodactylidae previously known only in European and African Late]urassic deposits. The discovery of the Haopterus It also represents the earliest occurrence as well as the recordrecord outside Brazil. outside Brazil. It also represents the earliest occurrence as well as the pean and African Late Jurassic deposits. The discovery of the Haopterus represents the first of theoffamily in Asia; it alsoitextends the distribution represents therecord first record the family in Asia; also extends the distribution of theoffamily to thetoEarly Cretaceous. Members of theofAnurognathidae were were Sinopterus has a has wingspan of about 1.2 m,1.2with the skull length aboutabout Sinopterus a wingspan of about m, with the skull length the family the Early Cretaceous. Members the Anurognathidae rarelyrarely known in theinpast; they they are mainly known from from the Solnhofen in in 170 mm. It is characterized by anby edentulous low and with with long long 170 mm. It is characterized an edentulous lowlong and skull, long skull, known the past; are mainly known the Solnhofen Germany and Karatau in Kazakhstan. The discovery of members of this The sagittal crest crest of theofpremaxilla and pointed rostrum and aand horny beak.beak. and pointed rostrum a horny The sagittal the premaxilla Germany and Karatau in Kazakhstan. The discovery of members of this most most complete skeleton of theoffamily. complete skeleton the family.
in Liaoning represents the first record of theoffamily in thein Early and dentary are low The posterior process of theofpremaxilla is family and dentary are and low small. and small. The posterior process the premaxilla is family in Liaoning represents the first record the family the Early Cretaceous. Sinopterus represents the first fossil record of the family Tapejaridae curved upward, separate from the skull, and parallel to the sagittal crest of the curved upward, separate from the skull, and parallel to the sagittal crest of the Cretaceous. Sinopterusrepresents the first fossil record of the family Tapejaridae outside Brazil. is large and long aboutabout 2.5 times parietal. The nasopreorbital fenestra parietal. The nasopreorbital fenestra is large and (length long (length 2.5 times outside Brazil. Two Two pterosaur assemblages appear to have existed in thein]ehol Group, of theofheight), exceeding 1/3 of1/3 theoftotal length of theofskull. The tibia about the height), exceeding the total length the skull. The is tibia is about pterosaur assemblages appear to have existed the Jehol Group, Formation. Among the the represented by the Formation and ]iufotang 1.4 times of theoflength of theoffemur. Metatarsal III is III about 22.1 % of theof the 1.4 times the length the femur. Metatarsal is about 22.1% represented byYixian the Yixian Formation and Jiufotang Formation. Among several dozens of pterosaur specimens known from from thesethese assemblages, most most of theofwing metacarpal, and metatarsal V is less the 1/5 theof the length length the wing metacarpal, and metatarsal V isthan less than theof1/5 several dozens of pterosaur specimens known assemblages, are pterodactyloids, and only a fewa of them are rhamphorhynchoids. The The length of metatarsal I. length of metatarsal I. are pterodactyloids, and only few of them are rhamphorhynchoids. from from the Yixian Formation, the lower assemblage, comprises Chaoyangopterus (Fig. (Fig. 136) 136) is a medium to large-sized pterodactyloid Chaoyangopterus is a medium to large-sized pterodactyloidassemblage assemblage the Yixian Formation, the lower assemblage, comprises is long and low, a pointed and Haopterus belonging to Pterodactyloidea, and Dendrorhynchoides with with a wingspan aboutabout 1.85 m long. The skull a wingspan 1.85 m long. The skull is long and with low, with a pointedEosipterus Eosipterus and Haopterus belonging to Pterodactyloidea, and Dendrorhynchoides andjeholopterus belonging to thetorhamphorhynchoid Anurognathidae. This This rostrum; it is edentulous. Its manual digitsdigits I ~ IIII---III are robust, and wing clawsclaws rostrum; it is edentulous. Its manual are robust, and wing andJeholopterus belonging the rhamphorhynchoid Anurognathidae. largelarge and curved. The wing digit digit comprises four four phalanges, progressively theofLate]urassic in Solnhofen showsshows somesome resemblance to that and curved. The wing comprises phalanges, progressivelyassemblage assemblage resemblance to of that the Late Jurassic in Solnhofen by the of members of the and and metacarpal and first of theofwing shorter toward the distal end. Wing shorter toward the distal end. Wing metacarpal and phalanx first phalanx the wing(Tithonian) (Tithonian) by sharing the sharing of members of Pterodactylidae the Pterodactylidae digit digit are relatively shortshort compared to Nyctosaurus gracilis from from the Upper The assemblage is associated with with the Confitciusornis avianavian are relatively compared to Nyctosaurus gracilis the UpperAnurognathidae. Anurognathidae. The assemblage is associated the Confuciusornis Associated dinosaurs are also including feathered theropods Cretaceous of western Kansas. Ratios of tibia to femur and tibia to humerus Cretaceous of western Kansas. Ratios of tibia to femur and tibia to humerusfauna.fauna. Associated dinosaurs are abundant, also abundant, including feathered theropods of forelimb (humerus + ulna + +Sinosauropteryx, Beipiaosaurus, Sinornithosaurus, Protarchaeopteryx and Caudipteryx, are 1.5 and the areand 1.5 2.2, and respectively, 2.2, respectively, and ratio the ratio of forelimb (humerus + ulna Sinosauropteryx, Beipiaosaurus, Sinornithosaurus, Protarchaeopteryx and Caudipteryx, wingwing metacarpal) to hind limblimb (femur + tibia + metacarpal III) isIII) 1.1. and the Liaoningosaurus, etc. etc. metacarpal) to hind (femur + tibia + metacarpal is 1.1.the iguanodontidjinzhousaurus, the iguanodontidJinzhousaurus, and ankylosaurid the ankylosaurid Liaoningosaurus, Chaoyangopterus not only represents the first record in Asia but also are mainly fromfrom the Jianshangou Bed,Bed, Dawangzhangzi Bed Bed and and Chaoyangopterus not only represents the such first such record in Asia but the also theTheyThey are mainly the Jianshangou Dawangzhangzi earliest record and most complete skeleton of theoffamily Nyctosauridae. SomeSome ]ingangshan Bed Bed of the Formation, withwith isotope datesdates ranging earliest record and most complete skeleton the family Nyctosauridae. Jingangshan of Yixian the Yixian Formation, isotope ranging of the family are made, such as having four wing digits and wellbetween 121 ~ 125 Ma. revisions revisions of the family are made, such as having four wing digits and well- between 121 --- 12 5 Ma. developed manual digitsdigits I ~ III. developed manual I---III.
The The Daohugou bed of Yixian Formation (note:(note: it is still Daohugou bedthe of lowest the lowest Yixian Formation it is still Liaoningopterus (Fig. (Fig. 137) 137) is referred to thetofamily Anhangureidae. It is It isdebatable as to whether this bed should be referred to the Yixian Formation) Liaoningopterus is referred the family Anhangureidae. debatable as to whether this bed should be referred to the Yixian Formation) the largest pterosaur ever discovered in China; its teeth also represent the producedjeholopterltS. Although currently therethere is no isprecise age for the largest pterosaur ever discovered in China; its teeth also represent the producedJeholopterus. Although currently no precise agethe forfossil the fossil
largest known fromfrom any any pterosaurs. Liaoningopterus is a is large-sized we estimate that that it cannot be older than than 139 Ma, recent 4°Arj39Ar age age largest known pterosaurs. Liaoningopterus a large-sizedbed, bed, we estimate it cannot be older 139 aMa, a recent 4°Ar/39Ar pterodaetyloid, withwith an estimated skullskull length of 610 mm mm and wingspan part part of the Formation, which is underlying the the pterodactyloid, an estimated length of 610 and wingspanof the of upper the upper of Tuchengzi the Tuchengzi Formation, which is underlying skullskull is lowis and Premaxilla and dentary are equipped aboutabout 5 m. 5The bed. bed. Our Our conclusion on the relationship between the the m. The low long. and long. Premaxilla and dentary are equippedDaohugou Daohugou conclusion onstratigraphic the stratigraphic relationship between The The teethteeth are only restricted to the part part of theof theDaohugou with with sagittal crest.crest. is based on the observation bed and Formation sagittal are only restricted to proximal the proximal Daohugou bed the andTuchengzi the Tuchengzi Formation is based onfield the field observation upper and lower jaws.jaws. Toothed portion of theofjaws, aboutabout half the of ofof their contact at the locality. upper and lower Toothed portion the jaws, halflength the length of their contact at Daohugou the Daohugou locality. the skull, does does not extend posteriorly to 1/3 theofnasopreorbital. Teeth near near The upper pterosaur assemblage comprises pterosaurs fromfrom the J the iufotang the skull, not extend posteriorly to of 1/3 the nasopreorbital. Teeth The upper pterosaur assemblage comprises pterosaurs Jiufotang are huge. The The fourth toothtooth of theofpremaxilla is theis theFormation. All pterosaurs collected so far from this formation are the rostral end of the rostral endtheofjaws the jaws are huge. fourth the premaxilla Formation. All pterosaurs collected so far from this formation are largest; the first and third are much smaller than the second and the fourth pterodactyloids. Among themthem Sinopterus, Chaoyangopterus and Liaoningopterus largest; the first and third are much smaller than the second and the fourth pterodactyloids. Among Sinopterus, Chaoyangopterus and Liaoningopterus ones.ones. havehave beenbeen described. No rhamphorhynchoid has been discovered among over over described. No rhamphorhynchoid has been discovered among
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a dozen specimens known from the Jiufotang Formation. The upper assemblage shows more resemblance to that of the Early Cretaceous Santana
Yanornis, Sapeornis and Jeholornis. Associated dinosaurs include the feathered dromaeosaurids Microraptor zhaoianus and M. gui. Although no radiometric
(Aptian/Albian) in comprising only pterodactyloids the dateclpeomi has beenand obtained directly from thedinosaurs beds in which jeb%mis. A ociated in ludethe chepterosaurs feachered were a dozen Formation pecimen known from che Jiufotang Formation. The upper as such em- as YOllomis, Tapejaridae the Anhangueridae. TheEarly age of the Jiufotangamana Formationdromaeo is collected, basalt overlying theand Jiufotang dated as 110 Ma aurid the ,\llcroraplor zhaoiollllS M. glli.Formation Although was n radiometric blage how more reand emblan e co that of the retaceou slightly older than in thecomprising Santana Formation. This pterosaur is hain been Inner obtained Mongolia,directly and therefore, thebed temporal distribution this formation dace from che in which rhe preroofaur \ ere Formation (Aptian/Albian) only pterodaccyloid uchassemblage the associated with the CathayornisThe avianage fauna thatJiufocang includes birds such asiSinornis, is estimated 110-'- 120cheMa. colleered, che bas Itasoverlying Tapejaridae and the Anhangueridae. of the Formation Jiufotang Formation was dated a ItO Ma lightly older than the amana Formation. Thi ptero aur a semblage i ......
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associated with the Cotbayornis avian fauna thac in tude bird
u has
IIIO/1llJ,
in Inner M ngolia, and rherefore, che temporal di tribution of chi formation i e cimated
ltO-120 Ma.
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136 Holotype of Chaoyangopteruszhangi (wingspan about 1.85 m), a nyctosaurid pterosaur, from Gonggao locality (]iufotang Formation), in Chaoyang, Liaoning. (Photo. IVPP) 136 Holotype of Chaoyangopterus z/lOngi (wing pan about I. 5 mI. a n ceo aurid prero aur. from Gon gao locality Oiuforang Formation). in Chaoyang, Liaoning. (Phoro: IVPP)
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137 Holotype ofLiaoningopterus gui (estimated wingspan about 5 m, skull length 61 cm), a large pterodactyloid with the largest tooth known from any pterosaurs, from locality (Jiufotanggui Formation) in wing Chaoyang, Liaoning. IVPP)61 em). a large pterodactyloid with the largest tooth known from any ptero aurs. 137 Xiaoyugou Holotype of Liaonillgopteru (estimated pan about 5 Ill. (Photo: skull length from Xiaoyugou locality Uiufotang Formation) in Chaoyang. Liaoning. (Photo: IVPP)
~ a 139 Close-up view of the integumelats ~)fSi~>sat~ropte~3,,vp~i~no. The filamentous integuments ~ei)resent p~-imiti\,e Feathers ~acl this discovery gives us a ~olimpse ~lt the o~-igin and eaHy evolution offeathers. (Photo: Da-jian l.i/CAS)
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z~cz~138 HolotYl)e of Silloso~loVlel-),x p~in~o, cl ~ooste>sized the~-{)t)()d dinosaur, fl-om Sihetun locality {lowel l)alt ofYixian l-:onnc/ti{)~l}, Beipiao, l.iaoning. {Photo: Da-jian Li/CAS}
Xing Xu
D
100
aur are among the best known prehistOrical animals. They
including auropod
theropod ornithopods, ceraropsians and ankylosaurs.
were the world' conqueror during the [e ozoic repre enting
The following paragraphs briefly describe some important dinosaur taxa from
one of the mo t successful vertebrate 'roup. That i why the
the Jehol Biota \ ith comments on their e olutionary implications.
hina i exrraordinarily
Coeluro aur are a group of derived therop d dino aurs, including the
rich in dinosaur fo sits, the rec rd of which span almost the entire age of
giant tyranno aurids, the long-clawed therizino auroids the deep- kulled
dino aur . Recently, the dino aur remains from the Low r Cretaceous Jehol
oviraproro aur , and the volant bird and their clo e relative -
Group have aroused worldwide arrention. The feathered dino aurs, in
dromaeo aurs and the large-brained rroodontids. In the Jehol Biota the
parricuJar, have challenged the conventional idea on both dinosaur' appearance
non-avian coeluro aurian are repre ented by 12 pecie in 10 genera.
Me oz ic Era i at
0
called the" Age of Dino aur ".
and their life hisrory. Mo t major dino aur gr up are represented in the Jehol Group,
the agile
illosclllroptelJ'x primo (Figs. 13 - L 0) the first named coeluro aurian
from theJehol Biota was found in western Liaoning in 1996.lt is about the
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140 Recon truction of Sinosouropteryx prima. Although a few artist re tored orne non-avian dino aurs with feathers or featherlike integuments as early as 1970s. the discovery of Sino ouropteryx prima provide the first fossil evidence for such a restoration. (Art: Xiao-lian Zeng/ KIB)
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a 141 complete A complete skeleton Caudipteryx zoui, emu-sized 141 keleton of ofaudipteryx ;01/;, anan emuized theropod dina aur, from Zhan 'Iagou localn (low r theropod dinosaur, from Zhangjiagou locality (lower parrpart of ofI Yixian ian F rmatlon). Formation),Belpiao, Beipiao,Liaoning. Liaoning,red red arrow denoting th the gizzard PP) arrow denoting gizzardtone, stones.(Photo: (Photo:I IVPP)
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142 Reconstruction of Coud'pleryx feather but
ZOIll.
imilar to it ovirapt rid relative. ol/dipleryx i a cursorial animal. with
uld not fly, (Art: Ander on Yang)
Reconstruction of Caudipteryxzoui. Similar to its oviraptorid relatives, Caudipteryxis a cursorial animal, with feathers but could notnfly. (Art: Anderson izc of a roo ter, with a large kull and hMp reeth, ve h rr.um and. unable to fl Yang) . Proltlrcb"wpw)x h harp reeth and long arm , and i 142
imilar
unu ually long tail. Alrh ugh ir i a primitive coeluro aunan, u reologi all}' size of a rooster, with a large skull and sharp teeth, very short arms and an quire differenr from bird compared r ad\'anced non-a ian oeluro aurian , unusually long tail. Although it is a primitive coelurosaurian, osteologically I1lfJ tlllmpl I)X oc upi an importanr po irion in under randing rhe rigin of quite different from birds compared to advanced non-avian coelurosaurians, bird be au e of rhe pre ence f a hairlike covering n ir bod}'. fan' Sinosauropteryx occupies an important position in understanding the origin of paleonr logi rs re ard d the hairlike tructure primitive fearher u ed for birds because of the presence of a hairlike covering on its body. Many in ulation bur oth r di a reed, nd b Lie ed they are unrelared ro fe rher . paleontologists regarded the hairlike structures as primitive feathers used for The following year, r 0 Other feather d dina au were ound from the insulation, others disagreed and believedrolJ/l.JltI they are to feathers. ame area. nebut was named Prol rchaeoplel)X andunrelated rhe other Call lipll'I).\
ro, bur Joe nor belong to, the dromaeo aurid . Dromaeo aurid are one of unable to fly. Protarchaeopteryxhas sharp teeth and long arms, and is similar rhe rna r birdlike dina aur 'r up rhar al a include rhe mall- ized elocirllplor, to, but does not belong to, the dromaeosaurids. Dromaeosaurids are one of Igured in rhe rna ie "Jura ic Park", Uldlpl r)x ha h rt arm like the most birdlike dinosaur groups that also include the small-sized Velociraptor, CfJJ11P fJgtltllhllJ but Other feature u 'ge tit i an oviraptoro urian din auf. figured in the movie "Jurassic Park". Caudipteryx has short arms like It has a tall and h rr kull atypic I of rh t in thero dino aur ,I ng leg, Compsognathus but other features suggest it is an oviraptorosaurian dinosaur. and horr rail. The gizzard tOne (Fig, 1 1) indi are thar Ctmdiplel)X ZOIll i It has a tall and short skull atypical of that in theropod dinosaurs, long legs, likely a herbi are, Both Proltlrrhtl Oplel)X and Gal/dip! ryx hav unque tionand The gizzard (Fig.d 141) that Caudipteryx zouivane. is ableshort rrue tail. feather th t restones compo fa indicate pr minent hafr and flat
TheI following year,etwo dinosaurs were more found imilar from the ;:ol/i (Figs. 1 1 2. The tw other dino feathered aur are com ararively ro
likely a herbivore. Both of Protarchaeopteryx Caudipteryx unquestionnJike flighr fearher bird, the Ion and feather on thhave arm and tail of able true feathers that are composed of a prominent shaft and flat vanes.that Pro! trr/}(leOple1)',\ and I/dip! I)'X have ymmetrical vane uggesring
same was named as Protarchaeopteryx robusta andprmJt; the other bird in area. rermOne of bony tru rure th n is ll1oJllf(rOp,el)'X yer Caudipteryx borh were zoui (Figs. 141,142). These two dinosaurs are comparatively more similar to birds in terms of bony structure than is Sinosauropteryxprima; yet both were
Unlike flight feathers of birds, the long feathers on the arms and tails of
Protarchaeopteryx and Caudipteryx have symmetrical vanes, suggesting that
m 143 Holotype of Beipiaosaurus inexpectus, the largest feathered dinosaur found from the Jehol Biota to date with estimated total length more than 2 m, from Sihetun locality (lower part of Yixian Formation), Liaoning. Different typical Beipiaosaurus has numerous teeth, andmore broad feet2 suggestive a slow 143 Holotype of Beipiao aurus inexpectu Beipiao, . the large t feathered dinofrom aur f,theund fromtheropods, theJehol Biota to date with e timatedtiny totallengrh than m. from ihofrun locality IVPP) (I lifestyle. wer part(Photo: ofYixian Formation). Beipiao, Liaoning. Different from the typical therop d . Beipiao auru ha nllmerOll tiny teeth, and broad feet sligge tive of a low lifestyle. (Photo: IVPP)
apparently more complex than those of Sinosauropteryx in having a branched structure. tho e of inosallrOplel)'X in having a branched complex chan Inapparencly 2000, the more sixth and seventh feathered dinosaur species were reported cruccure. from western Liaoning. One is a new species of an established genus In 2000 che150, sixch151), and seveoch feachered species, ere reported Caudipteryx dongi (Figs. which helps clarifydino someaur morphological wegenus. cern Liaonin Oneis ia tiny a new pecie less of than an escablished featuresfrom of the The otherJ. one dinosaur, 40 cm long.genu Callt!iplel)'X dongi 150, zhaoianus 151), which helps morphological This dinosaur, named as (Fig. Microraptor (Figs. 152clarify N 154),orne represents feature of che genu. The ocher one is a tiny dinosaur, less than 0 cm long. the smallest adult dinosaur found to date. Like Sinornithosaurus, it is also a This dino aur, named a 1icyomplol' zhaoiall/ls (Figs. 152 - 154) r preseoc dromaeosaurid. Microraptor is the most birdlike among all known dinosaurs. che smalle t adult dino aur found to dace. Like illomilhosallrlls, ic i al 0 a It is small and has an expanded braincase like birds, long arms that could flap, dromaeo aurid. i\1icl'oraplor i che mo c birdlike among all known dinosaur. and feet that show adaptations to dwelling in trees. This finding suggests that [c i mall and has an expanded braincase like birds long arms chac could flap, some small-sized non-avian dinosaurs moved into the trees in order to escape and feec chat show adaptation co dwelling in cree. Thi finding ugge c thac larger hunters or chase small prey, and gradually evolved flight capability orne mall-sized non-avian dino aur moved ioco che crees in order co e cape living in the trees. A halo of featherlike structures surrounds the fossil skeleton larger hunter or chase mall prey, and gradually evolved flight capabilicy of Microraptor zhaoianus, some of which bear the central shaft. Although the living in the trees. A halo offeatherlike scructure urround che fo il kelecon preservation does not allow us to draw a conclusion that Microraptorzhaoianus of 1icroraplor zhaoitl1lllS, some of which bear the ceocral shaft. Alchough che had vaned feathers, it is most likely that it did. preservation doe not allow u CO draw a conclu ion that Microraplor zhaoianlfS In 2002, three new coelurosaurian theropods have been reported from had aned feacher , ic i mo c likely chat ic did. western Liaoning and adjacent Inner Mongolia. Two of them, namely In 2002, chree new coeluro aurian rheropod have been reported from m 144 Close-up view of integuments ofBeipiaosaurus inexpectus.These filamentous Sinovenator changii (Figs. 155, 156) and Incisivosaurusgauthieri (Figs. 157, 158) we tern Liaoning and adjacene Inner Mongolia. T, 0 of them, namely structures probably represent the primitive feathers and may have wider were from the same fossil the156) Lujiatun Bed of thegalllhieri lowest (Figs. YixianL57, L5 ) 144 Close-up view ofinreguments of Beipiaosauru inexpectu . These filamentous im)l'tnalor changii (Fig bed, . 155, and lncisivosa/lrlls distribution among non-avian theropods. (Photo: IVPP) structure probably represent the primitive feathers and may have wider Formation Chapter 1 forilstratigraphic information). Although no Yixian were (refer from tothe arne fos bed, the Lujiarun Bed of rhe lowest distribution among non-avian theropods. (photo: IVPP) feathers were found on to thesehapcer specimens, due toinformacion). the nature ofAlchough the they are not suitable for flight. Formacion (refer 1 for maybe cracigraphic no
d e p o sfeacher i t s - coarse sandstone, most probably had feathers body of the were found onthey chese specimens, maybe due on to their the nature duringdeposic life. Sinovenator a troodontid, the most had birdlike dinosaur non-avianThis animals extinct, it was the lasthad time, fortunately. - coar eis and cone, cheyone mo of c probably feacher on cheir body was mtheliving fir t orcime in hibut tory thatnot feather been di co ered on groups and very similar in some features to the dromaeosaurids. Again, In 1999, two other feathered dinosaurs were again reported from western during life. in01 1 nalol' is a croodoneid, one of che most birdlike dinosaur non-avian animal -Ii ing or e tincc, but it was noc che lasc cime, forcunacely. Sinovenator is aand small animal less inthan onefeatures meter long. Its dromaeo braincase aurids. is very Again Liaoning. Beipiaosaurus (Figs. is the biggest theropod groups very imilar orne to the In 1999, cwo ocherinexpectus feathered dino143, aurs144) \ ere again reported from yet we tern similar in01'enalol'i to early birds,a indicating a relatively high level of intelligence. Despite i very discovered from Liaoning. It is more (Fig than . 2143, meters with tiny teeth, a yet mall animal Ie s than one mecer long. Its braincase Liaoning. Beipiaosalll'lfS illexpectlls 1 long, ) i che biggesc cheropod Thisarewas first time history that feathers had been discovered on they notthe suitable for in flight.
a theropod dinosaur usually regarded as carnivores, is De pice bulky body, andfrom a short tail. Although some features of Beipiaosaurus similar to early birds, indicating a relatively high level Incisivosaurus of intelligence. di covered Liaoning. Ic i more chan 2 meter long, with suggest ciny teech,being a in having teeth similar to uthose of regarded the plant-eaters. it perhaps a plantand eater, Beipiaosaurus has unusually long, curved, sharp claws being a theropod dinosaur ually a carnivores, IllCish'osa/lrils i bulky body, a hort cail. AIchough orne feacure f Beipiaosam71s suggeunusual t on ititsperhap hands aasplane in meat-eating dinosaurs. eater, Beipiaosalll'lls has Sinornithosaurus unu ually long, millenii curved, (Figs. harp claw
The reported in 2002 small arboreal dinosaur, unu third ual intheropod having teeth imilar co thowas e ofa che plane-eater.
Epidendrosaurus (Figs. 159, 160), and itswas holotype was collected on it hand a in meac-eacing dinosaurs. mi//enii Theningchengensis chird theropod reported in 2002 a small arboreal dinosaur, 145--149), the second species, is a close relative illornilhosallrlls of Velociraptor but much(Figs. Epidendrosllmls ningrhengmsis 160), andlong ic holotype was collecced from southeastern Inner Mongolia. (Figs. It has L59, an extremely third manual 1 5-1 9),dagger-like che secondteeth, species, a clo e relacive of and lIe/ociniplor smaller. It has long iarms that could flap, a rodlike,buc stiffmuch from oucheastern It Sinornithosaurus has dagger-like represents teeth longone arms couldbirdlike flap, anddinosaurs, a rodJike, sciff Inner 10ngolia. Ic unreported ha an excremely chird manual digit, indicating a type of adaptation previously from thelong Mesozoic. tail.smaller. Actually of chat the most Actually one of che mo c birdlikeabove. dinosaurEpidendrosaurus digic, indicacing a cype ofadapcaci n previou unreported che Me ozoic. has some features previously unknownIy in non-avianfrom dinosaurs, andcail. is more closelyiliomilhosallf'flS related to birdsrepre than ene the other species mentioned and is more clo el feathers relaced cowere birds found chan che meneionedorabove. Epidendrosam'/ls ha hallux. orne feature previou Iy unknown in non-avian dino aur , including a fully reversed Its precise phylogenetic position is not yet Although no vaned onocher eitherpecies Beipiaosaurus in luding a fully re ersed hallux. Icsregarded preci e phylogenecic icion of is noc yec Alchough no aned the feacher werestructures found on eicher Beipi 10SallJ'llS clear though Epidendrosaurus is tentatively as the closest po relative Sinornithosaurus specimens, featherlike on the two dinosaurs are or
illomilhosmmls pecimen the featherlike cruccure on che two dino aur are
clear chough Epic!ellc!l'ostl/lI'llS is ceneacively regarded as the closesc relative of
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145 Holot pe of of S;norn;tlJosaurus 1 145 Holotype Sinornithosaurus
millen;;, a small theropod dinomillenii, a small theropod dinoaursaur (estimated length (estimated length1.11.1mi.m), from ihetun locality (lower part
from Sihetun locality (lower part
of Yixian Formation). Beipiao.
of Yixian Formation), Beipiao,
Liaoning. It has feathered fore-
Liaoning. It has feathered fore-
limbs that can move the wa
limbs that can move the way birds' wings do, indicating that
birds' wings do. indicating that
non-avian theropod were pre-
non-avian theropods were pre-
adapted for evolving nying
adapted for evolving flying
capacity. (Photo: IVPP)
capacity. (Photo: IVPP)
~
146 Skull of Sinornithosaurus millenii, showing the sharp and serrated teeth of a
ferocious animal.howing (Photo: the IVPP) 146 Skull ofseemingly Sinornithosaurus millenii, sharp and serrated teeth of a
seemingly ferocious animal. (photo: IVPP)
~,~ 148 Wishbone of Sinornithosaurus millenii, a structure previously thought to be 148 Wi hbone of Sinornithosauru millenii, a structure previou Iy thought to be present only in birds, but recently found to be widespread among dinosaurs. pre ent only in bird . but recently found to be wide pread among dinosaurs. It is morphologically identical to that of Archaeopteryx, the most primitive It i morphologically identical to that of Archaeoplery , the most primitive bird. (Photo: IVPP) bird. (Photo: IVPP)
~;~ 147 The rodlike tail bones ofSinornithosaurus millenfi, unique to the dromaeosaurids, stiff tail. (Photo: IVPP) millenii, unique to the dromaeosaurids. 147 Theindicating rodlike taila bones ofSinomithosaurus
indicating a stiff tail. (Photo: IVPP)
..,
~ 149 Close-up view of the integuments of Sinornithosaurus millenii, showing 14g Clo e-up view of the integument of Sinornithosaurus millenii, showing the branched filamentous integuments, a unique feature in feathers. the branched filamentous integuments. a unique feature in feather. (Photo: IVPP) (Photo: IVPP)
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,150 Holot pe of Caudlpteryx dongi. 150 Holotype of Caudipt imilar in size to C. zoui. from
similar in size to EC, Zhangjiagou locality (I wer part
of Vi ian Formation). Beipiao. Liaoning. (Photo; I PPJ Liaoning, (Photo: IVpP), ;:
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1151 The remige pre erved on the
Caudipteryx dongi pecimen. the flight 151 TheDifferent remiges from preserved on feathers the in volanr bird . the remige Caudipteryx dongi specimen. on the forelim of Caudipteryx Different from thebsflight feathers have symmetrical vane • dongi in volant birds, the remiges on indicating a lack of aerodynamic the forelimbs of Caudipteryx function. (Photo: IVPP) dongi have symmetrical vanes,
indicating a lack ofaer function. (Photo: IVPP)
mic
152 Holotype of Mlcroraptor zhaoianus, the mallest known adult non-avian
153 The pes of icroraptor zhaoianus, orne of it pedal features uggestive of
dino aur (e timated total length les than 40 em), from Lallgshan locality
an arboreal habit, indicating that the theropod ancestors of birds might
152 Holotype of Microraptor zhaoianus, the smallest known adult non-avian Uiufotang Formation) in Chaoyang, Liaoning. (Photo: IVPP) dinosaur (estimated total length less than 40 cm), from Langshan locality (Jiufotang Formation)in Chaoyang, Liaoning. (Photo: IVPP)
153passed The pes of Microraptor zhaoianus, of its pedal features suggestive of IVPP) have through an arboreal pha e.some (Photo: an arboreal habit, indicating that the theropod ancestors of birds might have passed through an arboreal phase. (Photo: IVPP)
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154 Recon trllction of Micro154 Reconstruction of Microraptor (Art: raptor zhaoianus. zlJaoianus. (Art: Rong-shan Rong-shan Li/IVPP) LilIVPP)
'2_157 iirJ 157
Holotype Holotype of of Incisivosaurus Incisivosaurus gauthieri gauthieri (skull (skull length length about about 1III Clll), em), aa sillall small oviraptorosaur with peculiar tooth Illorphology, fronl Lujiatull locality oviraptorosaur with peculiar tooth morphology. from Lujiarun locality (lowest (lowest part part ofYixian of Yixian Fornlation) Formation) inin Beipiao, Beipiao. Liaoning. Liaoning. (Photo: (Photo: IVPP) IVPP)
[he the birds. birds. A new feathered feathered dinosaur, dinosaur, i\tlicrorclptor Mirrorap/or glli glli (Figs. (Figs. 161, 161, 162), 162), \vas was dedeA ne\v scribed scribed early early in in 2003. 2003. It It isis the the second econd species species of of "'licrorc/ptor, Microrap/or, and and aa small small light\veight lightweight aninlal animal of of 77 7 em cm long, long, \vith with aa rodlike rodlike long long tail. tail. This This dinosaur dinosaur probably probably lived lived in in the the trees. trees. Surprisingly urprisingly itit has has long long pennaceous pennaceous feathers feathers not not only only on on its its forelimbs forelimbs and and tail, tail, but but also also on on its its hind hind limbs limbs (the (the so-called so-called "four"four\vinged winged dinosaur"). dinosaur"). Furthermore, Furthermore, the the feathers feathers \"ere were almost almost identical identical to to those those of of living living birds, birds, \vith with asynlmetrical asymmetrical vanes, vanes, aa feature feature associated associated \vith with flight flight or or gliding glli isis aa gliding gliding in in extant extant birds. birds. It It isis very very likely likely that that )vlicroraptor MiCl'orap/orglii gliding animal, animal, representing an intermediate stage bet\veen the flightless dinosaurs and representing an intermediate stage between the flightless dinosaurs and the the volant volant birds. birds. Anlazingly, Amazingly, as as many many as as 12 12 coelurosaurian coelurosaurian theropod theropod species species have have been been reported reported from from aa small small "area area in in \vestern western Liaoning Liaoning and and the the adjacent adjacent Inner Inner -j
. . 155 Partial .155 Partial skull skull of of the the holotype holotype of ofS;novenalor Sinovenator chang;; changii. aa small small theropod m, theropod dinosaur dinosaur with with estimated estimated body body length length less less than than 1I m,
Mongolia Mongolia in in last last aa fe\v few years. years. More More surprisingly, surprisingly, these these discoveries discoveries have have provided provided us us ,vith with much-needed much-needed information information on on important important issues issues such such as as the the
Yixian Formation) from from Lujiatun Lujiatun locality locality (lowest (lowest part part of ofYixian Formation) in in
origin origin of of feathers feathers and and bird bird flight flight that that ,vas was unavailable unavailable from from all all other other kno\vn known
Beipiao, Beipiao, Liaoning. Liaoning. (Photo: (Photo: IVPP) IVPP)
dinosaur dinosaur specimens specimens collected collected world\vide worldwide over over the the last last century century and and aa half. half. Before Before these these discoveries discoveries from from ,vestern western Liaoning, Liaoning, there there ,vas was little little fossil fossil
t
120 120
evidence evidence for for studying studying the the origin origin of offeathers. feathers. Due Due to to their their structural structural complexity, complexity, feathers feathers are are very very distinctive distinctive from from all all other other integumentary integumentary structures, struCtures, and and their their abrupt abrupt appearanc~ appearance in in the the fossil fossil record record has has been been perplexing. perplexing. No,v ow that that the the hypothesis hypothesis of ofdinosaurian dinosaurian origin origin of of bird bird has has been been ,veIl well established, established, itit theoretitheoretically cally fo11o,vs follows that that feathers feathers should should be be present present on on some some birdlike birdlike theropods. theropods. Ho\vever, no intermediate structures that could be feather precursors However, no intermediate structures that could be feather precursors ,vere were preserved preserved in in the the related related theropods. theropods. On On the the contrary, contrary, most most evidence evidence suggested suggested that that dinosaurs dinosaurs ,vere were scaled scaled animals, animals, including including some some theropod theropod dinosaurs. dinosaurs. For For example, example, some some psittacosaurid psirracosaurid specimens specimens from from the the same same localities localities as as the the feathered feathered theropod theropod specimens specimens preserved preserved beautiful beautiful skin skin impressions, impressions, sho\ving showing that that these these animals animals had had scaly scaly skin. skin. The The feathered feathered dinosaurs dinosaurs from from Liaoning Liaoning are are just ,vhat many paleontologists ,vere anticipating, and thus provide direct just what many paleontologists were anticipating, and thus provide direCt fossil fossil evidence evidence for for the the hypothesis hypothesis that that feathers feathers are are not not unique unique to to birds birds and and can can be be traced traced back back into into the the dinosaurian dinosaurian ancestors ancestors of of birds. birds. The The featherlike featherlike
_156 .156 Reconstruction Reconstruction of of S;novenalor Sinovenator chang;;. c1rangii.
structures structures on on these these different different dinosaurs dinosaurs are are diverse diverse in in morphology, morphology, but but display display an an evolutionary evolutionary trend trend in in complexity complexity approaching approaching the the origin origin of of birds. birds.
(Art: (Art: Michael Michael W. W. Skrepnickl Skrepnickl FMNH) FMNH)
SiJlOS{lIlroptelJ'x inosallrop/eryx has has simple simple feathers; feathers; Beipic/osallrllJ Beipiaosallms and and SillorllithosCtllrllJ Sinornithosallms have have ~ 158 Reconstruction .158 Reconstruction of of Incisivosal/rus Incisivosaurus gal/thieri. gauthieri.
(Art: (Art: Portia Portia Sloan). Sloan).
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branched branched feathers feathers but but bear bear no no vanes; vanes; Protc/rchc/eopteIJ'x PI'O/arrhaeop/el)'x and and Cetllclipteryx Calldip/eryx have have vaned feathers. The lack of vaned feathers on Beipic/osclllrlls developed and developed vaned feathers. The lack of vaned feathers on Beipiaosallms and
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_ . 160 Reconstruction of Epidendrosaurus Epidrndro auru n;ngchengensis. ningclJrngrn i . showing its it . feature . (An: (Art: RongRang-shan han LilIVPP). L' IVPP). arboreal features.
inomilhos UlrIIS are mo likel due [0 to preservations. pre ervarion . ]n In fact, the most mo t recent discovery di over suggests uggest that rhat SinornithoJallrtlJ mostt likely inor1lilhoStlllrlls and Microraplor fiero/'ap,or have h ve vaned feathers fearhers as Protarchaeoptery·x PrOIa,., ha OPIflJX and Catldipleryx alldiplflJx do. SinornithoJatlrtlJ Here is i the rhe current urrenr picture of feather fear her evolution: evolurion: most mo r dinosaurs, din saurs including all ornithischian ornithi chian u
("bird-hipped" dinosaurs), dino aur ), all prosauropods pro auropods and sauropods, auropods, and some primitive primirive theropods, rheropod ,are are similar imilar ("bird-hipped ro typical rypical reptiles reptile in that rhar their rheir bodies are covered co ered with wirh scales cale or tubercles; rubercle ; the rhe first firsr feather evolved evol ed in to rhe early earl stages off coelurosaurian coelur aurian dinosaurs, dino aur , and was a simple, imple, hairlike structure, rrucrure, probably used u ed to ro the rhe animal warm; more complex feathers fear her evolved e olved later, and display di playaa branching structure; rru rure; long, keep the fearher evolved e olved in some orne maniraptoran maniraproran dinosaurs, dinosaur, probably including therizinosauroids, rherizino auroids, vaned feathers o iraptoro aurs, troodontids rroodonrids and dromaeosaurids, dromaeo aurids, and these rhese complex feathers fearher were likely for display; display' oviraptorosaurs, a ymmerrical flight flighr feathers fearhers evolved e olved and are used to ro generate lift during flight. Therefore, Therefl re, finally, asymmetrical fearher appeared before the rhe origin of birds bird (Fig. 163) and early feathers fearhers functionally fun rionally had nothing norhing to ro feathers irh flight. In the rhe future, furure, if we find a feathered fearhered fossil fos il animal, we need to ro be careful areful when v hen do with derermining what \ har kind off animal it ir is i - it ir could be a bird, but bur it ir could also al 0 be a flightless flighrles dinosaur. din aur. determining mpared to ro the rhe highly publicized theropod rheropod dinosaurs dino aurs from the rhe Jehol Fauna, the rhe ornithischian ornirhi chian Compared dino aur remain little lirrle known, knov n, despite despire the rhe fact facr that rhar the rhe first fir r dinosaurian dino aurian discovery di c very from the rhe Jehol Jeh I dinosaurs
Fauna was an ornithischian. ornirhischian. To date, dare three rhree major omithischian ornirhi chian groups group have been reported reporred from the rhe Jehol Biota, Bi ra, i.e., ankylosaurs, ankylo aur ,ornirhopod ornithopods,, and ceratopsians. ceratop ians. ceratop ian are a late lare group of herbivorous herbi orou dinosaurs, mostly mo rl restricted resrricted to the rhe Cretaceous rera e u The ceratopsians orne cranial ranial modifications, modificarion including in luding the rhe unique rostral ro rral bone that rhar period. They are characterized by some i otherwise orh rwi e unknown in any other orher dinosaurs. dino aur . In 19705 1970 some orne ceratopsian cerarop ian dinosaur din aur specimens pecimen were is collected from the rhe Jiufotang Jiuforang Formation Formarion of western wesrern Liaoning. Identified Idenrified later larer as a a new species pe ie of
psittacosaurid psirraco aurid dinosaur, din aur, PsjllaCOJaNrNJ Psilla OSt/II/'IIS mei/eyingenJiJ llIeile)'iflgensis has a relatively relarively tall rail and rounded skull kull (Fig. 164). P irra 0 aurid , known only from the rhe Early Cretaceous reraceous of Asia, A ia, are a basal lineage of the rhe ceratopsian ceratop ian Psittacosaurids, dino aur . They are facultative faculrarive biped, different differenr from the rhe more derived, quadrupedal neoceratopsians. neoceratop ians. dinosaurs.
The other orher ceratopsian cerarop ian from theJehol rhe Jehol Fauna is the rhe goat-sized goar- ized liaOCeralOpJ Litlow' tlOPS yanzigollenJjJ )'al1zigolleflsis (Figs. 165, 166), neo erarop ian, which belongs belong to ro the rhe second e ond major lineage linea 'e of the rhe ceratopsians. cerarop ian. It Ir has ha only a neoceratopsian, rudimenrar horns and a frill, different from the rhe more derived neoceratops neocerarop that rhar usually weigh much mu h rudimentary mas ive horns horn and a wide frill. Lia«eratopJ Liaocrralops yanzigoNenjjJ ),tl1lzigollensis is i the rhe smallest, malle r, oldest olde rand and heavier and have massive mo r primitive primirive neoceratopsian ne eratop ian ever found. most ornirhopod represent repre enr the rhe most mo r diverse diver e group of ornithischian ornirhi chian dinosaurs, dino aur ,includin rhe The ornithopods including the primirive small-bodied forms form such u h as Heterodontosauridae, Hererodonr auridae, the rhe intermediate inrermediate representatives repre enrarive such uch as primitive famou 19t1anodon, [gllt/1Iodo1l, and the derived large-bodied forms form such u h as a duck-billed dinosaurs. dino aur . The first fir t the famous rhe Jehol Fauna F una isjeholoJaNrtlJ i jeholostlltr!IS JhanytlanenJiJ Sht/1I)lItll1 1Isis (Fig. 167). 167 . It is i a small mall dinosaur dino aur ornithopod found from the Ie than one meter long, Ion' and appears to ro be very primitive in a number of features feature despite de pire being a less retaceou ornithopod.jeholoJatlrtlJ ornithop d.jrholos 1ImlS is i placed pia ed in the Ornithopoda, Ornirhopoda, but displays di play a few features feature similar imilar Cretaceous
to ro those tho e of ceratopsian eratOP ian dinosaurs. dinosaur. It is i a potentially porentially important importanr taxon for understanding under randing the early Epidendro auru ningchengensis, ningclJrngrn is. a small mall theropod with the size of . . 159 Holotype of Epid~ndroSQurus a house sparrow. from Daohugou locality localit in Ningcheng, Ningcheng. Inner Mongolia. (Photo: IVPP)
evolution e olution of the ornithischian ornirhi chian dinosaurs.jinzhoJisallrNJ dino aurs.ji1lzhollstlllrtlS yangi ytll1gi (Fig. 168) 16 ) represents repre enrs the rhe second econd
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ornithopod pecies found from the Yixian Formation of we tern Liaoning. About seven co eight meters long, ir repre ents the largest named dino aur from the Jehol Group. jinzhollJall.,./ls i
imilar to
Igllanodon, one of the earliest named dinosaurs in the world, in many features, such as the pike-like pollex. Interestingly, it has a combination of primitive and derived charaCters, some features more primitive than the cOntemporary iguanodontids but others imilar co those of derived hadrosaurs. Ankylosaurs are a group of highly pecialized ornithischian dinosaurs and easily recognized by their extensive body armor. Liaoningosa/lrtls paradox/ls (Fig. 169) represents the only known ankylosaur from the Jehol Biota. It is a beautifully preserved juvenile pecimen less than 0 cm long. Liaoningosalll71s has a large, somewhat shell-like bony plate under its belly. Thi di covery is the first record of such a tructure among dinosaurs thu adding to our kno~
ledge of the morphological diver ity of dino aur . Though a cladistic
analysis has placed Liaoningosallrtls in
odosauridae, a number of distinCt
ankylo aurid features eem to bridge the morphological gap ben een the two ankylosaurian familie . lr i also po sible that Liaoningosallrtls is neither
Pterosauria Ornithischia Sauropodomorpha Cerato auria Carno auria Comp ognathus Sinosauropteryx Tyranno aurus Beipiaosaurus Therizinosauru lncisivosaurus Caudipteryx Oviraptor Protarchaeopteryx Sinovenator Troodon Microraptor Sinornithosaurus Deinonychus Archaeopteryx Modern birds 163 The phylogenetic positions of the Jehol non-avian
theropods (in red) among the related groups.
ankylosaurid nor nodosaurid, but rathet a ba al ankylo aur. o far 17 new dinosaurian species have been reported from the Jehol Biota. The disco eries of feathered dino aurs are significant because they pro ide the most compelling e idence upporring the hypothesi that birds 161 Holotype of Microraptor gui (total length 77 em). a small dromaeosaurid and
the so-called "four-winged dinosaur" characterized by feathers on all it limbs. from Dapingfang locality Uiufotang Formation) in Chaoyang. Liaoning. (Photo: IVPP)
were descended from dinosaurs, and also improve our under tanding of the origin and early evolution offeathers and the origin of bird flight by indicating that feather evolved before powered flight, and that flight probably evolved through a gliding tage. Other di co eries of non-feathered dino aurs from western Liaoning are also very important becau e they have changed dinosaurian phylogenetic patterns proposed by previous tudie and ignificanrly advanced the study on character e olution for many dinosaurian lineages. The recent dino aurian disco eries from the Jehol Group have provided the most comprehensive evidence yet of dinosaurian oft tissues and will undoubtedly produce many more insights in the future.
162 Reconstruction of Microraptor gui. (Art: Portia Sloan)
164 Skull of the holotype of Psittacosaurus meileyingensis
(estimated body length I-2m) from Meileyingzi locality Uiufotang Formation). Chaoyang. Liaoning. Psittacosaurs are a group of herbivorous dinosaur living in the Early Cretaceous of Asia and a distant relative of Triceratops.
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less than l m), a Small p imlile Llaoceratops yanzigouensl part of • • r "tire c • ~s(tot Yixlan Formation)in D_. e.ratopslan, from Yank:_ al body length oe~plao, Liaonin~ ¢n~ gou lOcality (lower ~," ~rlloto: ompl Ie kull of a juvenile Liaoceralop yanzigouensi (totalIVPp) body length •
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167 A skull ofjeholo auru slwngyuanen i (e timated bod len th Ie
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167 A skull °fJeholosaurus shanffO'Uanensis (estimated bodylength less than 1 m), a small ornithopod dinosaur. from Lujiatun I cality (I wer part Small °rnithopod dinosaur, from(Photo: LujiatunIVPP) locality (lower part ofYi1 m), ianaFormation). Beipiao. Liaoning.
parr of Yi ian Formati n) in B ipiao. Liaonin . (Photo: IVPP)
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m 169 Holotype of Liaoningosaurus paradoxus (body length less than 40 cm), the smallest ankyiosaur 169 Holot pe of Liaon;ngo auru paradoxu (bod found to date, from Wangjiagou locality (middle length Ie than 40 em). the malle t ankylo aur part of Yixian Formation), Jinzhou, Liaoning. found to date, from Wangjiagou loeali
(Photo: IVPP)
(middle
part of Yixian Formation). Jinzhou. Liaoning.
168 Skull of the holotype ofJinzhousaurusyangi (skull about 50 cm long and 28 cm high), an iguanodont, fromkull Baicaigou partollru of Yixian IVPP) 168 ofrhe hlocality I type(middle ofJ;nzholl yang; Formation), (skull ab utJinzhou, 50 em I Liaoning. ng and 2 (Photo: em hi h), an iguanodont. fr m Baicaigou loeali
(middle part fYi ian Formation). Jinzhou, Liaoning. (Phot : IVPP)
(Photo: IVPP)
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BRD~ Zh Il 7_1 r bably m re than
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pe pIe all over the world. Largelr Imitating avian fli 'ht, .. e have cen our dr am of flying c me eru thr u 'h the inventl n of irplane and pa e raft.
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tern
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Mod I of Archoeoplery , the old t bird, whi h link d reptil to bird . U I.' ting a reptile-bird r I ti nhip. Ie uny: Larry D. Martini J
f thou and
171 A complete keleton of COnfuClu om;
po, a primitive beaked bird, fr m Yixian formation) in Beipia . Liaoning, howin well pre erved impre feath r'. te the t\ 0 P cial feature 0 thi bird jaw beaked (t othl . Upper); humeru trian ular with elliptic Ii ne tra t it I.' panded pro imal end (Lower). (Pilot : IVPP)
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.t opt IJX have been found over the past pa t 140 1 0 years. year. Although Alth ugh some orne other opte'J.\" Mesozoi birds bird were also al 0 discovered di overed in other ocher regions region of the world, they are all Mesozoic
little diversity. much younger than Archaeopteryx, with litde Thi situation icuation has significantly signifl andy changed since in e the 1990s 1990 with the This dis overies of abundant Mesozoic Me ozoic birds bird from Liaoning, Liaonin Hebei, Inner Mongolia, discoveries handong, Ningxia ingxi and other areas area of China. hina. These The e new fossils fo il have Shandong, e olution and radiation of birds. bird revolutionarized our view of the early evolution Me ozoic bird bitd from Liaoning was actually collected by a farmer The first Mesozoic haoyang in 1987 19 7 and then sent to the from the Meileyingzi locality in Chaoyang atural History Hi cory Museum for study; tudy; this thi bird was w later named SinorniJ. inomiJ. Beijing Natural eptember of 1990, we '\ e discovered di 0 ered three fossil fo il bird skeletons skeletOn from the In September I cality in Chaoyang, ha ang. Liaoning Province Pro ince (one one of them was later Boluochi locality
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172 A Confuciusornis "couple" buried together. The one on the left with a Confuciu ornis 'couple' long tail feather is assumed a umed as a a male. (Photo: Da-jian Lil LiI CAS) CA )
Beipiao. Liaoning. liaoning. The num.. numofYi ian Formation) in Beipiao, part ofYixian
ha surpassed urpassed the total number ber of Confuciusornis specimens has of other Mesozoic Me ozoic birds bird combined. ~Photo: (Photo: IVPP)
di tal ends of central tail feather ofCon!uciusomis. of Confuciu ornis. (Photo: IVPP) . . 174 The distal
named
Cathayornis). These fossils were all from the Early Cretaceous Jiufotang Formation, and fit right in the transition form the Late Jurassic Archaeopteryx
Confuciusornis(Figs. 171--- 175) is the earliest known bird without teeth. Unlike most other early birds, Confuciusornishad a horny
to the Late Cretaceous birds. However, their discovery was only the beginning
beak, just like the birds in your backyard. The loss of teeth and the appearance
Confuciusomis
of a series of exciting fossil findings in the area that have lasted to this day. For of horny beak may also indicate the reduction of the cutting function of jaws, named II!Jfl}Orlm). The e fI it Wl're all from the: Earl rl'taceous J IUfocang Confuciusornis IIfJji/IIIWmJiJ 'lrd. The 10 0 teeth ,ind the appearance: In 1993, the first skeleton of Confuciusornis in a fossil collector's home Confuciusornis is about the size of Archaeopteryx. The postorbital of of a ene of exciting fo il tindi;1 ,. In the area that ha e I ted to tim day. For o horn be. k may al 0 Indl ate the reduction of the curring function of jaw. drew the attention of some colleagues from the IVPP. It was later named after Confuciusornis is large, and ventrally connected with the jugal, suggesting that in tance, ove:r
t\ 0
doze:n of bird keletOn were dl co\ered from thi local it
a renowned Chinese saint, Confucius, who lived more than 2,500 years ago, alone dunng the follo\ ing leld ea on in the early 1990 .
and the hiftlng of the m
tlcatOry proce
m, inl to it gizzard. for which,
Archaeopteryx may also have a postorbital, thus a typical diapsid skull. ho\ e cr. we ha e not had an ' direct fos il c idcnce.
and recognized as the earliest known bird with a horny beak. A large number girdle bones, scapula and coracoid, are firmly connected, In 199. , the fi r t kcletOn of fJ11jiICIIISIJl'lm In a fo if colleCtor' home The pectoral ol1ji/cil/SonJi i ab ut the ize of /lrc!Jfl 0pll1l).\. The po corbit, 1 of of Confuciusornis specimens have since been excavated from the Yixian and appear more primitive in Archaeopteryx. Confudusornis also drew the arre:ntion 0 orne colleague rom the I PP. It w later n.tmed after rJlJjilf//lSorm I large:.than and ventrall ' connected WIth rhe jug. I. has uggea ting that Formation at several localities in Beipiao and Chaoyang, western Liaoning. a reno\ ned
hine e aint,
primitive hand similarmay to that Archaeopteryx. Except thepical pectoral onfuciu, who Ii cd more than 2,500 year ago, rlnhrILoplel)" al 0 ofhave a rororbital. thuforat diapgirdle id kull.
Many birds havethe also beent collected from a younger formation, thenumber and forelimb,The most of other features however, moreconnected, andfossil recogntzed' earlle known bird with a horn' beak. A hlrge pe:ctoral girdle bone.of Confudusornis apula and c are, ra oid, arc firml}' Jiufotang Formation in pecimen that area. Hundreds specimens of Mesozoic derived,and suggesting a moreI powerful capability. For example, the distalal h a of rmji/(/lIsr)"l/i ha e ince:ofbeen excavated rom birds the Yixian appear more rimiti\'e flight than in /In!Juopltl)x. CrllJflll"illJ(JnJlJ haveFormati been discovered fromlocalitie the JeholInBiota sinceand Sinornis was described, tail vertebrae of Confuciusornis into one solid bone, pygostyle, n at everal Beipiao h oyang. we tern and Liaoning. primltivc hand imilarare to fused that of Ilr(!Jtltflpltrp. E cept r thewhich pectoral girdle western Liaoning has increasingly the hottest areaaounger for the study of the fany fo il bird ha e al 0 become been ollected from formation. the and forelimb. m t 0from othe:r of (l11jiIClIISQnUS are. caudal ho\ ever. more is remarkably distinguishable that eacure of Archaeopteryx that has a long origin and early evolution of that birds.area. Hundre:d of pelime:n f 1e ozoic bIrd Jiufotang Formati n in denved, uggevertebrae. ting a more werful pabdlty. For example, tails with 22 unfused A short and flIght steadycbody of Confuciusornis is, the di tal have been di
0
ered from thc Jeh I Biota ince
l1I'mm w
de cribed. and
tad ertebrae of fJ1Jjim/lSIJl'J1i are fu ed into one olid bone, p go tyle which
Uji 176 A specimen of Sapeornis chaoyangensis, the largest bird (about as large as Archaeopteryx), from Shangheshou (]iufotang Formation) we tern Lla nin' h· in re ingl' be orne the hknown tte t Early area Cretaceous for the cudy f the twiceremarkably dl tin UI habit: fr m that oflocality /lnh(/t:Op/~'JX that h a long caudal in Chaoyang, Liaoning. Its elongated forelimbs are longer than those of Longipteryx in proportion; however, its short and robust coracoids are comparable to those t d \ ith 2_ unfu ed vertebrae. hoft and teady body OlJjiI of Archaeopteryxand theropod dinosaurs. (Photo: IVPP)
ongin and early evolution of bird.
176 A P cim n of apeomi cllao 'angen i , rhe large I kn wn Earl in Chaoyang. Lia nm . II
reta ou bird lab UI twi e a large a Arcllat'optery ). fr m han he h Ulocali
longaled for limb are I nger than th
of Archaeopteryx and Iherop d dino aur . (pholo: I PP)
e of Longipt ry in pr portion; h w ver. it
hort and robu I
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ra oid are c mparable I Ih
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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
..
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.
175).
o
tail.
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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183 YHolotype of Liaoxiomis delicate. a ju enile or ubadult oppo ite bird ,lJlgzl JOl ,JIll ~ (mlClcl/ l ' p,lfl oj Jw,rngzh l.rnl'Y onl rn litl. rorrnall 1,111 ab ut a Great 10 ali (middle part of (IXize 11,lOnrng (1)1 1I,1I1.Dawangzhangzi '"from lotn: J\ PP)
Yi ian Formation) in Lingyuan. Liaonin . (photo: I PP) _184 Rl-C. ()n~1 flU. t Inn of /10 184 dt'i,( R con II.~ 1m III' a /<" lrueti n of Liaoxiomi Andc.-rson Y,tn~) de/irat .(An (An: Andl.'rs n Yang)
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of Boluochia is that the most anterior end of premaxilla is curved into a hook
at that time. It is distinguishable from other birds by its relatively short
as in some extant raptorial and passerine birds. No tooth has been discovered
rostrum and deep skull. Eoenantiornis is smaller than Confuciusornis, but larger
from the premaxilla. was therefore thati Boluochia probably a that i di tingui hablc from other birds relatively hart size of BO/1I0c/JlfI i that the mostItanterior end ofproposed premaxilla curved inro a hookhad at thantime. otherIt enantiornithine birds, consistent with by theittendency of body similar lifestyle as some raptorial passerine birds.disco ered : in ome extanr raproriaJ and extant pas erine birds. and 0 tOOth has been
ro trum and deep kull.evolution EOU/fll/liol'1l1J i smaller but larger are reduction in the of early birds.than The ol/jllrillsomis, teeth of Eoenantiornis
than other enaoriornithine birds con i teor with the tendency of body ize from the premaxilla. It wa therefore cd thatis BO/llorh", probably had aclaws, Another significant featurepropo of Boluochia its long and curved foot characteristic of all known toothed birds in having a constriction at the base imilarsuggesting life tyle a climbing ome extanr raprorial ability. and I' Aterine birds.end of the tarsometa-reduCtion in the evolution of early bird. The teeth of EOCl/flllliol'n1J are and perching the distal of the crown. Another notable feature of Eoenantiornis is the preservation of chara teri tic of all known roothed bird in haVing a constriction ar rhe ba e Another BO/"Oc/JllI IS it on long curved claw, tarsus theignificanr trochleae feature for the of digits are nearly theand same level, foot further proving
bastard wing as in Protopteryx. The bastard wing was not present in Archae-
ugge that ting it dimbing ability. Atcapability. the di tal end of the tar ometais a birdand withperching strong perching
of rhe crown. Another norable feature of Eotl101l11ll1'/1IJ is the pre ervation of
opteryx and Confuciusornis. It is probably present in all other enantiornithines
tar us the trochleae for the digit are(Figs. nearly on 184) the same eI, further provin ' ba tard " ing as in PmloplelJ.\. The bastard \ ing \ as nor pre enr in ,Irrh({/:Liaoxiomis Liaoxiornis 183, is theIesmallest known Mesozoic and ornithurines (all extant birds are ornithurine birds). that it bird i a reported bird withfrom (fang capability.in Lingyuan, western Liaoning in OpltlJx and onj"cillJornis. Ir i probably pre enr in all other enaoriornirhine theperching Yixian Formation
Liaoxiornis Liflo.\lornis (Fig. 184) i the mallest kno\ n Me azoic 1999. It is about the size of a183. sparrow.
Protopteryx Protopteryx (Figs,
186, 187) is about the size of a gray
and ornithurine ( II extaor bird are ornirhurine birds).
bird reponed from the Yixian Formation in Lingyuan, \ estern Liaoning in
Liaoxiornis has a large and deep skull. Both upper and lower jaws are
starling. As its name implies it is "A bird with primitive feathers". The two Protopteryx PrOIOpll:lJX (Fig, 1 6, 1 7) i abour the ize of a gray central tail feathers of Protopteryx
rarling. As ir name implie it i "A bird with primiri e feather ". The twO 1999. It i abour the ize of a sparrow. toothed as in most other Mesozoic birds. Both primitive and derived features are most distinctive Liao.\/fimis has a large and deep skull. Barh upper and lower jaw are
ceorral tail feather of Prolopll:ryx
can be found in Liaoxiornis. One of distinctive features of this bird is that the
roothed as in mo t other 1e ozoic bird. Both primiti e and derived feature
are mo t di tinctlve
thighbone is longer than the humerus; the pygostyle is longer than the neck. can be found in Liaoxiornis. One of di tinerive fe ture of thi bird i that the The derived features of Liaoxiornis include" the proximal ends of thighbone i Ion ter than the humerus; the pygostyle i longer than the neck.
metacarpal bones being fused together; the number of
The derived feature of Lltloxiomis include: the proximal end of
phalanges of the hand less than that of
metacarpal bone being fused rogether' the number of
more primitive birds such as
phalange of the hand Ie
Archaeopteryx
more primitive bird
than that of
uch a
.tI rrhaI:Opll:r).\
among early birds. birds. Its distal among end is n oearly differt much
m 187 Reconstruction of Protopteryx, with a long, central tail feather. This unbranched feather 187 Recon truction f Pr%p/eryx. with a long. may represent the ancestral type of feather, central tail feather. Thi unbranched feather providing evidence for a scale-feather ma represent the ancestral type of feather. relationship. (Art: IVPP) providing evidence for a cale-feather relationship. IArt: IVPP)
di ral nor mu h different fromIt that of end otheri birds in comprising eorshaft from or that of other compri a central rachis, andbird barbsin on both ing sides; a ceorral shaft orregion rachi ,isand barb on both ide; however, its proximal unbranched, that is, lacks howeverbarbs it proximal region i unbranched, rharproximal i , lack tail differentiated on either side of the rachis. Thus the differeoriated barb either idescalelike of the structure, rachi . Thu thewas proximal tail as feather is more like on an elongated which interpreted
feather i more like anstage e10ngared caJelike crucrure, whi avian h wasfeather. inrerpreted an intermediate between reptilian scale and Aftera the and Confuciusornis; the wishbone having a long hypocleideum; and thean inrermediate rage ben een reptilian cale and a ian fe cher. After che discovery of Protopteryx, similar primitive tail feathers have also been recogelongated. dis overy of the ProIOPlnJX, imilar primitive cail feather have also been recogand coracoid ol/jllrilfJomis; the \Vi hbone ha ing a long hypocleideum; and the nized in specimens of Confuciusornis and some other enaritiornithine birds. The juvenile and enantiornithine features of Liaoxiornis indicate that itnized in rhe specimens of on/t,ci"sol1lis and ome Other enanciornithine bird. coracoid elongated. Protopteryxalso represents the most primitive of the known enantiornithine is a juvenile bird, yet more materials are needed clarify PIVIOPIt:IJX aJso repre eors the rna c primiti e of the known enanciornichim: The juvenileenantiornithine and enanriornithine feature f UlIOXlOl'f1lJ indicatetothat it its birds. Some of its hand bones are longer than those of other enantiornithine phylogenetic position in early bird. orne of it hand b ne are longer than tho e of orher enantiornithine a juvenile enaoriornithine bird, yetavian moreevolution. material arc needed ro clarify it birds. On the other hand, it develops the initial procoracoid on the coracoid, Eoenantiornis Eoenantiornis (Fig. 185) was so named because when itbird. On the other hand, it develop the initial procoracoid on the coracoid, phylogenetic po it ion in early avian evolution. indicative of the presence of triosseal canal, which is a derived and key was published inEOUltlllliomis 1999 it represented most 0primitive enantiornithine Eoenantiornis (Fig. 1 the '5) \Va named because when It birdindicarive of rhe pre cnce of trio seal canal, which is a derived and key structure adapted for flapping flight of modern birds. tructure adapted for flapping fljghr of modern birds. wa publi hed in 1999 it repre eored the most prinllti e enanriornithine bird
Protopteryx also preserves bastard wing or alula, which is important for
to the ring-necked pheasant. This is an ornithurine bird from the Jiufotang
balance during the slow flight and take-off of birds. It again is a derived
Formation. Long beak, elongated snout and densely distributed teeth are
feature of birds, suggesting the capacity of more skillful flight in this early bird
three of the most marked features of Yanornis. The neck vertebrae are long and
thanPrOIIJPlelp in Archaeopteryx Confuciusornis. al 0 preand erves bastard wing or alula, which i importane for Longipteryx Longipteryx (Fig.take-off 188) is aoflong-winged bird,i which was balance during the low flIght and bird. It again a derived
saddle-shaped, suggesting thatThi Yanornis head and bird neckfrom maneuverability to the ring-necked pheasant. i an has ornithurine the] iufotang
discovered from the ting samethelocality of of Sapeornis in Chaoyang. feacure of bird, u 'ge capacity more killful flight in Its thi forelimb early birdis
claws of aretherelatively short.feature Like modern plover, Yanornis might have spentand most marked of)' 11101711 . The neck vertebrae arc long three
markedly longer thanand theCon/I/cil/somis. hind limb; the ratio of forelimb to hind limb is about than in /t rc!Jfleoplel).\
most of hared, their time on the catching mollusks, fishesmaneu or arthropods addlesu 'J!,e tingwaterside, that )'dnomiJ ha head and neck erability
1.4.Longipteryx About nine cervical vertebrae saddle-shaped vertebral LOlIgipleI)x (Fig. show 1 8) ideveloped a long-winged bird, which wa
as food. The to long and The flexible long neckare arerelatively well adapted thisthe comparable m mouth dern bird. pedal digit long,tobut
comparable to modern The pedal are relatively long, but theare Formation. Lon' beak,birds. elongated snoutdigits and densely di tributed teeth
relativel~'
hort. Like modern plover, )'l1/omis might ha e pene
body. This is anthe indicator of high of agility of thein head and neck. discovered from ame locality apl!onm Chaoyang. It forelimb is
claw are lifestyle.
markedly longer than the ratio processes oHorelimb hindside; limbthis i about Longipteryx hasthe at hind least limb; four uncinate oncoeach is the
most of their time on the water (Figs. ide, catching mollu ,fishe or from arthropod Yixianornis Yixianornis 191, 192) wask collected the
J. first . About nineofcer vertebrae ho\ developed addlehaped vertebral evidence suchical structure in enantiornithine birds. In some modern birds
aJiufotang food. The long mouth and flexible neckCity, are Liaoning well adapted co thi Formation in Yixian County, long Jinzhou Province.
body. is anprocesses indicator high agility the head andofneck. the Thi uncinate areofattached to theofcaudal border the ribs, providing
lifestyle. Compared with
the uncinate proce has e are arrached to the caudal border of the rib, providing Longipteryx long thoracic limb and beak, and it also has strong
Yanornis, Yixianornis has a relatively short head, less teeth in Yixianornis Ylxlel1/ornis (Figs. the jaws, and thinner long bones. Like191, 192) wa collected from the ]iufotang in Yixian ouney, ]inzhou ity, Province. Yanornis, itFormation is an ornithurine bird with '~. . . Liaoning .. ~ pared with )~71lOnm, ha a relatively hort head, Ie teeth in a om well-developed sternalYixianol"flis keel.
connection between neighboring ribs. The reinforced rib provide effecti e
the jaws, and thinner long bone. Like
Lollp,iptel'Yx has at neighboring least four uncinate proreinforced e se on each side; thieffective i the connection between ribs. The ribs provide firstattachment e idence offoruch enaneiornithine bird.and In respiration. ome modern bird sometructure musclesin responsible for flight
perching capability. Therefore, it is suggested that the bird probably had a
arrachmene for ome mu cle re ponsible for flight and respiration.
lifestyle comparable to kingfisher. The discovery of Longipteryx also suggests Longiplel)X has long thoracic limb and beak, and it at 0 ha trong that early enantiornithine birds had undergone a significant radiation in the
perching capability. Therefore, it is sugge ted that the bird probably had a
Early Cretaceous.
lifestyle comparable co kingfi her. The discovery of Longlplel)X al
0
ugge t
Liaoningomis Liaoningornis (Fig. 189) was collected from the Sihetun
that early enaneiornithine bird had undergone a ignificane radiation in the
locality, a site famous for producing the bird Confuciusornis, the feathered
Early
retaceous.
dinosaur Sinornithosaurus and many other important vertebrates. Liaoningornis Liaoningornis Liaoningomis (Fig. I 9) wa collected from the ihetun is the only ornithurine bird known from the Yixian Formation. Among locality, a site famous or produ ing the bird on/I/cil/somis, the feathered ornithurine birds, Liaoningornis is a small-sized bird, comparable to the gray dino aur inol7lilhosallYIIs and many other importane vertebrate. Liaoningornis starling. It has a nearly completely fused tarsometatarsus. It also has sharp is the only ornithurine bird kno\ n from the Yixian Formation. Among
and curved pedal claws, indicating strong perching capability.
ornithurine bird, Linoni/lgomis i a mall- ized bird, comparable to the gray
Another distinctive feature of this bird is the well-developed keel of the
tarling. Ie has a nearly completely fu ed tar ometatarsus. 1t also has harp
sternum, suggesting that Liaoningornis has more power of flight than
and curved pedal claw, indi ating strong perching capability.
e n a n t i o r n i t h i n e s and other basal birds. It is also notable that Another distinctive feacure of thi bird i the well-developed keel of the
Liaoningornis'sternum is thicker than that of Yanornis and Yixianornis, both of ternum, ugge ting that Llooningornis ha more I o\! er of flight than which are more advanced and younger found in western enantiornithine and other ba al ornithurine bird. It birds is alalso 0 notable that Liaoning (seeternum bellow).I The sternum Lia0ning0rnis lateral processes. Liaoningomis' thicker than of that of )'01l0,.1IISalso andlacks }~ixi&l11omis, both of Yanornis 190)ornithurine was derivedbird from an which areYanomis more advanced and(Fig. younger aI "Yan", 0 found name in \ e of tern ancient( country withThe Chaoyang It is a aI largesized bird, comparable ternumthe of capital. Liooningomis 0 lacks lateral proce e. Liaoning ee bellow). Yanornis Yanom/S FIg. 190) \ a derived from "Yan", name of an m 189 Holotype ofLiaoningornis (Slab A), the only ornithurine anciene counery \! ith haoyang the capital.longidigitus It i a large ized bird, comparable bird (size about a sparrow) found in Sihetun locality (lower part of Yixian Formation)in Beipiao, Liaoning. (Photo: IVPP) 189 Holorype of Uaoningornis Jongldigiw ( lab A), the only ornithurine bird ( ize about a sparrow) found in Sihetun locality (lower part ofYixian Formation) in Beipiao, Liaoning. (photo: IVPPj
Yixianornis also preserved wellY01lomis, it i an ornithurine bird with a well-developed
Yixio1l0111/S al
0
(ernal keel.
pr erved well-
147
190 Holotype of Yanornismartini, a large ornithurine bird (size about a ring-necked pheasant), from Dapingfang locality (Jiufotang Formation) in Chaoyang, Liaoning. (Photo: IVPP)
190 Holotype of Yanornis martini, a large ornithurine bird (size abollt a ring-necked pheasant), from Dapingfang locality Uiufotang Formation) in Chaoyang. Liaoning. (Photo: IVPP)
developed uncinate processes, indicating the presence of a strong ribcage.
other hand, it also retains a long pubic symphysis comparable to more
Chaoyangia Chaoyangia 193) was the the pre firstence ornithurine birdribcage. primitiveother birds. hand, it also retains a long pubic ymphysi comparable to more developed uncinate proces(Fig. es, indicating of a trong described from the Jehol Biota. It was (Fig. collected of numerous fossils, as well as a high taxonomic Chaoyangia Chaoyangia 193from wasthe thesame first locality ornithurine birdWith primitive birdwell-preserved . diversity by both very primitive and fos quiteil advanced the taxonomic Boluochia and Cathayornis, Boluochi is anthe incomplete de cribed from the the JehoI Biota. site It ~ofasChaoyang. colleered Itfrom same locality of represented With numerous well-preserved as well forms, as a high Jehol birds have important on and the phylogeny and form, the specimen, with and onlyCathayorniJ, partial postcranial bones preserved. UncinateItprocesses Boillochia the Boluochi site of haoyang. i an incomplete diver ity provided repre enred by both information very primitive quite advanced evolution of early birdshave (Fig.pro 194).ided Though still debatable concerning various are wellpecimen, preserved in the of Chaoyangia. It has least nine sacralproce with onlyholotype partial postcraniaI bone pre aterved. ncinate se Jehol bird important information on the phylogeny and hypotheses, some consensus has (Fig. been reached. vertebrae compared to eight in enantiornithines and sevenJtinhas Confuciusornis. are well preserved in the holorype of Chaoyangia. at least nine sacral evolution of early bird 19 ). Though till debatable concerning various It alsovertebrae has a well-developed cnemial at the proximal and tibiotarsus. the compared co eight crest in enanriornithines seven inOn Conf"ciuJorniJ. hypotheses, some consen us ha been rea hed. It a!
0
has a well-developed cnemial crest at the proxima! tibiotarsus. On the
~ 191
Holorype of Yixianornis grabaui. an ornithurine bird (slightly smaller than Yanornls) with ,,,ell· preserved skeleton and feather inlprints. from
Qianyang locality Uiufotang Fonnation) ill Chaoyang, liaoning. (Photo: IVPP)
~ 192 Wing
~.tI~
. - - - - - - - - - - - - - Archaeopteryx ,.......--------jeholornis ,.....------- Sapeornis ,.....------ Confuciusornis ......- - - Protopteryx ......- - Eoenanliornis Cathayornis Longipteryx Yanornis beishanensis. an Yixianornis
feather of Yixianomi." grabaui, same as
to those of extant flying birds. (Photo: IVPP)
I
j i
11\~1-",9' 1.
I
-
-
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I I
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~193 Holotype of C/rao.yangia ornithurine bird \vith uncinate processes
... '
i
.'
l'
..'# '"
;lor.' jJ'p,J'
(denoted by a red arrow) on ribs and developed cnemial crest (denot~d by a blue arrow) at the proxilnal tibiotarsus. frorn Reish,}n locality Uiufotang fonllation) in Chaoyang. Liaoning. (Photo: IVPP)
Cllaoyangia
Modem birds ~ 194 Cladogram showing the phylogenetic
relationship of the Jehol birds and to Archaeopteryx and modern birds.
//
Yt,t(llT,-qitTg
Yao-min g Hu, Chuan-kui Li Yuall-qing Wang, Yao-lning Hil, Chl1017-klii Li
M
~
M
ammalia is biologically the most differentiated group of
Zhangheotherium quinquecuspidensin 1997. Since then, six other mammalian
ammalia isconsisting biologically thethe most differentiated group of vertebrates, of all living mammals, their quinquecmpiclens in 1997. then, to sixmany otherother mammalian speciesZhangheotherium have been reported from the Jehol Biota. Since Compared vertebrates, consisting all the living mammals, have been reported fromtothe Biota. Compared lineage to many other common ancestor, and their of extinct allies. Mammals have their kinds species of animals in the fauna, especially theJehol feathered dinosaur-bird
their extincr allies.theMammals in the fauna, especially to at theallfeathered predominated thecommon modern ancestor, terrestrial and ecosystems throughout Cenozoic have and a kinds varietyofofanimals invertebrates, mammals were not that rich.dinosaur-bird Despite this, lineage and a variety modern throughout the to Cenozoic of Jehol invertebrates, were notintegrative at ail thatpart rich.ofDespite this, Erapredominated since some 65 the million yearsterrestrial (Myr) ago,ecosystems which is thus often referred as fossil mammals of the Biota aremammals an important and the since some 65 million yearsthe (Myr) which of is mammalian thus often referred as fossilmammalian mammals ofhistory. the Jehol Biota are an important and integrative part of the theEra "Age of Mammals". However, first ago, two-thirds history to Mesozoic The exceptionally well-preserved materials the "Age of Mammals". However,Era thethat fir extended t two-thirds ofroughly mammalian histOry The exceptionally Mesozoic occurred in the preceding Mesozoic from 250 to provide critical mammalian evidence for histOry. understanding the transitionwell-preserved of mammalianmaterials occurred provide critical evidence for understanding the transition of mammalian in before the preceding Mesozoic extended ftOmLate roughly 250 to about 65 Myr present. After theirEra firstthat appearance in the Triassic characteristics and reconstructing the phylogeny of early mammals. These about 65 Myr before present. After their first appearance in the Late Triassic characteristics and reconstructing the phylogeny of early mammals. These (about 220 Myr ago) and throughout the remainder of the Mesozoic, fossils have shown a fairly high diversity of Mammalia in eastern Asia during (about 220 Myr ago) and throughout the remainder of the Mesozoic, fossils have shown a fairly high diversity of Mammalia in eastern Asia during mammals shared the earth with some great reptiles such as dinosaurs. During the Early Cretaceous and represent four major mammalian groups: mammals shared the earth with some great reptile such as dinosaurs. During the Early Cretaceous and represent four major mammalian groups: all that time, mammals were small and scarce, and lived in the shadow of triconodonts, multituberculates, symmetrodonts, and eutherians. In the all that time, mammals were small and scarce, and lived in the shadow of criconodonts, multituberculates, symmetrodonts, and eutherians. In the dinosaurs. Mammals, however, outsmarted dinosaurs by surviving through following paragraphs, they will be briefly introduced in systematical sequence. dinosaurs. Mammals, however, outsmarted dinosaurs by surviving through following paragraphs, they will be briefly introduced in systematical sequence. the mass extinction at the end of the Cretaceous and became dominant on the Triconodonts Triconodonts consist of the most primitive known the mass extinction at the end of the Cretaceous and became dominant on the Triconodonts Triconodonts consist of the most primitive known earth's landmasses afterwards. mammals and some of their derived relatives. As their name implies, their earth's landmasses afterwards. mammals and some of their derived relatives. As their name implies, their Mammals lived in the Mesozoic Era for about 155 Myr, but their fossil basic dental structure comprises three major cusps anteroposteriorly aligned, Mammals lived in the Mesozoic Era for about 155 Myr, but their fossil basic dental structure comprises three major cusps anteroposteriorly aligned, record is rather poor and fragmentary. Unlike the uniform teeth of reptiles, whereas some advanced forms have an enlarged fourth cusp on the distal record is rather poor and fragmentary. Unlike the uniform teeth of reptiles, whereas some advanced forms have an enlarged fourth cusp on the distal mammalian teeth differentiated into four kinds: incisor, canine, premolar, margin. The triconodont lineage spans the entire Mesozoic mammalian margin. The triconodont lineage spans the entire Mesozoic mammalian mammalian teeth differentiated into four kinds: incisor, canine, premolar, and molar, with each kind performing a different function. Based mainly on history from the Late Triassic to Late Cretaceous. Although abundantly history from the Late Triassic to Late Cretaceous. Although abundantly and molar, with each kind performing a different function. Based mainly on
thethe dental morphology, Mesozoic mammals are grouped as Triconodonta, represented by teeth and some and postcranial materials, fully fully dental morphology, Mesozoic mammals are grouped as Triconodonta, represented by teeth and cranial some cranial and postcranial materials, Docodonta, Multituberculata, Symmetrodonta, Eupantotheria, Monotremata, articulated skeleton of triconodonts was yetwas unknown until theuntil discovery of arriculated skeletOn Docodonta, MuJtituberculata, Symmetrodonta, EupantOtheria, Monotremata, of triconodonts yet unknown the discovery of Marsupialia, andand Eutheria. Among them, onlyonly the last groups are still triconodonts in the in Jehol Marsupialia, Eutheria. Among them, the three last three groups are still triconodonts the Biota. Jehol Biota. living on earth today. Before the recovery of mammals from the Jehol Biota, The first triconodont reported from the Jehol The first triconodont reported from theBiota JeholisJeholodensjenkinsi Biota is]eholoclens jenkinsi living on earth tOday. Before the recovery of mammals from the Jehol Biota, tenten localities had been reported to yield Mesozoic mammal fossils in China. (Figs. 195,196). It was represented by a nearly complete skeleton consisting (Figs. 195, 196). It was represented by a nearly complete skeletOn consisting localities had been reported to yield Mesozoic mammal fossils in China. OfOf them, only thethe Early Jurassic Lufeng of Yunnan Province in southwestern of a partial skull skull and the postcranial skeleton preserved as two as twO them, only Early Jurassic Lufeng of Yunnan Province in ourhwestern of a partial andwhole the whole postcranial skeleton preserved China produced some well-preserved specimens, including a number of skulls counterparts and was by Q. Ji others 1999.Jeholodens is China produced some well-preserved specimens, including a number of skulls counterparts anddescribed was described by and Q. Ji and in others in 1999.]eholoclens is distinctive from the mammals in having diagnostic features:features: of of triconodonts, such as Sinoconodon andand Morganucodon. Fossils fromfrom the other distinctive fromother rhe primitive other primitive mammal in having diagnostic triconodonts, such as Sinoconodon Morgctnl/.coclon. Fossils the other dentaldental formula (i.e., number of incisors, canine, canine, premolars, molars on each on each sites were generally represented by fragmentary jaws.jaws. FromFrom the Middle formula (i.e., number of incisors, premolars, molars sites were generally represented by fragmentary the Middle side of theofupper/lower jaw) 4"jaw) 1"2-3/4" 1"2"4; buccolingually compressed Jurassic coal-bed of of northeastern China was was reported the first Chinese side the upper/lower 4·1·2·3/4·1·2-4; buccolingually compressed Jurassic coal-bed northeastern China reported the first Chinese molarsmolars with three a straight alignment, and possessing spoon- spoonMesozoic mammal, Manchurodon. Later, twotwo other mammals, Endotherium with main three cusps main in cusps in a straight alignment, and possessing Mesozoic mammal, Manchlllwlon. Later, other mammals, Endothel·i1lm and Liaotherium, were discovered in that area.area. and LiaotherillJll, were discovered in that
shaped incisors that isthat uniquely derivedderived among among triconodonts. shaped incisors is uniquely criconodonts.
1992, mammal of the Jehol Biota unearthed a small In In 1992, thethe firstfirst mammal of the Jehol Biota was was unearthed near near a small
The articulated type specimen ofJeholodens jenkinsijertkinsi shows a hows mosaica of The articulated type specimen of]eholoclem mo aic of
derived, therian-like characters forparts mostofparts of the shoulder girdle derived, therian-like characters for most the shoulder girdle and the and the illage,Jianshangou Jianshangou western Liaoning Province. lr was named village, of of western Liaoning Province. It was named
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B 195ofjeholodens Holotype ofJeholodensjenkinsi, a triconodont cm), from Sihetun 195 Holorype jenkinsi, a triconodont (skulliengrh(skull aboLitlength 2 em).about from 2Sihetun locality partFormation) of Yixian Formation) in Beipiao,(Photo: Liaoning. CMNH) locality (lower part (lower ofYixian in Beipiao. Liaoning. eM (Photo: H)
196 Reconstruction of jenkinsi. Jeholodensjenkinsi. (Art: Mark A. 196 Reconstruction of jeholodens (Art: Mark A. Klingler/ eMKlingler/CMNH) H)
15l contacting nasals; retainment a large septomaxilla; humeri, but verycharacters primitive characters for thecolumn, vertebralpelvic column, pelvic contacting girdle, the nasals;the retainment of a large of septomaxilla; short and short low and low humeri, but very primitive for the vertebral girdle, sagittal crest; well-developed and theoccipital sloping occipital limbs and Q. Ji and others interpreted mosaic of and crest; sagittal well-developed lambdoid lambdoid crest; andcrest; the sloping hind limbshind and pedes. Q. Jipedes. and others interpreted the mosaicthe of derived andderived surfaceinexposed in dorsal of the skull. features in Jeholodens as either theshoulder derived girdle shoulder and exposed of theview skull. surface dorsal view as either the derived andgirdle primitive primitive features in]eholodens most feature strikingoffeature of these Repenomamus is the forelimbs representing convergence those of multituberculates-therianThe most The striking these Repenomamus specimensspecimens is the forelimbs representing convergence with thosewith of multituberculates-therian presence of abone separate bone to attached to the medial side ofThe dentary. The clade, or the shoulder primitivegirdle shoulder and in forelimbs in monotremes being of a separate attached the medial side of dentary. clade, or the primitive andgirdle forelimbs monorremes being presence is the single lower-jaw all the most extinct atavistictoreversals to the conditions ancestral conditions in distantly the more relared distantly related dentary isdentary the single lower-jaw bone of allbone the of living andliving most and extinct atavistic reversals the ancestral in the more non-mammalian non-mammalian cynodonts.cynodonts.
In the mammallike and mostmammals, primitive mammals, mammals. mammals. In the mammallike reptiles andreptiles most primitive however, however,
the together dentary, with together with postdentary bonesinclude (usuallyarticular, include articular, after the report ofJeholodens, J.-1. and her colleagues the dentary, postdentary bones (usually One year One after year the report of]eholodens, ].-1. Li and herLicolleagues angular, and surangular), the lower jaw. describedtriconodont, another triconodont, Repenomamus robustus, a fairlyprearticular, well- prearticular, angular, and surangular), constitutedconstituted the lower jaw. During theDuring the Repenomamus robustus, based on abased fairly on welldescribed another some associated bones 198).evolutionary early evolutionary stage of mammals, the postdentary bones greatly reduced stage of mammals, preserved preserved skull and skull some and associated postcranialpostcranial bones (Figs. 197,(Figs. 198).197,early the postdentary bones greatly reduced Several additional werefrom collected fromlocality the same locality either into the ear region (becoming the ear Several additional specimens specimens were collected the same after the after and the either and shifted intoshifted the ear region (becoming the ear ossicles andossicles part ofand part of report of this It is the largestknown mammal known from the Mesozoic bulla) toor the fused to the The dentary. The bone peculiar bone on the first reportfirst of this animal. It isanimal. the largest mammal from the Mesozoic bulla) or fused dentary. peculiar on the lower jawlower of jaw of in the 108 in mm in theand holotype andin114 Repenomamus mm in Repenomamus is rodlike, with anterior a pointedtipanterior tip and a flaredend posterior end Era in the Era world. Theworld. skull isThe 108skull mmislong thelong holotype 114 mm is rodlike, with a pointed and a flared posterior another specimen. dentalofformula of Repenomamus robustus is 3" 1.2" 4/3197). Its anterior portioninis alodged in a depression that another specimen. The dentalThe formula Repenomamus robustus is 3·1·2' 4/3' is lodged depression that appears to appears be an to be an (Fig. 197). (Fig. Its anterior portion 1.2.5 198). On molars, its upper onemain of three cusps posterior expandedportion posterior of the meckelian groove.there Because 1· 2· 5 (Figs. 198).(Figs. On its upper themolars, middlethe onemiddle of three cuspsmainexpanded of portion the meckelian groove. Because is no there other is no other large and two the are other two without are weak,distinct without distinct ectocingulum girdleon theit dentary, it can be inferred that all postdentary bones in Repenomamus is large andisthe other weak, ectocingulum (a girdle- (a scar on the scar demary, can be inferred that all postdemary bones in Repenomamus like on structure on labial of the tooth).theBesides the and largethe size and been from detached from theStructures dentary. Structures of the ear of margin the tooth). Besides large size like structure labial margin havethe been have detached the demary. of the ear region andregion their and their dentalRepenomamus features, Repenomamus further othermammals Mesozoic mammals small dimensions indicate that intorelation to of thethe sizes of and the skull and further differs fromdiffers otherfrom Mesozoic in smallindimensions dental features, in relation the sizes skull indicate that the character following combination: character combination: short dorsal of premaxilla not mandible, the ear in Repenomamus beeninreduced of process premaxilla not the following short dorsal process Repenomamus must havemust been have reduced size. in size. mandible, the ear ossicles in ossicles
n 197 Skull (108 mm long) of the holotype of Repenomamus 197 Skull (108 mm long) of the holotype of Repenomal11l1s robustus, the largest Mesozoic mammal to date, in robustlls. the largest Mesozoic mammal to date. in dorsal (Left) and ventral (Right) views, from Lujiatun dorsal (Left) and ventral (Right) views, from Lujiatun locality (lowest Yixian Formation) in Beipiao, Liaoning, locality (lowest Yixian Formation) in Beipiao. Liaoning, red arrow denoting the ossified Meckel s cartilage on red arrow denoting the ossified Meckel s cartilage on the lower jaw. (Photo: Jin Meng/AMNH) the lower jaw. (Photo:Jin MengiAM H) I:
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Therefore, it is reasonable to conclude that the bone attached to the to the Therefore, it i reasonable to conclude thatrodlike the rodlike bone attached dentary is notis any theofpostdentary bones.bones. Embryological studiesstudies of living dentary not of any the po tdentary Embryological of living mammals have have demonstrated the whole processproce fromsthe posterior part ofpart the of the mammals demonstrated the whole from the posterior Meckel's cartilage theossicles, ear ossicles, and anatomical research the Meckel's cartilage to thetoear and anatomical research revealsreveals the Meckel's cartilage an internal the dentary in prenatal Meckel's cartilage lyinglying in an in internal groovegroove of the of dentary in prenatal and and postnatal extant mar upials, monotremes, and eutherians. The of shape of somesome postnatal extant marsupials, monotremes, and eutherians. The shape the separate andrelation it relation the skull in Repenol77alfl/lS are closely the separate bonebone and its to thetoskull in Repenomamus are closely comparable to those the Meckel's cartilage the aforementioned comparable to those of theofMeckel's cartilage in the inaforementioned living living mammals. We identified thi peculiar the ossified of mammals. We identified this peculiar bone bone as the asossified middlemiddle portionportion of the Meckel's cartilage in 2001. This finding, thetime, fir t documented time, documented the Meckel's cartilage in 2001. This finding, for theforfirst the Meckel's cartilage os in ified in fos il record and provided evidence for the Meckel's cartilage ossified fossil record and provided direct direct evidence for its persistence in adults the common ancestor of living mammals. its persistence in adults of theofcommon ancestor of living mammals. bas been strengthened by another kullboth withlower both lower ThisThis has been strengthened yet byyet another partialparcial skull with the arne locality thatpreserves also pre erves the ossified Meckel's cartilage. jawsjaw fromfrom the same locality that also the ossified Meckel's cartilage. specimen represent new pecies of Gobiconodon, a triconodont TheThe specimen represents a newa species of Gobiconodon, also a also triconodont (Fig. (Fig. genu is pbylogenetically clorelated ely related to Repenol77r11f1/1S and has 199).199). ThisThis genus is phylogenetically closely to Repenomamus and has previouknown Iy known the Early Cretaceous ofNorth both America orth America beenbeen previously fromfrom the Early Cretaceous of both and and Irs dental formula i 2·1· ·4/1'1·4'5. In 2003, C.-k. Li and his colleagues Asia.Asia. Its dental formula is 2" 1.4.4/1" 1"4"5. In 2003, C.-k. Li and his colleagues named it GobiCO/loclon zofiae. 198 Skull of Repenomamus robustlJs in anterior (Upper) and lateral views (Lower).named it Gobiconodonzofiae. Discovery of the os ified Meckel's cartilage helps interpret the function 198 SkullofRepenomarnus robustus in anterior (Upper) and lateral views (Lower), Discovery of the ossified Meckel's cartilage helps interpret the function showing the tooth morphology. (Photo: Jin Mengl AM H) showing the tooth morphology. (Photo: Jin Meng/AMNH)
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of the internal groove on dentaries of many early mammals. The groove has been variably of the internal groove on dentaries many and/or considered as holding dentalofnerves early mammals. The groove has been variably arteries, or the Meckel's cartilage, or even as considered as holding dental nerves and/or facets for the postdentary bones. Because of the arteries, or the Meckel's cartilage, or even as presence of such a groove in a number of facets for the postdentary bones. Because of the phylogenetically derived mammalian forms, presence of such a groove in a number of the interpretation for the persistence of the phylogenetically derived mammalian forms, postdentary bones in these taxa argues for the interpretation for the persistence of the multiple origins of the definitive triossicular postdentary bones in these taxa argues for mammalian middle ear. The discovery of the multiple origins of the definitive triossicular ossified Meckel's cartilage from the internal mammalian middle ear. The discovery of the groove of the dentary in Repenomamus and os ified Meckel's cartilage from the internal Gobiconodon clearly shows the function of the groove of the dentary in RepenomamltS and groove as holding the cartilage. This in turn Gobiconodon clearly shows the function of the suggests early mammals a Meckel's groove as holdingthat the in cartilage. This in turn whether ossified aorMeckel's not, should be suggestscartilage, that in early mammals as a potential, even be primary, cartilage,considered whether ossified or not, or should occupant for the internal groove. Thus, this considered as a potential, or even primary, no longer callsThus, for multiple orioccupantextrapolation for the internal groove. this gins ofnothelonger triossicular mammalian ear. extrapolation calls for multiple middle ori-
Multituberculates Multituberculates first mammalian middle ear. gins of the triossicular appeared in theMultiruberculates latest Triassic and Multituberculates firstbecame the lateTriassic Eocene and of thebecame Cenozoic Era. appearedextinct in theinlatest are Eocene frequently "rodents extinct inThey the late of thecalled Cenozoic Era. of the duecalled to their"rodents superficially rodentlike They areMesozoic", frequently of the features, such as the pair rodentlike of enlarged and Mesozoic", due to their uperficially lower In laterand forms, the features,procumbent such as the pairincisors. of enlarged first lower pair ofinci lower incisors is chisellike procumbent ors. In later forms, theand only fir t pair the of lower incisors chisellike and onlywhich is ventral side is iscovered by enamel, is But, the ventral side is covered especially similar by to enamel, those ofwhich rodents. especially similar to those of rodents. usually But, have unlike rodents, multituberculates unlike rodents, multiruberculates usually In have more than one pair of incisors. the midmore than one pair of incisors. In the mid-of most portion of the lower dentitions portion multituberculates, of the lower dentitions of modified most into premolars are
m 199 Holotype of Gobiconodonzofiae (skull length 45 mm), showing cranium in lateral (Upper) and ventral (Middle) multituberculates, premolarsTheir are modified intovarying _ 199 Holotype of Gobiconodon zojiae (skulilengrh 45 mm). in lateral (Upper) andYixian ventral (Middle) in bladelike structure. molars have views and lower jaw in labial and lingual views showing (Lower), cranium from Lujiatun locality (lowest Formation) lowerLiaoning, jaw in labial and lingual views Lujiatun locality (lowest Yixian Formation) in views and bladelikenumber structure. Their molars havethan varying Beipiao, red arrow denoting the(Lower), ossified from Meckers cartilage. (Photo: IVPP) (from several to more a dozen) of number (from several
to
more than a dozen) of
Beipiao, Liaoning, red arrow denoting the ossified Meckel's cartilage. (Photo: IVPP)
~200
Holotype of Sinobaatar lingyuanensis (Slab A, skull length 26.6 mm), a multituberculate from Dawangzhangzi locality (middle 200 Holorype of Sinobaalar Jingyuanensis (Slab A. skull length 26.6 partmm), of Yixian Formation)in Lingyuan, Liaoning. (Photo: IVPP) a multiruberculate from Dawan zhangzi localiry (middle part of ixian Formation) in Lingyuan. Liaoning. (Photo: IVPP)
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201 A Abladelike bladelikepremolar premolaron onthe thelower lower jawofof Sinobaatar m 201 law Sinobaatar Jingyuanen is (ea t). (Photo: Jin Mengl AM lingyuanensis {cast). {Photo: Jin Meng/AMNH)H)
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203 The upper and lower tooth rows and the lower jaw ofZhangheotherium m 203 The upper and lower tooth rows and the lower jaw of Zhangheotherium quinquecu pidens.
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204 Reconstruction of the ear region of Zhangheotherium quinquecuspidens,
~204
showing the finger-shaped promontorium (denoted by a red arrow) that Reconstruction of the ear region ofZhangheotherium quinquecuspidens, houses the cochlea, implying its auditory system not very sensitive to showing the finger-shaped promontorium (denoted by a red arrow) that high frequency sound.
houses the cochlea, implying its auditory system not very sensitive to high frequency sound.
the on buccal side on The the lowers. earliest symmetrodonts tuberclelike lowwhich cusps, they fromderived which they their name. Distinctuppers, changeswhileuppers, tuberclelike low cusps, from theirderived name. Distinct changes at the while buccalatside the lowers. earliest The symmetrodonts in the latest Triassic to earliest Jurassic of Wales, while their latest in morphology their dental morphology time are characterized by the in inwere in their dental through timethrough are characterized were found the found latest Triassic to earliest Jurassic by the decrease in decrease of Wales, while their latest record from the Late of Cretaceous of North America. thepremolars, number ofthe premolars, thethe increase in of theserrations number of serrations ridges the number of increase in number and ridges and North America. record was from thewas Late Cretaceous Y.-m. and his reported colleaguesa symmetrodont reported a symmetrodont on the last lower and in the number of molar cusps. Y.-m. Hu and of molar cusps. on the bladelike lastbladelike lower premolar andpremolar in the number his Hu colleagues mammal, mammal,
quinquecuspidens, fromBiota the Jehol Biota in 1997 (Figs. date, onlyofone species of multituberculates has been recovered from Zhangheotherium multituberculates has been recovered from To date, onlyTo one species Zhangheotherium quinquecuspidens, from the Jehol in 1997 (Figs. 202"-'205). is the first mammal from and is represented the Jehol Y.-m. HuWang, and Y.-q. Wang, in 2002, named it Sinobaatar the Jehol Biota. Y-m.Biota. Hu and Y-g. in 2002, named it Sinobaatar 202-205). It is the first Itmammal known from known the Biota andtheis Biota represented by a well-preserved skeletonofconsisting of a partial skullofand (Fig.type 200). The type specimena represents a subadult. Its abody lingyuanensis lingyuanensis (Fig. 200). The specimen represents subadult. Its body a partial skull and most the most of the by well-preserved skeleton consisting postcranial skeleton, all cervical thoracicforelimbs vertebrae, forelimbs from the tiptoofthe rostrum the hip, about 10.3 cm. Its dentalpostcranial formula skeleton, including allincluding cervical and thoracic and vertebrae, length, from length, the tip of rostrum hip, is to about 10.3isem. Its dental formula pectoral girdle, as well impressions as excellent of impressions of the pelvic hind limbs, pelvic is 3"?'5"2/1"0"3"2. Differing from later multituberculates, the first as well as excellent the hind limbs, andlower pectoral and girdle, is 3' ?'5' 2/1,0-3-2. Differing from later multituberculates, the first lower girdle, ribs. The dental formula of Zhangheotherium is 3" 1"2-5/3-1.2.6 (Fig. incisor ofisSinobaatar is conical and completely covered by The of Zhangheotherium is 3·1·2'5/3·1·2·6 (Fig. girdle, and ribs. Theand dentalformula conical and completely covered by enamel. Theenamel. incisor of Sinobaatar 203). isThis animal isbydiagnosed by having blunt trigonid cusps, weak cristids, bladelike middle of the lower dentition three premolars 203).(Fig. This animal diagnosed having blunt trigonid cusps, weak cristids, bladelike middle portion of theportion lower dentition involves threeinvolves premolars (Fig. large cingulidand cuspules, andand nolingual labial and lingualBefore cingulids. 201). the formula same dental formula and similar dental morphology, large cingulid cuspules, no labial cingulids. the Before the 201). Sinobaatar hasSinobaatar the same has dental as and similarasdental morphology, of Zhangheotherium, symmetrodonts were only from the especially of the to cheek to Early that ofCretaceous some Earlymultituberculates Cretaceous multituberculates discovery ofdiscovery Zhangheotherium, symmetrodonts were only known fromknown the cheek teeth, that teeth, ofsome especially ofthe from Mongolia. from Mongolia.
fragments of lower upper jaws. and lower jaws. The nearlyskeleton completeof skeleton of fragments of upper and The nearly complete
some valuable morphological for this holotype of Sinobaatar lingyuanensis is the most complete Zhangheotherium pre-Late Zhangheotherium reveals somereveals valuable morphological information information for this The holotypeThe of Sinobaatar lingyuanensis is the most complete pre-Late Cretaceous multituberculate It also new morphological Cretaceous multituberculate specimen. It specimen. also provides newprovides morphological group.
group.
of postcranial skeleton previously unknown One of the most cranial interesting cranial featurestoisauditory related to auditory informationinformation of postcranial skeleton previously unknown for early for early One of the most interesting features is related multituberculates. In the contrast to the changes significant changes of their dental Although apparatus.fractured Althoughand fractured in preservation, multituberculates. In contrast to significant of their dental apparatus. distortedand in distorted preservation, the petrosal the petrosal available information change in the postcranial bone of Zhangheotherium is complete. Its promontorium, which the cochlea features, thefeatures, availablethe information shows little shows changelittle in the postcranial bone of Zhangheotherium is complete. Its promontorium, in which the in cochlea of multituberculates during their This may that hasisahoused, has and a cylindrical fingerlike shapesimilar (Fig. 204), similar morphology morphology of multituberculates during their history. This history. may suggest that suggest is housed, cylindrical fingerlikeand shape (Fig. 204), to those of to those of the patterns locomotory patternsmultituberculates in different multituberculates didmuch not change much triconodonts and multituberculates. kind of promontorium is closely the locomotory in different did not change triconodonts and multituberculates. This kind ofThis promontorium is closely duringTheir evolution. Their morphologically generalized skeletons postcranial skeletons correlated either straightcurved or slightly curved (but uncoiled) during evolution. morphologically generalized postcranial correlated with either awith straight or aslightly (but uncoiled) cochlea, a cochlea, a theAninner ear. An uncoiled was, therefore, thatonthey lived on a variety of substrata. they lived a variety of substrata. indicate thatindicate spiral-shapedspiral-shaped cavity of thecavity inner of ear. uncoiled cochlea was,cochlea therefore, specimen of far reveals most morphological informa- for inferred for Zhangheotherium. that the hearing of Sinobaatar bySinobaatar far revealsby most morphological informa- inferred The specimenThe Zhangheotherium. This means This that means the hearing function of function of tion on multituberculate hand.structure The wristof structure is very Zhangheotherium was not as well-developed as more in most of more derived tion on multituberculate hand. The wrist SinobaatarofisSinobaatar very Zhangheotherium was not as well-developed as in most of derived to that ofKrypt0baatar, a Latemultituberculate Cretaceous multituberculate similar to thatsimilar of Kryptobaatar, a Late Cretaceous ofMongolia, of Mongolia, mammals. mammals. and that ofZhangheotherium, a symmetrodont from the Sinobaatar Jehol Biota. Sinobaatar shoulder girdle andofforelimbs of Zhangheotherium and that of Zhangheotherium, a symmetrodont from the Jehol Biota. The shoulderThe girdle and forelimbs Zhangheotherium are the best-are the besthas well-preserved feet and resembles Ptilodus,multituberculate a Paleocene multituberculate and most parts informative parts in itsskeleton. postcranial has well-preserved feet and resembles Ptilodus, a Paleocene preserved andpreserved most informative in its postcranial Theskeleton. clavicle The clavicle of North in having large tibial and asymmetrical but of Zhangheotherium rodlike and in similar to that ofand marsupials and of North America, in America, having a large tibiala condyle, andcondyle, asymmetrical but of Zhangheotherium is rodlike andis in similar shape to thatshape of marsupials widely movable astragalotibial joint, on etc.theBased foot and ankle But, placentals. But, unlikeand marsupials anda placentals, a separate interclavicle is widely movable astragalotibial joint, etc. Based foot on andthe ankle placentals. unlike marsupials placentals, separate interclavicle is structure, long andtails, prehensile tails, F. Jr. A. Jenkins, Jr. and D. W.anterior structure, plus present anterior to the sternum (breastbone) the longplus andthe prehensile F. A. Jenkins, and D. W. present to the sternum (breastbone) and betweenand twobetween clavicles.two In clavicles. In Krause reconstructed an arboreal animal, a lifestyle may also and marsupials andinterclavicle placentals, interclavicle does exist asbone, a separate bone, Krause reconstructed Ptilodus as anPtilodus arborealasanimal, a lifestyle that may alsothat marsupials placentals, does not exist as anot separate shared by Sinobaatar. Sinobaatar. be shared bybe it in is very large in monotremes and acts whereas it is whereas very large monotremes (e.g. platypus(e.g. andplatypus echidnas)and andechidnas) acts Symmetrodonts Symmetrodonts were mammals, shrew-sizedcharacmammals, charac- as primary supporting structure. Unlike monotremes where the clavicleSymmetrodonts Symmetrodonts were shrew-sized as primary supporting structure. Unlike monotremes where the clavicleterized by the triangular, imperfectly symmetrical cusp arrangement of botfi interclavicle joint is fixed, Zhangheotheriumhas a mobile clavicle-interclavicle terized by the triangular, imperfectly symmetrical cusp arrangement of both interclavicle joint is fixed, Zhangheotherium has a mobile clavicle-interclavicle uppermolars. and lower The of orientation of the the upper and upper and lower The molars. orientation the triangles on triangles the upperon and joint, which allowstothemove clavicle moverange in a and widerfurther range results and further results joint, which allows the clavicle in a to wider molars- is the reversed--the is at side the lingual on the in free movement of forelimbs. is reversed central cusp central is at thecusp lingual on the side lower molarslower in free movement of forelimbs.
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m 206 Holotype of Eomaiascansoria(Slab A, skull length about 3 cm)0an eutherian, from Dawangzhangzi locality (middle part of Yixian Formation) in Lingyuan, Liaoning. 206 Holotype of Eomaia can aria (Slab A. sklllliength about 3 em), an elltherian, from Dawangzhangzi locality (middle parr of Yixian Formation) in Lingyuan. Liaoning. (Photo: CMNH) (Photo: CMNH)
In extant land mammals, marsupials and placentals move in a parasagittal probably suggests that Zhangheotherium has a forelimb posture intermediate In extant land mammals, marsupials and placentals move in a parasagittal probably suggests that Zhangheotherium has a forelimb posture intermediate between that of the sprawling monotremes and that of the parasagittal posture, i.e., the fore and hind limbs straight under the body during between that of the sprawling monotremes and that of the parasagittal posture, i.e., the fore and hind limbs straight under the body during therians. movement, while monotremes in a sprawling posture with legs extending therians. movement, while monotremes in a sprawling posture with legs extending Early in 2003, G. W. Rougier and others reported another symmetrodont, laterally in certain extent. Comparative anatomical study shows that different Early in 2003, G. W. Rougier and others reported another symmetrodont, laterally in certain extent. Comparative anatomical study shows that different postures are related to morphology of limb bones. The forelimbs of Maotherium sinensis, from the ]ehol Biota. It is represented by a fully postures are related to morphology of limb bones. The forelimbs of Maotherium sinensis, from the Jehol Biota. It is represented by a fully Zhangheotherium have some features similar to those of therian mammals with articulated skeleton and closely related to Zhangheotherium. Although Zhangheotherium have some features similar to those of therian mammals with articulated skeleton and closely related to Zhangheotherium. Although differing in some details, they share many morphological features of cranium, parasagittal posture. However, they also retained some primitive characters differing in some details, they share many morphological features of cranium, parasagittal posture. However, they also retained some primitive characters dentition, and postcranial skeleton, and thus are grouped into the same that are common in the known non-therian mammals having a sprawling dentition, and postcranial skeleton, and thus are grouped into the same that are common in the known non-therian mammals having a sprawling family. posture. The combination of both primitive and derived features of forelimbs family. posture. The combination of both primitive and derived features of forelimbs
207 Recon truction of Comaia can aria. (Art:
ark A. Klin lerl M H).
207 Reconstruction ofEomaia scansoria. (Art: Mark A. Klingler/CMNH).
Eutherians Eutherian are a roup of mammals on i ting of extant placental
Tritylodontidae
nd Eutherians their extinct cl e relative . Theyof mammals h re the mo t recent Eutherians are a group consisting of extant
comm n placentals ance cor with . The r of mol di ringui he recent and mar their upial extinct closenumb relatives. Theyr share the most eutherians (three molar on e ch ide for both upper and lower dentition) common ancestor with marsupials. The number of molars distinguishes fr m mar upiaJ (; ur up r and four low r molar . The arli t record w eutherians (three molars on each side for both upper and lower dentitions) previou Iy d cumented in the lac arly retace u (ca. 110 Myr a 0). In from marsupials (four upper and four lower molars). The earliest record was 2002, Q. Ji and his colla rae rs reponed an eutheri 0 m mmal, Eomllia previously documented in the late Early Cretaceous (ca. 110 Myr ago). In scans(),-ia, fr m th Jehol Bioea (i .206,207). Ie is ehe arli St r pr m tiv 2002, Q. Ji and his collaborators reported an eutherian mammal, Eomaia of eurheri os 0 far kn wn in th rid. The animal h peciaJized limb nd scansoria, from the Jehol Biota (Figs. 206, 207). It is the earliest representative fi Ot featuI that Ie oly known fr m an orial (climbing) and arb real of eutherians so far known in the world. The animal has specialized limb and (cree-dwellin ) exeant mammal , quite differ m from the eerre trial or foot features that are only known from scansorial (climbing) and arboreal cur orial cunnin ) £i acur f other retaceou eueherians. This u rs chat (tree-dwelling) extant mammals, quite different from the terrestrial or the earlie t eurh rian lineage d el ped different I m cory adaptation , cursorial (running) features of other Cretaceous eutherians. This suggests that fa ilieating their prea co dl r e ecolo ical niche in the reeaceous. earliest different The the ph 10 enecieutherian r laeion lineages hip f e developed ely mammal re oflocomotory great inter adaptations, r co
Tritylodontidae 5inoconodon Sinoconodon Morganucodon Morganucodon Ha/danodon
'-
Haldanodon Hadrocodium
Hadrocodium Jeh%dell
l
Priacodon Jeholodens Gobiconodoll Priacodon
~H
RepenomalTlu Gobiconodon
facilitating their spread tofodiverse niches in are the either Cretaceous. pal omammaJogists. ammalian iJ fr ecological m the J hoI Biota the phylogenetic of early within mammals are re of great interest to be c-pre erved The or ehe eaelie t crelationships urred pccimen cheir pective
Monotremata Repenomamus (e.g., Sinobaatar) Multituberculate Monotremata
paleomammalogists. from Jehol Biota aretheither the roup. Therefore, th pro Mammalian a new in i fossils he into th the relation hip of or the within kelecon. their respective major linebest-preserved e of mammal ndearliest the ev occurred lution of specimens the mammalian Differ or phylogen eic an I they provide a new insight into • butthe therelationships r ult ar of the group. Therefore, the e of olutionary tr nd skeleton. f n t eeongly onfli t d with h oth and r andthehow major lineages of mammals evolution the mammalian
ZhangheotileriulTl Multituberculate (e.g., Sinobaatar)
l
t
om m jor mammali n ch raceer . fr mew rk f th but e r theultresults are Different phylogenetic analysis shows some uncertainties, ),not ie strongly eem afeconflicted to make with the folio in on ion:the (1)evolutionary The ab encetrends of each other andlushow othtrillmcharacters. i best conGiven id c dthe primici e, i.e., ofcoil d results in Zhangh some major mammalian framework these c chle y (Fig. t to be ev itIved. The r athe ittfollowing I fi relimb p cure of(1)Ii The in absence 208), seems2safe to pmake conclusions: th nOt pre em in monotreme, multiruber ulate, nd man of a coiled cochlea in Zhangheotheriumis best considered primitive, i.e., a coiled m mmal u has Zhllngheorh rimn, Hmkelofh -illm and juce/estes. Ie cochlea yet to be evolved. (2) The parasagittal forelimb posture of living r babl ar e in later ta e f m. mmali n e luti n. ( ) The pr enc of therians was not present in monotremes, multituberculates, and many definitive rri icul r mamm lian middle e r in R 'Penofnafnll nd Gobicono Ion Mesozoic mammals such as Zhangheotherium,Henkelotheriumand Vincelestes.It provid e id 0 e fl r it id di tribud n in rh mammals. probably arose in later stage of mammalian evolution. (3) The presence of
definitive triossicular mammalian middle ear in Repenomamusand Gobiconodon provides evidence for its wide distribution in the mammals.
MaotheriulTl Zhangheotherium Henke/otherium Maotherium Vincelestes Henkelotherium
1 208 Clad
Metath ria Vincelestes Eutheria (e.g.• [omaial
t4
Metatheria
ram h wing mammalian phyl gen tic
relation hip ,combining reEutheria ult of differ analy(e.g., or Eomaia) e r lated to the Jehol mammals (in red).
/~208 Cladogram showing mammalian phylogenetic relationships, combining results of different analyses related to the Jehol mammals (in red).
Qiff i wang, HI,t[-HTH. Lu, jlin.g-lin Yci,ng Qi-.fei \Vallg. Hili-nan Lu, ling-lin Yang ~ , ~ h e charophytes (or stoneworts), a kind of algae, grow in fresh or
T
Ihc
lit
of 18 m, compact or bushy. The structure of charophytes is relatively simple
he charophytes stonewores), kind ofponds, algae, lakes, growlagoons, in fre hfiords or and of 1 m, compact ororgans, bushy.with The no strucrure charophytes is relatively simple brackish(or waters including astreams, in vegetative vascularofbundles. However, their reproductive brackish have waters including streams, ponds, lakes, lagoons, fjords and except in vegetative with no vascular a wide geographical distribution on all continents organs organ, are rather complex. Peoplebundle. usually However, place themtheir in a reproductive separate division geographical distribution on all exceptand rich have a wide organsofare usually place them in a separate division Antarctica. Most of them live in the waters that are continents clear, transparent therather plant complex. kingdom, People the Charophyta.
Antarctica. of them livewith in the thatfrom are clear, and rich highof the planeThe kingdom, the Charophyta. in Most calcium, mostly pHwaters ranging 7.0 totransparene 8.0, e.g., alkaline, charophyte thallus is green, ranging from 0.5 to 200 cm in height mostly with pH ranging from 7.0 to 8.0, e.g. alkaline, highin calcium, The charophyte thallus is green, ranginghas from to 200 cm axis in height calcium lakes. Charophytes are present in running water only where move(15---25 cm in average), and usually an 0.5 erect, central or stalk, calcium lakes. Charophytes are presene in running water only where move(15-25 cm in average), and usually has an erect, cenrral axis or stalk, ment is slight. They grow submerged in the water from shorewards to a depth regularly whorled branches arising from the very short nodal cells and rhizoids ment is slight. They grow submerged in the water from shorewards co a depth regularly whorled branches arising from the very shore nodal cells and rhizoids at the base to attach to the muddy or sandy substrate (Fig. 209). The plant at the base to attach to the muddy or sandy substrate (Fig. 209). The plane cells contain small and elliptic chloroplasts that carry out photosynthesis and cells contain small and ellipric chloroplasts that carry out photosynehesis and convert inorganic compounds to organic nutrients. The plants commonly are convert inorganic compounds co organic nutrienes. The plants commonly are monoecoius, i.e., a single plant producing both types of gametes. Both male monoecoius, i.e., a single plane producing borh types of gametes. Both male gametes (antheridia) and female gametes (oogonia) occur on the branches. gametes (aneheridia) and female gametes (oogonia) occur on the branches. Oogonia in living charophytes are elliptic to spherical, covered with five Oogonia in living charophytes are elliptic to spherical covered with five .. "@]:; sinistral enveloping cells, whereas antheridia are spherical, protected with !,iJ l/;.~:i.., h sinistral enveloping cells, whereas antheridia are spherical, prorected with four to eight shield cells. Charophytes reproduce both vegetatively and four to eight shield cells. Charophytes reproduce both vegetarively and sexually. Vegetative reproduction may occur by means of bulbils or fragments sexually. Vegetative reproducrion may occur by means ofbulbils or fragmenes ......
.....
4 _ _-+
J
~ ~ 2
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from mature Greater or lesser of calcification of calcification takestakes placesplaces in thein the from mature plane. plant. Greater or lesser degreedegree plant during the growth of charophytes, especially the enveloping of charophyres, especially withinwithin the enveloping cells cells plane during the growth
r~
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of oogonia, the preservation good preservation the shells. lime shells. The term of oogonia, which which enablesenables the good of theoflime The term gyrogonite is commonly used to designate the fossilized oogonium. is commonly used to designate the fossilized oogonium. gyrogonite Most charophyte are remains of gyrogonites as shown in the Most charophyte fossils fossils are remains of gyrogonites as shown in the (Figs. 210---223). The geological occurrence of fossil charophytes figures figures (Figs. 210-223). The geological occurrence offossil charophytes may may tracetoback the Late Silurian, 400 million ago. Quite the to Late ilurian, nearlynearly 400 million years years ago. Quite a fewa few trace back havegeographic wide geographic but limited geologic but limited geologic range,range, whichwhich allowallow their their species species have wide indicesindices for freshwater and brackish sedimenes as well as as use as stratigraphic use as stratigraphic for freshwater and brackish sediments as well tools intools environment reconstruction. The diagnoses in fossil charophytes ' in environment reconstruction. The diagnoses in fossil charophytes' the size of gyrogonites, classification and identification include classification and identification include theand sizeshape and shape of gyrogonites, characters of calcification and utricle, apicalapical structures and depressions, characters of calcification and utricle, structures and depressions,
and coronular cells, etc. rosettesrosettes and coronular cells, etc. 209 Sketch showing living charophyte plant (Left. x I;(Left, I, rhizoid; 209map Sketch mapashowing a living charophyte plant x 1" 1, rhizoid; The Jurassic charophytes are ab are eneabsent in oreheast China.China. 0 charophytes The Jurassic charophytes in Northeast No charophytes 2, axis; 3.2,branch) its reproductive organs (Right, 40; 4, × 40; 4, axis. 3,and branch) and its reproductive organs x (Right, have been reported from the upper part of the Yixian Formation and antheridium; 5, oogonium; 6, coronular cells). (After Nordstedt, antheridium. 5, oogonium; 6, coronular cells). (After Nordstedt, have been reported from the upper part of the Yixian Formation the andlowet the lower 1891; redrawn Yu-gao by Ren/ NIGP)Ren/NIGP) 1891.byredrawn Yu-gao the Jiufotang Formation at present. The fossil charophytes obtained pare ofpart of the Jiufotang Formation at present. The fossil charophytes obtained
iii
from the lower part of the ixian Formation of northern Heb i and,; e tern from the lower part of the Yixian Formation of northern Hebei and western Liaonin are predominant \ irh Mesochartl, Peckisphaerc/ and Mirlbechara. Liaoning are predominant with Mesochara, Peckisphaera and Minhechara. The upper part of the Jiufotang Formation to the Fuxin Formation The upper part of the Jiufotang Formation to the Fuxin Formation yields rich char phyte including mainly Flabellorhara bebeie1/Sis F. barrisi, yields rich charophytes, including mainly Flabellochara hebeiensis, F. harrisi, Il/opocbam tril'OIvis rriqllelm Aclistochara 'flllIl IlIltl A. hllihllibaoensis, Mesochara
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Atopochara triquetra, Aclistochara mundula,The A. huihuibaoensis, Mesochara in the sripitata,trivolvis 'f. voillta, Peckisphaera verticil/ara. e specie are common stipitata, voluta, Peckisphaera verticillata. These are commonhina. in the middlM.part of the Lo~ er Cretaceous d po species it in northern Among
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middle of the Lower Cretaceous deposits in trit'ol northern China. Among them,part Flabellochara beheiemis and Atoporhara is triquetra have pecial them, Flabellochara hebeiensis i.e., and bAtopochara trivolvis triquetra have in special tratigraphic ignificance, th are typical to the Barremian the world. stratigraphic significance, i.e.,Jiufotang, both are typical to the in the world. an be harophytes from the hahai andBarremian Fuxin Formation included infrom the the third la atoraShahai eae Biozone of Formations hina, namely Charophytes Jiufotang, and Fuxin can the be late Barremian Afoporhara trivo/t'is rriqllerra-Flabel/ochartl ieusis Biozone. included in the third Clavatoraceae Biozone of China,heb namely the late
Cretaceous.
Cretaceous. M rochm'a prodllcta Fig. 216): gyr gonite ooidal, apex br adly round According to trivolvis the charophyte di tributionhebeiensis the geologi al age f the Barremian Atopochara triquetra-Flabellochara Biozone. producta and (Fig. forming 216): gyrogonites ovoidal, round, ba e contracted a taLklike projeapex tion,broadly 250-3 0 J..lrn long, lower part of to thethe Yixian Formation can be attributed to the According charophyte distribution, the geological age early of the Early Mesochara base contracted and wide forming-9 a stalklike projection, Cretaceou the Yixian upper pan of the Jiufotang Formati to n as as Early the hahai 200-250 J..lm conv lution vi ible 250N380/am in lateral view'long, found in the lower part of. the Formation can be attributed thewell early 200---250 wide, 8N9ofconvolutions visible in lateral view; found in the and Fuxin Formation are regarded as the I te Barremian. Lower~m Cretaceous northern hina. Cretaceous; the upper part of the Jiufotang Formation as well as the Shahai The figure h '; some fos it charophyte from Inner Mongolia, HebLower Cretaceous Peckisphaera IIIlIltispira (Fig. 217): gyrogonite broadly ovoidal apex of northern China. and Fuxin Formations are regarded as the late Barremian. and Li oning province . Their diagno e are briefly tated as folJow . round, ba e multispira narrowly (Fig. round217)" or contracted a stalk ovoidal, 3 0- apex 0 J..lrn long, Peckisphaera gyrogonitestobroadly The figures show some fossil charophytes from Inner Mongolia, Hebei ~ wide, 10convolution isible300N400 in lateral view; found in the Aclistochal'a IJllihllibaoemis (Fig. 210): gyrogonites 0 al to prolate, round,200-300 base narrowly round or L1 contracted to a stalk, lam long, and Liaoning provinces. Their diagnoses are briefly stated as follows. u Nof Mongolia and inHebei. o -550 ~m long, 350-5 0 ~m wide, -9 convolution vi ible in lateral200 NLower 300 gtmreta wide,e 10 11Inner convolutions visible lateral view; found in the Aclistochara huihuibaoensis (Fig. 210)" gyrogonites oval to prolate, view piral cell c ncave intercellular ridge narrow and harp; found from Peckisphaera paragranlllifera (Fig. 21 ): gyrogonite ovoidal or 400N550 ~m long, 350N500 ~m wide, 8N9 convolutions visible in lateral Lower Cretaceous of Inner Mongolia and Hebei. the Lower retaceou of hina. ub pheroidal, apex round, ba e r und and protruding, ba al end flat Peckisphaera paragranulifera (Fig. 218)" gyrogonites ovoidal or view, spiral cells concave, intercellular ridges narrow and sharp; found from Aclislochara IIIlIndllla (Fig. 2 LL): gyro onite urn-shaped, 370-600 ~1 50-600 ll-m long, 50-65 ll-m wide 11-12 convolution visible in subspheroidal, apex round, base round and protruding, basal end flat, the Lower Cretaceous China. lateral view' found in the Lower retaceous of hina. long, 25000 ~ of wide, 10-12 convolution isible in lateral view; found 450N600 gtm long, 350N465 ~m wide, 11N12 convolutions visible in Aclistochara mundula (Fig. 211)" gyrogonites urn-shaped, 370N600 ~m from the Lower retaceou 0 er th world. Atopoche/ra trit'ol is triquetra (Fig. 2 L9-2_1): urricles _0-10 0 llm lateral view; found in the Lower Cretaceous of China. long, 250N400 ~tm wide, 10 N 12 convolutions visible in lateral view; found P rkispha ra verticillaltl (Fig. 212, -13): gyrogonite pheroidal, apex ng, 750-1000 J..lm wide, Ii rmed with 36 units" hich are divided into three Atopochara trivolvis triquetra (Figs. 219 N 221): utricles 820 N 1080 gm from the Lower over the round, world. 50-750 llm long 00-600 ~m group, ba al vie, rriangular; di rributed in the Barremian over th world. br adly round Cretaceous and ba e narrowly long, 750 N 1000 lam wide, formed with 36 units, which are divided into three verticillata 212, in 213): gyrogonites spheroidal, wide,Peckisphaera 10- Leon olution(Figs.i ible lateral view' found in theapex Lower Flabello ham bth iensis Fig. 222): utricle ov idal 620- 00 ll-m long, groups, basal view triangular; distributed in the Barremian over the world. broadly roundover andthe baseworld. narrowly round, 450N750 l~m long, 400N600 ~m retaceou 520-700 llm wide, lateral unit con i t of5 or 6 ceLIs' found in the Barremian Flabellochara hebeiensis (Fig. 222): utricles ovoidal, 620N800 ~tm long, wide, Mesochara 10N12 convolutions view; found the ooidal Lower to over the world. xlla1lziemis visible (Fi . 21in ):lateral gyr gonite very inmalJ 520N700 ~m wide, lateral units consist of 5 or 6 cells; found in the Barremian 1inhechara sp. (Fig. 22 ): gyro onite ub pheroidal, apex and base ubovoidal,over 230-350 Cretaceous the world.J..lm long, 200-25 llm wide, 6- convolution over the world. Mesochara xuanziensis (Fig.in214)" gyrogonites veryussmall, ovoidalhaanxi, to vi ible in lateral view' found the Lower Cretace f Hebei, round, apical cemer pr jected, forming columeUa 750-1050 J..lm long Minhechara sp. (Fig. 223): gyrogonites subspheroidal, apex and base Anhui, Henan, ansu and hina; northern north we tern 50-610 J..lffi wide, 10-11 convolution visible in lateral iew; found in the subovoidal, 230N350 ~m Xinjiang long, 200~250 ~tm wide, pain 6N8 and convolutions round, center projected, forming a columella, 750 N 1050 gtm long, ermany in Eur view; pe. found in the Lower Cretaceous of Hebei, Shaanxi, Lo apical er reta eous of Lia ning and Qinghai province. visible in lateral ~mfowide, visible found the il 10 h N wn11inconvolutions chi hapter are the in fo lateral ilized view; oogonia i.e. in gyrogonites, so hm'aGansu voillta . 215):China; gyrogonite m II and conically ovoidal 450N610(AU Anhui, Henan, andFiXinjiang, northern Spain northwestern Cretaceous of Liaoning Qinghai provinces. Mao and Zhou-qing Chen 300- 00 ~m long, 210- 0 ~ wide, 8-9 convolution vi ible in lateral Lower of the planes. Photo wereand taken by Yong-qiang Germany in Europe. shown in this chapter are the fossilized oogonia, i.e. gyrogonites, of(All the fossils I P) view; Mesochara having a\voluta orldwide tribution during small, the Late Jura icovoidal, and Early (Fig.di215)" gyrogonites conically of the plants. Photos were taken by Yong-qiang Mao and Zhou-qing Chen 300N400 lum long, 210"--330 lain wide, 8"-"9 convolutions visible in lateral view; having a worldwide distribution during the Late Jurassic and Early of the NIGP)
"'1
:63
_
. . 210 Aclistochara huihuibaoensis.lateral view (550 Jlrn long,
.211 Aclistochara mundula, lateral view (550 J.1rn
. . 212 Peckisphaera verticillata, apical view (550 Jlrn
.-. 21 3 Peckisphaera verticillata, lateral view (575 J..lm long,
... 214 Mesochara xuanziensis lateral view (290
.-. 215 Mesochara vo/uta, lateral view (390 Jlrn long,
440 Jlm wide), from Halawusu drilling sample (Shahai
long, 250 Jlrn wide), from Pijiagou locality
wide), from Dongliang drilling sample (Fuxin
525 J..lffi wide), from Dongliang drilling sample
300 Jlrn wide), from Sanguanrniao locality
Formation) in Horqin Zuoyi Houqi, Inner Mongolia.
Uiufotang Formation) in Yixian, Liaoning.
Formation) in Fuxin, Liaoning.
(Fuxin Formation) in Fuxin, Liaoning.
Jlm long, 230 J..lrn wide), from Dadianzi locality (lower part ofYixian Formation) in luanping, HebeL
216 Mesochara producta, lateral view (360 Jlrn long, 200 Jlm wide), from
Dadianzi locality (lower part ofYixian Formation) in Luanping. HebeL
_
217 Peckisphaera rnuftispira, lateral view
_
218 Peckisphaera paragranulifero, lateral
_
Liaoning.
.-. 220 Atopochara trivolvis triquetra, lateral
.-. 221 Atopochara trivo/vis triquetra basal view
.-. 222 Flabellochara hebeiensis, lateral view
... 223 Minhechara sp .. lateral view
view oftile same fossil in Fig. 219 (900 J.1m long. 860 J..lm wide). from Halawusu drilling sample (Shahai Formation) in Horqin Zuoyi Houqi. Inner Mongolia.
of the same fossil in Fig. 219 (860 Jim wide), from Halawusu drilling sample (Shahai Formation) in Horqin Zuoyi Houqi, Inner Mongolia.
(750 Jlrn long. 660 J.lm wide), from
(1030 J.lrn long, 610 flrn wide),
Qijiawopeng drilling sample (Shahai
from Sanguanmiao locality (lower part ofYixian Formation) in Kazuo, liaoning.
Dadianzi locality (lower part ofYixian
from Pijiagou locality Uiufotang
Formation) in Luanping, HebeL
Formation) in Yixian, Liaoning.
Formation) in Horqin Zuoyi
view (450 J.1rn long. 350 Jlrn wide),
(lower part ofYixian Formation) in Kazuo,
219 Atopochara trivalvis triquetra, api-
cal view (860 J..lrn wide). from Halawusu drilling sample (Shahai
(370 Jlm long, 300 J..lm wide) , from
I
HouqL Inner Mongolia.
1
Formation) in Kangping, Liaoning.
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II
plants constitute an important component of the Jehol Biota. They and are plants recordedonmainly from the Yixian Formation of West tiwte an important component of the Jehol Biota. Liaoning, almost all major of land Formati plants known Theyand arerepresent recorded mainly fromgroups the Yixian n of esr
L
Liaoning from the Mesozoic Era including free-sporing plants (Bryophyta, Lycopsida, and represent almosr all major groups ofland plants known from rhe Filicopsida) Mesozoic Era including free-sporing plantsCzekanowskiales, (Bryophyta, Lycop ida, Sphenopsida, and seed plants (Ginkgoales, phenopsida Filicopsida) andandeecl planr ( inkgoales, Czekan w Coniferales, Bennettitales, Gnetales Angiospermae). This presentation is kiale , Bennerritale , Gnerales permae). Thi entarion i basedConiferale mainly on , the study of plant fossils and fromAngio the Jianshangou Bedpre of the on rhe wdy ofatplant fos il from Beipiao rhe JianCity, hangou Bed of the lowerbased part ofmainly the Yixian Formation Huangbanjigou, western lowerAparr rhe54Yixian ar Huangbanjigou, B ipiaoAmong ity, \ e rem Liaoning. totalofof speciesFormation in 34 genera have been identified. Liaoning. A roral of 5
pecie in 3 genera have been identified. Among
these, 28 species and nine genera are new taxa. Below is a brief characteriza-
tion of rhe major plant groups and where po ible, rheir inferred pale eco-
..'"
logical condirion will be aJ
oCIl
rhe e, 2 specie and nine genera are new raxa. Below is a brief characteriza-
tion of the major plants groups and where possible, their inferred paleoecological conditions will be also discussed. 0
di cu ed.
Bryophytes Bryophytes (Bryophyta) are the most basal group among Bryophytes Bryophyte (Bryophyra) are rhe
010
t basal group among
the land plants. They include small green plants of simple construction with
rhe land plants. They include smaJl green plant of simple con rrucrion wirh
unlignified conducting tissue and without roots. They are distinguishable
unlignified conducring ri ue and withour roOt. They are di tinguishable
from from all other land plants in having a dominant gametophyte and a very all other land plant in having a dominant gamerophyre and aery simple sporophyte that isthat bornis on theon gametophyte (in other plants simple sporophyte born rhe gamerophyte (inland ocher land the plant the sporophyte is thei dominant and the tissue is The The porophyte the dominant andconducting the conducting ti lignified). ue is lignified). gametophyte is typically upright with dichotomous branching and small gamerophyte is rypically uprighr with dichoromou branching andleafmall leaflike appendages, or in or liverworts mayit bemay flattened and thalloid. The The like appendage, in liverwit rt be flarrened and thalloid. unbranched porophyte born the gameroph and produce unbranched sporophyte is bornis on theongametophyte and re produces a singlea ingle terminal sporangium. Bryophytes hadyhumid and humid terminal sporangium. Bryophytes mostlymostly inhabitinhabit shady and en ironment. the Jehol rhey are represented four species environments. In theInJehol Biota Bi theytaare represented by fourby species in two in (';
0
genera. and thaJloid are pre ent 224, (Fig.225). 22 ,225). genera. Both Both leafy leafy and thalloid types type are present (Figs.
Lycopods Lycop(Lycopsida) ids (Lycopsida) ba al poamong ition among the /yoopods Lycopsids have ahave basala position the va cular plant. lycopod all herbaceous They are characvascular plants. ExtantExtant lycopods are allare herbaceous plants. plants. They are characrerized by ha upright ing upright dichoromou Iy branching terns growing terized by having dichotomously branching stems growing from from horizontal rhizomes and bearing densely The plants are horizontal rhizomes and bearing densely spacedspaced leaves.leave. The plants are hetero porou or homo and roussporangia and p rangia grouped mall cone heterosporous or homosporous either either grouped in smallincones or cattered along the axe, born on the adaxial urface of leaf-like or scattered along the axes, born on the adaxial surface of leaf-like p rangiophylls. Lycopod ha e a wide ecological ran e
ith repre entative
sporangiophylls. Lycopods have a wide ecological range with representatives
in wet and humid environment as well a in arid regions. The rhree extant
in wet and humid environment as well as in arid regions. The three extant 224 Muscites tenellus. a bryophyte. 3.15 cm long.
224 Muscites tenellus, a bryophyte, 3.15 cm long.
lycopod familie ,Lycopodiaceae, elaginellaceae and Isoetac ae, \ ere al
lycopod families, Lycopodiaceae, Selaginellaceae and Isoetaceae, were also
0
%
,..o
present in the Cretaceous, but so far only members of the Lycopodiaceae were pre ent in thefrom retthe ceou , but so far onlas member of the Lycopodiaceae recorded Jehol Biota, such Lycopoditesfaustus, a homosporouswer<: plant recorded theJehol Biota,insuch as L)'copodih.rfil/llllJ, a homo porou pl. nt with from sporangia arranged a strobilus (Figs. 226, 227). with porangia arranged Sphenopsids in a strobiluor(Fig. nG, (Sphenopsida) 22 ). Sphenopsids horsetails are characterSphenopsids phenop ids or horsetail ( phenop ida) are charaCter-
ized by upright and distinctly articulate and typically hollow stems arising
ized by upright and di tincdy < niculate and typically hollow tems ari ing
from horizontal rhizomes. Branches and leaves are born in whorls at the
from horizontal rhizomes. Branche 'lOd leave arc born in whorl at the
nodes. The plants are heterosporous or homosporous with terminal, mostly
nodes. The plant arc hetero porous or homosporous with terminal, mo dy
abaxial sporangia, born on peltate, scaly sporophylls that are arranged in
abaxial porangia, born on peltate, scaly porophyll that are arranged in
terminal, ellipsoid cones. The only extant genus, Equisetum (Equisetaceae) is
terminal, ellip oid cone. The anI extant genu, Eqllistllllll (Equi etaceae) is
herbaceous and homosporous. It has very small leaves that are more or less
herbaceou and homosporou . Ir ha
ery mall lea es that are more or Ie
fused to form a dentiform leaf sheath at the node. The surface of the internodes
fu cd t form a dentiform leaf heath at the node. The urface of the internode
shows distinct longitudinal ridges from the vascular bundles. Equisetum
h \ s di tinct I ngitudinal ridge from rhe va cular bundle. EqllistllllJ/
predominately occurs in wet environments, and the hollow stems, charac-
predominately occur in wet environment
and the hollow tem
charac-
teristic of many sphenopsids, indicate it was adaptedto to teristic also also of many cxtinextinct t phenopsid, indicate thatthat it wa adapted 225 Thallites a bryophyte, cmround long, round print probably 225 TllOl/ite nccioite riccioites, . a bryophyte. 1.1 cm 1.1 long. prim probabl
in moist areas. Remnants of Equisetum-like plants, grO\ growth th in moi r area. Remnant of EqlliJellIlI/-like plant, de described cribed a as
representing the regetative-reproductive organ gemma repr senting the regetative-reproducti e organ gemma cups. cups.
226 Lyropodites!austu. a lycopod. m 226 Lycopoditesfaustus,a lycopod, 5.95 CI11 long.
5.95 cm long.
227 Enlargemems of the rip of the specimen shown in Fig. 226, showing a 227 Enlargements of the the specimen shown in Fig. 226, showing sporangia. the trobili wirh two rowtipofofround
the strobili with two rows of round sporangia.
Equisetites, are very common in the fossil-bearing strata of the Yixian Formation. The most abundant sphenopsid in the Biota is Equisetites Equisetites, are very common in the fossil-bearing srrata of the Yixian longevaginatus (Fig. 228), characterized by very small stem and rather long leaf
Formation. The most abundant sphenopsid in the Biota is Eql/isetites sheath. longevaginattls (Fig. 228), characterized by very small stem and rather long leaf Filicopsids Filicopsids or ferns (Filicopsida) are the largest group of the sheath. free-sporing vascular plants. The plants have distinctly differentiated root, Filicopsids Filicopsids or ferns (Filicopsida) are the largest group of the stem and leaves. They are woody or herbaceous and some attain tree habit, free-sporing vascular plants. The plants have distinctly differentiated root, characterized by having an upright growth with a crown of large leaves. The stem and leaves. They are woody or herbaceous and some attain tree habit, leaves, often referred to asanfronds, aregrowth simple or more commonly pinnately characterized by having upright with a crown of large leaves. or The
dichotomously compound. Most ferns are homosporous whereaspinnately a few are or leaves, often referred to as fronds, are simple or more commonly heterosporous. are Most terminal born the abaxial surface of anormal dichotomouslySporangia compound. ferns areon homosporous whereas few are or specialized leaves. Their position on the leaves highly variable. They may heterosporous. Sporangia are terminal born on isthe abaxial surface of normal be scattered over the surface of the lamina variously aggregated in They circular, or specialized leaves. Their position on theorleaves is highly variable. may elongate or linear Ferns mainly inor humid and moist areas, but some be scattered over sori. the surface of the grow lamina vatiously aggregated in circular, elongate or linear sori.arid Ferns mainly grow humid moist areas, some are adapted to more conditions. Fernsin are veryand common in the bur Yixian are adaptedand to so more arid conditions. Ferns very common Particularly, in the Yixian Formation far more than 5 species haveare been recognized. Formation and so far genera more than 5 species have been recognized. Particularly, species of the extinct Botrychites (Figs. 229, 230), Eboracia (Fig. 231), species of the extinct genera Bot1ychites (Figs. 229,assemblages. 230), Ehoracia (Fig. 231), and Coniopteris are conspicuous in the Jehol plant and Coniopteris conspicuous in the comprise Jehol plant assemblages. Ginkgos are Ginkgos (Ginkgoales) large trees characterized by Ginkgos Ginkgos (Ginkgoales) comprise large trees characterized by long shoots and short shoots. They have wood with small pith, proliferate long shoots and short shoots. They have wood with small pith, proliferate helical branching and leaves born in a helical arrangement. Leaves on long helical branching and leaves born in a helical arrangement. Leaves on long shoots are typically widely spaced while those on the short shoots are clustered shoots are typically widely spaced while those on the short shoots are clustered around the apical part. They are unique, fan-shaped with dichotomously around the apical patt. They are unique, fan-shaped with dichotomously branching veins, bilobed or sometime strongly dissected. The only extant branching veins, bilobed or sometime strongly dissected. The only extant species, Ginkgo biloba, is dioecious with female and male trees. The ovules and species, Ginkgo hi/oha, is dioecious with female and male trees. The ovules and microsporophylls are terminal and born on the short shoots. The ovules are microsporophyJls are terminal and born on the short shoots. The ovules are borne in small cups on a ovulate axis. Of typically two ovules in extant Ginkgo, borne in small cups on a ovulate axis. Of typically two ovules in extant Ginkgo, only one reaches maturity. In some ancestral ginkgos ovules were more only one reaches maturity. In some ancestral ginkgos ovules were more numerous on the ovulate axis. The ovules and mature seeds are relatively large numerous on the ovulate axis. The ovules and mature seeds are relatively large with an outer fleshy layer. Ginkgos were much more abundant and diverse with an outer fleshy layer. Ginkgos were much more abundant and diverse in the Mesozoic, particularly in the Late Jurassic and Early Cretaceous. in the Mesozoic, particularly in the Late Jurassic and Early Cretaceous. Ginkgos declined declined drastically Ginkgos drastically both both inin diversity diversityand andgeographic geographicdistribution di tribution
during the the Late Late Cretaceous during Cretaceous and and Tertiary. Tertiary. During Duringthe theQuaternary Quaternaryititbecame became restricted to small areas restricted ro small areas in in eastern eastern China. China.The Theonly onlyknown knownwild wildpopulation populationofof
Ginkgo hi/oha biloba isis in Ginkgo in aa temperate temperate moist moist forest forest ofofZhejiang. Zhejiang.Ginkgo Ginkgobiloba hi/ohais is ~ 2 2228 8 Equisetites Equisetiteslongevaginatus, !ongevaginatus,aasphenopsid, sphenopsid,3.7 3.7cm emlong. long.
deciduous and the abundant occurrence of leaves in certain geological horizons indicates that at least some extinct ginkgos were also deciduous. In deciduou :lnd rhe abundanr
ccurrence of leave in cenain geological
the Yixian Formation a single species of Ginkgo was reported recently by
horizon indicare rhat at leasr ome extincr ginkgos \ ere also deciduous. In
Z.-y. Zhou and S.-1. Zheng (2003) (Fig. 232). It is intermediate in the ovulate
rhe Yixian Formarion a ingle pecies of Ginkgo was reponed recently by
structure between modern Ginkgo and Jurassic ginkgos with many ovules.
Z.-y. ZhoLl and .-1. Zheng (_DO ) (Fig. 232). Ir is inrermediate in rhe ovulate
The flora also comprises several taxa belonging to extinct ginkgoalean genera
rructure berween modern Ginkgo and Jura ic ginkgos with many ovule.
such as Baiera (Fig. 233), Ginkgoites, and Sphenobaiera.
The flora al
0
compri e
everal raxa belonging
to
exrinct gink,goalean genera
Czekanowskialeans Czekanowskialeans (Czekanowskiales) represent uch as Baiera Fig. _ >, GiJlkgoiles, and phOlobaiera. anCzekanowskialeans extinct group of seed zekanow plants of kialean unknown( zekanowskiale) systematic affinity. was repreIt enr an extinct group of eed plane of unknown s
m 2 2 9 Botrychitesreheensis, a filicale, 6.8 cm long, showing the heteromorphic 229 Botrychites i , a filicale, sterilereheen and fertile pinna. 6.8 sterile and fertile pinna.
COl
long, showing the heteromorphic
temaric affiniry. Ir wa
~230 Pinna ofBotrychites reheensis,a filicale, 4 cm long. 230 Pinna of Botrychites reheensi , a filicale, 4 Col long.
~
CO
d
Ginkgobiloba
c
~
CD ...... CO
::J
a
'Ginkgoadiantoides ca. 56 Myr BP
~~Ginkgonew species ca. 121 Myr BP
Ginkgoyirnaensis
()ca.
en en
Ginkgo yimaensilt yr BP
~i~~
,=
~
..., ::J
Ginkgo apodes. a ginkgophyte, ~ 232232Ginkgo apodes, a ginkgophyte, fromfrom 231 Sterile pinna of Eboracia lobifolia, a filicale. 1.8 cm ~ 2 3 1 Sterile pinna ofEboracia lobifolia, a filicale, 1.8 cm long. Note the shape of the variant pinnule at the long. Note the shape of the variant pinnule at the right of the base of pinna. (Photo: Shi-wei Zhao/ right of the base of pinna. (Photo: Shi-wei Zhao/ NIGP) NIGP)
233 Baiera borealis, a ginkgophyte, 7.1 cm long. (Photo: 233 Baiera borealis, a ginkgophyte, 7.1 cm long. (Photo: Shi-wei Zhao/ NIGP) Shi-wei Zhao/NIGP)
. 234 Solenites murrayana, a ginkgophyte, 4.55 cm long. m 234 Solenitesmurrayana, a ginkgophyte, 4.55 cm long.
Toudaohezi locality (Zhuanchengzi Toudaohezi locality (Zhuanchengzi Bed ofYixian Formation) in Yixian, Bed of Yixian Formation)in Yixian, Liaoning. All scale bars = 5 mm. a, Liaoning. All scale bars = 5 mm. a, A juvenile ovulate organ with 6 A juvenile ovulate organ with 6 collars and very short pedicels; b. collars and very short pedicels; b, Associated leaf; c, An ovulate orAssociated leaf; c, An ovulate organ bearing one nearly mature (red gan bearing one nearly mature (red arrow) and probably one aborted arrow) and probably one aborted ovule (blue arrow) and 1-2(7) ovule (blue a r r o w ) a n d 1N2(?) empty collars; d, Sketch reconstrucempty collars; d, Sketch reconstruction of the species and some other tionGinkgo of the taxa, species and some other of showing evolution Ginkgo taxa, showing evolution of this group in geological history. this group in geological history. (Courtesy: Zhi-yan Zhoul NIGP) (Courtesy: Zhi-yan Zhou/NIGP)
Cone ofSchizokpis beipiaoensis. a conifer. 9.15 an long.
Samara of Schizo/~p;s b~;p;Qo~ns;s. a conifer. 1.15 an long.
included in the Ginkgoales, but their leaves and their reproductive structures,
structures that may be unisexual or bisexual. The ovules and mature seeds are
particularly the ovulate structures, are distinct from those of Ginkgo, and they m i n u t e and terminal densely spaced on a conical or spherical receptacle included in the Ginkgoale , but theit leave and their reproductive tructure struCture that may be unisexual or bi exual. The ovule and mature eeds are are now usually placed in a separate order. The leaves are borne in clusters on together with small interseminal scales. The microsporangia are borne on minute and terminal den ely spa ed on a conical or spherical receptacle particularly the ovulate tructures, are distinct from those of Ginkgo, and they short shoots. They are very narrow and supplied by only a single vein, in simple or branched (pinnate, bipinnate) microsporophylls. The bennettites are now usually placed in a separate order. The leave are borne in clu ters on rogether with mall interseminal ales. The microsporangia are borne on contrast to the two veins that enter the leaves of Ginkgo. The leaves are were c o m m o n in Triassic, Jurassic and Early Cretaceous strata but became short hoots. They are very narrow and supplied by only a ingle vein, in imple or branched (pinnate bipinnate) micro porophylls. The bennenites typically dichotomized into several sections, each with a single vein. The extinct by the end of the Cretaceous. In the Yixian Formation four forms have contrast to the twO veins that enter the leaves of Ginkgo. The leaves are were common in Triassic, Jurassic and Early Cretaceous strata but became ovulate structure called Leptostrobus is characterized by having elongated axes been reported (Figs. 238---240). extinct by the end of the reta eou . In the Yixian Formation four forms have rypi ally di hotomized into everal eetion, each with a ingle ein. The with bivalved ovule bearing units in a spiral arrangement. Each bivalved unit, G n e t a l e s Gnetales comprise three extant genera (Ephedra, Gnetum, been reported (Figs. 238-2 0). ovulate tructure called Leptos/I'oblfS is characterized by having el ngated axe often referred to as a capsule, contains several ovules. The microsporangiate Welwitschia) that are morphologically very distinct. The phylogenetic posiwith bivalved ovule bearing unit in a spiral arrangement. Each bivalved unit, Gnetales Gnetales compri e three extant genera (Ephedra, ne/II11l, structure, Ixostrobus, is a simple axis with pairs of pollen sacs also arranged in Welwi/schia) that are morphologically very distinct. The phylogenetic posioften referred to as a capsule, contains several ovules. Th microsporangiate a spiral pattern. The Czekanowskiales were widespread and abundant during structure, IxostroblfS, i a simple axis with pairs of pollen acs also arranged in the Jurassic and Cretaceous, particularly in floras of the Northern Hemisphere. a piral pattern. The zekanowskiale were wide pread and abundant during In the Jehol Biota species of Solenites (Fig. 2_34) and Sphenarion are common theJuras i and 'rera eous, parri ularly in nora (rhe rrhern llemi phere. Czekanowskialean elements. In the Jehol Biota pecies of oi nite.r (Fig. 23 ) and phel1arion are common Conifers Conifers (Coniferales) comprise trees or more rarely shrubs zekanowskialean elements. characterized by long shoots and short shoots. W o o d is massive with small Conifers onifers ( oniferales) comprise trees or more rarely shrubs pith. Branching is proliferating and phyllotaxis is helical or decussate. Leaves characterized by long hoot and short shoots. Wood is rna ive with small are h o m o m o r p h i c or h e t e r o m o r p h i c , typically small needlelike, scaly or pith. Branching i proliferaring and phyllotaxi i helical or decu sate. Leave conical, supplied by a single vein or more rarely two veins. Most conifers are are homomorphic or heteromorphi , typically mall needlelike, scaly or monoecious. The ovules and microsporangia are terminal borne in cones. coni al, supplied by a single vein or more rarely twO veins. Most conifers are Ovulate cones are c o m p o u n d with ovules born on seed scale that is fused to monoecious. The ovules and microsporangia are terminal borne in cones. the supporting cone scale. Mature seeds are usually rather small with a hard Ovulate cones ate compound with ovules born on seed scale that is fused ro seed wall. Microsporangiate cones are simple and the pollen sacs are born on the upporting cone cale. Mature seeds are usually rather small with a hard the abaxial surface of the microsporophylls. Conifers were a b u n d a n t in the eed wall. Micro porangiate c ne are simple and tht: pollen sac are born on Mesozoic with many extinct forms occurring well into the Cretaceous. In the the abaxial urface of the micr p rophylls. onifer were abundant in the Yixian Formation all conifers so far recorded appear to belong to extinct Mesozoic with many extinct form occurring well into the retaceous. In the genera and families. The most abundant conifer is Schizolepis (Figs. 2_35, 2_36), Yixian Formation all onifers 0 far recorded appear ro belong ro extinct but Elatodadus (Fig. 2_37) and Brachyphyllum also constitute an important part genera and families. The m t abundant conifer is chizolepis (Figs. 2 5,2 6), of the Jehol plant assemblage. but El.1/odatllfS (Fig. 2 7) and BrachyphylllllfJ also con titute an important part Bennettites Bennettites (Bennettitales) are extinct seed plants with a of the Jehol plant as emblage. habit resembling that of the cycads both in the stature and in the morphology Bennettites Bennettite (Bennettitales) are extinct e d plant with a of leaves. B e n n e t t i t e s are r e c o n s t r u c t e d as small, sparsely b r a n c h e d or habit resembling that of th cycads both in the stature and in the morphology unbranched trees or small shrubs. The trunk is slender or in some bennettites of lea e . Bennerrites are recon tructed as small, sparsely branched or barrel-shaped. Leaves are large, simple (entire) or pinnately divided, very unbran hed tree or mall hrubs. The trunk i lender or in some bennenites similar in gross m o r p h o l o g y to some cycad leaves. The two groups can, barrel- haped. Leave are large imple (entire) or pinnately di ided, very however, easily be distinguished by their epidermal features when present. imilar in gros m rphology to some cycad leave. The twO groups an, The ovules and microsporophylls are crowded in rather large flowerlike m 238 Wiiliamsoniabella, a bennettite, 10.1 cm long. howe er, easily be distingui hed by their epidermal feature when pre ent.
The ovules and microsp rophylls are crowded in rath r large flowerlike
238 Williamsonia bella. a benn nile, 10.1 em long.
... "'%,..o Cl
173
l::
;:
lit
tion of the g r o u p is uncertain. It was sometimes placed close to the
174
W u , 2000), and Chaoyangia liangii (Fig. 242) assigned to the extinct genus
Gurvanella (Sun et al., 2001). They are all closely similar to Ephedra in their angiosperms, but has recently been associated with the conifers. Ephedra and tion of the group is uncertain. It was sometimes placed close to the Wu, 2000), and Chaoyangia liangii (Fig. 2 2) assigned to the extinct genu vegetative morphology, but some (e.g., Gurvanella) are distinguishable from Gnetum include trees, shrubs, vines and climbers with proliferate branching angio perm, but has recently been associated with the conifers. Ephedra and GlIrvanelia ( un et al. 2001). They are all closely similar to Eph dra in their Ephedra in the ovulate structures. and decussate or whorled phyllotaxis. Welwitschia is unusual in having a very Gne/llm include tree, shrubs, vines and climbers with proliferate btanching vegetative morphology, but orne (e.g., GlIrvanella) are distinguishable from Angiosperms Angiosperms (Angiospermae) are the most diverse of all condensed, unbranched stem and two persistent leaves that grow for the Ephedm in the ovulate tructures. and decus ate or whorled phyllotaxi . Welwi/schia i unu ual in having a very plant groups and exhibit an enormous range in vegetative and reproductive entire life of the plant. The plants are mostly dioecious, rarely monoecious. Angiosperms Angio perms (Angio permae) are the mo t diver e of all condensed unbranched stem and two persistent leaves that grow for the features. Probably the most distinct feature that distinguishes angiosperms The ovules and microsporophylls are terminal borne in small compound, plant groups and exhibit an enormous range in vegetative and reprodu tive entire life of the plant. The plants are mostly dioecious, rarely monoecious. from other plants is the enclosure of the ovules in carpellary tissue and pollen unisexual cones. The mature seeds are small to large. The microsporangia are The ovules and microsporophylls are terminal borne in small compound, features. Probably the most distinct feature that distinguishes angiosperms tube growth via the carpellary tissue to the ovules. Several angiosperms have borne in synangia. Ephedra is xerophytic and Welwitschia extreme xerophytic, uni exual cones. The mature eeds are small to large. The microsporangia are from Other plants is the enclo ure of the ovules in carpellary tissue and pollen been described from the Yixian formation but most of them are dubious (Figs. while Gnetum is tropical genus with a wider ecological range inhabiting moist borne in synangia. Ephedra is xerophytic and Welwi/schia extreme xerophytic, tube growth via the carpellary tissue to the ovules. everal angiosperms have 243---250). Some have already been placed in the Gnetales, while the features to rather dry environments. The fossil history of Gnetales is poorly known but been described from the Yixian formation but mo t of them are dubious (Fi s. while Gne/llm i tr pi al genu with a wider ecological range inhabiting moi t of others such as Archaefructus is still debated (see the Chapter "Angiosperms"). Gnetales pollen occurs abundantly in Early Cretaceous sediments. In the to rather dry environments. The fossil history of Gnetale is poorly known but 2 3-250). orne have already been placed in the Gnetale ,while the feature A more distinct angiosperm was described recently by Qin Leng and Else M. Yixian Formation remnants of Gnetales are relatively common and several of others such as Archae/rlletlls is still debated (see the hapter' Angiosperms"). Gnetales pollen occurs abundantly in Early Cretaceous sediments. In the Friis and will be treated in the next chapter together with a discussion on the taxa have been described. Among these are Eragrosites changii and Liaoxia Yixian Formation remnants of Gnetales are relatively common and several A more distinct angiosperm was described recently by Qin Leng and ELse M. chenii (Fig. 241) that was assigned to the extinct genus Ephedrites (Guo and Jehol angiosperms. Friis and will be treated in the next chapter rogether with a di cussion on the taxa have been de cribed. Among these are Eragrosites changii and Liaoxia (Unless otherwise chenii (Fig. 241) that was a signed ro the extinct genus Ephedrites (Guo and Jehol angiosperms. stated, fossils shown in (U nle otherwi e this chapter are collected stated, fo il hown in from the Jianshangou Bed this chapter are collected of the lower part of the from the J ianshangou Bed Y i x i a n F o r m a t i o n at of the lower part of the Huangbanjigou, Beipiao, Yixian Formation at Liaoning, photo by Da-jian 1 uangbanjigou, Beipiao, Li/CAS) Liaoning, photo by Da-jian Lil A)
239 Rehezamites anisolobus, a possible bennettite, 10.1anisolobus, cm long (Left). 239 Rehezamites a possible bennenite, )0.1 cm long (Left).
i 2 4 0 Tyrmia acrodonta, a bennettite, 6.9 cm long
(Right). 240 Tyrmia acrodonta, a bennettite, 6.9 em long (Right).
~242
Chaoyangia liangii, a gnetale, 0.85 cm long.
~243
Erenia stenoptera, Plantae incertae sedis, 0.5 cm long. (Photo: Dong-xing Deng/NIGP}
244 Typhaerafusiformis, Piantae incertae sedis, 1.8 cm long. (Photo: Shi-wei Zhao/NIGP)
~ 2 4 1Liaoxia Liaoxia chenii, a gnetale, long. (Photo: 241 chenii, a gnetale, 8.9 8.9 cm cm long. (Photo: Dong-xing Deng/NIGP) Dong-xing Dengl IGP)
Q245 2 4 5 Leaf Leaf shootofofLilites Lilitesreheensis, reheensis,Plantae Plantae incertae incertae sed sedis, 9.8 cm cm long, shoot is, 9.8 long, showing opposite and amplexicaul leaves, and arching veins. showing opposite and amplexicaul leaves, and arching veins.
246 sedis, 4.54.5 cmcm 246 Fruit Fruitspur spurofLilites of Lilitesreheensis, rel1eensis,Plantae Plantaeincertae incertae sed is, long, long, showing showingterminal terminalfruit fruitand andopposite oppositeleaves. leaves.
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m 247 Carpolithussp., Plantae incertae sedis, 0.6 cm 247Dong-xing Carpolithus sp.• Plantae ineertae sedis. 0.6 em long. (Photo: Deng/NIGP) long. (Photo: Dong-xing Deng/ IGP)
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IGP)
250 Antholithussp. 2, Plantae incertae sedis, 3.5 cm long. (Photo: Shi-wei 250 Antholitlru sp. 2, Plantae ineertae edi Zhao/NIGP) ,3.5 em long. (Photo: Shi-wei Zhao/ NIGP)
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ngiosperms ngi p rm (flowering (fl wenn plants) plant are ecologically ec 10 i ally dominant d man ne In most m t terrestrial terre eri J vegetation ve etation today, t day, accounting a oumin for fi r the bulk of the primary productivity pr du ti icy on land. They constitute con ticuc the ch most m c
species-rich pe ie -ri h group r up off land plants pI m that thac ever eXisted, exi ced, with wich an overwhelming ov r\ helming
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oceans. De pic their cheir prominent pc minem position p icion In in the che modern m dern world, the che anan . Despite
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179
ecological placeeo over IntervalI dUring e 10 i I dominance d min nettookk pia er a relatively r laci el short hore time cime imerv durin the th Cretaceous, ret u more mor than chan 300 00 millton milli n years after fcer plants' plancs' first fir t Invasion in i n onto nc the land land. The earhesc arlie c uneqUivocal unequi al angiosperm an i erm fossils fo il are pollen poll n grains grain from fr m Valanginian-Hauterivian Ian in ian- Haucerivian strata erata of JIsrael rael (Brenner, Brenner, 1996) and nd southern ouchern England (Hughes, 1994), about 130-135 -1 5 millton milli n years ear old. Angiosperms An i perm In che these earltest are rrare ar re in earlie c Cretaceous r ca eou strata ceaca but buc by the Barremian-Aptian, Barremian-Apcian,
10-15 ffiliiton afterr cheir their first occurrence, milli n years year aft fir curren e, they were already aJre dy Widely widely establtshed, the eran transition the ancient MeSOZOIc cabli hed and th icion from ch anciene Me ozoi vegetation, e cati n, dominated (cycads, bennettttaleans, d minated by b gymnosperms ymn perm (cy ad, ginkgos, inkgo seed d ferns, fern bennetcicalean, the modern conifers) onifer and nd pteridophytes pteridophyte (lycopods, (Iycop d hhorsetails, retail , ferns), fern ), cto che m dern ecosystems was well under way. ec cern w Plum ere trees, tea bu bushes, walnut cree trees, sunflowers, h unflo er rice, n e and the th crops r p that among che the familiar famlltar an angiosperms thac give leuus our breakfast br akfa t cereals cere I are re amon io perm essential features enti J to our ur eeveryday ryday Iilives. The first angiosperms shared hared many feature their present-day descendants, but th they were also different How With ~ ith cheir re ene-day de c ndam , buc ere al 0 differ m. H w different, we do nOt not fully kno know . Th The ori origin diverSification differenc, in and early di er ificacion of angiosperms intenSively more years, buc but an i p rm have been incen i Iy studied tudied for m r than 100 I year, questions unresolved regarding time of their Origin, their many que ti n remain une olved re ardin the cime ri in, th ir
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251 Archaefructus sinensis, an aquatic (preserved 15long), cm long), Dawangzhangzi locality 251 Archaefructu sinensis, an aquatic plantplant (preserved mainmain axis axis 15 cm fromfrom Dawangzhangzi locality (Dawangzhangzi Bed of Yixian Formation) in Lingyuan, Liaoning 125122 or 122 picture Bed ofYixian Formation) in Lingyuan, Liaoning (ca. (ca. 125 or Ma).Ma). The The left left picture (Dawangzhangzi showsshows the clothee-up view of a reproductive axis. axis. (Photo: Yvonne Arremo/ RM) close-up view of a reproductive (Photo: Yvonne Arremo/NRM)
ancestral ancestralfeatures, features,and andtheir theirclosest closestrelatives relativesamong amongother otherplant plantgroups. groups.So Sofar far there are neither clear signals from molecular studies nor universally accepted there are neither clear signals from molecular studies nor universally accepted theoretical theoreticalmodels modelson onangiosperm angiospermevolution. evolution.Therefore, Therefore,information informationfrom fromthe the fossil fossilrecord recordisiscrucial crucialtotoour ourunderstanding understanding of ofangiosperm angiospermevolution evolutionand andfor for testing testingvarious various models modelsfor fortheir their phylogenetic phylogenetic relationships. relationships. The TheJehol Jehol Group Group sediments sediments were weredeposited deposited during during the the early earlyphases phases of of this this Cretaceous Cretaceous vegetational vegetational revolution revolution and and provide provide aa window window to to study study changes and innovations in Early Cretaceous floras. The deposits preserve changes and innovations in Early Cretaceous floras. The deposits preserve aa biota biotathat that either eitherlived livedininor orwas wastrapped trapped ininaasystem systemof oflow-energy low-energylakes. lakes.There There isis no no or or little little indication indication of of transportation transportation prior prior to to fossilization. fossilization. Some Some of ofthe the plant fossils are preserved as whole plants with roots, stems, leaves and plant fossils are preserved as whole plants with roots, stems, leaves and reproductive reproductive organs organs in in organic organic connection, connection, and and some some of of the the plants plants were were obviously obviously aquatic. aquatic. Whole Whole plant plant preservation preservation isis very very rare rare in in the the fossil fossil record record and and has has been been reported reported only only from from aafew fewplaces places around around the the world. world. Fossil Fossil remains remains of whole water plants are even more unusual because they typically have of whole water plants are even more unusual because they typically have very very delicate delicate structures structures with with low low fossilization fossilization potential. potential. The The Jehol Jehol plant plant assemassem-
blage thus provides insight into a part of the vegetational history that is poorly blage thus provides insight into a part of the vegetational history that is poorly known. Despite the deposition that resulted in whole plant preservation and known. Despite the deposition that resulted in whole plant preservation and excellent information on gross morphology, plant fossils from the Jehol excellent information on gross morphology, plant fossils from the Jehol sediments have experienced rapid tissue decay during the very early stage of sediments have experienced rapid tissue decay during the very early stage of fossilization, which destroyed most of the cell-level structures (Leng and fossilization, which destroyed most of the cell-level structures (Leng and Yang, 2003). This, along with the highly specialized features of water plants, Yang, 2003). This, along with the highly specialized features of water plants, sometimes makes interpretations and systematic inferences very difficult. sometimes makes interpretations and systematic inferences very difficult. History of the search for Jehol angiosperms The first Jehol plants History of the search for dehol angiosperms The first Jehol plants assigned to angiosperms were Potamogeton jeholensis Yabe et Endo and assigned to angiosperms were Potamogeton jeholensis Yabe et Endo and Potamogeton? sp. reported by two]apanese botanists H. Yabe and S. Endo in Potamogeton? sp. reported by two Japanese botanists H. Yabe and S. Endo in 1935, but an intensive search for angiosperms did not start until about 60 1935, but an intensive search for angiosperms did not start until about 60 years later and several putative angiosperms were recently described (rom the years later and several putative angiosperms were recently described from the Yixian Formation. The Late Jurassic age assignment of the lower Yixian Yixian Formation. The Late Jurassic age assignment of the lower Yixian Formation that was adopted by some authors gave the fossils an older age than Formation that was adopted by some authors gave the fossils an older age than angiosperms from other places and certainly contributed to the excitement of angiosperms from other places and certainly contributed to the excitement of the search. The mid-Early Cretaceous age that is now widely accepted (Zhou the search. The mid-Early Cretaceous age that is now widely accepted (Zhou
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. . 253 An almosr complete plant (13.4 em long) of Archaefructus sinensis, from Dawangzhangzi locality (Dawangzhangzi
Bed ofYixian Formation) in Lingyuan, Liaoning (ca. 125 or 122 Ma). (Photo: Yvonne Arremo/ NRM)
953 An almost complete plant (13.4 cm long) ofArchaefructussinensis, from Dawangzhangzi locality (Dawangzhangzi Bed of Yixian Formation) in Lingyuan, Liaoning (ca. 125 or 122 Ma). (Photo. Yvonne Arremo/NRM) er al., 2003) places rhe fossils in another conrext as coeval with early
Dilcher Zheng et Zhou ( un et al., 1998) and A. sinensis un, Dilcher, Ji et
angiosperm floras from other regions. Among rhe fossil planes that have been Nixon (Sun et aI., 2002); Beipiaoa pOl'va Dilcher, Sun et Zheng, B. rotunda et al., 2003) places the fossils in another context as coeval with early Dilcher, Zheng et Zhou (Sun et al., 1998) and A. sinensis Sun, Dilcher, Ji et recently related to the angiosperms are Chaoyangia liangii Duan (Duan, Dilcher, Sun et Zheng, and B. spinosa Dilcher, un et Zheng (Sun et al., 2001); Nixon (Sun et al., 2002); Beipiaoa parva Dilcher, Sun et Zheng, B. rotunda angiosperm floras from other regions. Among the fossil plants that have been 1997); E"agrosites ehOlzgii Cao et Wu, Liaoxia ehenii Cao et Wu, and "Mono- and Sinoearpm deClfssatm Leng et Friis (Leng and Friis, 2003). Other putative Dilcher, Sun et Zheng, and B. spinosa Dilcher, Sun et Zheng (Sun et al., 2001); recently related to the angiosperms are Chaoyangia liangii Duan (Duan, cotyLedon Leaf' (Cao et al., 1998); Erenia stenoptera Krassilov, Lilites reheensis angio perms were described based on very fragmentary or poorly preserved and Sinocarpus decussatus Leng et Friis (Leng and Friis, 2003). Other putative 1997); Eragrosites changii Cao et Wu, Liaoxia chenii Cao et Wu, and "MonoWu, Orehidites linearifoLim Wu, Orchidites laneifolills Wu, Trapa? sp., and material (e.g. several taxa described under the heading "Problematic Ancotyledon Leaf' (Cao et al., 1998); Erenia stenoptera Krassilov, Lilites reheensis angiosperms were described based on very fragmentary or poorly preserved Typhaera filsi/om/is Krassilov (Wu, 1999); Archaefmet1tS liaoningensis Sun, giosperms" by Wu, 1999) and are nOt discussed further here. material (e.g. several taxa described under the heading "Problematic AnWu, Orchiditeslinearifolius Wu, Orchidites lancifolius Wu, Trapa? sp., and
Typhaera fusiformis Krassilov (Wu, 1999); Archaefructus liaoningensis Sun,
giosperms" by Wu, 1999) and are not discussed further here.
The rapid decay prior to fossilization and possible morphological specialization of some of the plants to an aquatic environment make it difficult to interpret the morphology and structures. Interpretation and systematic assignment have subsequently been questioned for many of these fossils. Chaoyangia liangii,
Eragrosites changE, Liaoxia chenii and Potamogetonjeholensis(later reassigned to Ranunculusjeholensis by Miki, The rapid decay prior co fos ilization and possible morphological specialization of some of the plant
1964) show distinct affinity with members of the relict seed plant group Gnetales and two of them,
ro an aquatic environment make it difficult co interpret the morphology and structures. Interpretation and
Eragrosites changii and Liaoxia chenii were transferred to the extinct genus Ephedrites (Guo and Wu, 2000). haO)'Clngia liangii, Potamogenton? sp., the "Monocotyledon Leaf', Lilites, and the two species of Orchidites were re-interpreted E1'agrosites changii, Liaoxia chenii and Potamogeto71 jeholenJis (later reas igned ro Ram(7/mlm jeholensis by Miki as conifers (Sun et al., 2000, 2001), an assignment that is also problematic (see below). Archaefructus was
sy tematic assignment have ubsequenrly been que tioned for many of these fossils.
em
1964) how distinct affinity with members of the relict seed plant group Gnetale and twO of them
originally described as bearing multiparted bisexual and naked flowers and was resolved as sister to all Eragrosites chcmgii and Liaoxia chmii were transferred co the extinct genus Ephe Irites (Guo and Wu, _000). extant angiosperms (Sun et al., 1998, 2002). This interpretation has also been questioned and an Potamogento71? sp., the "Monocotyledon Leaf', Lilites, and the twO pecie ofOrrhidites were re-interpteted alternative explanation offered (Friis et al., 2003), and the possibility that Archaefructus is more closely as conifers ( un er aI., 2000,2001), an as ignment that j al 0 problematic (see below). Archae/mctlls was related to other seed plants was discussed by Zhou and others (2003). In the following sections we originally described as bearing multiparred bisexual and naked flower and
as re olved as sisrer to all
comment briefly on (fossil plants that still remain of interpretation interest in the has discussion on Early Cretaceous extant angiosperms un et al. L99 2002). This also been questioned and an angiosperm structure and diversity. alternative explanation offered (Friis et aI., 2003), and the po sibility that Archae/mc/IIS i more closely Archaefructus--a submerged water The extinct genus(2003). Archaefructus wasfollowing established by Sun we related co other eed plants was discus edplant by Zhou and other In the sections
m 2 5 6 Reconstruction ofSinocarpus decussatus. (Art: Qin Leng and Yu-gao Ren/NIGP)
and others briefly (1998). on Thefo original description single species, A. liaoningensis, which wasCretaceou known comment sil plants that stillincluded remain aof interest in the discu sion on Early angiosperm structure and di er ity.
256 Reconstruction of Sinocarpus demssatus. (Art: Qin Leng and Yu-gao Ren/ NIGP)
submerged water plant The extinct genus Archae/metlls was establi hed by un m254Archaefructus-a Sinocarpusdecussatus (5 cm long), an angiosperm with advanced features, 255 A close-up view of the partly united carpels of the specimen shown and others (1998). The original description included singleFormation) species, A. h was kno nXiao-yi Fan/NIGP) in Fig. 254 (Photo: from Dawangzhangzi locality (Dawangzhangzi Bed ofa Yixian in liaoningemis, whi
from Dawangzhangzi locality (Dawangzhangzi Bed ofYixian Formation) in Lingyuan. Liaoning (ca. 125 or 122 Ma). (Photo: Yvonne Arremo/ NRM)
m
:z: o
1II 1II
;:
Lingyuan, Liaoning (ca. 125 or 122 Ma). (Photo: Yvonne Arremo/NRM) 254 Sinocarpus decussatus (5 em long). an angiosperm with advanced features.
.."'i . tl 1II
255 Aclose-up view of the partly united carpels of the specimen shown in Fig. 254 (Photo: Xiao-yi Fan/ NIGP)
83
only from fragments of reproductive axes. Later more complete specimens arrangement in pairs or clusters along the axis suggests that each reproductive only from fragments of reproductive axes. Later more complete specimens arrangement in pairs or clusters along the axis suggests that each reproductive were found and an additional species, A. sinensis, was described (Sun et al., axis is an inflorescence with many laterally arranged flowers (Friis et al., were found and an additional species, A. sinensis, was described (Sun et al., axis is an inflorescence with many laterally arranged flowers (Friis et al., 2002). Archaefructus includes small herbaceous plants with axillary branching 2003), rather than a single flower as originally described. Each flower in the 2002). Archae/ructus includes small herbaceous plants with axillary branching 2003), rather than a single flower as originally described. Each flower in the and elongated terminal reproductive axes (Figs. 251"253). Their general inflorescence is simple and unisexual with only few parts. The female flowers and elongated terminal reproductive axes (Figs. 251-253). Their general inflorescence is simple and unisexual with only few partS. The female flowers above consist of one or two carpels and male flowers below one to four habit and morphology, particularly the strongly dissected leaves, indicate habit and morphology, particularly the strongly dissected leaves, indicate above consist of one or two carpels and male flowers below one to four (typically two or three) stamens. The flowers are naked, which may be that the A rchaefructus plants were aquatic. The reproductive axes have that the A rchae/ructus plants were aquatic. The reproductive axes have (typically two or three) stamens. The flowers are naked, which may be ovulate (female) organs above and microsporangiate (male) organs below interpreted as a secondary loss. This is compatible with a submerged aquatic ovulate (female) organs above and microsporangiate (male) organs below interpreted as a secondary loss. This is compatible with a submerged aquatic (Fig. 252). The ovulate organs are often borne in pairs and the microsporannature inferred for the flowers, since flowers underwater do not require (Fig. 252). The ovulate organs are often borne in pairs and the microsporannature inferred for the flowers, since flowers underwater do not require giate organs in clusters of one to four (Figs. 251, 253). The presence of critical perianth parts for protection against desiccation or for attracting pollinators. giate organs in clusters of one to four (Figs. 251, 253). The presence of critical perianth parts for protection against desiccation or for attracting pollinators. Similar simple and naked floral structures are typical for modern angiosperms angiosperm features of Archaefructus has not been fully documented, and angiosperm features of Archae/ructus has not been fully documented, and Similar simple and naked floral structures are typical for modern angiosperms specialized for flowering under water. The position of the inflorescence axes many of the reproductive features are still poorly understood. The morpholmany of the reproductive features are still poorly understood. The morphol- specialized for flowering under water. The position of the inflorescence axes ogy of the reproductive organs indicates that the ovulate organs might be at different levels of the plants (Figs. 251,253) also supports the interpretaogy of the reproductive organs indicates that the ovulate organs might be at different levels of the plants (Figs. 251, 253) also supports the interpretation of submerged flowers as do the lax appearance of some of the reproductive carpels and the microsporangiate organs stamens as in angiosperms. Their tion of submerged flowers as do the lax appearance ofsome of the reproductive carpels and the microsporangiate organs stamens as in angiosperms. Their
IIJ4
258 OrclJidites /ancifo/ius, an uncertain angiosperm (2.2 cm long). from 257 Orchidites an uncertainanangiosperm (6.7 cm long). ! 2 5 7 /inearifolius. Orchiditeslinearifolius, uncertain angiosperm (6.7 cm long),
m 2 5 8 Orchidites lancifolius, an uncertain angiosperm (2.2 cm long), from of Yixian Huangbanjigou locality Uianshangou Bed of Yixian Formation) in Huangbanjigou locality (Jianshangou Bed of Yixian Formation) in Formation) Formation) in Beipiao. in Liaoning (ca. 125 Mal. (photo: Xiao-yi Beipiao, Liaoning (ca. 125 Ma). (Photo: Xiao-yi Beipiao, Liaoning (ca. 125 Ma). (Photo: Xiao-yi Fan! NIGP) Beipiao, Liaoning (ca. 125 Ma). (Photo: Xiao-yi Fan/NIGP) Fan! NIGP) Fan/NIGP)
from Huangbanjigou locality Uianshangou Bed of Yixian from Huangbanjigou locality 0ianshangou Bed
axes (Fig. axes 252).(Fig. 252).
Sinocarpus--an Sinocarpus angiosperm - an angiosperm with advanced with advanced features features Sinocarpus Sinocarpus decussatuscomprises a single fragment an infructescence axis (Figs. axis 254 (Figs. N 256). decussatus comprises a single of fragment ofan infructescence 254~ 256). The specimen is fossilized in the fruiting withstage fruitswith born on long, The specimen is fossilized in the stage fruiting fruits born on long, slender stalks in an opposite 254, 256).254, The256). fruitsThe are fruits from are from slender stalks in an pattern opposite(Figs. pattern (Figs. hypogynous flowers and composed of (3 N) 4 of carpels (3 ~) 4 that carpels are partly that are united. partly united. hypogynous flowers and composed Although Although the fossil is and strongly it is clearlyit from fossil is incomplete and compressed, strongly compressed, is clearly from theincomplete an angiosperm mainly because its united of itscarpels. united United carpels.carpels Unitedare carpels are an angiosperm mainly of because generally generally regarded regarded as a more asderived angiosperms and charactera morefeature derivedinfeature in angiosperms and characteristic for most "higher" plants called eudicots. systematic of the flowering "higher" flowering plants called The eudicots. The systematic isticof forthe most position of Sinocarpus has not been established with certainty, but the fossil of Sinocarpus has not been established with certainty, but the fossil position shows particular to livingplants eudicot have a combination shows particular similaritysimilarity to living eudicot thatplants have that a combination
of derivedofand more and primitive features offeatures angiosperms as seen in as members of derived more primitive of angiosperms seen in members of the Ranunculaceae, Buxaceae,Buxaceae, and Myrothamnaceae. the Ranunculaceae, and Myrothamnaceae.
Other possible The affinity of affinity other putative Other angiosperms possible angiosperms The ofotherangiosperms putative angiosperms in the Jehol Flora is still uncertain. Since many of them fragmentarily ]ehol Flora is still uncertain. Since many are of them are fragmentarily in the preservedpreserved or lack reproductive organs, we can only speculate until more until more or lack reproductive organs, we can only speculate information is available. Among these uncertain angiosperms are the vegetainformation is available. Among these uncertain angiosperms are the vegetative branches of Potamogeton? sp., the "Monocotyledon and the two tive branches of Potamogeton? sp., the "Monocotyledon Leaf", andLeaf", the two 259 A spiny fruit (seed) ofBeipiaoa spinosa, an uncertain angiosperm (1 cm long), of Orchidites (Figs. 257, Allhave of them have that species of species Orchidites (Figs. 257,258). All 258). of them leaves thatleaves in size andin size and _ 259 A spiny fruit (seed) of Beipiaoa spinosa. an uncertain angiosperm (1 em long). from Huangbanjigou locality (Jianshangou Bed of Yixian Formation) in from Huangbanjigou locality Uianshangou Bed of Yixian Formation} in shape, venation arrangement are very similar to similar each other andother and Beipiao, Liaoning shape, pattern venationand pattern and arrangement are very to each (ca. 125 Ma). (Photo: Xiao-yi Fan/NIGP) Beipiao, Liaoning (ca. 125 Ma). (Photo: Xiao-yi Fan! NIGP) may be conspecific. They all display linear andlinear lax leaves with parallel may be conspecific. They alllong, display long, and lax leaves with parallel
venation, venation, and in general they resemble monocotyledons and and inappearance general appearance they some resemble some monocotyledons and aquatic and these and organs may also be produced by ancientby water aquatic angiosperms, these organs may also be produced ancient water also some also modern They wereThey all transferred to the conifers someaquatic modernplants. aquatic plants. were all transferred to theand conifers and angiosperms, plants. so farBut theysohave not been in organicinassociation with plants. far they have discovered not been discovered organic association with boii Sun, et Mei et But included in thetaxon extinct taxon Liaoningocladus included in the extinct Liaoningocladus boii Sun, Zheng et Zheng Mei (Sun et (Sun vegetative structuresstructures and thus their full their naturefull hasnature yet tohas be yet clarified. any vegetative and thus to be clarified. 2001, 2000). supporting Features supporting thewith affinity withare conifers are notany clear al., 2001, al., 2000). Features the affinity conifers not clear
angiosperms in a global angiosperms Jehol angiosperms in context a global Although context Although angiosperms and the special (syndetocheilic) type ofdescribed and the special (syndetocheilic) type of stomata for Liaoningocladus stomata described for LiaoningocladusThe JeholThe are relatively in the arethat relatively rare in therare Jehol Biota and Biota different plantfrom fossils described ]ehol and from different plant fossils described is characteristic for angiosperms It is thusthat possible is characteristic for angiosperms but absentbut in absent conifers.inItconifers. is thus possible otherthey regions, theyinare also interms broader termsagreement in good agreement with from otherfrom regions, are also broader in good with these fossils arefossils indeedare angiosperms as first suggested, but their but systematic as first suggested, their systematic these indeed angiosperms plants from other Barremian-Aptian floras. Small,and simple and few parted plants from other Barremian-Aptian floras. Small, simple few parted affinity stilltoremains to beinstudied in more The angiosperm putative angiosperm affinity still remains be studied more detail. Thedetail. putative
unisexual unisexual flowers asflowers known as forknown Archaefructus are common contemporaneous for Archae/ructus are in common in contemporaneous Liliteswas reheensis was also re-interpreted as and a conifer and Podocarpites renamed Podocarpites Lilites reheensis also re-interpreted as a conifer renamed fromand Europe North and America andwere eudicots were established floras fromfloras Europe Northand America eudicots established by this by this reheensis (Wu) Sun(Wu) et Zheng (Sun et al.,(Sun 2001). However, for a full documenreheensis Sun et Zheng et al., 2001). However, for a full documentimeinatAfrica, least in Africa,and Europe North America. The of presence time at least Europe Northand America. The presence aquaticof aquatic tation of its systematic position, more tation of its systematic position, more details of details venation of pattern, venationepidermal pattern, epidermal plants early in the history angiosperms has also been inferred on based on of angiosperms has also been based inferred plants early in theofhistory and reproductive are needed. structure, structute, and reproductive organs areorgans needed. fossils from otherbut places, so far whole plant evidence for an early fossils from other places, so farbut whole plant evidence for an early Some of the isolated organs from the from YixiantheFormation Some of thereproductive isolated reproductive organs Yixian Formation specialization aquatic environment has only been discovered in the ]ehol specialization to aquatictoenvironment has only been discovered in the Jehol suchdescribed as those described Trapa? sp., Beipiaoa parva, B.and rotunda and B. spinosa such as those as Trapa? assp., Beipiaoaparva, B. rotunda B. spinosa Biota. Biota. have sturdy spines (Fig. 259),(Fig. a feature seenoften in fruits seeds and of extant have sturdy spines 259), often a feature seenand in fruits seeds of extant
..... :r
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,85
SPORfS B~~ POllf~ ~
S
Wen-ben Li Wen-ben Li
pores and pollen are productive organs of plants. Pteridophytes and
some of thearemore primitive plants, such asPteridophyres bryophytes, algae pores and pollen productive organs of plants. and and fungi, produce spores byplants, which such new generation of plant is reproduced some of the more primitive as bryophytes, algae and sexually or nonsexually. (spermatophytes), gymnofungi, produce spores by Seed whichplants new generation of plant isincluding reproduced
sexuallysperms or nonsexually. Seed plants (spermatophytes), gymnoand angiosperms, produce pollen. Sperms including of pollen (male spores) sperms combine and angiosperms, pollen. pollen spores) with ova ofproduce female ovule andSperms developof into seeds.(male The reproduction female ovule and develop seeds. The The study reproduction combine ova of depends ofwith seed plants on germination ofinro the seeds. of spores and
2BO Densoisporites microrugulatus, spore of a selaginellales. "260 Densoisporites microruguJatus, spore of a selaginellales.
of seed pollen plantsisdepends on germination thepollen seeds.are The srudy and called palynology. Sporesof and very smallofinspores size, commonly called200 palynology. pores pollen are very small in size, commonly pollen isunder microns, but veryand large in quantity. Because of their organic antimicrons, but very large they in quantity. Because of their organic antiunder 200 decomposition wall (exine), can be well preserved together with mineral decomposition wall (exine), they can be well pre erved rogether with mineral
clastics in the sediments through the geological history. In order to extract
clastics in the sediments through the geological hisrory. In order ro extract
spores and pollen from the sediments, the rock containing spores and pollen
spores and pollen from the sediments, the rock containing spores and pollen
must be generally macerated in nitric and hydrofluoric solutions to separate
~G
must be generally macerated in nitric and hydrofluoric solutions ro separate
the spores and pollen from mineral clastics. After collected from the rock
the spores and pollen from mineral clastics. After colleered from the rock
18fJ
residue, the spores and pollen are mounted with glycerine jelly on glass slides.
residue, the spores and pollen are mounted with glycerine jelly on glass slides.
We can observe and identify them under microscope at power from 600 to
We can observe and identify them under microscope at power from 600 ro
1000 enlargements. Generally speaking, the spores and pollen from different 1000 enlargements. Generally speaking, the spores and pollen from different plant taxadifferent show different morphology, including ofexinal their exinal structure plant taxa show morphology, including that ofthat their strucrure m261 Tenuangulasporis microverrucosus, spore of a possible selaginellales. . . 261 TenuanguJasporis microverrucosus, spore of a possible selaginellales.
and ornamentation. By means of palynological taxonomic wewell can well By means of palynological taxonomic srudy,study, we can and ornamentation. reconstruct the vegetation, and environment, and the dateage thewhen age when the vegetation, climateclimate and environment, and date reconstruct theygrowing. were growing. they were The Yixian Formation cropping at Jianshangou and Huangbanjigou The Yixian Formation cropping Out at out Jianshangou and Huangbanjigou of Beipiao, Zaocishan of Yixian and Sunjialing, Sanguanmiao and Gujialing ofBeipiao, Zaocishan ofYixian and unjialing, anguanmiao and Gujialing of Harqin western Liaoning abundant and pollen of Harqin Zuoyi, Zuoyi, western Liaoning yields yields abundant fossil fossil sporesspores and pollen with about 70 known speciesspecies assigned ro 40 to genera. MostMost (Figs. 260-268), (Figs. 260---268), with about 70 known assigned 40 genera. the whole specie species in the spore-pollen assemblage are common taxa through in the spore-pollen assemblage are common taxa through the whole Mesozoic era, forera, exampIe, the spores ofStereisporite.r antiqllasporites ofbryophytes, Mesozoic for example, the spores ofStereisporites antiquasporites ofbryophytes, Lycopodiltmspol'ite.r austroc!avatidite.r, Leptolepidite.r verrltcosus, Neol'aistrickia equalis, Lycopodiumsporites austroclavatidites, Leptolepidites verrucosus, Neoraistrickia equalis,
Densoisporites micromgulatlts of lycopodiales, C)'athidites minor,minor, Osmundacidites Densoisporites microrugulatus of lycopodiales, Cyathidites Osmundacidites wellmanii, Klukisporites psettc/oretiCillatlts and BaCillatispo1'ites comalllllemis of of wellmanii, Klukisporites pseudoreticulatus and Baculatisporites comaumensis filicales, and the grainsgrains of Classopollis anmtlatltS, GinkgoC)'cadophytltS filicales, andpollen the pollen of Classopollis annulatus, Ginkgocycadophytus
262 Osmundacidites wellmanii, spore of a filicales.
. . 262 Osmundacidites wel/manii, spore of a filicales.
nitidllS,nitidus, Ephedripites sp., Caytonipollenites pallidllS, Perinopollenites elatoides, Ephedripites sp., Caytonipollenites pallidus, Perinopollenites elatoides, PinltSpollenites divltlgatltS, Abietineaepollenite.rpectinelllts, Ceclripite.r pmillLtS, Cedripite.r Pin uspollenites divu lgatus , A bietineaepollenitespectinellus , Cedripitespus illus , Cedripites
nj263 Cicatricosisporitespacificus, ~264 Schizaeoisporitescertus, spore of a filicales. 264 Srhizaeoi porites rertus. spore of a filicales . ... 263 Cicatricosisporites parifiCII • spore of a filicales.
aju265 Pinuspollenitesdivulgatus, pollen of a gymnosperm. .265 Pinuspollenites divu/gatus, pollen of a gymnosperm.
spore of a filicales.
microsaccoides,Piceaepollenitesspp., Abiespollenites spp., Podocarpiditesmultesimus, we have not found a single reliable angiosperm pollen grain from the whole mirrosaeeoides, Pieeaepollenites spp., Abiespollenites spp., Podocatpidites mllitesirr//(s, we have not found a single reliable angiosperm pollen grain from the whole Podocarpidites ornatus, Callialasporites dampieri, Bicestopollis wulanensis, Yixian Formation. In consideration of the infrequent appearance of the Podoealpidites omatm, CatliaLaspol'l'tes dampieri, Bicestopotlis wlILctnemis, Yixian Formation. In consideration of the infrequent appearance of the Quadraeculina limbata, Protoconiferus funarius, Protopinus sp., Pseudopicea genera Cicatricosisporites, Schizaeoisporites, Tenuangulasporis andJiaohepollis, the QlladraeClilina limbata, Protoeonifems fimarim ProtopinllS sp., Pselldopicea genera Cieatrirosisporites, rhizaeoisporites, TenllangllLasporis andJiaohepollis, the variabiliformis and Pseudopicea rotundiformis of gymnosperms. But some others age of the Yixian Formation should be of the earliest Cretaceous (Berriasian), 7/ariabilifomlis and Pselldopleea rotlllldiformis of gymnosperms. But some others age of the Yixian Formation should be of the earliest Cretaceous (Berriasian), are are species rising fromfrom the Cretaceous period, such such as Cicatricosisporites 13 5"-- 131 millionmillion years before species rising the Cretaceous period, as Cieatriwsisporites l35-l3l year present. before present. australiensis, Cicatricosisporites pacificus, Schizaeoisporites certuscerllls of lygodiaceae, of xerophytic cheirolepidiaceae, usually usually amountsamounts to cmstraLiewis, ieatl'irosisporites paeifiCIIs, Sehizaeoisporites of Iygodiaceae, The Classopollis The ClassopoLLis of xerophytic cheirolepidiaceae, to Cyclocristella senticosa, Tenuangulasporis qiuchengensis, Tenuangulasporis 15---91% in content, dominating the spore-pollen assemblage of the latest Cyc!oC1'istella sentieosa, Tenllalzglliasporis qillehengemis, Tenllanglliasporis l5 -9l % in content, dominating the spore-pollen assemblage of the latest microverrucosus of selaginellales andJugella claribaculata,Jiaohepollis flexuosus Jurassic Houcheng Formation in Xuanhua and Wanquan, northernnorthern Hebei Hebei micrwermroslls of selaginellales andJlIgella claribaCIILata ,Jiaohepollis j1exlloS/ls Juras ic Houcheng Formation in Xuanhua and Wanquan, andJiaohepollis aml/ilatltS of gymnosperms. Province. In contrast, ClassopoLiis andarid-tolerating other arid-tOlerating taxa, Sehizaeoisporites andJiaohepollis annulatus of gymnosperms. Province. In contrast, Classopollis and other taxa, Schizaeoisporites Palynological demonstrates that during the accumulation time and of Ephedripites, and Ephed,-ipites, in the Yixian Formation, only2% under 2% in content. Palynological studystudy demonstrates that during the accumulation time of are rareare in rare the Yixian Formation, only under in content. deposition ofYixian the Yixian Formation, the main part the vegetation in western Thi indicates faCt indicates the climate from arid Late Juras ic into the deposition of the Formation, the main part of theof vegetation in western This fact the climate changechange from the aridthe Late Jurassic into the composed of conifers that occupies over % of the Liaoning Province relatively Early Cretaceous in northern Hebei and western Liaoning Province was was composed of conifers that occupies over 90 % 90 of the relatively humidhumid Early Cretaceous in northern Hebei and western LiaoningLiaoning Provinces. whole vegetation. Under the coniferous t some bryophytes, pteridoProvinces. whole vegetation. Under the coniferous forestfore some bryophytes, pteridophytes cycadophytes sparAngiosperm ely. Angio perm megafossil plant inocClIpfIJ (All specimens this chapter thepart upper part of the (All specimens shown shown in this in chapter are fromarethefrom upper of the phytes and and cycadophytes grewgrew sparsely. megafossil plant Sinocarpus deC/matm and several other purported ones were recorded from different Yixian Formation in anguanmiao, Harqin Zuoyi Mongolian Autonomous Yixian Formation in Sanguanmiao, Harqin Zuoyi Mongolian Autonomous decussatus and several other purported ones were recorded from different horizons of the Yi.."ian Formation (see the
hapter" Angio perms") but so far
horizons of the Yixian Formation (see the Chapter "Angiosperms"), but so far
266 Podocarpidites ornatus, pollen of a gymnosperm.
266 Podocarpiditesornatus, pollen of a gymnosperm.
County, Liaoning Province, and photOgraphed by Wen-ben Lil
County, Liaoning Province, and photographed by Wen-ben Li/NIGP)
267 Proloroniferusfullarius, pollen ofa gym no penn.
IGP)
268 jugeJla c1aribnrulata. pollen ofa gymnospeml.
267 Protoconiferusfunarius,pollen of a gymnosperm. ~ 268 Jugellaclaribaculata,pollenof a gymnosperm.
I
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Ul
187
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Anlmana Phylum Mollusca Cla Ss Ga stropod a . . . . .
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Animalia ou ' ~" ~-permmfl,, lng'? onioid-"-acea :
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Nakamuranaiaehingshanensis uperfamily Trig nioida ea (Grabau, 1923) ~
CiaFamily Ga trop a Cyclophoridae 1847
....
ubcLa Pro obranchia Pseudarinia yushugouensis Zhu, 1976 Cycl phMuller, rida 1847 r ~Family l y Valvatidae 1774
. . . .
Family VaJvatida Fischer, Muller,1885 1774 Family Hy&obidae Amplovalvata Reesidetla . . sp. . . p.
baya hi, Family Plicatqingquanensis um nida K --Wang, Weichangella i982196' Weichangel/a angulari Wang, 19 21980 .... Weichangella shalingouensis Yu et Yao; Weichan ella qin quanen i Wang, J9 2 Subclass Heterodonta Weichan ella halingouen i Yu et Yao 19 0 Order Cyrenodonta "
....
~
Famil Hydr bida
Family Micromelaniidae Brusina, 1874 ... Ree idella p. Probaicalia gerassimovi (Reis, i 9 1 0 ) : i r m laniid e Bru
Family ippononaiidae hi 1976 196 Nakamuranaia subrotundaKGubaya et Ma, ching hanen i1968 Grabau, 192 Family Nakamuranaia Plicatounionidae Kobayashi, ...... akamuranaia uhrotunda Gu et Weichangella angularisWang, 1982 Ma, 1976 .....
•
P eudariniasp'ushugouen is Zhu 1976 amplovalvata
Family
.
Family N ippo nona iidae Kobayashi' 1 9 6 8
SubclassMollu erosobr~chia Phylum ca
.
~
.
in~
.
ubcla
SuperfamilyC0rbiculacea
L874
.
.
.
Het rodonta
....
Ord r Cyrenod nta
Probaicalia vitimensis Maainson~ 1949, Probai alia g rassimovi Rei , 1910) Subclass P~monata . . . . . Probai alia vitimel i Martin on 1949 F a ~ l y Ellobiidae Bolten, 1 7 9 8
Family: Pisidiidae Gray,: 1857 uperfamily
...
~
orbicula ea
Sphaerium anderssoni (Grabau, 1923)
.
Family Pi idiidae Gra , I 57
Sphaerium jeholense (Grabau, 1923) ubcl Pulmonata phaerium andel's oni Grabau, 192 ) P~chos~lus philippii (Dunker, 1 8 4 6 ) S p h a e r i u m pujiangense et Ma, 1976 phaerium jeholenseGuGrabau, 192 ) Family Ell biidae Bolten, 179 P~chos~lus 1837) .... Ptycho tyluharpaeformis(Koch philippii Dunker et1 Dunker, 46) phaerium pujian en e Gu el Ma, 1976 Zap~chius sp, harpaefonni K h et Dunker, I 7) .... Phylum A ~ o p o d a . Pty ho tylu Family Lymnaeidae Zoprychiu p. Brodefip, 1839 Galbasphaira Pan, 1983 Family Lymnaeida Broderip, 1839
Class Cmstacea Phylum Arthropoda Subclass BranChiopoda I
Galba sphaira Pan, 19 1767 Family Planorbidae Geffroy, Gyraulus sp. rbida G ffro , 1767 Family Plan Gyraufuloryi p. Coquand, 1855 Gyraulus Gyraulus fory'i
.
.......
Class BiValvia ..... ....
......
uperfamiJy ni Fle~ng, nac a Family Unionidae 1828 Family Unionidae Heming,(Grabau, I 2 Mengyinaia mengyinensis 1 9 2 Mengyinaia meng ,in 11 i Grabau, 1923)
....
. . . . Order ub Conchostraca la Bran hiopoda Family Eosestheriidae Order Concho traca Zhang et Chen, 1 9 7 6
oquand, I 55
Cia Bi Palaeoheterodonta al ia SubClass Palae h Stoliczka ler nta Order Unionoida Order licz Superfamily Unionacea a
......
3
Mengyinaia shifoensis Yu, Dong et Yao, 1989 Mengyinaia hifo 11 is Yu, D ng et Yao, J 9 Mengyinaia tugrigensis (M~inson, 196I) Mengyinaia tu rigensi (Martin n 1961 Family Sibireconchidae Kolesnikov, 1977 Family ibire on hidae Kole niko 1977 Arguniella lingyuanensis (Gu, 1976) Arguniella ling 'uanen is (Gu 1976) Arguniella yanshanensis (Gu, 1976) Arguniella yan hanen i (Gu, 1976)
)
~ ~
Family Eo erotunda theriidae Zh ng Abrestheria Wang, 1981then, 1976 Allestheria striata Shen et Chen, 1982 hrestheria rolllnda Wang, 1981 llestheria h n et(Jones, hen, I862) 19 2 Eosestheria aft.striata middendo~i Eo e theria aff. middendorfii EosestheriafuxinensisChen, 1976(lone, I 62) Eo e theria figujialingensis inen i hen, 1976 i987) Eosestheriopsis (Wang, Eo e theriopsi ujialingen i Eosestheria jingangshanensis Chen,Wang, 1976 1987 Eo e theria jin an hanen hen, 1976 Eosestheria lingyuanensis Chen;is1976 Eo e theria Jin 'uanen1976) i hen 197 Eosestheria ovata(fhen, Eo e theria ovata ( hen, 1976 Eosestheria peipiaoensis (Kobayashi et Kuzumi, 1953) Eo e theria peipiaoen is K bay hi el Kuzumi, 195 Eosestheria subrotunda Chen, 1976 Eo estheria subroumda hen, 1976 Yanjiestheria jiufotangensis (Chen, 1976) Yanjiestheria jiufotan en i hen, 1976) Yanjiestheria beipiaoensis Chen, 1999 Yanjiestheria beipiaoen i h n, 1999 Yanshania xishunjingensis "Wang, 1981 Yan hania xi hunjingen is Wang 19 I
197
Yumenestheria delicatula Shen et Chen, 1982 Family Diestheriidae Zhang et Chen, 1976 Diestheria jeholensis(Kobayashi et Kuzumi, 1953) Yumene Theria delieatu/a hen et Chen, hen, 19 2 Diestheria longinqua 1976 Family Die th Diestheria riida Zhang et Ch n, Chen, J976 1976 yixianensis DiesTheriajeholen is (Kobaya hi et Kuzumi, I 53 Family Loxomegaglyptidae Novojilov, 1950 Die theria longinqua hen, J976 Ambonella lepida Wang, 1981 Die theria yixianen i hen 1976 Nestoria dabeigouensis Wang, 1981 Family Lo om gaglyptida 0 ojilo . 1950 Nestoria pissovi Krasinetz, 1962 Ambonella /epida ang, 198 J Family Sinoestheriidae Chen et Shen, 1982 Ne IOria dabeigouensi Wang, 19 1 Sentestheria banjietaensis Wang, 1981 Ne IOria pi ol'i Kr inetz, 1962 Sentestheria weichangensis Wang, 1981 Family inoe theriidae hen et Sh n. 19 2 Family Ipsiloniidae Novojilov, ente Theria banjietaen is Wang, 1981 1958 Wang, 1981 ente theria Keratestheria weiehan en i gigantea Wang, J981 Keratestheria Family Ip iloniidae 0 ojilo longa , J95 Wang, 1981 Family KeraTe theria i Palaeolimnadiidae anTea Wang, 1981 Tasch, 1956 KeraTestheriaJibeilimnadia /onga Wang, ovata 19 I Wang, 1981 Family PaJa olimnadiidae Ta ch, 1956
!
lg8 198
Jibeilimnadio ovalO Wang, 19 1 Subclass Ostracoda
Order Podocopida ubcla 0 tra oda Superfamily Cypridacea Order Podocopida Family Cyclocyprididae Kaufmann, 1900 uperfamily ypridacea Subfamily Cyclocypridinae Kaufmann, 1900 Family
yclocyprididae Kaufmann 1900
Damonella extenda Wu et Yang, 1980 Damonella formosa Cao, 1999 Damonella extenda Wu t Yang, 19 0 Damonella subsymmetraca Zhang, 1985 Damol/ellaformo a ao, 1999 Ziziphocypris cosdata (Galeeva, 1955) Damonella ub 'mllletroea Zhang, L9 5 Ziziphocypris linchengensis Suet Li, 1981 Ziziphocypri co daTa (Galeeva, 195 Ziziphocypris simacovi (Mandelstam, 1955) Ziziphoc pri linchengen is u t Li, 19 Family Cyprididae Baird, 1845 Zhphocypris imacovi (Mandel tam, 1955 Subfamily Cyprideinae Martin, 1940 Family ypridida Baird, 1845 Cheilocypridea trapezoidea Zhang, 1985 ubfarnily yprideinae Manin, 1940 Cypridea (Cypridea) altidorsangulata Pang, 1984 hei/ocypridea Trapezoidea Zhang, 19 5 (Cypridea) dabeigouensis ypridea C Cypridea prideo) a/Tidor angu/aTa Pang. 19 4Yang, 1981 (Cypridea) fingangshanensis Zhang, 1985 ypridea (C Cypridea 'pridea) dabeigouen i Yang 198J Cyprideajingangshal/el/si (Cypridea) liaoningensis Zhang, 1985 ypridea C)pridea) Zhang, 19 5 (Cypridea)isluanpingensis Cypridea C Cypridea 'pridea) /iaoningen Zhang, 19 5 Pang,1984 Cypridea (Cypridea) obliquoblonga Cypridea Cypridea) /uanpingen is Pang,19 4 Pang, 1984 (Cypridea) prognata Lubimova, 1956 Cypridea C Cypridea pridea ob/iquob/onga Pang. 19 4 (Cypridea) sihetunensis ypridea (C Cypridea 'pridea) prognaTa Lubimova, J956Cao, 1999 ypridea C Cypridea pridea) ihetune/lSi a. 19 9 Pang, 1984 (Cypridea) subgranulosa C,}pridea (C Cypridea 'pridea) ubgra11ulo Pang, 19 4 1985 (Cypridea) atersa Zhang, ubfamily Cy locypridinae Kaufmann, 1900
Cypridea C pridea) ter a Zhang, 198
Cypridea (Cypridea) tubercularis Pang, 1984 Cypridea (Cypridea) unicostata Galeeva, 1955 Cypridea (Cypridea) vitimensis Mandelstam, 1955 Cypridea ( Cypridea ypridea) (Tuben u/arizaocishanensis Pang, 19 4 Zhang, 1985 Cypridea) pridea ( Cypridea ypridea) (unieo ToTa beipiaoensis Galee a, 1955 Ulwellia) Cao, 1999 pridea ( }prideo) viTimen i Mandel tam, 1955 Cypridea ( Ulwellia) koskulensis Mandelstam, 1958 Cypridea ( )1Jridea zao ishonensi Zhang 19 5 Cypridea (Ulwellia) muriculata Zhang, 1985 C pridea (U/wellia) beipiaoensi Cao. L999 Cypridea ( Ulwellia) regia Lubimova, 1956 Cyprideo (U/wellia) ko ku/ensi Mandel tam, 195 Cypridea (Ulwellia) subeIongata Zhang, 1985 ypridea (Ulwellia) murieulaTa Zhang 19 5 Djungarica camarata Zhang, 1985 ypridea (Ulwellio) regia Lubimova, 1956 Djungarica circulitriangula Zhang, 1985 ypridea Ulwellia) ubelongaTo Zhang, 19 5 procurva 1985 DjungaricoDjungarica camaraTa Zhang, 19 Zhang, 5 1956 DjungoricaLimnocypridea circu/iTriangu/aabscondida Zhang 19 Lubimova, 5 grammi DjungaricoLimnocypridea procurva Zhang, 19 5 Lubimova, 1956 Limnocypridea Zhang, 1985 Linmoc)pridea ab condidaposticontracta Lubimo a, 1956 Limnocypridea rara Zhang, LiIlIllO 'pridea grammi Lubimova, L9561985 Limnoc 'pridea posTicontra redunca Ta Zhang, 19 5 1985 Limnocypridea Zhang, Limno pridea rara Zhang, 19 Limnocypridea tulongshanensis Zhang, 1985 Limlloc 'pridea redLIII a Zhang, 198Yang, 1981 Luanpingella postacuta Limnoc 'prideo TU/ongshanensi Zhang, 1985 Mongolianella palmosa Mandelstam, 1955 Luanpingella po Ta uTa Yang, 19 1 Mongolianella subtrapezoidea Yang, 1981 Mongo/ial/ella pa/mo a Mandel tam, 1955 Yanshanina dabeigouensis (Yang, 1981) Mon o/ianella ubTrapezoidea Yang. 19 1 Yumenia casta Zhang, 1985 Yanshanina dabeigouensi Yang, 19 I) Yumenia jianchangensis (Su et Li, 1985) Yum nia asta Zhang, 19 5 Subfamily Cypridinae Baird, 1845 Yumenia jian hangensis ( u et Li, 19 ) Lycopterocypris infantinis Lubimova, 1956 ubfamily ypridina Baird I 45 Mantelliana sp. Lyeoptero pri infanTini Lubimova, 1956 Torinina obesa (Pang, 1984) Mall1elliana p. Torinina tersa Sinitsa, 1992 Torinina obesa Pang, I 4) Yixianella Zhang, 1985 Torinina Tersa ini a marginulata 1992 Ilyocypridae 1900 Y'uianellaFamily mar i11ulaTa Zhang,Kaufmann, I 8 Subfamily Ilyocyprinae Kaufmann, 1900 Famil lIyocypridae Kaufmann, 1900 Rhinocyprisjurassica ubfamily Ilyo yprinae Kaufmann. (Martin, 1900 1940) Superfamily Rhinoeypri jurassieaDarwinulacea (Martin, 1940) Darwinulidae Brady et Norman, 1889 up rfamily Family Dan inulacea amily Darwinulidae Brady et orman, I 89 1855) Darwinula leguminella (Forbes, Dam'inu/a /eguminella (Forbe ,(Roemer, I 55) 1839) Darwinula oblonga Dam'inu/a ob/ongo (Roemer, I 39) Superfamily Cytheracea lip rfamily Family ytheracea Limnocytheridae Klie, 1933 Family Limno Timiriasevia ytherida Klie, 1933Zhang, 1985 eminula Timiria eviaTimiriaseviajianshangouensis eminu/a Zhang, 19 5 Zhang, 1985 Timiria eviaTimiriasevia jion hangoupolymorpha IlSis Zhang,Mandelstam, 19 5 1955 Till/iria el'ia po/ymorpha Mand I tam, 1955
Liaonemobius tanae Ren, 1998
Subclass Malacostraca
Pseudacrida costata Lin, 1982
Order Decapoda
Sinohagla pleioneura Ren, 1995
Superfamily Astacoidea Family Cricoidoscelosidae Taylor, Schram et Shen, 1999 (emend.) ubel ala 0 tra a Cricoidoscelosus aethus Taylor, Schram et Shen, 1999 Order D capoda
Palaeocambarus licenti (Van Straelen, 1928) Taylor, Schram et uperfamily id Shen, 1999 Famil ri oid
c I
idae Ta I r
brarn et h n, 1999 (emend.)
celo u a tllU Tayl r, chrarn et hen, 19 9 Order Cricoido Hemicaridea Palaeo ambaru Iicenti ( an 1957 traelen, 192 ) Ta I r, chram et Family Spelaeogriphidae Gorden, hen, I 99 Liaoningogriphus quadripartitus Shen, Taylor et Schram, 1998 Order Hemicaridea
Class Family Arachnidapela ogriphidae G rd n, 1957 Order Liaoningo Araneida riphus quadripartitu hen, Taylor et
bram, 199
Family Araneidae Koch et Berendt, 1854 rachnida Araenidae indet. Order Aran ida ran ida K h t Berendt I 54 Famil Class Insecta Araerudae indet. Order Ephemeroptera la
Ephemeropsis trisetalis Eichwald, 1864
las In ta Order Odonata Order Ephem roptera
Aeschnidium Ephemerop heishankowense i tri etalis Eich (Hong, ald, I 1965) 64 Chrysogomphus beipiaoensis Ren, 1994 Order Odonata Congqingia rhora 1992 Ae c1midiwn hei Zhang, hankowen e (Hong, I 65) Liogomphus yixianensis Ren iet Ren, Guo, 1994 1996 Chrysogomphu beipiaoen Mesocordulia boreala Ren et1992 Guo, 1996 Congqingia rhora Zhang, Rudiaeschna limnobia Ren et 1996 Liogomphu ixianen i RenGuo, et Guo 1996 Me ocordulia bor. ala Zhang Ren et et Guo, I 96 Stylaeschnidium rarum Zhang, 2001 Iimllobia Ren et Guo, 1996 OrderRudiaesclma Blattaria tylae ehnidium Blattula exetenuataramm Ren, Zhang 1995 et Zhang, 2001 OrderBlattula Blattaria platypa Ren, 1995
Blattuladelicatula exet nuata Ren1995 199 Blattula Ren, Blattula plarypa Ren, 19Ren, 1995 Karatauoblatta formosa Blattllla delicawla n, 1995 Laiyangia delicatulaRZhang, 1985 Kararalloblattaformosa Ren, J 995 Nipponoblatta acerba Ren, 1995 Lai 'an ia delieatula Zhang, 19 Parablattula cretacea (Hong, 1982) ipponoblatta a erba Ren 19 5 Order Dermaptera Parablattula creta ea (Hong, 19 2 Archaeosoma serratum Zhang, 1993 Ord r Dermaptera
Longicerciata mesozoica Zhang, 1993 Archaeo oma erratum Zhang, 1993 Sinostaphylina nanligezhuangensis (Hong et Wang, 1990) Longicerciata m a-oiea Zhang, 199 Order Orthoptera iTlo taph lina nanligevzuan en i (H ng et Wang, I 90) Falsirameus ravus Zhang, 1985 Order Onh ptera Habrohagla curtivenata Ren, 1995 Fal irameu rawls Zhang, 19 5 Habrohagla curtivenata Ren, 1995
Order Phasmatodea Liaonemobius ranae Ren, 19 Aethephasma megista Ren, 1997 P eudacrida 0 tara Lin 19 2 Hagiphasma paradoxa Ren,R 1997 inohagla pleioneura n, I 9
Orephasma eumorpha Ren, 1997 Pha matodea Order Order Hemiptera Aethephasma megisra Ren, 1997 Ha ipha rna parado_ a Ren, 97 1998 Anomoscytina anomala Ren, Yin etI Dou,
Orephasma eUlILorpha 97 1998 Anthoscytina aphthosa Ren,Ren, Yin etI Dou, Order Hemjpt ra Caudaphis spinalis Zhang, Zhang, Hou et Ma, 1989 Anolllo cytina anomala Ren Yin et D u, 199 Clypostemma xyphiala Popov, 1964 Anrho decorus C) tina aphtho a Ren, Yin et D u, 199 Lapicixius Ren, Yin et Dou, 1998 audaphis pinalis Liaocossus hui Ren, Yin Zhang, et Dou,Zhang, 1998 Hou et a, 19 9 Cl 'po temma • phiala P po , I 64 Liaocossus beipiaoensis Ren, Yin et Dou, 1998 Lapieixiu decort Ren, Yin et D u, 199 Liaocossus exiguus Ren, Yin et Dou, 1998 Liaoeo u IlUi Ren, Yin et Dou, 199 Liaocossus fengningensis Ren, Yin et Dou, 1998 Liaoeo u beipiaoensi R n, Yin et D u, 199 Liaocossus pingquanensis Ren, Yin et Dou, 1998 Liaoeos u exi uu R n Yin et Dou, 199 Mesanthocoris brunneus Hong et Wang, 1990 Liaoeo u fengningeTl i Ren, Yin et Dou, J99 Mesolygaeus laiyangensis Ping, 1928 Liaoeo su pillgquanen i Ren, Yin et D u, 199 Miracossus ingentius Ren, Yin et Dou, 1998 Me aflfho ori brullneu H ng et ang. 1990 Paroviparosiphum opimum Zhang, Zhang, Me olygaeu laiyangen i Ping, 192 Hou et Ma, 1989 Pauropentacoris macrurata Ren, Zhu Lu, 1998 1995 Mira 0 us in entiu Ren, Yin et etDou, Schizopterax shandongensis Hong, 1984Zhang, Hou et Ma 19 9 Parol'iparo iphwn opimum Zhang Sinaphis epichare Zhang, Zhang, Ren Hou et Ma, Pauropenraeoris maerurara Zhu et 1989 Lu 19 5 Sinojassus brevispinalis Zhang, 1985 ehi-opterax handongen i Hong, 19 4 Sinoviparosiphum lini Ren, 1995 inaphi epiehare Zhang, Zhang H u a, 19 9 Yanocossus Ren, 1995 Zhang 19 5 inojas guoi u brevispinali Order Coleoptera inoviparo iphum lini R n, 19 5
Coptoclava Ping,1995 1928 Yano 0 longipoda u guoi Ren, Ord r 01 opt chifengensis ra Cretihaliplus Ren, Zhu et Lu, 1995
Coptoclava longipoda Ping, Cretihaliplus sidaojingensis Ren, 192 Zhu et Lu, 1995 Crerihaliplu chifengen n, Zhu Geotrupoides fortus Ren, Zhui etR Lu, 1995et Lu, 199 retihaliplu idaojin en1984 i Ren, Zhu et Lu 1995 Glypta qingshilaensis Hong, Geotrupoides fortus Ren Zhu Lu 1992 199 Hesterniasca obesa Zhang, Wang etetXu, Gl 'Pta qingshilaen i H ng, Holcoribeus evittatus Zhang, 1992I 4 Hesternia a obe Zhang, ang et u, I 92 Notocupes laetus (Lin,a1982) Holeorib u evittatu Zhang, 92 Notocupes tuanwangensis (Hong etI Wang, 1990) 010 upe laew Lin, 19 2) Ovidytes gaoi Ren, Zhu et Lu, 1995 010 upe tuanwangen i (Hong tang. 1990) Palaeoendomychus gymnus Zhang, 1992 Ovid 're gaoi R n, Zhu et Lu, 1995 Tetraphalerus lentus Ren, 1995 Palaeoendomychu gymnu Zhang, 1992 Sinosornia longiantenna Zhang, 1992 Tetraphaleru lentu Ren, 19 inosomia Ion iantenna Zhang 1992
Igg
Order Neuroptera
Allopterus luianus Zhang, 1990 Choromyrmeleon othneius Ren et Guo, 1996 Ord r europlera sinica et Hong, 1990 1I0pferu Drakochrysa luianLi Zhang, 19 Yang 0 Kalligramma liaoningensis et Guo, 1996 horolllyrmeleon ot/lIleiLi Ren et Gu Ren . \996 Lasiosmylus newi Drakochrysa ini a Yang el Ren Honget Guo, 1990 1996 Lembochrysa Kalligramma liaonin ellminiscula i Ren etRen Gu et. \Guo, 6 1996 La io m 'luLembochrysa newi Ren etpolyneura Guo, 199Ren et Guo, 1996 miratRen, LembochryLimnogramma a miniscula Ren Guo,2003 \996 yangi Ren,, 1995 LembochryMesascalaphus a polynellra Ren el Gu 199 Lilllnogralllma mira Ren, 200Ren et Guo, 1996 Oloberotha sinica Me a alap/ILI yan i Ren, 1995Ren, 2003 Oregramma gloriosa OloberOfhaSiniphes sini a Rdelicates n t Gu Ren ,I et6Yin, 2002 Oregramma Iorio a R n.eucalla 200 Ren et Guo, 1996 Sophogramma iniphe deli ates Ren t papilionacea Yin, 2002 Ren et Guo, 1996 Sophogramma opho ralllma eLicalla R n t Gu , 19 Ren 6 et Guo, 1996 Sophogramma plecophlebia opho ramma papilionacea Ren etRen, Gu 1995 , 199 Yanosmylus rarivenatus Sophogralllma plecoplzlebia Ren et Guo, 1996 Order Raphidioptera Yallo m 'Ius rari\'ellafll R n, 1995
Alloraphidia anomala Ren, 1997 Alloraphidia longistigmosa Ren, 1994 1I0raphidia anomala Ren, 1997 Baissoptera grandis Ren, 1995 1I0raphidia Lon i ti mo a Ren, 1994 Baissoptera euneura Ren, 1997 Bai sopfera randis Ren, 1995 Caloraphidia glossophylla Ren, 1997 Bai soptera eunellra Ren 1997 Mesoraphidia amoena Ren, 1997 aloraphidia lo oph.vLla R n 1997 Mesoraphidia heteroneura Ren, 1997 Me oraphidia amoena R n, 1997 Mesoraphidia sinica Ren, 1997 Me oraphidia heteroneLira Ren, 1997 Mioraphidia furcivenata (Ren, 1995) Me oraphidia inica Ren, 1997 Phiradia myrioneura Ren, 1997 Mioraphidiafurcivenata Ren, 199 Rudiraphidia liaoningensis (Ren, 1994) Phiradia m 'rioneLira Ren, 1997 Siboptera 1994) Rudiraphidia liaonillfornicata ell i R (Ren, n, 1994) Xynoraphidia (Ren, 1994) Siboptera fomicata Ren.polyphlebia 1994) Xynoraphidia shangyuanensis (Ren, 1994) X 'noraphidia polyphlebia (Ren, 1 4 Yanoraphidia gaoii Ren, X '1lOraphidia shangyLianen (R n,1995 I 94)
Ord r Raphidioptera
Order Mecoptera Yanoraphidia aoi Ren, 19 5 e
pteraLiaobittacus longantennatus Ren, 1993
Megabittacus colosseus Ren, 1997 LiaobitfacLis Ion anfellnatLI Ren, 1993 Megabittacus beipiaoensis Ren, 1997 Megabittacu colo ells Ren, 1997 liaoningensis MegabittacuOrthophlebia beipiaoen is Ren, 1 7 Ren, 1997 Sibirobittacus Orthophlebia liaollin en atalus is R n,Ren, 1 971997 Yanorthophlebia hebeiensis Ren, 1995 ibirobittacLis ataLu Ren, 1997 Order Trichoptera Yanorthophlebia hebeien i R n, \ 9 Ord r Trich ptera
Multimodus dissitus Ren, 1995 Multimodus stigmaeus Ren, 1995 Multimodus ? elongatus Ren, 1995 MLiltimodu dis itLi Ren, \995 Tuanwangica aethoneura Zhang, 1985
MLiLtimodu
tigmaeLis Ren \99
Order Diptera
MLiLtimodu , elon atLi Ren, 19 5
xingi Ren, 1995 TuanwanAlleremonomus ica aethoneLira Zhang, 19 5 Ord r Dipt raAlleremonomus liaoningensis Ren, 1995
AllomyiaxiII ruderalis lIeremonomLi i R n, Ren, 199 1998 Archisolva cupressai Zhang, Zhang et Li, 1993 lIeremononw Liaoningen Ren, 1995 Ren, 1995 ALtomyiaAtalosciophila ruderali R n,yanensis 199
Basilorhagio venustus Ren,et1995 Archi oll'a cupre a Zhang, Zhang Li, \993 Chironomaptera talo ciophila yanen i Rgregaria n, 1995(Grabau, 1923) Ba ilorha io venu tLi Ren, 1995 Chironomaptera vesca Kalugina, 1976 hironomaptera regaria 1 2 Eopangonius pletus(Grabau Ren, 1998 hironomaptera ve ca Kalugina, 1976Ren, 1998 Florinemestrius pulcherrimus Eopangonius pletueucalla R n, IRen, 9 1998 Helempis Florineme trillS puLcherrimLi Ren, 1998 199 Helempis yixianensis Ren, Helempi Lepteremochaetus eLicaLta Ren, 19 lithoecius Ren, 1998 HeLempi Lichnoplecia yixianen is R n, 199Ren, 1995 kovalevi LepteremochaetLi litllOe ill Ren, 199
Manlayamyia dabeigouensis Zhang, 1991 Oiobrachyceron limnogenus Ren, 1998 Malllayam ia dabeigoLien i Zhang, 199\ Opiparifungivora aliena Ren, 1995 Oiobra hyceron limnogellLl Ren. 19 Orsobrachyceron chinensis Ren, 1998 Opiparifimgivora aliena Ren, 1995 Palaepangonius eupterus Ren, 1998 Or obm hyceron chillen i Ren, 1 9 Pauromyia oresbia Ren, 1998 Palaepall olliu eLipteru R n, I Pleciomimella perbella Zhang, Zhang, Liu et Shangguan, 1986 PaLirom 'ia ore bia Ren, 199 Protapiocera megista Ren, 1998 Pleciolllimella pel'bella Zhang, Zhang, Liu et hangguan, 19 6 Protapiocera ischyra Ren, 1998 Protapio era megista Ren 199 Protempis minuta Ren, 1998 Protapio era i chyra R n, 199 Ren, 1998 Profemp'Protonemestrius minwa Ren, I beipiaoensis 9 Protonemestrius jurassicus Ren, 1998 Protonemestriu beipiaoen i Ren, 199 Lichnople ia kovaLe i Ren, 1995
Order Hymenoptera Protonemesfriu jura i
LI
Ren, 199
ovata Ren, 1995 Ord r Hymen Allogaster plera Alloserphus Ilogaster ovafa R n,saxosus 19 5 Zhang et Zhang, 2001 Alloxyelula Ren, 1995 Lto erphu axo LI lingyuanensis Zhang el Zhang, 200 1 Angaridyela endemica Zhang et Zhang, 2000 ALloxyelula lingyLianen i Ren, \995 Angaridyela Zhang et Zhang, n aridyela endemi aexculpta Zhang et Zhang, 2000 2000 Angaridyela robustaelZhang et Zhang, ngaridyela ex ulpfa Zhang Zhang. 2000 2000 n aridyela robu ta Zhang et Zhang, Angaridyela suspecta Zhang et2 Zhang, 2000 n aridyela suspecta ZhangRen, et Zhang, Baissodes grabaui 1995 2000 Bai sode Beipiaoserphus rabaui Ren, 199 elegans Zhang et Zhang, 2000 Beipiao erp/lLIs ele an Zhang et Zhang, 2000
Ceratoxyela decorosa Zhang et Zhang, 2000 Sinoxyela viriosa Zhang et Zhang, 2000 Chengdeserphus petidatus Ren, 1995 Spherogaster coronata Zhang et Zhang, 2001 Crephanogaster rara Zhang, Rasnitsyn et Zhang, 2002 Steleoserphus beipiaoensis Zhang et Zhang, 2001 inoxyela virio a Zhan t Zhang, 2000 Cerato 'eLa decoro a Zhang et Zhang, 2 Eopelecinus similaris Zhang, Rasnitsyn et Zhang, 2002 Stemmogaster celata Zhang, 1985 Chen de erphu petidatu Ren, 1 95 ph roga ter coronata Zhang et Zhang, 200J Eopelecinus vicinus Zhang, Rasnitsyn et Zhang, 2002 Tanychora beipiaoensis Zhang et Rasnitsyn, 2003 Crephano a ter rara Zhang, R ni yn t Zhang, 2002 teLeo erphu beipiaoen i Zhang et Zhang, 200 I Eophelecinus shangyuanensis Zhang, Rasnitsyn et Zhang, 2002 Tanychora exquisita Zhang et Rasnitsyn, 2003 temmo a ter elata Zhang, 19 EopeLe inus imilari Zhang, Ra ni yn et Zhang, 2002 Gurvanotrupes exiguus Zhang et Zhang, 2001 Tanychora sinensis Zhang, 1991 EopeLecilw vi iI/us Zhang, Ra nit n et Zhang. 2002 Tan)' hora beipiaoensis Zhang t Ra nit n, 2003 Gurvanotrupes liaoningensis Zhang et Zhang, 2000 Tanychora spinata Zhang et Rasnitsyn, Eop/zelecinu /zan )'liQnen i Zhang, Ra nit n t Zhang, 2002 Tony hora exqui ita Zhang 2003 et Ra nil n 2003 Gurvanotrupes stolidus Zhang et Zhang, 2001 Tanychorella dubia Zhang et Rasnitsyn, 2003 Gurvanolrupe exi Wi Zhang t Zhang, 2001 Ton)'ehora sinen i Zhan 1991 Heteroxyela ignota Zhang et Zhang, Trematothoracoides liaoningensis Zhang, et Wei, GlirvanOlrupe liaonin en is2000 Zhang et Zhang, 2000 Tonychora pinata Zhang et RZhang nit yn, 20032001 IsoxyelaGurvanotrupe. rudis Zhang et Zhang, Tuphephialtites zherikhinidubio Zhang, Rasnitsyn Zhang, 2002 toLidus2000 Zhang et Zhang, 200 I Tanyehorella Zhang et Raetnit yn, 2003 Jeholoropronia pingi ignota Ren, 1995 Xyelites lingyuanensis Zhang et Zhang, 2000i Zhang. Zhang et Wei, 2001 Heteroxyela Zhang t Zhang, 2000 Trematothoracoides /iaoningen Lethoxyela excurva Zhang et Zhang, 2000 Yanocleistogaster canaliculata Ren, 1995 I oxye/a rudi Zhang et Zhang 2000 Tupltephialtite -herikltini Zhang, Ra ni yn et Zhang, 2002 Lethoxyela vulgata Zhang 2000 Jeholoropronia pinet iZhang, Ren, 1995 yefite lingyuanen i Zhang el Zhang, 2 Liadoxyela chengdeensis Ren, 1995 et Zhang, 2000 Phylum Chordata Let/zo 'eLa excurva Zhang Yanoclei to aster canaLiculata R n, I 9 Liaoropronia leonina ZhangZhang et Zhang, 2001 2000 Subphylum Vertebrata Lethox)'ela vulgata t Zhang, Liaoropronia et Zhang, Class Osteichthyes Liado regia 'ela Zhang eh n deensi R 2001 n.1995 Ph fum hordata Liaoropronia leonina Zhang et Zhang, 2 ubph lum ertebrata Subclass Actinopterygii Liaoserphus perrarus Zhang et Zhang, 2001 Liaoropronia ia Zhang t Zhang, 2001 la 0 tei hthye Order Acipenseriformes Liaotoma linearis Ren,r 1995 Liao erp/zus perranl Zhang t Zhang, 2 ubcl clin Liu pterygii Family Peipiaosteidae et Zhou, 1965 Liaoxyela antiqua Zhang et Zhang, 2000 Liaotoma Lineari R n, J995 Ord r fengningensis ipen eri~ rmBai, 1983 Peipiaosteus Manlayaflexuosa (Ren, 1995) Liaoxyela antiquo Zhang Familpani Peipia leida 1965 Liu t Zh u, 1965 Peipiaosteus Liu et Zhou, Mesaulacinus rasnitsyni Ren, 1995et Zhang 2000 Manlayaflexuo a R n. J99 ) Peipiao teu fen nin en i et Bai, 19 1995 Yanosteus longidorsalis Jin, Tian, Yang Deng, Ocnoserphus sculptus Zhang et Zhang, 2001 Me aula iTW rasnil yni Ren, Peipiaosteu pani Liu Family Polyodontidae Bonaparte, 1838t Zh u, 1965 Palaeathalia laiyangensis Zhang, 19851995 Gcno erp/zu eulpw Zhang et Zhang, 2 Yano liui teu Lu, Lon1994 idorsali Jin, Tian, Yan o et Deng, 1995 Protopsephurus Pompiloperus sp. PalaeQ//zaLia laiyangensi Zhang 19 5 amil Pol d ntida Bonaparte, 1 3 Order Amiiformes Procretevania pristina Zhang et Zhang, 2000 Pompiloperu p. Protop plzurus liui Lu, 1994 Family Sinamiidae Berg, 1940 Protocyrtus validus Zhang et Zhang, 2001 Ord r miif rm Pro retel'ania pri tina Zhang et Zhang, 200 Sinamia zdanskyi Stensi6, 1935 Protoscolia imperialis Zhang, Rasnitsyn et Zhang, 2002 Protoeyrtus validu Zhan
t Zhang, 200 I
Protoscolia normalis Zhang, Rasnitsyn et Zhang, 2002
Proto eolia imperia/is Zhang, Ra nit yn et Zhang, 2002
t'rotoscolia Zhang, Rasnitsyn et Zhang, Protosinensis olia normali Zhang Ra ni yn 2002 et Zhang, 2002 Saucrotrupes et Zhang, Proto decorosus olia sinenZhang is Zhang, Ra nj2001n et Zhang, 2002 Scalprogasterfossilis Zhang et Zhang, au rotrupe decorosll Zhang2001 el Zhang, 200 1 Scolichneumon 1995 t Zhang, 200 1 Scalpro rectivenius asterfo iIiRen, Zhang Scorpiopelecinus versatilis Zhang, coLi 11I1eumon re ti,'eniu Rasnitsyn Ren, 1 9 et Zhang, 2002 eorpiopelecinu veT alili Shandongodes lithodes Zhang, 1985Zhang, R nits n I Zhan ,2002 handon od lithodes 19 et Zhang, 2002 Sinopelecinus delicatus Zhang, Zhang, Rasnitsyn Sinopelecil/u d Zhang, licat Rasnitsyn Zhang, Raetni~ n I2002 Zhang, 2002 Sinopelecinus epigaeus Zhang, inop lecinlls aeu Rasnitsyn Zhang, R et ni yn et2002 Zhang, 2002 Sinopelecinus magicusepi Zhang, Zhang, inop lecinus iel Rasnitsyn Zhang, R etnit n I2002 Zhang. 2002 Sinopelecinus viriosusma Zhang, Zhang, inopelecinu viriosus Zh ng, Ra nil n et Zhang, 2002 Sinosepulca giganthoracalis Ren, 1995 inosepul a i al1lllOracaLi 1995 Sinowestratia communicata Zhang etRen, Zhang, 2000 inolte tratia ommuni ata Zhang et Zhang, 2000
Family inamiidae B rg, 1940
Superorder Osteoglossomorpha inamia -dan fryi ten i", 19 5 Family Lycopteridae Cockerell, 1925 uper rder 0 teogl Lycoptera davidi (Sauvage, 1880)rpha amily Ly pt rid Lycoptera fuxinensis Zhang, 2002ckerelJ, 1925 L 'coptera dal'idi ( au age, 1 0 Lycoptera muroii (Takai, 1943) L 'copt ra fiainen i Zhang, 200_ Lycoptera sankeyushuensis (Ma et Sun, 1988) L 'coptera l1luroii Takai. 194 ) Lycoptera sinensis Woodward, 1901 L 'coptera ankeyu Iwen i a et un, J9 Lycoptera tokunagai Saito, 1936 L 'coptera ineT i 00<1\ ard, I 1 Family Kuyangichthyidae Liu, Ma et Liu, 1982 L 'coptera lokuno ai ail , 19 Jinanichthys longicephalus (Liu, Su, Huang et Zhang, 1963) Family Kuyangichthyidae Liu,
a
I
Liu, 19 2
Teleostei incertae sedis Jinaniehthy Longicepltalu Liu, u Huang et Zhang, 1963) Longdeichthys luojiaxiaensis Liu, 1982 Tel
0
lei in erta
edi
Class Amphibia Linnaeus, 1758 Longdeiehlh 'S luojiaxiaen i Liu, 19 2 la
mphibia Linna u , 17
Subclass Lissamphibia Haeckel, 1866 Superorder Salientia Laurenti, 1768 Li amphibia Ha ekel, I 66
Family Discoglossidae Gtinther, 1859 Callobatrachus sanyanensis Wang et Gao, 1999
Super rder alientia Laurenti, 176 nura Rafine que, 18 I5
Order
Famil Di eoglo id e Gunther, I 59
Liaobatrachus grabaui Ji et Ji, 1998 Callobatrachu sanyanell i Wang et Gao 1999 Mesophryne beipiaoensis Gao et Wang, 2001 edi
Superorder Caudata Scopoli, 1777
Liaobatrachu
rabaui Ji et Ji, 199
Order Urodela Dum6ril, 1806
Me oplzryne beipiaoell i Ga et
ang,2001
Family Cryptobranchidae Fitzinger, 1826
uperorder Caudata
opoli 1777
Chunerpeton tianyiensis Gao et Shubin, 2003
Order Urodela Dumeril, 1806
incertae sedis Family Family ryptobranehidae Fitzinger, I 26
Jeholotriton paradoxus Chunerpeton tian ien i Gao etWang, hubin2000 200 Laccotriton subsolanus Gao, Cheng et Xu, 1998 Family in erta edi Liaoxitriton zhongjiani Dong et Wang, 1998 Jeholorriroll paradoxu Wang, 2000 Sinerpetonfengshanensis GaoetetXu, Shubin, Laccotriton ub o/anu Gao, Cheng 199 2001 Liaoxirriron zhongjiani Dong et Wang, 1998
Class Reptilia lumen i Gao et hubin,2001 inerperonfeng Order Chelonia la
ReptiliaSuborder Cryptodira
Ord r h Ionia Family Sinemydidae Yeh, 1963 uborder Manchurochelys ryptodira liaoxiensis Ji, 1995 Family in Manchurochelys mydida Yeh, 1963 manchoukuoensis Endo et Shikama, 1942 Manchurochely liao 'ien i Ji 1995 Manchurochel s mcmchoukuoen i Endo et hikama, 1942 Subclass Diapsida
Order Choristodera DiapFamily ida incertae sedis
ub I
Order Chori todera Hyphalosaurus lingyuanensis Gao, Tang et Wang, 1999 Family ineenae edi (=Sinohydrosaurus lingyuanensis Li, Zhang et Ji, 1999) Hyphalosauru lingyuanen i Gao, Tang et tWang, 1999 lkechosaurus gaoi Lti, Kobayashi Li, 1999 (=Sinoh 'dro aLIrUS ling 'uanensiEndo, Li, Zhang Monjurosuchus splendens 1940 et Ji 1999) IkecllO auru gaoi Lii, Kobaya hi et Li, 1999 MOl/jurosu plendel/s End 1940 Infraclass hu Lepidosauromorpha
Order Squamata Infraela
Lepido Suborderauromorpha Lacertilia
Order quamata Infraorder Gekkota ub rder La erti1ia Family Ardeosauridae Camp, 1923 Infraorder Gekkota Yabeinosaurus tenuis Endo et Shikama, 1942 FamilyInfraorder rdeo auridae amp 1923 Scincomorpha Yabeino aurus renui Endo t 1825 hikama 1942 Family ?Lacertidae Gray,
Infra rder cine morpha Family ?La ertidae Gray, I 25
Jeho/acertaformo a Ji et Ren, 1999
Dalinghosaurus longidigitus Ji, 1998
Infra rd r ineertae edi Famil in en
Subclass Archosauromorpha
Family incertae sedis
Family in ena
Infraorder incertae sedis Family incertae sedis
Order Anura Rafinesque, 1815 ubcla
Jeholacerta formosa Ji et Ren, 1999
Da/illgho aUrLt longidigitLC Ji 1998
Order Pterosauria Suborder Rhamphorhynchoidea ubcla reho auromorpha Family Anurognathidae Kuhn, 1937 Order Ptero auna
Dendrorhynchoides curvidentatus (Ji et Ji, 1998) Jeholopterus ningchengensis Wang, Zhou, Zhang et Xu, 2002
uborder Rhamphorhynehoidea
Famil Anurognathidae Kuhn, 1937
Suborder Pterodactyloidea Ji et Ji, 199 ) Family Anhangueridae Campos et Kellner, 1985
Dendrorhyn Iwide curvidentatu
Jeh%pteru ningchengensi Wang Zhou, Zhang et
u, 2002
Liaoningopterus ub rder Pter a tyloidea gui Wang et Zhou, 2003 Nyctosauridae Nicholson et Lydekker, et Kellner, 19 5 1889 FamilyFamily nhangueridae Campo
Chaoyangopterus zhangi Wang2003 et Zhou, 2003 Liaollingopterus gui Wang et Zhou, FamilyFamily etoPterodactylidae auridae ieholBonaparte, on et Lyd 1838 kker, 1889
(Ji Wang et Ji, 1997) hao Eosipterus angopreruyangi -hangi et Zh u, 200
Haopterus gracilis Wang et 18 Ltirt, 2001 Fami! Pteroda tylidae Bonapane, Family Tapejaridae Eosipteru yangi Ji et Kellner, Ji 1997)1989
Sinopterus Wang et Zhou, Haopreru racilidongi Wang et Liin, 20012002 Famil Tapejarida Kellner, 19 9 illopreru dongi Wang et Zh u, 2002 Order Saurischia
Suborder Theropoda Order auri hiaCompsognathidae Family ubord r Theropoda Sinosauropteryx prima Ji et Ji, 1996 Family omp gnathidae
Family Dromaeosauridae Matthew et Brown, 1922
illo auropleryx prima Ji et Ji, 1996
Sinornithosaurus milleni Xu, Wang et Wu, 1999 0 auridae atth et Brown, 1922 Microraptor gui Xu, Zhou, Wang, Kuang, Zhang et Du, 2003 inomirho auru milleni u, Wang et u, 1999 Microraptor zhaoianus Xu, Zhou et Wang, 2000
amily Dr rna
Microrapror gui Xu, Zhou, Wang. Kuang, Zhang et Du, 2003
Family Troodontidae Gilmore, 1924
u, Zhou tWang 2000 Norell, Wang, Makovicky et Wu, 2002 Family TrSinovenator dontidae changii Gilmore,Xu, 1924 Microrapror zhaoianu
Superfamily Therizinosauroidea orelJ, ana, Mako i ky t Wu, 2002 Family incertae sedis uperfamily Therizin aur idea inovenalor chan ii Xu,
Beipiaosaurus inexpectus Xu, Tang et Wang, 1999 Family ineertae edi Infraorder Beipiao auruOviraptorosauria inexpectLC Xu, Tang et Wang, 1999 Familyirapt Caudipteridae Infra rder r auria Zhou et Wang, 2000 amily Caudipteryx audipt ridaezoui Zh Ji, u etCurrie, Wang,Norell 2 0 et Ji, 1998 audipteryx ';.oui Jidongi urrie et Ji,2000 1998 Caudipteryx Zhou orell et Wang,
Familyryx incertae audipt dongi sedis Zhou et Wang 2000 di gauthieri Xu, Cheng, Wang et Chang, 2002 Family ineertae Incisivosaurus Inci iva auru gaulhieri u, Cheng, Wang et Chang, 2002
Theropoda, Family incertae sedis
Family Liaoxiornithidae Hou, Zhou, Zhang et Gu, 2002
Liaoxiornis delicatus Hou et Chen, 1999
Epidendrosaurus ningchengensis Zhang, Zhou, Xu et Wang, 2002 Protarchaeopteryx robusta Ji et Ji, 1997 Ther p da Family incena edi Yixianosaurus et Wang, 2003 Zh u, u et Wang, 2002 Epidendrolongimanus Guru ningXuhengen i Zhang,
Order Sinornithiformes Family Liaoxiomithidae Hou, Zh u Zhang et Gu 2002 Family Sinornithidae 1997 Hou et hen, 1999 Liaoxiomi Hou, delicatus
Sinornis Sereno et Rao, 1992 Ord r santensis in rnithiform
Order Ornithischia Protarchaeoptery robu ta Ii et Ii, I 97 Suborder Ankylosauria Yixiano auru longimanu u et ang,2003 Family incertae rd r Omithisedis hia
Order Cathayomithiformes Famil inornithid
Eocathayornis Order atha walkeri rnithi~Zhou, rrne 2002 Cathayornis Hou, Zhou, Gu, 2002 atha omithidae Zh u,Zhang lin et et Zhang, I 92 Famil aberransis Cathayornis caudatus Hou, 1997 Eo athayorni walkeri Zhou, 2002 athayorni u Zhou, Zhang t Gu 2002 Cathayornis yandicaaberran Zhou, Jini etHZhang, 1992 athayorni caudatu Hou, Hou,1997 1997 Longchengornis sanyanensis
Liaoningosaurus paradoxus Xu, Wang et You, 2001 ub rder n Suborder Ceratopsia Famil in na
edi
Liaoningosauru Osborn, paradoxus Family Psittacosauridae 1923 u,
ang et ou, 200 I
ub rder eratop houi ia You, Xu et Wang, 2003 Hongshanosaurus aurida 0 born, 1923 Family P ittac Psittacosaurus meileyingensis Sereno, Zhao, Cheng et Rao, 1988 Hon /zano aurus houi You, u et Psittacosaurus mongoliensis Osborn, 1923 ang, 2003
athayorni yandicaHou, Zhou, lin et Zhang, 19 2 Family Cuspirostrisornithidae 1997 Long hengornis allyallen Cuspirostrisornis houi Hou, 1997 i H u, 1997 pir tri mithida H u 19 7 Famil u sexdentornis Largirostrornis Hou, 1997 u piro tri omi houi Hou, I 7 Order Longipterygiformes
P ittacosauru meileyingen i ereno, Zha, heng et Rao, 19 Neoceratopsia P ittacosauru mon olien i 0 born, 1923 Liaoceratops yanzigouensis Xu, Makovicky, Wang, Norell et You, 2002 e
eratop ia
Suborder Omithopoda [jaoceratop yan-igouen Family incertae sedis
u, Mako icky,
ang,
orelJ et Y, u, 2002
uborder Omith poda Family in ena
Order L
edi
Infra rder 19uan
'UO/leIl
i
u, Wang et You, 2000
nanti rnithe , Ord r and Family ind t.
Jibeinia luanhera Hou, 1997 Boluo /zia zh Il i Zh u, 19 Otogornis genghisi Hou, 1994 libeillia Ilianhera H u, 19 7 Protopteryxfengningensis et Zhou, 2000 Otogorni gen hi iZhang H u, 1994
di
an
u, 200 I
t
Class Aves
Subclass Ornithurae Protopteryxfen nin el i Zhang et Zhou. 2 Order Liaoningornithiformes ubcla Ornithura
Subclass Sauriurae Order and Family indet ubcla auriura
Family Liaoningornithidae Hou, 1996 Order Liaoning rnithiforme
Jeholornis prima Zhou Order and Family indetet Zhang, 2002
Liaoningornis longiditris Hou, 1996 H u 1996 Famil Liaoningornithidae
Sapeornis chaoyangensis et Zhang, 2002 leholomi prima Zh Zhou u et Zhang, 2002
Order Chaoyangiformes Liaollingorni Ion iditri H u, 19 6 Family Chaoyangidae nne 1997 Ord r ha angi~Hou,
Order Confuciusomithiformes apeorni chaoyan ensis Zhou et Zhang, 2002 Family Hou, Zhou, Gu et Zhang, 1995 rderConfuciusornithidae onfu iu omithiforrn
Chaoyangia beishanensis et 1997 Zhang, 1993 Family Cha angida Hou H u,
Changchengornis Ji, Zh Chiappe 1999 1995 mithida Hou, u, GuetetJi,Zhang, Famil Confu iuhengdaoziensis C/zang h 11chuonzhous omi hengdaoziensi Confuciusornis Hou, 1997 Ii, hiappe t Ii, 19 9 COllfu iu omi huo1l':.hou H u, et 1997 Confuciusornis dui Hou, Martin, Zhou Feduccia, 1999 Confuciu omi dlli Hou, Hou,Zhou, aninGuZh u t Fedu cia, 1999 Confuciusornis sanctus et Zhang, 1995 Confuciu omi sanctu H u, Zh u, Gu t Zhang, 19 5 Confuciusornis suniae Hou, 1997 on/II iuyixianensis omi suniae H Zhou, u, 1997 Jinzhouornis Hou, Zhang et Gu, 2002 lilrhouomi yL ianen Hou, is Hou, Zhou, Zhang et Gu, 2002 Jinzhouornis zhangjiyingia Zhou, Zhang et Gu, 2002 Jin-llOuomi Walker, -/lQllgjiyingia Subclass Enantiornithes 1981 H u, Zh u, Zhang et Gu, 2
Family Songlingornithidae Hou, 1997 hao.. . an ia bei hallen i H u et Zhang. 199 Songlingornis linghensis Hou, 1997 Famil nglingornithida Hou. I 7
onglillgornis Family incertae sedis. ling/zen i Hou, I
7
incenae Zhou di.et Zhang, 2001 Famil grabaui Yixianornis
YlXianomi Zhou graballi Zhou et2001 Zhang, 2001 Order Yanomithiformes et Zhang, OrdYanornithidae r an rnithi~Zhou rm etZh u et Zhang, Family Zhang, 2001 2001 2
alk r, 19 I ubcla nanti mithe Order Eoenantiornithiformes rder Eoenantionithidae E nanti mithif" rrne. Family Hou, Martin, Zhou et Feduccia, 1999 Famil E nanti nithidae H u, Manin, Zh u et Fedu ia, 19 Eoenantiornis buhleri Hou, Martin, Zhou et Feduccia, 1999
Eoenantiorni buhleri H u, Order Liaoxiornithiformes rder Lia i rnithiforrn
rrne
Family L ngipterygidae Zhang, Zhou Hou et Gu 2000
ntia
linzhou aunt yan
ngipterygi~
Enantiornithes, Order and Family indet. Longipter)'x chaoyallgen i Zhang, Zh u, H u et Gu, 2000 Boluochia zhengi Zhou, 1995
Jinzhousaurus yangi Wang et Xu, 2001 Family in ena
LAr iro tromi dentomi H u, 1997 Family Longipterygidae Zhang, Zhou, Hou et Gu, 2000
Longipteryx chaoyangensis Zhang, Zhou, Hou et Gu, 2000
Jeholosaurus shangyuanensis Xu, Wang et You, 2000 Infraorder Iguanodontia le/zolosaul"Us /zan Family incertae sedis
H u, 1997
Family Cathayornithidae Jin etet Zhang, 1992 inorni antenZhou, i ereno Ra , 1992
anin, Zh u t edu cia, 1999
Zh u et Zhang, 200 I Familmartini Yan rnithidae Yanornis Zhou et Zhang, 2001
Yallomi Hou martini Zh 1984 u et Zhang 2001 Order Gansuiformes et Liu, OrdGansuidae r Gan uiforrne Liu, 19 4 Family Hou et Hou Liu, et 1984 Gan uidaHouH etuLiu, et Liu, 19 4 Famil Gansus yumenensis 1984
Gall Class Mammalia Cia
1I
ywnenel i Hut Liu, 19 4
Mammalia
~03
Sinobaatar lingyuanen i Hu et Wang, 2002
Order Tricon d nta Family incertae
Order ymmetr donta
di
leh%den jenkin i Ji, Luo elJi, I 99 Family Repen mamidae Li,
ang,
nlidae
Gobi onodon zofiae Li,
h
ang el Li, 2000
el Rich, I
lh riida Mar h, I
pal
7
Maotheriwn inen i R ugier, Ji et
ang el Li, 2000
Repenomamu robu ru Li Wang, Famil G bic n
Famil
0
a ek,2003
Zhan heorherium quinquecu pidens Hu,
4
Infr la
ang, Hu l M ng, 2003
Euth ria
Order and Family incerta
edi
Eomaia scan oria Ji, Luo, Yuan,
Order Mullituber ulata
ang, Lu el Li. I 97
ible, Zhang et G
rgi.2002
Family Eobaatarida Kielan-Jaw r w ka Da hzeveg l Trofimo , 19 7
Plantae Divi i n Char phyta la
Thallites d yphylLu
Charoph
e
ThaLlites ric ioite
haracea L. CI. Ri hard, I 15
ubfamiJy
eli t
har ideae Madler, 1952
A Ii rochara huihuibaoen i ubfamily Charoid 0
~'oLLlta
it lIoideae
oniopreri bllrejensi (zaJe ky) eward, 1904 oniopteris pectabi/is Brick, I 5 Eboracia Lobifolia (Phillip) Thom ,1 29
1. Braun et Migula, 1890
Pe ki phaera mu/tispira Lu et Yuan, 1991) Pecki phaera vertici/lara Pe k, I 37 Famil Cia
ang, I 6
l ra ea Pia, 1927
Atopo hara trh'oll i triquetra L. Gramb t 19 FLabellochara harri i Peck, 1941 FLabelLochara hebeien i Lu, Zhang et Zha . 19 Family Porochara cae L. Gramba t, 19 2 ubfamil
uneat
Botrychite reheen i Wu, 19
u, 19
Pe ki phaera paragranuLifera
har idea Z. Wang et Huang, 197
Minhechara p.
u, 1999
P ida
(Peck, 1937)
Me ochara 'llanzien is Yang, 19 3 ubfamily
Equi etites longel'aginaru iii
Le nhardi, I 63
hara producra Liu [
Me ochara
L' opodite faustu Wu. 1999
phen p ida
. Wang, 1965
A Ii to hara IIILllldula Pe k, 1941 Me
u 199
Lyc p ida
Ord r CharaJ Family
u 19
eed planl Ginkgoal
Baiera boreali Wu, I
9
Baiera gra iii (B en M ) Bunbary, 1 51 Gink
0
apode Zheng l Zhou, _003
Gink oite
p.
phenobaiera p. zekan w bale
-ekanolVskia? debi/i Wu, 19 Solenites murrayana Lindley l Hutton
4
phenarion parili Wu, 19 9 oniferaJe
Fr - p ring plant
BrachyphyLLum cfjaponi mn Yok
Bry phyta
BrachyphyLLum rhombicu/11
Mu cite drepanophylLu Wu. 1999
Cupre ino ladu
MilS it
C pari idium p.
ten lIu Wu, 1999
p.
u,19
ma Oi hi, I
4
yathidite minor ouper, 1953
Elatocladu leptophylh Wu, 199 Pityo ladus den ijoliu Wu, 1999
Leptolepidite verruco u
Pityospermum p.
o mundacidile
chi-olepi beipiaoell is
Baculati pOI·ire comaumen is (
u, 1999
Neorai tri kia equali (0
chizolepi jeholensi Yabe el End , 1934 Tyrmia acrodoflta Wu, 1999 William onia bella,
Pu el
u, 19
n, 1953) P t nie, 1956
ouper, 195
i atrico i porite australien i ( ook on, 195 ) Ba1me, 1957
u, 1999
ifi u
i atricosi porite pa
Boleho ilina 1961) Zhang, 1965
chi-aeoi porites certu (B I h
Rehezamite anisolobu Wu 19 9
ilina, 19 6) Gao
I
Zhao, 1976
Tenuangula pori qiuchen en i (Wang et Li, 19 I) ]ia, 19 6
p.
Tenuangula pori microl'erruco u (Zhang, 19 4) ]ia, 19 6
Gnetale
Den oi porite microrugulatu Brenner, 1963
Liaoxia changii ( a el Wu) 1997 Liaoxia ellenii a
I
yclo ristel/a enti
Wu, 1997
Izaoyallgia liangii Duan 1997
0
a Phillip et Felix, 1971
Gymno permou pollen
Perinopollenite elalOides ouper, 195 Cia opolli aflflulatu (Verbit kaya, 19 2 Li, 19 4
perma
ino arpus decus atu Leng t Frii , 2003 ngi
on et Dettmann, 195
Kluki pOl·ite p eudoreticulaw
Bennettitale .
ngi
k n, 195 ) POlonie, 1956
Lycopodiun poriles au lro lavalidite (
Benneltitale
Rehe amite
up r, 1953
wellmanii Coup r, 1953
permae?
Ginkgo 'cadoph ·tli nitidu (Balme, 19 7) de ]er ey, 1 62
Archaefructus liaoningen is un, Dileher, Zheng el Zhou, 199
lugel/a clariba ulata Ml hedli hili
Archaefructu
Ephedripile
inen is un. Dileber, Ji el
ion, 2002
hakhmunde, 1973
t
p.
Beipiaoa parva Dil her, un el Zheng, 2001
liaolzepolli annulaw Yu et Mia , 19 4
Beipiaoa rotunda Dil her, un et Zheng, 2001
liaohepol/i j7exllo u (Mia , 19 2)
Beipiaoa pino a Dilch r, un el Zheng, 200 I Liaoniflgocladus boii un, Zheng el Mei, 2000 (PotamogefOn? p.; Orchidites lancijoliu Wu, 1999; Orchidile linearijoliu
ia el Yu, 19 4
al/ialasporites dampieri (Balm , 1957) D v, 1961 u. 1999
Trapa? p.
Bice topolli wulanen i Li, 19 3 a ·tonipolleniles pallidu (Rei inger, 1950) ouper, 195 Quadraeculina anellae/ormi Maljawkina, 1949 Quadrae ulina limbata
Planta Incertae edi
alja kina, 1949
Antholithu ovatu Wu, 1999
Protoconiferu funariu
Carpolithus p.
Protopinus p.
Erenia telloptera Kra ilo, 19 2
P eudopicea variabilijormis (Maljawkina, 1949 Bolcho ilina, 1956
Lilite reheen i
P eudopicea rotundiformis (Maljawkina, 1949 Bol ho ilina, 1956
u, 1999
aumo a, 19 7) Bol ho ilina, 1956
Poly onite planus Wu, 1999
Pinu pollellires divulgafU (B Ieh vilina, 19 6) Qu, 19 0
Poly onite polyclonus Wu, 1999
Abietineaepol/el/ite pectineI/us (Ma1ja kina, 1949) Liu, 19 2
Rhi<.oma el/ipti a Wu 1999
Abie pollenite
Typhaerafu iformi Kra ilo
Piceaepollenile
19 2
pp. p.
Cedripile pll illll (Zauer 1954) Krutz ch 1971 Cedripite microsa coide
p re and poUen
Plerid phyte por
1
Podocarpidire multe imu (B Ieho itina, 1956) Poe
Bryophyte p re
terei porites antiqua porite (W I n el Web
ong el Zh ng, 1
I
r, 1946) Dettmann, I 63
Podocarpidite onzatus Poco k 1962
k, 1962
A COMPOSITE PICTURE OF THE JEHOL FOSSILS rl:
Ilder
Yall
011
F sh Fish /I)j{~1IJ p(1/1/ I. PUPI 1. PeipiaoJteuJ pani
16. IProtopteryx /"()ff)/JttlJ.\ fll/!,IIIIIf,W IS Itngningensis
·opt. /. I 11/111'01/ 2. L) Lycoptera mumii
Itbll)III"I/I )(111 it I 17. Calhayornis yandica
Ampfl Amphibian
1 'IPWJX tb(lII) lIIp,t'n I lB.> LOll Longipteryx chaoyangensis
tlillbltri bm Jall)'antnJiJ IU) IIWI IS 3. CallobalrachuJ 11"1 ",XIIS 4. jlbolotl"/toll jtholotriton Pparado"·/IJ
Turle Turtle
20. Zbt/l1gheotbn'ill/// 711/1/lfllf 'lIs/Jllon Zhangheothtrium quinqlltcuJpidens
70.\11:1111 5. AllI/tblll'lItbd)s Manchuroche/)'J Ii /iaoxienJiJ
Cho to) r Choristodere 6.
19. YanorniJ )'U/O/7/1 Irtlrtllll 19. martini
Mlmma Mammal
Insect 21. Ephnneropsis Ephulltro/J I tristta/is triS tt/IIS
II)pbllloJ(IIII"IIJ lingYIlI/tllJlj Hypha/oJauruJ /ingyuanenJis
Lizal Lizard )' Ib~lI/oJ //1m tU/l1t 7. YabeinoJaurtlJ tenNis
p . ~ Pterosaur ,8.>. HaOpltrllS l/t/oPtt rll/!,I"I tlis gr«i/is 0 S Dinosaur
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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B AMNH CAS CIB
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American Mu eum of atural Hi tory Chine e Academy of Science American Museum of Natural History AMNH Chengdu In 6tute of Biology, Chine e Academy of Science
Chinese Academy of Sciences
CMNH CAS CNU
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Carnegie Mu eum of Natural Hi tory
Institute of Biology, Chinese Academy of Sciences CIB Capital Chengdu ormal Univer ity (Beijing)
of Hi Natural Mu eumMuseum of atural tory History FMNH CMNHThe FieldCarnegie
GPH
Capital University (Beijing) CNU Geological Pub Ii Normal hing Hou e (Beijing)
IHB
FMNHIn titute of The Field Museum of Natural Historyof Science Hydrobiology, Chine e Academy
IVPP
VertebratePublishing Paleontology and(Beijing) Paleoanthropology Chine e Academy of Sciences Geological House GPH In titute of
KIB
In tituteof ofHydrobiology, Botany, ChineChinese e Academy of Science Academy of Sciences IHB Kunming Institute
KU
ofKan of a Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences IVPP Univer ityInstitute
NIGP
titute ofInstitute Geologyofand Palaeontology, Chine eof Academy KIB Nanjing InKunming Botany, Chinese Academy Sciencesof Science
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I01
NRM PKU SAPE ZMUA
Naturhi tori ka riksmuseet (Swedi h Mu eum of Natural Hi tory)
KU
University of Kansas
NIGP NRM
PKU
Peking Univer ity
Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences
Society of Avian Paleontology and E olu6on
Naturhistoriska riksmuseet (Swedish Museum of Natural History)
Zoological Mu eum, Univer ity of Am terdam
Peking University
SAPE
Society of Avian Paleontology and Evolution
ZMUA
Zoological Museum, University of Amsterdam
