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Vertebrate diversity of the Jehol Biota as compared with other Lagerstätten

  • Paleontological Museum of China, Beijing, China

Abstract and Figures

In the last twenty years, the extraordinary discoveries of vertebrate fossils from the Jehol Biota not only have important implications for studying the evolution of major Mesozoic vertebrate groups, their paleobiostratigraphy and paleoenvironmentology, but also provide critical evidence for understanding the biodiversity changes of the Early Cretaceous ecosystem. Currently, the Jehol Biota in a narrow sense (i.e., distribution limited to western Liaoning, northern Hebei, and southeastern Inner Mongolia) comprises a vertebrate assemblage of at least 121 genera and 142 species. Among them are 13 genera and 15 species of mammals, 33 genera and 39 species of birds, 30 genera and 35 species of dinosaurs, 17 genera and species of pterosaurs, 5 genera and species of squamates, 5 genera and 7 species of choristoderes, 2 genera and species of turtles, 8 genera and species of amphibians, 7 genera and 13 species of fishes as well as 1 genus and species of agnathan. All these known 121 genera are extinct forms, and only a small percentage of them (e.g., agnathans, some fishes and amphibians) can be referred to extant families. The Jehol vertebrate diversity already exceeds that of the contemporaneous lagerstätten such as Santana Fauna from Brazil and the Las Hoyas Fauna from Spain, and is nearly as great as that of the Jurassic Solnhofen Fauna and the Eocene Messel Fauna from Germany. Therefore, The Jehol Biota undoubtedly represents a world class lagerstätte in terms of both fossil preservation and vertebrate diversity. The success of the Jehol vertebrate diversity had a complex biological, geological, and paleoenvironmental background. Analysis of the habitat and diet of various vertebrate groups also indicates that the habitat and dietary differentiation had played a key role in the success of the taxonomic diversity of vertebrates of various ranks. Furthermore, the interactions among vertebrates, plants, and invertebrates as well as the competitions among various vertebrate groups and some key morphological innovations also contributed to the success of the Jehol vertebrate diversity. KeywordsJehol Biota-Early Cretaceous-vertebrate-diversity-lagerstätten
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Earth Sciences
© Science China Press and Springer-Verlag Berlin Heidelberg 2010
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RESEARCH PAPER December 2010 Vol.53 No.12: 1894–1907
doi: 10.1007/s11430-010-4094-9
Vertebrate diversity of the Jehol Biota as compared with other
ZHOU ZhongHe* & WANG Yuan
Key Laboratory of Evolutionary Systematics, Institute of Vertebrate Paleontology and Paleoanthropology,
Chinese Academy of Sciences, Beijing 100044, China
Received March 19, 2010; accepted August 18, 2010
In the last twenty years, the extraordinary discoveries of vertebrate fossils from the Jehol Biota not only have important impli-
cations for studying the evolution of major Mesozoic vertebrate groups, their paleobiostratigraphy and paleoenvironmentology,
but also provide critical evidence for understanding the biodiversity changes of the Early Cretaceous ecosystem. Currently, the
Jehol Biota in a narrow sense (i.e., distribution limited to western Liaoning, northern Hebei, and southeastern Inner Mongolia)
comprises a vertebrate assemblage of at least 121 genera and 142 species. Among them are 13 genera and 15 species of mam-
mals, 33 genera and 39 species of birds, 30 genera and 35 species of dinosaurs, 17 genera and species of pterosaurs, 5 genera
and species of squamates, 5 genera and 7 species of choristoderes, 2 genera and species of turtles, 8 genera and species of am-
phibians, 7 genera and 13 species of fishes as well as 1 genus and species of agnathan. All these known 121 genera are extinct
forms, and only a small percentage of them (e.g., agnathans, some fishes and amphibians) can be referred to extant families.
The Jehol vertebrate diversity already exceeds that of the contemporaneous lagerstätten such as Santana Fauna from Brazil and
the Las Hoyas Fauna from Spain, and is nearly as great as that of the Jurassic Solnhofen Fauna and the Eocene Messel Fauna
from Germany. Therefore, The Jehol Biota undoubtedly represents a world class lagerstätte in terms of both fossil preservation
and vertebrate diversity. The success of the Jehol vertebrate diversity had a complex biological, geological, and paleoenviron-
mental background. Analysis of the habitat and diet of various vertebrate groups also indicates that the habitat and dietary dif-
ferentiation had played a key role in the success of the taxonomic diversity of vertebrates of various ranks. Furthermore, the
interactions among vertebrates, plants, and invertebrates as well as the competitions among various vertebrate groups and some
key morphological innovations also contributed to the success of the Jehol vertebrate diversity.
Jehol Biota, Early Cretaceous, vertebrate, diversity, lagerstätten
Citation: Zhou Z H, Wang Y. Vertebrate diversity of the Jehol Biota as compared with other lagerstätten. Sci China Earth Sci, 2010, 53: 18941907,
doi: 10.1007/s11430-010-4094-9
1 Introduction
The Jehol Biota represents a terrestrial and freshwater fau-
nal and floral assemblage in the middle Early Cretaceous
(131–120 Ma [1, 2]) that was mainly distributed in East Asia.
It currently comprises invertebrates such as gastropods, bi-
valves, crustaceans (conchostracans, tadpole shrimps, os-
tracods, and shrimps), spiders and insects, vertebrates in-
cluding agnathans, fishes (chondrichthyans, acipenseriforms,
amiiformes and osteoglossomorphs), amphibians (anurans
and urodeles), turtles, choristoderes, squamates, pterosaurs,
dinosaurs (saurischians and ornithischians), birds (enantior-
nithines, ornithurines and more basal forms), and mammals
(triconodonts, multituberculates, symmetrodonts, metathe-
rians and eutherians), as well as algae, lichens, ferns, gym-
ZHOU ZhongHe, et al. Sci China Earth Sci December (2010) Vol.53 No.12 1895
nosperms and angiosperms. Abundant fossils, exceptional
preservation, and the unique geological background of the
region open a window into the evolution of various bio-
logical groups and the Early Cretaceous terrestrial ecosys-
tem [3–5]. The Jehol Biota in a broad sense had a distribu-
tion in northern China, Mongolia, Kazakhstan, Siberia, Ja-
pan, and the Korea Peninsula, but in a narrow sense it only
refers to the biota in northern Hebei, western Liaoning, and
southeastern Inner Mongolia where the fossils are best pre-
served and represented.
In the 1860s, the French missionarier Père Armand
David started to collect fish fossils in Lingyuan County of
Chaoyang City, western Liaoning Province. In 1880 the
French ichthyologist H. E. Sauvage studied these fossils,
but misidentified them as a Tertiary fish and referred them
to a new species of Prolebias (P. davidi), which was re-
named Lycoptera davidi by the British ichthyologist A. S.
Woodward in 1901. These earlier work marked the begin-
ning of the scientific study of the Jehol Biota.
In the 1920s, the American geologist A. W. Grabau did
some pioneering work on the Mesozoic stratigraphy and
paleontology in western Liaoning, and for the first time
proposed the name of “Jehol Fauna”. In the 1930s and
1940s, the paleontological work in this region was done
mainly by some Japanese paleontologists, and a number of
well known reptilian fossils such as the lizard Yabeinosau-
rus, the choristodere Monjurosuchus, and the turtle Manchu-
rochelys were described [6].
In the 1950s and 1960s, Chinese geologists and paleon-
tologists conducted a comprehensive stratigraphic and pa-
leontological survey of the region. In 1962, Z. W. Gu ush-
ered in the concept of the “Jehol Group” and the “Jehol Bi-
ota”. Liu and colleagues [7] published a systematic study of
the fishes from the biota. Shortly after, another most com-
mon fish of the Jehol Biota, Peipiaosteus pani, a fossil
acipenseriform, was reported [8].
In the 1970s and 1980s, many comprehensive papers or
books were published on the biostratigraphy of the Jehol
Group, but comparatively little progress was made on the
study of the vertebrate fossils. This situation has dramati-
cally changed since the 1990s with many important verte-
brate fossils discovered and reported, and a new era in the
study of Jehol vertebrate assemblages has begun. At present,
the Jehol vertebrate diversity is already greater than any
contemporaneous counterpart in other areas of the world.
Recently in the study of the Jehol Biota, significant pro-
gresses have been made not only in the origin of birds, their
flight and feathers, and the early evolution of various im-
portant biological groups such as birds, mammals, ptero-
saurs, amphibians, insects and angiosperms [9–16], but also
in the biostratigraphy, geochronology, paleomagnetism, and
paleoenvironments [17–21]. These studies provided impor-
tant clues to understanding the major features and mecha-
nism of the Jehol vertebrate diversity.
It is noteworthy that the important vertebrate fossils from
the late Mesozoic terrestrial deposits in northeastern China
were from a long geological duration, ranging from the
Middle Jurassic to the Middle Cretaceous, and belong to
different biotae. For instance, the Middle and Late Jurassic
Yanliao Biota (also known as the Daohugou Biota) is well
known for producing many important groups of vertebrates
such as primitive mammals, pterosaurs and feathered dino-
saurs in addition to abundant insects and salamanders
[22–25]. The Yanliao Biota bears some resemblance to the
Early Cretaceous Biota; however, its fossil assemblage dif-
fers significantly from that of the latter. In addition, the
Fuxin Biota represents a middle Cretaceous biota that is
slightly younger than the Jehol Biota and also has a unique
vertebrate assemblage. In this paper, we restrict our discus-
sion of fossils from the Jehol Group (i.e., from bottom up,
the Dabeigou Formation, Yixian Formation, and Jiufotang
Formation), and the vertebrate taxa in the analysis of this
paper only refer to those from the Jehol Biota in a narrow
sense, i.e., only from northern Hebei, western Liaoning, and
southeastern Inner Mongolia. Although some of the Jehol
taxa are also known from other areas of northern China,
such as Xinjiang, Gansu, middle and western parts of Inner
Mongolia, Shaanxi, Jilin, and Shandong, they are not in-
cluded in the statistics of this paper.
2 Analysis of the Jehol vertebrate diversity
Currently, the Jehol birds comprise 33 genera and 39 spe-
cies, which constitute approximately one third of the known
Mesozoic avian species globally, or about 40% if only the
species published since 1990 are counted [26]. Chinese ma-
terials are represented mostly by complete skeletons and
often with preservation of feathers, and thus the information
they provide on the early avian evolution is greater than
from other contemporaneous regions.
Most of the Jehol birds are recognized as arboreal forms.
They demonstrated significant differentiations in morphol-
ogy, size, flight, diet, and habitat, representing the first ma-
jor radiation in avian evolutionary history.
The Jehol dinosaurs currently comprise 30 genera and 35
species. The number of dinosaur species in China is now
greater than from any other countries (Hailu You, pers.
comm.), and more than two thirds of them were described
since 1990. The Jehol dinosaurs constitute about 20% of all
dinosaur species from China, and both its species diversity
and number of individuals exceed that of any other fauna.
Among the dinosaurs from the Jehol Biota, theropods are
most diversified, comprising 21 genera and 24 species and
constituting about two thirds of the total dinosaur diversity
of the biota. The Jehol theropods represent nearly all major
Cretaceous lineages, and many are feathered, characteristic
of the appearance of many arboreal or herbivorous forms
The Jehol pterosaurs comprise 17 genera and species,
1896 ZHOU ZhongHe, et al. Sci China Earth Sci December (2010) Vol.53 No.12
and can be referred to at least 10 families or superfamilies.
All except one genus and species belong to the pterodacty-
loids, and the only exception represents a member of the
short tailed rhamphorhynchoid family Anurognathidae.
China now has the richest pterosaur fossil record largely
due to the recent discoveries from the Jehol Biota. The Je-
hol pterosaur assemblage already exceeds that of the Late
Jurassic Solnhofen in both abundance and the generic di-
versity. The Late Jurassic Solnhofen now only comprises 8
genera and approximately 16–19 species (Helmut Tischlin-
ger, pers. comm.).
The Jehol pterosaur discovery has remarkably changed
our view on the ecological niches of pterosaurs. Previous
discoveries were mainly from the sea shores whereas the
pterosaurs from the typical terrestrial or lake shore envi-
ronments in the Early Cretaceous of northeastern China
have expanded their distribution to more diverse areas
Although about 300 genera of mammals are known from
the Mesozoic globally, and the Jehol Biota only comprises
13 genera and 15 species, the information about early
mammalian evolution as inferred from the Jehol mammals
is extraordinary as they are almost all represented by com-
plete skeletons while about 100 species of mammals from
over 50 different localities in other areas are mostly frag-
mentary teeth or partial skeletons [26]. The Jehol mammals
can be referred to five major groups of Mesozoic mammals:
Triconodonta, Multituberculata, Symmetrodonta, Metathe-
ria, and Eutheria. Among them, triconodonts, multitubercu-
lates, and symmetrodonts are all important Mesozoic
mammalian groups, and the earliest members of the
metatherians and eutherians represent the beginning of their
later evolution and radiation [13, 34–36].
In addition to birds, dinosaurs, pterosaurs, and mammals,
the Jehol fishes, amphibians, turtles, choristoderes, and
squamates also demonstrated distinctive radiations.
The only known agnathan from the Jehol Biota is a lam-
prey genus and species that represents its earliest freshwater
record worldwide [37]. Fishes comprise 4 orders (or higher
ranks), 6 families, 7 genera and 13 species, as well as a few
fishes of undetermined family or species. Among these
fishes are a chondrichthyan (gen. et sp. undet.), acipenseri-
formes that comprise two families, three genera and four
species, and amiiformes represented by one genus and spe-
cies. Teleostei are most diversified, with two families, two
genera and seven species belonging to osteoglossomorphs,
and one genus and species of undetermined order and fam-
The Jehol amphibians all belong to lissamphibians, in-
cluding four genera and species of frogs and four genera
and species of salamanders, as well an advanced frog be-
longing to a species of undetermined order and family [38].
These fossils represent the earliest radiation of lissamphibi-
ans in East Asia [39], which is characteristic of abundant
individuals and marked species differentiation as well as the
appearance of representatives of extant families such as the
Discoglossidae [40, 41]. Although currently only one genus
is referred to an extant family, seven genera have not yet
been referred to any family. Some of them such as Meso-
phryne possibly belong to an extinct family [39, 42] while
others such as Liaoxitriton and Regalerpeton seem to be
closely related to the origin of some extant taxa [11, 14, 43].
The Jehol turtles currently comprise two genera and spe-
cies belonging to an extinct family. Despite the abundance
of individuals in some horizons, their taxonomic differen-
tiation is less significant than other reptiles.
Choristoderes all belong to an extinct reptilian lineage,
and are referred to three families, five genera and seven
species. Among them, two genera and three species are re-
ferred to the Simoedosauridae, one genus and two species to
the Hyphalosauridae, and two genera and species to the
Monjurosuchidae. These aquatic or semi-aquatic animals
represent the top level predator in the food web of Jehol
lake ecosystem.
The squamates from the Jehol Biota comprise five genera
and species of lizards, and none of them has been referred to
a known family. Among them, two genera and species (Ya-
beinosaurus tenuis [44], Liushusaurus acanthocaudata [45])
are the relics of the Jurassic lineage from Laurasia, and one
genus and species (Xianglong zhaoi [46]) belongs to Iguania,
and the rest belong to Scleroglossa [14, 43, 47].
To sum up, the Jehol vertebrate assemblage now com-
prises 121 genera and 142 species. All of them are extinct
genera and species. Most of them can only be referred to
extinct families, with a few exceptions in agnathans, some
fishes and amphibians which have been referred to extant
families (Table 1, Figure 1). We estimate that the count of
the Jehol genera and species will certainly increase with
many new discoveries to be reported in the near future. We
did not count the total number of families mainly because
many published genera and species have not been referred
to any familial category, and more morphological and
taxonomic work of these materials is needed. The species
number will also most likely increase with more detailed
morphological and comparative studies of the known Jehol
vertebrate fossils.
It is notable that we made a rough estimate of the number
of at least 180 vertebrate species of the Jehol Biota in a
broad sense, i.e., from Xinjiang, Gansu, middle and western
parts of Inner Mongolia, Shaanxi, Jilin, and Shandong in
addition to western Liaoning, northern Hebei and south-
western Inner Mongolia. This number does not include
those outside of northern China due to incomplete data and
the difficulty of precise stratigraphic correlations.
3 Dietary and habitat differentiation of the Je-
hol vertebrates
The Jehol vertebrate diversity is characterized not only by
ZHOU ZhongHe, et al. Sci China Earth Sci December (2010) Vol.53 No.12 1897
Table 1 List of vertebrates of the Jehol Biota
Order or higher taxa Family Genus Species
Agnatha: one order, one family, one genus and species
Petromyzontiformes Petromyzontidae Mesomyzon M. mengae
Pisces: four orders, 6 families, 7 genera and 13 species (one additional genus and species undetermined)
Elasmobranchii Hybodontoidea indet. indet.
P. fengningensis Peipiaosteus
P. pani
Yanosteus Y. longidorsalis
Polyodontidae Protopsephurus P. liui
Amiiformes Sinamiidae Sinamia S. zdanskyi
L. davidi
L. fuxinensis
L. muroii
L. sankeyushuensis
L. sinensis
Lycopteridae Lycoptera
L. tokunagai
Kuyangichthyidae Jinanichthys J. longicephalus
Teleostei indet. Longdeichthys L. luojiaxiaensis
Amphibia: two orders, 1 family and 8 genera and species (one additional genus and species undetermined)
Discoglossidae Callobatrachus C. sanyanensis
Liaobatrachus L. grabaui
M. beipiaoensis
(=Dalianbatrachus mengi)
Yizhoubatrachus Y. macilentus
indet. indet.
Laccotriton L. subsolanus
Liaoxitriton L. zhongjiani
Regalerpeton R. weichangensis
Urodela indet.
Sinerpeton S. fengshanensis
Chelonia: one order, one family, two genera and species
Ordosemys O. liaoxiensis
(=Manchurochelys liaoxiensis)
Chelonia Sinemydidae
Manchurochelys M. manchoukuoensis (type lost)
Choristodera: one order, 3 families, 5 genera and 7 species
I. gaoi Ikechosaurus
I. pijiagouensis
Liaoxisaurus L. chaoyangensis
H. lingyuanensis
( =Sinohydrosaurus lingyuanensis)
Hyphalosauridae Hyphalosaurus
H. baitaigouensis
Monjurosuchus M. manchoukuoensis
Philydosaurus P. proseilus
Squamata: one order, 5 genera and species
indet. Yabeinosaurus Y. tenuis (type lost)
(?=Jeholacerta formosa)
indet. Xianglong X. zhaoi
indet. Dalinghosaurus D. longidigitus
indet. Liaoningolacerta L. brevirostra, nomen dubium
indet. Liushusaurus L. acanthocaudata
Pterosauria: one order, 10 families, 16 genera and species
Anurognathidae Dendrorhynchoides D. curvidentatus
?Gallodactylidae Feilongus F. youngi
Anhangueridae Liaoningopterus L. gui
Pteranodontidae Chaoyangopterus
C. zhangi
(=Jidapterus edentus;
=Eopteranodon lii;
=Eoazhdarcho liaoxianesis)
Eosipterus E. yangi Pterodactylidae
Haopterus H. gracilis
Tapejaridae Sinopterus S. dongi
(=Huaxiapterus jii;
=Sinopterus gui)
Cathayopterus C. grabaui
Elanodactylus E. prolatus
Gegepterus G. changae
(To be continued on the next page)
1898 ZHOU ZhongHe, et al. Sci China Earth Sci December (2010) Vol.53 No.12
Order or higher taxa Family Genus Species
Dsungaripteroidea Nemicolopterus N. crypticus
Nurhachius N. ignaciobritoi
Hongshanopterus H. lacustrisi
Liaoxipterus L. brachyognathus
Istiodactylus I. sinensis
Ornithocheiridae Boreopterus B. cuiae
Ornithischia: one order, 3 families, 7 genera and 19 species
Heterodontosauridae Tianyulong T. confuciusi
P. lujiatunensis
P. meileyingensis
P. mongoliensis
P. gobiensis
Hongshanosaurus H. houi
Neoceratopsia fam. indet. Liaoceratops L. yanzigouensis
Ornithopoda fam. indet. Jeholosaurus J. shangyuanensis
Iguanodontia fam. indet. Jinzhousaurus J. yangi
Saurischia: one order, 8 families, 23 genera and 25 species (one additional species indertermined)
Titanosauroidea Euhelopus E. sp.
Titanosauriformes Dongbeititan D. dongi
Dilong D. paradoxus
Raptorex R. kriegsteini
Sinotyrannus S. kazuoensis
Protarchaeopteryx P. robusta Coelurosauria fam. indet.
Yixianosaurus Y. longimanus
Sinosauropteryx S. prima
Huaxiagnathus H. orientalis
Sinocalliopteryx S. gigas
S. changii
Mei M. long
Sinusonasus S. magnodens
Jinfengopteryx J. elegans
S. haoiana Sinornithosaurus
S. millenii
M. gui Microraptor
M. zhaoianus
Graciliraptor G. lujiatunensis
Tianyuraptor T. ostromi
Oviraptorsauria fam. indet. Incisivosaurus I. gauthieri
Caudipteryx C. dongi
C. zoui
Similicaudipteryx S. yixianensis
Therizinosauroidea Beipiaosaurus B. inexpectus
S. orientalis
Aves: 13 orders, 14 families, 33 genera and 39 species
Jeholornithiformes Jeholornithidae Jeholornis J. prima
(= Shenzhouraptor sinensis;
= Jixiangornis orientalis)
S. chaoyangensis Sapeornithiformes Sapeornithidae Sapeornis
S. angustis
Eoconfuciusornis E. zhengi
C. sanctus
C. dui
Confuciusornithiformes Confuciusornithidae
C. suni
(To be continued on the next page)
ZHOU ZhongHe, et al. Sci China Earth Sci December (2010) Vol.53 No.12 1899
Order or higher taxa Family Genus Species
C. chuonzhous Confuciusornis
C. feducciai
Changchengornis C. hengdaoziensis
J. yixianensis
Confuciusornithiformes Confuciusornithidae
J. zhangjiyingia
Zhongjianornis Z. yangi Order indet. indet.
Zhongornis Z. haoae
Protopterygiformes Protopterygidae Protopteryx P. fengningensis
Eoenantiornithiformes Eoenantiornithidae Eoenantiornis E. buhleri
Longipterygithiformes Longipterygithidae Longipteryx
L. chaoyangensis
L. hani
R. pani
S. cooperorum
B. zhengi
Eocathayornis E. walkeri
Cathayornis Y. yandica
(= Cathayornis caudatus;
=Cathayornis aberransis;
= Longchengornis sanyanensis;
= Cuspirostrisornis houi;
= Largirostrornis sexdentornis)
Cathayornithiformes Cathayornithidae
Sinornis S. santensis
Liaoxiornithidae Liaoxiornis L. delicatus
(= Lingyuanornis parvus)
Jibeiniaithiformes Jibeiniaithidae Jibeinia J. luanhera
Pengornis P. houi
Vescornis V. hebeiensis
Paraprotopteryx P. gracilis
Shenqiornis S. mengi
Enantiomithes indet.
Liaoningornithiformes Liaoningornithidae Liaoningornis L. longidigitris
Chaoyangornithidae Chaoyangia C. beishanensis Chaoyangornithiformes
Indet. Songlingornis S. linghensis
Yanornithiformes Yanornithidae Yanornis Y. martini
(=Aberratiodontus wui)
Yixianornithiformes Yixianornithidae Yixianornis Y. grabaui
Hongshanornis H. longicresta Ornithurae
Order indet.
indet. Longicrusavis L. houi
Archaeorhynchus A. spathula Order indet.
Order indet.
indet. Jianchangornis J. microdonta
Mammalia: three orders, 5 families, 13 genera and 15 species
Gobiconodon G. zofiae Gobiconodontidae
Meemannodon M. lujiatunensis
R. robustus Repenomamidae Repenomamus
R. giganticus
Jeholodens J. jenkinsi
Yanoconodon Y. allini
Multituberculata Eobaataridae Sinobaatar S. lingyuanensis
Akidolestes A. cifellii
M. sinensis Maotherium
M. asiaticus
Symmetrodonta Spalacotheriidae
Zhangheotherium Z. quinquecuspidens
Metatheria indet. Sinodelphys S. szalayi
Eomaia E. scansoria Eutheria indet.
Acristatherium A. yanensis
Mammalia indet. Juchilestes J. liaoningensis
1900 ZHOU ZhongHe, et al. Sci China Earth Sci December (2010) Vol.53 No.12
Figure 1 Comparison of the generic and specific numbers of major groups of the Jehol vertebrate assemblage. Numbers in the parenthesis after each taxon
represent known genus and species.
the great taxonomic differentiation among various groups
but also by the significant differentiation in their diets and
habitats. Among the Jehol vertebrates, the agnathans, fishes,
amphibians, turtles, and choristoderes were mostly dwelling
in aquatic or semi-aquatic environments, constituting the
top level of food web of the Jehol lake ecosystem. For in-
stance, the lampreys were parasitical; the acipenseriforms
were the largest fishes in the Jehol lakes, with some reach-
ing over one meter long, and probably mainly filtering food
items [48]. The teleosts were the most abundant fishes, and
probably mostly carnivorous. Amphibians were all car-
nivorous, presumably mainly predating on insects or worms
by living near the water. Turtles were also abundant and the
known species had some aquatic features. Choristoderes
represent extinct aquatic or semi-aquatic reptiles. Among
them, Monjurosuchus lived a semi-aquatic life [49]; hypha-
losaurids with a significantly elongated neck and tail were
piscivorous and lived in the water. They currently comprise
two species, one can be as long as one meter, and the other
is smaller in size [50] and is represented by hundreds of
individuals in fossil record. The third group of choristoderes
is represented by Ikechosaurus, which is characterized by a
large body size with elongated rostrum and believed to be as
ferocious as crocodiles [51].
Unlike fishes and other aquatic or semi-aquatic verte-
brates, the Jehol lizards were typical terrestrial forms. The
newly discovered Yabeinosaurus material showed a robust
body, in contrast to a previous view of slender body which
is now recognized as a juvenile feature. Its habitat is com-
parable to some extant larger terrestrial lizards [44], and one
newly discovered fossil actually preserved in its stomach
some fish remains, suggesting at least some of them were
piscivorous (Y. Wang, pers. observ.). The recently reported
Dalinghosaurus possesses a hindlimb with scansorial adap-
tation [52]. Xianglong was probably a perching lizard and
could glide with its unique membranes, representing a spe-
cialized clade of extinct lizards [46]. It seems that the forest
environment had played a key role in the differentiation of
the Jehol lizards.
The taxonomically diversified Jehol dinosaurs also dem-
onstrate great differentiation in habitats. Although most
dinosaurs were terrestrial animals, arboreal dinosaurs have
been recognized (e.g., Microraptor), which not only sig-
nificantly changed our view of the life of some theropods
but also provided further evidence supporting the arboreal
hypothesis on the origin of avian flight. The Jehol dinosaurs
vary greatly in body size. For instance, among the smallest
dinosaurs are the herbivorous Psittacosaurs and the car-
nivorous Microraptor zhaoianus, some of which are even
smaller than the contemporaneous birds. The similar sized
dromaeosaurid Sinornithosaurus was recently reported to
have possessed venomous glands [53]. Tyrannosauroids
from the Jehol Biota are larger, but still small compared to
their Late Cretaceous relatives [29, 54, 55]. Together with
compsognathids [56–58], tyrannosauroids constitute the
most ferocious predators on the top level of the food web of
the Jehol terrestrial ecosystem. The iguanosaurid Jin-
zhousaurus is gigantic with an estimated body length of 5–6
m. And the herbivorous titanosauriforms probably represent
the largest animal in the Jehol Biota [59, 60].
The herbivorous ornithischian dinosaurs such as Psitta-
cosaurus and Jinzhousaurus are also well represented in the
Jehol Biota despite a lower diversity compared to sauris-
chians. It is notable that among the theropods, some of them
are raptatorial such as Sinosauropteryx that often preserved
mammal or lizard remains in their stomach, whereas some
other taxa are herbivorous and often preserved gizzard
stones in the stomach, such as the oviraptorosaur Caudip-
teryx and the ornithomimosaurid Shenzhousaurus [61]. The
oviraptorosaur Incisivosaurus even possessed a pair of in-
cisor-like premaxillary teeth, suggesting a highly special-
ized herbivorous diet [62]. These discoveries have enriched
ZHOU ZhongHe, et al. Sci China Earth Sci December (2010) Vol.53 No.12 1901
our understanding of the dietary differentiation of theropods,
and the presence of many herbivorous theropods also seems
to be characteristic of the Jehol dinosaur assemblage.
As the first flying vertebrates, pterosaurs are mainly ar-
boreal, as in the case of the Jehol pterosaurs. Although
many of them were piscivorous, other dietary adaptations
such as filtering or herbivory have also been recognized.
For instance, Sinopterus is probably a seed eater (Z. Zhou,
pers. observ.). The differentiation of the Jehol pterosaurs in
body size is also significant and probably greater than that
in birds, with the smallest (e.g., Nemicolopterus) as big as a
swallow [33], and the largest (e.g., Liaoningopterus) with a
wing span of about 5 m, definitely the giant of the sky at the
time [32].
In extant birds, most species are arboreal. Arboreal forms
were also predominant in early birds. In fact, the evolution
of a reversed hallux and the possession of arboreal capabil-
ity represent a major event in the evolution of birds and
avian flight. Current study indicates that arboreal forms
most likely represent the most primitive ecological types in
avian history. Arboreal birds are dominant in the Jehol
avian assemblage. Among the Jehol birds, the ornithurines
are mainly terrestrial dwellers, but nearly all enantiornithi-
nes and more basal birds (e.g., Jeholornis, Sapeornis and
Confuciusornis) are arboreal [63–65].
Due to limited preservation, our knowledge of the diets
of the Jehol birds remains poor. Although it is generally
presumed that most enantiornithines were probably insec-
tivorous or feeding on other invertebrates, there is yet no
direct evidence for this hypothesis. However, some other
birds actually preserved direct evidence for their dietary
preference. For instance, several specimens of Jeholornis
and Sapeornis preserved seeds in their stomach, suggesting
they were seed eaters [66]. Their dietary reconstruction was
also consistent with analysis of their habitat and the mor-
phological features of the jaws. Although a few Confuciu-
sornis preserved piscivorous evidence, it is uncertain
whether it mainly fed on fishes [67]; it might have as well
been omnivorous. Among basal ornithurines, Yanornis and
Jianchangornis were obviously piscivorous as several
specimens preserved fish remains in their stomach [4, 68],
and Archaeorhynchus was almost certainly herbivorous as
several referred specimens preserved gizzard stones [69].
Some enantiornithines such as longipterygithids possess an
elongated rostrum and teeth restricted to its rostral end,
suggesting a probing diet [70, 71]. Although Longipteryx
was presumed to be piscivorous, none of the Jehol enantior-
nithines preserved any direct fossil evidence of their diet
[72]. Given the arboreal life of most enantiornithines, in-
sects could well be their major food resources. It seems cer-
tain that dietary differentiation was significant in various
lineages of birds in the Jehol Biota.
Most of the Jehol mammals probably fed mainly on in-
sects or worms based on their dentitions. Multituberculates
were omnivorous. Repenomamus was carnivorous as evi-
denced by the preservation of dinosaur remains in one of its
specimens although it remains a mystery whether it was an
active predator or scavenger [36, 73]. The Jehol mammals
also show remarkable differentiation in body size. Although
most of them, e.g., metatherians and eutherians, are small,
some taxa such as Repenomamus already represent one of
the largest mammals in the Mesozoic. Symmetrodonts such
as Zhangheotherium were probably mainly terrestrial in life
with a parasagittal posture [74]; however, the earliest
known members of both metatherians and eutherians
showed scansorial adaptation [35].
In sum, the Jehol vertebrate diversity is shown not only
by its great taxonomic differentiations at generic or higher
levels but also by its remarkable differentiations in diets and
habitats. The dietary and habitat differentiations had un-
doubtedly played an important role in the evolution of the
Jehol vertebrate diversity at various taxonomic ranks.
4 Mechanism for the evolution of the Jehol
vertebrate diversity
The discoveries and studies of several major vertebrate
groups of Jehol fossils are critical to our understanding of
the evolution of the biodiversity of the Early Cretaceous
terrestrial ecosystem. The great vertebrate diversity of the
Jehol Biota must have had a complex paleoenvironmental
and ecological background. In particular, among them are
the unique paleoenvironmental, paleogeographic changes
and the evolution of the lake ecosystems in East Asia in the
late Mesozoic.
East and Central Asia were generally believed to be iso-
lated from Europe during the Middle Jurassic and Early
Cretaceous [75]; however, recent fossil discoveries seem to
indicate that East and Central Asia were never completely
separated, and by the middle and late Early Cretaceous
when the Jehol Biota was most flourishing, the interconti-
nental biological changes were already frequent [75]. Even
before the appearance of the Jehol Biota, in the Middle and
Late Jurassic when the Yanliao Biota lived in an area simi-
lar to that of the Jehol Biota, there existed some faunal ex-
changes between East and Central Asia and Europe, e.g.,
mammals and pterosaurs [25].
The Jehol Biota not only contains some relics and re-
gional fossil taxa [76, 77], but also comprise many taxa
suggesting that the Jehol area was the cradle and evolution-
ary center for many biological groups [75], which was fur-
ther supported by the study of the pterosaur assemblage of
the Jehol Biota [31]. By the early Early Cretaceous with the
disappearance of the paleogeographic barriers between
Europe and Asia, many of the vertebrates that originated
from Jehol spread to Europe and other continents, and dur-
ing the same interval there also existed bilateral exchanges,
which together account for the presence of many cosmo-
politan elements in the Jehol Biota [5], such as dinosaurs
1902 ZHOU ZhongHe, et al. Sci China Earth Sci December (2010) Vol.53 No.12
(e.g., iguanodontids, dromaeosaurids, and ankylosauroids),
birds (e.g., enantiornithines), pterosaurs (tapejarids and an-
hangurids), and mammals (e.g., multituberculates and go-
biconodontids) [78, 79]. Obviously, the paleogeographic
changes and the formation of the Eurasia continent in the
Early Cretaceous had a significant impact on the origin and
radiation of the Jehol vertebrates.
The East Asia had a complex and active tectonic back-
ground in the Early Cretaceous. For instance, the destruc-
tion of the North China Craton and the mantle thinning was
probably the result of the active global tectonic activity (e.g.,
the subduction of the Pacific Plate, large scale volcanic
eruptions, and the beginning of the Cretaceous Normal Su-
perchron) [80]. In contrast to northeastern region of China,
the western areas of China were much less affected. It was
also proposed [20] that northeastern China was affected by
several transgressions from the Pacific. During the Early
Cretaceous, the global temperature was generally warm [81],
which could have resulted in the flourishing of the flora. On
the other hand, against this warm background, there proba-
bly also existed some short periods of cold temperature. A
recent study of the oxygen isotopes from vertebrate teeth
from Jehol and other areas suggests that at the time of the
Yixian Formation, the average atmospheric temperature in
East Asia was slightly lower than that of today (Romain
Amiot, pers. comm.). Furthermore, seasonal climatic
changes were probably present in the Early Cretaceous [4,
82]. Finally, the impact of the frequent volcanic eruptions in
this region on the speciation rate and the turn-over of the
paleoecosystems should not be ignored. Evidence of mass
mortality of life due to volcanic eruptions has been well
documented [83–85]. We estimate that the frequent volcanic
eruptions must have played a key role in the pattern of the
Jehol vertebrate radiation.
Undoubtedly, the diverse and abundant Jehol invertebrate
assemblages, particularly the insect assemblage, as well as
the Jehol flora have greatly contributed to the evolution of
the Jehol vertebrate diversity as they were inseparable parts
of the whole Jehol ecosystem. The invertebrates and plants
in the middle or lower level of the food web provided not
only food resources for the Jehol vertebrates but also the
paleoenvironmental background for vertebrate diversifica-
It seems that the insect radiation in the Early Cretaceous
of northeastern China represents one of the greatest radia-
tions in the evolutionary history of insects. Liu et al. [86, 87]
showed that the insect assemblage in the Yixian Formation
comprises about 16 orders, 95 families, 201 genera and 271
species. The lakes at this time were relatively deep and ex-
tensive, and the lacustrine ecosystem was relatively stable.
The temperature was generally humid, warm, with seasonal
arid or semiarid niches. Liu et al. [86, 87] also predicted the
presence of high mountains nearby with an altitude of over
800 meters surrounded by lakes, swamps, and ponds, suit-
able for diverse animals and plants to thrive [87].
The Jehol Flora was dominated by gymnosperms and
ferns, and the angiosperms were only slightly differentiated
[88, 89]. There already exists evidence of the co-evolution
between insects and angiosperms [90], and the aquatic ad-
aptation of early angiosperms [91, 92]. The forests were
vital to the evolution of various vertebrate groups as shown
by the presence of a high percentage of arboreal and her-
bivorous forms in the vertebrate assemblages, such as birds,
pterosaurs, dinosaurs, lizards, and mammals.
The Jehol fishes became important food source for many
birds, pterosaurs, and aquatic reptiles. For instance, the ter-
restrial ornithurines Yanornis and Jianchangornis most
likely were mainly piscivorous as fish remains were often
preserved in their stomach [68]. The diet of the mainly ar-
boreal confuciusornithids is still unclear although a few
specimens actually preserved evidence of fish eating [67,
93]. Although the dietary evidence for Jehol pterosaurs is
generally lacking, it is generally believed that most of them
were piscivorous based on their jaw morphology [32]. Fur-
thermore, some theropod dinosaurs (e.g., Sinosauropteryx)
often preserved remains of lizards and mammals in the
It should be pointed out that the success of the Jehol ver-
tebrate diversity is related to the evolution of each major
vertebrate group. For instance, since its first appearance in
the Late Jurassic, birds by the Early Cretaceous had pos-
sessed a number of important features relating to their flight
such as a pygostyle, a well-developed wing, and the pecto-
ral girdle and sternum nearly identical to that of modern
birds. In pterosaur evolution, the Early Cretaceous assem-
blage was dominated by short-tailed pterodactyloids with
several morphological innovations (e.g., elongated wing
metacarpal and wing digit) beneficial to more powerful
To sum up, the success of the Jehol vertebrate diversity
had a complex geological and paleoenvironmental back-
ground; it was also closely related to both the interactions
among vertebrates, invertebrates and plants in the Jehol
ecosystem, and the appearance of many major morphologi-
cal innovations of each vertebrate group.
5 Jehol vertebrate diversity as compared to
that of other lagerstätten
The Jehol Biota in a narrow sense (i.e., distribution limited
to western Liaoning, northern Hebei, and southeastern Inner
Mongolia) currently comprises a vertebrate assemblage of
121 genera and 142 species (Figure 1, Table 1). The number
of vertebrate species of the Jehol Biota in a broad sense, i.e.,
geographically including those from other areas of northern
China (e.g., Xinjiang, Gansu, middle and western Inner
Mongolia, Shaanxi, Jilin, Shandong), is estimated to be
about 180. Currently, as many of the Jehol vertebrate genera
have not been referred to a family or higher ranks, it is dif-
ZHOU ZhongHe, et al. Sci China Earth Sci December (2010) Vol.53 No.12 1903
ficult to calculate and compare the Jehol vertebrate diversity
at familial or higher level, and thus the faunal comparison in
the paper is restricted to generic and species level. The
Santana Fauna in Brazil is among the most famous terres-
trial Early Cretaceous vertebrate faunae outside China. It is
well known for preserving abundant and diverse fishes and
pterosaurs. No mammal or bird skeleton has been reported
so far [94]. It currently comprises about 30 fish species
(Alexander Kellner, pers. comm.), which include chon-
drichthyans, actinopterygians, and sarcopterygians [94]; all
four known dinosaurs are theropods [95], and there are two
frogs and two crocodilians (Alexander Kellner, pers.
comm.), as well as six turtles [96]. Besides fishes, ptero-
saurs are the most diverse vertebrate group among the
Santana Fauna. Kellner and Campos [94] estimated the
presence of about 14 species, and most are represented by
incomplete individuals and belong to the families of An-
hangueridae and Tapejaridae. It is notable that members of
the two families have also been reported from the Jehol Bi-
ota. Sayão and Kellner [97] provided a list of the Santana
pterosaurs that comprises 11 genera and 23 species. Ac-
cording to A. Kellner (pers. comm.), two more species can
now be added to this list. Therefore, besides fishes and
pterosaurs, the Santana vertebrate groups are generally not
as diverse as the Jehol vertebrates at species level, and the
Jehol pterosaurs are more diverse at familial level.
The Las Hoyas Fauna from the La Huérguina Forma-
tion (Upper Barremian) in Cuenca, Spain is another impor-
tant Early Cretaceous terrestrial fauna, which is well known
for its exceptional preservation of abundant animal imprints,
ferns, cycads, conifers, insects, birds, fish, crocodilians,
dinosaurs, amphibians, and lizards, over 200 nearly com-
plete terrestrial vertebrate fossils in total (http://palaeo.gly.
The Las Hoyas Fauna currently comprises three genera and
species of birds, all belonging to enantiornithines [98]. Its
fish assemblage is more diverse, comprising about 12 gen-
era belonging to various higher ranks of taxa. Among the
four known amphibian genera, the anuran Eodiscoglossus
can be referred to the family Discoglossidae. One unnamed
species can be referred to pipiforms. There are also two
genera of urodeles, and one genus of albanerpetontids. The
latter group has not been discovered from China, and is re-
cently regarded to represent an extinct amphibian lineage
other than frogs, salamanders, and caecilians. Lizard fossils
comprise three genera and species. The best known dino-
saur is the theropod Pelecanimimus, and there are also some
fragmentary sauropods. The Las Hoyas vertebrate assem-
blage also comprises one species of turtle and four species
of crocodilians [99].
Las Hoyas is only the best known locality of the La
Huérguina Formation. Fossil-bearing deposits of this forma-
tion are also distributed in other areas such as Buenache and
Uña. In addition to abundant fishes from these two localities,
there are two other anurans and four lizard genera and spe-
cies, as well as fragmentary materials of lepidosauromorphs
and a few more turtles. Besides several fragmentary croco-
dilians, there are some crocodilomorphs referrable to at
least three families. Fragmentary pterosaurs cannot be re-
ferred to any definite taxon now. Dinosaurs and mammals
are best preserved in the Uña assemblage, including such
dinosaurs as Richardoestesia, Paronychodon, dromaeosaur-
ines, velociraptorines, and hypsilophodontids, and such
mammals as Crusafontia, Galveodon, Eobaartar, and an-
other multituberculate [99].
To add up the vertebrate taxa from the above-mentioned
three localities, the Early Cretaceous La Huérguina Forma-
tion has yielded about 20 genera and species of fishes, 8
amphibians, 3 turtles, 8 lizards, 9 crocodilians or croco-
dilomorphs, 7 dinosaurs, 3 birds, and 4 mammals (Hercu-
lano Alvarenga, pers. comm.). Thus, the total estimate of
the vertebrate species is probably no more than 70, ap-
proximately similar to that of the Santana Fauna, but sig-
nificantly less than that of the Jehol Biota. The La Huér-
guina vertebrate assemblage lacks agnathans or choristo-
deres that were present in the Jehol Biota, but the Jehol as-
semblage lacks albanerpetontids and crocodilians.
The Late Jurassic Solnhofen Fauna was preserved in ma-
rine or lagoon deposits; however, many of the vertebrate
fossils were from the continent. The Solnhofen fossils gen-
erally refer to those from the Solnhofen Lithographic Lime-
stone (Solnhofen Formation, Lower Tithonian, hybonotum
Zone); however, fossils from other Late Jurassic horizons
slightly older or younger have often been referred to this
fauna, such as the eighth example of Archaeopteryx that
was from the slightly younger Moernsheim Formation
(Helmut Tischlinger, pers. comm.). According to informa-
tion provided by H. Tischlinger, due to controversy
over the taxonomic treatment, the Solnhofen fish assem-
blage probably comprises about 74–90 genera and 107–130
species, there are 1 genus and 2–3 species of birds, 2 genera
and species of dinosaurs, 8 genera and 16–19 species of
pterosaurs, 7–9 genera and 8–10 species of crocodilians; 6
genera and 6–8 species of sphenodonts, 2–3 genera and 2–4
species of ichthyosaurs, 4 genera and species of squamates,
and 7–9 genera and 8–10 species of turtles. Thus, the total
estimate of the Solnhofen vertebrates is 111–132 genera and
155–190 species (Helmut Tischlinger, pers. comm.). Com-
pared to the Jehol Biota, The Solnhofen fishes are more
diverse and account for over half of the total vertebrate
genera or species. Other vertebrate groups from Solnhofen,
particular birds, are less diverse as compared with the Jehol
Biota. The Solnhofen fauna comprises the marine ichthyo-
saurs, but lacks the terrestrial amphibians and mammals.
The Middle and Late Jurassic Yanliao Fauna represents
another lagerstätte from the late Mesozoic of Northeast
China. It is well known for its beautiful preservation of
abundant insects and salamanders, as well as mammals,
pterosaurs and feathered dinosaurs. The most famous local-
ity is Daohugou in Ningcheng County, southeastern Inner
1904 ZHOU ZhongHe, et al. Sci China Earth Sci December (2010) Vol.53 No.12
Mongolia; other major vertebrate localities also include
Guancailiang in Jianpin County, Linlongta in Jianchang
County and Wubaiding in Linyuan City, western Liaoning,
and Gangou in Qinglong County, Hebei Province. Because
most of these vertebrate discoveries have been made in re-
cent years, and the scale of excavation is also smaller com-
pared to that of the Jehol Biota, the Yanliao vertebrate as-
semblage currently only comprises 19 genera and species,
as well as 3 unnamed species [25]. It comprises three genera
and four species of mammals, four genera and species of
dinosaurs, five genera and species of pterosaurs, one genus
and species of lizards (two additional juvenile individuals
probably represent two different genera and species), four
genera and species of urodeles, one unnamed juvenile indi-
vidual of anuran, and one genus and species of fish. No bird,
choristodere or turtle has been reported from the Yanliao
Biota. It is also notable that salamanders are most abundant
among all the Yanliao vertebrates and are often represented
by hundreds of completely articulated skeletons or impres-
sions, showing a variety of ecomorphs and differentiation
[100–103]. Overall, the vertebrate diversity of the Yanliao
Biota is much lower than that of the Jehol Biota.
The study of the fossils from the Eocene Green River
Formation in North America has a history of over 150 years,
and approximately 60 vertebrate species have been reported
including about 14 families, 19 genera and 26 species of
fishes (
river-fish) and 11 species of reptiles, as well as many birds
and mammals. It represents one of the most famous Ceno-
zoic fossil lagerstätte, whereas the number of its total
vertebrate species is fewer than half of the Jehol Biota.
The Eocene Messel shales in Germany have preserved
many exceptional fossils, including avian feathers, mam-
malian hairs, skin impressions, stomach contents, etc. The
fossil collecting history of Messel is also over one hundred
years. Up to now, about 142 species of vertebrates have
been known [104] (Gerald Mayr, pers. comm.), same as that
of the Jehol vertebrate assemblage. The Messel vertebrate
assemblage comprises 45 species of mammals, 52 species
of birds referable to at least 23 families, 32 species of rep-
tiles referable to 12 families (including 7 crocodiles, 20
squamates, and 5 turtles), 5 species of frogs and salaman-
ders referable to 4 families, and 8 species of fishes referable
to 6 families. Among this vertebrate composition, birds and
mammals have the highest species diversity, and reptiles are
second to them, well reflecting the successful radiation of
birds and mammals in the early Cenozoic that had replaced
the ecological niches left by extinct Mesozoic reptiles. On
the contrary, the Jehol reptiles are most diverse, with birds
being the second, and the diversity of mammals is much
lower than birds. The diversity of fishes, amphibians, and
turtles is similar in both biotae. On the other hand, the Mes-
Figure 2 Comparison of the species diversity of the Jehol Biota with
other lagerstätten over the world. Numbers in the parenthesis after each
biota name represent the minimum count at species level.
sel Fauna is also rich in squamates and crocodiles whereas
crocodilians have not yet been reported from the Jehol
Fauna and choristoderes that probably shared a similar
ecology are not found in the Messel.
From comparisons above, it seems clear that the Jehol
vertebrate diversity exceeds those from other contemporary
lagerstätten. It is undoubtedly among the best lagerstätten
even compared to some most famous biotae such as the Late
Jurassic Solnhofen or the Eocene Messel or Green River
(Figure 2).
This comparison, however, is based on published taxa
without considering the collecting history and other bias,
thus it may not completely reflect the actual paleobiodiver-
sity of each biota. For instance, the distribution and out-
crops of the Yanliao Biota-bearing sediments are much less
extensive compared to the Jehol Biota, and furthermore the
scale of fossil collecting of the former is also much smaller.
The collecting history of the Santana Fauna is long and also
similar to that of the Jehol Biota in the fact that most of the
vertebrate fossils were the result of commercial collectings.
Finally, the duration of the Jehol Biota had lasted for at least
11 Ma, which is probably also longer than some other bio-
tae, which might be part of the reason for a high vertebrate
Nevertheless, the Jehol Biota is undoubtedly now the
best widow into the Early Cretaceous terrestrial ecosystem;
it has preserved arguably the best evidence for studying the
evolutionary history of major Mesozoic vertebrate groups
[3, 105] as well as reconstructing their paleoecological his-
ZHOU ZhongHe, et al. Sci China Earth Sci December (2010) Vol.53 No.12 1905
We thank Alexander Kellner, Peter Wellnhofer, Gerald Mayr, Helmut
Tischlinger, Jesus Marugan and Herculano Alvarenga for help with in-
formation of the biotae in Brazil, Germany and Spain. We are grateful to
Xiaolin Wang, Xing Xu, Yuanqing Wang, Fucheng Zhang, Jiangyong
Zhang for discussions and helps. Two reviewers provided helpful sugges-
tions. Desui Miao improved the English. This study was supported by Na-
tional Basic Research Program of China (Grant No. 2006CB806400),
National Natural Science Foundation of China (Grant No. 40121202), and
Chinese Academy of Sciences.
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... T he Middle-Upper Jurassic Yanliao Biota and the Lower Cretaceous Jehol Biota in northern China have been the subject of intense paleontology research for over a decade, especially instances of unusually preserved soft tissues, such as dinosaur feathers, mammals with fur, and angiosperms with petals and other delicate structures (Zhou et al., 2010;Pan et al., 2013). The temporal gap between the record of the Yanliao Biota and the Jehol Biota is nearly equivalent to the age of the Tuchengzi Formation . ...
... The deinonychosaurian tracks are 8.4-10 cm long, and the estimated trackmaker length is 99.4-118.4 cm, similar to the typical bipedal deinonychosaurians from the Jehol Biota (Zhou et al., 2010). Bird tracks include Pullornipes aureus, Aquatilavipes isp. ...
... However, it is doubtful whether there is evidence of ornithopod in the Tuchengzi Formation because all possible ornithopod tracks are poorly preserved. No ornithopod body fossils have been recorded in Yanliao Biota, but ornithopods are known from the Jehol Biota (Zhou et al., 2010). A total of 19 dinosaur track sites have been found in the Tuchengzi Formation, with 2637 dinosaur tracks representing at least 2091 trackmakers. ...
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In northern China, mainly within Western Liaoning, Northern Hebei and Beijing, vertebrate skeletons from the Tuchengzi Formation (Upper Jurassic to Lower Cretaceous) are scarce. However, a large number of new dinosaur track discoveries have made in past 10 years. So far, a total of 19 dinosaur track sites have been found in the Tuchengzi Formation, collectively containing 2,637 individual tracks and representing at least 2,091 trackmakers. This track record has become an important sample for understanding the Tuchengzi dinosaur fauna. The abundant Tuchengzi track record includes six non-avian theropod ichnogenera (Asianopodus, Eubrontes, Grallator, Menglongipus, Therangospodus, and Velociraptorichnus), three bird ichnogenera (Aquatilavipes, Pullornipes, and Koreanaornis), two sauropod ichnogenera (Brontopodus and cf. Parabrontopodus isp.), and a possible ornithopod ichnogenus (cf. Dinehichnu). This assemblage is saurischian-dominated, which is consistent with most Jurassic and Cretaceous track sites in China. The presence of deinonychosaurian tracks is consistent with the rich skeletal record from the Yanliao Biota or Jehol Biota, and the bird tracks are consistent with the record of shorebirds in Jehol Biota. The existence of ornithopod tracks is doubtful and needs further discoveries to be confirmed.
... This is a typical Mesozoic continental coal-and oil-bearing sedimentary basin ( Fig. 1B; Wang et al., 1997;Jia et al., 2021). The western Liaoning Province area is crucial in that it reveals the evolution of the Jehol Biota and angiosperms, as well as the origination of birds in time with ecological change (Qiang et al., 1998;Zhou and Zhang, 2004;Zhou and Wang, 2010;Wan et al., 2017). The area is also considered to be critical for understanding the subduction boundary between the paleo-Pacific and Eurasian plates ( Fig. 1A; Guo et al., 2017;Zhang et al., 2019). ...
... The western Liaoning Province area underwent a major ecological transition across the early Aptian-late Aptian boundary during the middle Aptian (ca. 119 Ma; Zhou and Wang, 2010). The biodiverse Jehol Biota was gradually replaced by the Fuxin Biota described by Wan et al. (2017). ...
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Reconstruction of Aptian−Albian paleoclimate obtained from marine records remains a challenging topic, but studies on coeval terrestrial paleoclimate and trigger mechanisms have lagged substantially. In this study, new multiproxy data from mudrocks in the Fuxin Basin of NE China provide a high-resolution terrestrial climate record from East Asia. Here, we demonstrate the occurrence of terrestrial climate cooling during the late Aptian (118−113 Ma), which interrupted the mid-Cretaceous warming shown in global records. Nearly uniform long-term global climate trends attributable to tectonism, volcanism, and weathering occur in Early Cretaceous terrestrial and marine records. In the Fuxin Basin, the long-term terrestrial climate was characterized by increasing temperatures during the late early Aptian, gradual cooling during the late Aptian, and subsequent enhanced warming during the early Albian. Moreover, chemical weathering and humidity during these intervals were low, moderate to high, and then moderate, respectively. A markedly reduced high-elevation paleogeomorphology under strong continental weathering during the late Aptian increased the variability in chemical weathering fluxes as the Eurasian plate in NE China drifted SE during the Early Cretaceous and then NE during the Late Cretaceous. We suggest that a combination of enhanced continental weathering and weakened plate drift induced changes in atmospheric CO2, while the geographic setting ultimately led to cooling in the Fuxin Basin during the late Aptian. Our results illustrate the importance of exploring long-term tectonic-climatic-biotic feedbacks to improve our understanding of tectonic processes and ecological transitions across various spatiotemporal scales.
... Lockley et al. Cretaceous Research 135 (2022) 105190 Yixian and Jiufotang formations contain the famous Jehol Biota, and abundant bivalves, insects, ostracods, fishes, and reptiles (Zhou and Wang, 2010 Young (1960) and Shang (1986) suggested that the dinosaur tracks (IVPP V 2470, and 1986 specimen) came from the Haizhou Formation. The Fuxin Formation was first coined as the "Fuxin coal series" by Wang and Huang (1929), and then referred to as the "Fuxin coal layer" by Muroi (1940). ...
... Lockley and Xing (2015) explained these flattened footprints as forming in very thin mudstone and siltstone intervals in thick sanddominated sequences, that are significantly flattened due to the differential effects of overburden pressures on different lithologies. Wang (1987) named the biota that existed in the Shahai and Fuxin formations in the northern Hebei and western Liaoning area as the "Fuxin Biota," also known as the "K-Y-A Biota," which is represented by the fishes Kuyangichthys, the conchostraca Yanjiestheria-Pseudestherites-Neimongolestheria, and by the plant Acanthopteris, The Fuxin Biota represents a mid-Cretaceous biota that is slightly younger than the Jehol Biota and also has a unique vertebrate assemblage (Zhou and Wang 2010). The Fuxin Biota has yielded various mammalian remains, including eutridoconodontans and multituberculates (Shikama, 1947;Wang et al., 1995Wang et al., , 2018Hu et al., 2005;Li et al., 2005;Kusuhashi et al., 2009aKusuhashi et al., , 2009bKusuhashi et al., , 2010Kusuhashi et al., , 2016Kusuhashi et al., , 2019. ...
Dinosaur tracks have been known and reported sporadically from the coal-bearing beds of the Fuxin Formation (upper Aptian–Albian, upper Lower Cretaceous) of Liaoning Province, northeastern China for some time but have not been described in detail. Here available material is illustrated suggesting a diversity of theropod track morphotypes, including small grallatorid, eubrontid and possible Gigandipus-like forms, a Magnoavipes-like form and an isolated small ornithopod track. Some of the tracks show the characteristic distortion of flattened footprints. The ichnofauna supplements our knowledge of the Fuxin Biota by adding a more diverse dinosaur fanua, indicated by the footprint record.
... [210]), it also preserves several Lagerstätten. These deposits have received a large amount of attention because of the exquisitely preserved dinosaur and mammal fossils they have yielded [211][212][213][214], but they have also produced numerous, exceptionally preserved specimens of caudates (e.g. [37,40,42]). ...
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The disjunct geographical range of many lineages of caudates points to a complex evolutionary and biogeographic history that cannot be disentangled by only considering the present-day distribution of salamander biodiversity. Here, we provide a critical reappraisal of the published fossil record of caudates from the Palearctic and quantitatively evaluate the quality of the group's fossil record. Stem-Urodela and Karauridae were widespread in the Palearctic in the Middle Jurassic, suggesting an earlier, unsampled diversification for this group. Cryptobranchidae reached Europe no later than the Oligocene, but this clade was subsequently extirpated from this continent, as well as from western and central Asia. The relatively recent appearance of hynobiids in the fossil record (Early Miocene) is most likely an artefact of a taphonomic bias against the preservation of high-mountain, stream-type environments which early members likely inhabited. Salamandroids first appear in Europe, expanding into Asia by the Miocene. The apparently enigmatic and disjunct distribution of extant caudate lineages is therefore explained by a wider past geographical range, as testified by the fossil record, which was fragmented during the late Cenozoic by a combination of tectonic (i.e. the uplift of the Tibetan Plateau) and climatic drivers, resulting in regional extirpations.
... The Early Cretaceous Jehol Biota sensu stricto is a world-class terrestrial Lagerstӓtte, containing exceptionally well-preserved fossils, including feathered dinosaurs, early birds, mammals, pterosaurs, fishes, and amphibians, as well as abundant insects and early flowering plants, which significantly increased our understanding of the Mesozoic terrestrial ecosystem and the origin and evolution of a number of biological groups (Zhou et al., 2003(Zhou et al., , 2021Norell and Xu, 2005;Zhou and Wang, 2010;Zhou, 2014;Meng, 2014;Chiappe and Meng, 2016;Ren et al., 2019). Gu (1962) formally defined the Jehol Biota as the fossil assemblage containing Eosestheria (clam shrimp), Ephemeropsis trisetalis (insect) and Lycoptera (fish) (i.e., EEL assemblage). ...
The Early Cretaceous Jehol Biota in northeastern China significantly increased our understanding of the Mesozoic terrestrial ecosystem and the origin and evolution of a number of biological groups. The early evolutional phase of the Jehol Biota is best recorded in the Members 2 and 3 of the Dabeigou Formation (D-M2–3) in the Luanping Basin and their equivalent Huajiying Formation in the Senjitu-Sichakou Basin in northern Hebei Province. To date, however, no complete, high-resolution time scale has been established for these early Jehol Biota-bearing strata, hindering our understanding of the origin of the Jehol Biota. Here we carried out a cyclostratigraphic analysis using a high-resolution relative lake level variation record, called “depth ranks” (i.e., a series of numerically ranked sedimentary facies) in the ∼185-m-thick, continuously lacustrine D-M2–3 in the Yushuxia section, Luanping Basin. Our results suggest that the climate/lake level fluctuations in the Luanping Basin during the development of the early Jehol Biota were strongly paced by orbital cycles, including the 405 kyr and ∼ 100 kyr eccentricity and ∼ 34 kyr obliquity and ∼ 21 kyr precession. By anchoring the 405 kyr-tuned depth ranks to a relatively high-precise zircon UPb age of one tuff layer at the base of Member 3 (131.5 ± 0.4 Ma), we built a high-resolution astronomical time scale, spanning from 132.4 to 130.7 Ma, for the D-M2–3, indicating a ∼ 1.7 Myr duration of the early Jehol Biota and an average sedimentation rate of ∼11 cm/kyr of the D-M2–3 in the Luanping Basin. In addition, we suggest that the periods characterized by warm summer, enhanced hydrological cycle and expanded lake at high obliquity and low precession at eccentricity maximum were most conducive for both development and preservation of the early Jehol Biota.
... Although the Jehol Biota provides some of the most important palaeontological insights into early bird evolution (Zhou & Wang, 2010;Zhou, 2014;Xu et al., 2020), the anatomical and taxonomic knowledge of Jehol birds is far from complete. Few attempts have been made to provide comprehensive descriptions of the cranial morphology of Jehol birds, either 2D or three-dimensional (3D), with only some 2D descriptions for confuciusornithiforms and enantiornithines, and with limited 3D descriptions for Sapeornis and one juvenile enantiornithine (O'Connor & Chiappe, 2011;Elżanowski et al., 2018;Wang et al., 2019aWang et al., , 2021Hu et al., 2020a). ...
Jeholornis is a representative of the earliest-diverging bird lineages, providing important evidence of anatomical transitions involved in bird origins. Although ~100 specimens have been reported, its cranial morphology remains poorly documented owing to poor two-dimensional preservation, limiting our understanding of the morphology and ecology of the key avian lineage Jeholornithiformes, in addition to cranial evolution during the origin and early evolution of birds. Here, we provide a detailed description of the cranial osteology of Jeholornis prima, based primarily on high-quality, three-dimensional data of a recently reported specimen. New anatomical information confirms the overall plesiomorphic morphology of the skull, with the exception of the more specialized rostrum. Data from a large sample size of specimens reveal the dental formula of J. prima to be 0–2–3 (premaxillary–maxillary–dentary tooth counts), contrary to previous suggestions that the presence of maxillary teeth is diagnostic of a separate species, Jeholornis palmapenis. We also present evidence of sensory adaptation, including relatively large olfactory bulbs in comparison to other known stem birds, suggesting that olfaction was an important aspect of Jeholornis ecology. The digitally reconstructed scleral ring suggests a strongly diurnal habit, supporting the hypothesis that early-diverging birds were predominantly active during the day.
... In this warmer climate, various forms of life, namely the Early Cretaceous terrestrial Jehol Biota, are mainly distributed in North China and are at paleolatitudes of ~40°N latitude (Zhou 2014). However, in recent years, the records of Early Cretaceous abnormal climate fluctuation were found in North China at 30-40°N latitude: Cheng et al. (2002) believed that glacial debris flow deposits developed in the Lower Cretaceous at the northeastern margin of the Ordos Basin (Cheng et al. 2002); Wang et al. (1996) discovered red bed ice-rafted deposits in the Quantou Formation in the Songliao Basin (Wang et al. 1996); Xenoxylon, which represents a colder climate, is preserved in North China (Ding et al. 2016); Northeast China lacks thermophilic reptiles such as crocodiles (Zhou and Wang 2010); The pollen of cold and warm (conifers) plants coexist in North China (Zhong et al. 2019). The estimated average paleotemperature is approximately 10°C by Amiot et al. (2011), which is a present-day cool temperate climatic condition (e.g. ...
The Cretaceous, with relatively high atmospheric CO2 concentrations, temperatures, and sea levels, is a typical greenhouse period in the geologic history of the Earth. Recently, gradual changes in the understanding of the Early Cretaceous climate have led to the proposal of small-scale polar glaciers or cooling events in polar regions. However, the cooling events in North China at mid-latitudes are still poorly understood. In this study, 48 wedge structures that are 35-240 cm wide and from 120 cm to more than 300 cm in depth down through the bottom of Lower Cretaceous Luohe Formation conglomerate and penetrated vertically downward into Jurassic Anding Formation mudstone in the Ordos Basin, North China. According to findings, these sand wedges are distinguished by large-scale V shapes, vertical to steeply dipping laminations and pebbles, aeolian abrasion of sand grains, ventifacts, upturned host strata, and other features that are most likely the source of thermal contraction cracking in periglacial desert region. The sand wedges formed when thermal contraction cracks were filled with wind-blown deposits and other material in dry, cold, and windy regions. This record may support the hypothesis that the Ordos Basin, western North China, may have been at a high altitude and was cold during the Early Cretaceous. During the Early Cretaceous, the complex topography and climate of North China allowed the Jehol Fauna to coexist with permafrost and glaciers.
Community zonation of the Late Cretaceous Western Interior Seaway (WIS) has been suggested for bivalves, cephalopods, foraminifera, gastropods, and tetrapods. Most proposed WIS community zones consist of a northern and southern subprovince with a gradational boundary across central or south-central North America. Since it has been over three decades since the WIS community zonation hypothesis has been investigated for vertebrates, recent radiometric age determinations, taxonomic revisions, additional specimen discoveries, and recently available online museum specimen catalogues allow for an updated description of Manitoba (MB) escarpment faunal assemblages and testing of the community zonation hypothesis. Nine time bins were used to represent nine Upper Cretaceous lithostratigraphic units of the MB escarpment to test the zonation hypothesis consistency for nearly the entire Late Cretaceous (~71–95 Ma). Relatively high genus-level community similarity values (25–50%) of south-central WIS localities and low values (<20%) of localities furthest north and south support the existence of a central subprovince during late Cenomanian to early Turonian and late Coniacian to early Campanian times, when the gradational subprovincial boundary would have been furthest south between Kansas and Texas localities. Comparatively low genus-level community similarity values (<25%) of all localities south of MB during mid-Cenomanian and early to mid-Campanian times indicate the southern subprovincial boundary was farthest north between MB and South Dakota localities during these time intervals and had migrated throughout the Late Cretaceous. This work highlights significant fluctuations in vertebrate community zonation throughout the WIS through time and space and offers insight into the magnitude of compositional and palaeoecological changes that can occur in shallow marine vertebrate communities over an approximately 25 million year interval.
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Two Jurassic–Cretaceous anurans are described based on well-preserved specimens from the lower part of the Yixian Formation, western Liaoning Province, northeastern China. One specimen, from the Heitizigou site, documents a new genus and species, and the second, from the Sihetun site, is the holotype and only known specimen for the recently named Callobatrachus sanyanensis. Phylogenetic relationships of the major archaeobatrachian anuran clades are investigated with incorporation into the analysis of selected (well-established) early fossil taxa. The new taxon named and described in this paper is placed as the representative of a distinct archaic anuran clade, and Callobatrachus is considered to be an ingroup member of the Discoglossidae, constituting the earliest record of the family from Asia. The oldest known fossil anuran, Prosalirus from the Early Jurassic of Arizona, is grouped with Notobatrachus as sister taxa, and the two together form the most basal clade of Anura. Contradicting the widely accepted Leiopelmatidae–Discoglossidae sistergroup relationship, new evidence places the Leiopelmatidae as the most basal extant familial group and the sister group to other archaeobatrachian clades. The relationships and classification of the major archaic anuran clades are discussed, based on the phylogenetic results of this study.
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Recent coelurosaurian discoveries have greatly enriched our knowledge of the dinosaur-bird transition, but all reported taxa close to this transition are from relatively well-known coelurosaurian groups^1-3^. Here we report a new basal avialan, Epidexipteryx hui gen. et sp. nov., from the Middle-Late Jurassic of Inner Mongolia, China. This new species is characterized by an unexpected combination of characters seen in several different theropod groups, particularly the Oviraptorosauria. Phylogenetic analysis shows it to be the sister taxon to Epidendrosaurus^4,5^, forming a new clade at the base of Avialae^6^. Epidexipteryx also possesses two pairs of elongate ribbon-like tail feathers (ETFs), and its limbs lack contour feathers for flight. This finding shows that a member of the avialan lineage experimented with integumentary ornamentation as early as the Middle-Late Jurassic, and provides further evidence relating to this important aspect of the transition from non-avian theropods to birds.
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132 vertebrate species are known from the Messel Fossil Site. In this paper, all species and genera are listed, and for each of them the first report from Messel is cited. Moreover, recent discoveries and current research projects are mentioned. The list thus reflects the state-of-the-art knowledge on the present taxonomic status of all vertebrate species and genera of Messel. Kurzfassung 132 Wirbeltierarten sind derzeit aus der Fossilienfundstelle Grube Messel bekannt. Sie werden hier aufgeführt. Darüber hinaus werden jeweils erste Nachweise aus Messel, neue Funde sowie laufende Forschungsprojekte genannt, wodurch der aktuelle taxonomische Status der einzelnen Arten und Gattungen widergegeben wird.
A new species of enantiornithine bird from the Lower Cretaceous Yixian Formation of northeastern China is reported. The new taxon, Shanweiniao cooperorum, possesses several enantiornithine synapomorphies as well as the elongate rostral morphology (rostrum equal to or exceeding 60% the total length of the skull) of the Chinese early Cretaceous enantiornithines, Longipteryx chaoyangensis and Longirostravis hani. The discovery of this new specimen highlights the existence of a diverse clade of trophically specialized enantiornithines, Longipterygidae, for which we present phylogenetic support in a new comprehensive cladistic analysis of Mesozoic birds. Shanweiniao provides new information on the anatomy of longipterygids, and preserves a rectricial morphology previously unknown to enantior-nithines, with at least four tail feathers closely arranged. This supports the hypothesis that enantiornithines were strong fliers and adds to the diversity of known tail morphologies of these Cretaceous birds.