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An Updated Review of the Middle-Late Jurassic Yanliao Biota: Chronology, Taphonomy, Paleontology and Paleoecology

  • Paleontological Museum of China, Beijing, China

Abstract and Figures

The northeastern Chinese Yanliao Biota (sometimes called the Daohugou Biota) comprises numerous, frequently spectacular fossils of non-marine organisms, occurring in Middle-Upper Jurassic strata in western Liaoning, northern Hebei, and southeastern Inner Mongolia. The biota lasted for about 10 million years, divided into two phases: the Bathonian-Callovian Daohugou phase (about 168–164 million years ago) and the Oxfordian Linglongta phase (164–159 million years ago). The Yanliao fossils are often taphonomically exceptional (many vertebrate skeletons, for example, are complete and accompanied by preserved integumentary features), and not only are taxonomically diverse but also include the oldest known representatives of many groups of plants, invertebrates, and vertebrates. These fossils have provided significant new information regarding the origins and early evolution of such clades as fleas, birds, and mammals, in addition to the evolution of some major biological structures such as feathers, and have demonstrated the existence of a complex terrestrial ecosystem in northeast China around the time of the Middle-Late Jurassic boundary.
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1 Introduction
The Yanliao Area is a large region of northeast China,
situated north of the Yan Mountains, south of the
Daxinganling (Greater Khingan) Mountains, east of the
Hunshandak Sandland, and west of the Liao River.
Between roughly the middle of the Jurassic and the middle
of the Cretaceous, three major volcanic and sedimentary
cycles in the Yanliao Area successively produced the
Middle-Upper Jurassic Haifanggou/Jiulongshan and
Lanqi/Tiaojishan formations, the Upper Jurassic-Lower
Cretaceous Tuchengzi/Houcheng and Zhangjiakou
formations, and the Lower Cretaceous Dabeigou, Yixian
and Jiufotang formations. The famous and widely
distributed Early Cretaceous Jehol fossil assemblages
formed during the third cycle. However, strata deposited
during the first cycle contain a comparable fossil biota of
Middle-Late Jurassic age that is known as the Yanliao
Biota (Fig. 1) and is currently emerging as a source of
paleontological discoveries that may rival the Jehol Biota
in importance.
The Yanliao fossil assemblages were first noticed in
1983, when a rich insect assemblage was discovered from
the Middle Jurassic Jiulongshan Formation in the Yanliao
Area. The term “Yanliao Insect Fauna” was first
introduced to refer to this assemblage (Hong, 1983), but
was superseded by “Yanliao Fauna” after it became
apparent that a wide range of additional non-marine taxa
including conchostracans, bivalves, fish, reptiles, and
mammaliaforms coexisted with the insects (Ren et al.,
1995). Ren et al. (1995) further suggested that the fauna
should also be considered to include fossils from the
Mentougou, Tiaojishan, and Tuchengzi formations
exposed in the Yanliao Area, which were then believed to
be Middle Jurassic. The diagnostic elements of the
Yanliao Fauna were suggested to be the insect
Yanliaocorixa, the fish Liaosteus, and the lizard
‘Yabeinosaurus’ (Ren et al., 1995), which are known only
from the laterally equivalent Jiulongshan and Haifanggou
formations (Hoffstetter, 1964; Zhou et al., 1991; Jin, 1999;
Ren et al., 2010). It should be noted that ‘Yabeinosaurus
here refers toYabeinosaurus youngi, which is known
from the Jiulongshan/Haifanggou Formation of Lingyuan,
Liaoning. A recent study indicated that this species differs
markedly in limb proportions from the type species of the
An Updated Review of the Middle-Late Jurassic Yanliao Biota:
Chronology, Taphonomy, Paleontology and Paleoecology
XU Xing1, *, ZHOU Zhonghe1, Corwin SULLIVAN1, WANG Yuan1 and REN Dong2
1 Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology and
Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China
2 College of Life Sciences, Capital Normal University, Haidian District, Beijing 100048, China
Abstract: The northeastern Chinese Yanliao Biota (sometimes called the Daohugou Biota) comprises
numerous, frequently spectacular fossils of non-marine organisms, occurring in Middle-Upper
Jurassic strata in western Liaoning, northern Hebei, and southeastern Inner Mongolia. The biota
lasted for about 10 million years, divided into two phases: the Bathonian-Callovian Daohugou phase
(about 168-164 million years ago) and the Oxfordian Linglongta phase (164-159 million years ago).
The Yanliao fossils are often taphonomically exceptional (many vertebrate skeletons, for example, are
complete and accompanied by preserved integumentary features), and not only are taxonomically
diverse but also include the oldest known representatives of many groups of plants, invertebrates, and
vertebrates. These fossils have provided significant new information regarding the origins and early
evolution of such clades as fleas, birds, and mammals, in addition to the evolution of some major
biological structures such as feathers, and have demonstrated the existence of a complex terrestrial
ecosystem in northeast China around the time of the Middle-Late Jurassic boundary.
Key words: Yanliao Biota, Daohugou phase, Linglongta phase, Middle-Late Jurassic, Yanliao area
Vol. 90 No. 6 pp.2229–2243 ACTA GEOLOGICA SINICA (English Edition) Dec. 2016
* Corresponding author. E-mail:
© 2016 Geological Society of China
2230 Vol. 90 No. 6 Dec. 2016
genus, Yabeinosaurus tenuis from the Lower Cretaceous
Jehol Group, and its attribution to the genus is still
equivocal (Evans and Wang, 2012).
During 2009 and 2010, the term ‘Yanliao Biota’ was
independently introduced by multiple authors, but with
different definitions. Sun et al. (2009) defined the Yanliao
Biota as the entire Jurassic flora and fauna of western
Liaoning Province and neighboring areas (Sun et al.,
2009; Sun et al., 2011). Chang et al. (2009), by contrast,
used the term ‘Yanliao Biota’ to designate the fossil
content of the Haifanggou Formation of Liaoning
Province and the laterally equivalent Jiulongshan
Formation of Hebei Province, a definition concordant with
the original definition of the “Yanliao Insect Fauna”. Zhou
et al. (2010) considered the Yanliao Biota to include all
fossils from the Middle Jurassic Haifanggou/Jiulongshan
Formation and the Upper Jurassic Lanqi/Tiaojishan
Formation. Because the Jiulongshan/Haifanggou
Formation and the Lanqi/Tiaojishan Formation share
strong similarities in terms of lithology, taphonomy and
fossil content (Zhou et al., 2010; Sullivan et al., 2014), we
follow Zhou et al.’s (2010) definition of the Yanliao Biota.
The Yanliao Biota has sometimes been referred to as the
Daohugou Biota (e.g. Sullivan et al. 2014), after the
Daohugou Locality in southeastern Inner Mongolia, but
the term ‘Yanliao Biota’ is gaining currency among
researchers and is preferred here. Sullivan et al. (2014)
also defined the ‘Daohugou Biota’ more narrowly,
restricting its scope to sites where the abundant
salamander Chunerpeton tianyiensis is known to occur.
Over the last decade, numerous significant fossils have
been recovered from the Yanliao Biota, including plants
that may represent the oldest known angiosperms (Wang,
2010a), the earliest known members of several insect
clades (Huang et al., 2008a; Huang and Nel, 2009; Ren et
al., 2010; Gao et al., 2012; Huang et al., 2012; Huang,
2014; Huang and Cai, in press), the best-preserved crown
salamanders known from the Jurassic (Gao and Shubin,
2003; Gao and Shubin, 2012), transitional pterosaurs (Lü
et al., 2010a; Wang et al., 2010c), some of the earliest
known feathered dinosaurs such as the paravian
Anchiornis and the heterodontosaurid Tianyulong (Xu and
Zhang, 2005; Zhang et al., 2008a; Hu et al., 2009;
Godefroit et al., 2013a; Godefroit et al., 2013b), the
earliest known gliding and aquatic mammaliaforms (Ji et
al., 2006; Meng et al., 2006), and the earliest known
eutherian mammal (Luo et al., 2011). The Yanliao Biota
has thus emerged as a terrestrial fossil assemblage of the
greatest importance for understanding Mesozoic
ecosystems and the evolution of many major groups of
organisms, including the origins of some large modern
clades. The present paper provides an updated review of
this important biota, building on previous reviews written
from various perspectives (Liu et al., 2010; Zhou et al.,
2010; Guo et al., 2012; Zheng, 2013; Sullivan et al., 2014;
Xu et al., 2016).
2 Temporal and Geographical Distribution of
the Yanliao Biota
Like the Early Cretaceous Jehol Biota, the Yanliao
Biota is a collection of non-marine fossil assemblages
mainly preserved in lacustrine deposits rich in volcanic
ash. Major known Yanliao localities, many of which are
referred to by more than one name in current literature,
include: the Fangshen (Zhou et al., 1991), Yujiagou (Lu,
1995; Jin, 1999), Yipandaogou, Liaotugou (Wang et al.,
1989), Daxishan (or Daxigou), Yaolugou, Wubaiding (or
Reshuitang) (Zhang and Wang, 2004; Wang et al., 2005;
Wang and Evans, 2006), Guancaishan (or Guancailing)
(Liu et al., 2006b; Wang et al., 2010d; Gao and Shubin,
2012; Liu et al., 2012), and Haifenggou localities in
western Liaoning Province; the Fanzhangzi, Bawanggou
(or Mutoudeng) (Lü, 2009; Lü et al., 2012; Zheng et al.,
2013), and Zhuanshanzi (or Nanshimen or Gangou)
localities in northern Hebei Province; and the
Wuhuaxigou, Chentaizi, Jiangzhangzi, and Daohugou
localities in southeastern Inner Mongolia (Fig. 2). These
Fig. 1. Stratigraphic distribution of the Middle-Upper Juras-
sic Yanliao Biota and the Lower Cretaceous Jehol Biota.
Shading indicates the rock formations that contain the Yanliao and Jehol
Dec. 2016 Vol. 90 No. 6 2231
localities are within the core geographical range of the
Jehol Biota, and are widely accepted as sites where the
Haifanggou/Jiulongshan and/or Lanqi/Tiaojishan
formations crop out (Huang, 2015), but details remain
poorly known.
The Haifanggou/Jiulongshan Formation has been widely
accepted as Middle Jurassic in age based on
paleontological data (Ren et al., 1995; Shen et al., 2003;
Xu et al., 2003). Isotopic radiometric data have supported
this age assessment (Chen et al., 1997; Xu et al., 2003; Wu
et al., 2004) and further indicated that the formation is
partially Bathonian but mainly Callovian. For example, a
40Ar/39Ar age of 166.7+1.0 Ma was obtained from samples
collected from around the middle of the Haifanggou
Formation of Beipiao, western Liaoning Province (Chang
et al., 2013). However, the formation might extend into the
Upper Jurassic in some areas, such as in the Chengde Basin
of northern Hebei Province (Liu et al., 2006a; Zhang et al.,
2008b). The Lanqi/Tiaojishan Formation is probably
Oxfordian. Two recent 40Ar/39Ar dates for the Lanqi
Formation of western Liaoning are 159.5±0.6 Ma, based on
samples from the top of the formation (Chang et al., 2013),
and 161.8±0.4 Ma based on samples from the basal portion
of the formation (Chang et al., 2009; Chang et al., 2013).
The strata exposed at the Daohugou Locality belong
mainly to the Bathonian-Callovian Haifanggou Formation.
Radiometric dates based on samples collected from
volcanic rocks overlying the fossil-bearing layers at the
Daohugou Locality include two 206Pb/238U SHRIMP ages
of 162±2 Ma and 152±2.3 Ma (Liu et al., 2006b), an 40Ar-
39Ar age of 159.8±0.8 Ma (He et al., 2004; Liu et al.,
2006b), two 206Pb/238U SHRIMP ages of 166±1.5 Ma and
165±2.4 Ma, an 40Ar-39Ar age of 164±2.5 Ma (Chen et al.,
2004), and two 206Pb/238U SHRIMP ages of 164±1.2 Ma
and 165±1.2 Ma (Yang and Li, 2004). A 206Pb/238U
SHRIMP age of 165.0 ±1.2 Ma was obtained based on
samples collected from the fossil-bearing layers (Yang and
Li, 2008), suggesting that the Daohugou beds are mostly
Callovian. The beds at this locality have been informally
termed the ‘Daohugou Formation’ (Zhang, 2002), but this
usage has received little acceptance.
Other sites at which strata of the Haifanggou/
Jiulongshan Formation are considered to be exposed
include the Wuhuaxigou, Chentaizi, Jiangzhangzi,
Wubaiding, Guancaishan, Haifenggou, Fanzhangzi, and
Zhuanshanzi localities (Huang, 2015). Radiometric dates
are available for some of these exposures. The Callovian
Jiulongshan Formation is probably exposed at the
Wubaiding Locality, and samples collected from lavas
below the salamander-bearing beds at this site yielded a
206Pb/238U SHRIMP age of 164±4 Ma (Liu et al., 2006b).
However, the same sample also produced a 206Pb/238U
TIMS age of 168±4.4 Ma (Liu Yongqing, personal
communication). A minimum 206Pb/238U age of 157±3 Ma
has been reported for the salamander-bearing beds at the
Guancaishan Locality (Liu et al., 2006b). It is noteworthy
that controversy surrounds the stratigraphy at some of
these localities. For example, the beds at the Zhuanshanzi
Locality have been considered to pertain to the Tiaojishan
Formation by some studies (Xu et al., 2015), but to the
Haifanggou/Jiulongshan Formation by others (Huang,
2015). Similarly, either the Haifanggou/Jiulongshan
Formation (Huang, 2015) or Lanqi/Tiaojishan Formation
(Liu et al., 2006b; Gao and Shubin, 2012) may crop out at
the Guancaishan Locality.
The Daxishan Locality, another major Yanliao site,
exposes the Oxfordian Lanqi/Tiaojishan Formation.
Radiometric dates based on samples above, directly from,
or below the Anchiornis-bearing bed at the Daxishan
Locality in Linglongta, Jianchang, Liaoning Province
include U-Pb SHRIMP ages of 160.5±0.99 Ma and
Fig. 2. Geographic distribution of major Yanliao Biota lo-
calities (Daohugou Biota localities in red and Linglongta
Biota localities in blue).
Shaded area enlarged and shown in detail. 1, Wuhuaxigou Locality; 2,
Jiangzhangzi Locality; 3, Chentazi Locality; 4, Daohugou Locality; 5,
Guancaishan Locality; 6, Wubaiding (Reshuitang) Locality; 7, Fangshen
Locality; 8, Yujiagou Locality; 9, Liaotugou Locality; 10, Yipandaogou
Locality; 11, Fanzhangzi Locality; 12, Bawanggou (Mutoudeng) Local-
ity; 13, Zhuanshanzi (Gangou or Nanshimen) Locality; 14, Yaolugou
Locality; 15, Daxishan (Daxigou) Locality.
2232 Vol. 90 No. 6 Dec. 2016
161.0±1.44 Ma (Liu et al., 2012), of 160.7±3.2 Ma (Peng
et al., 2012), and of 160.7±1.7 Ma, 158.9±1.7 Ma, and
159.5±2.3 Ma (Wang et al., 2013), suggesting that the
Anchiornis-bearing bed is Oxfordian in age. Exposures at
several other sites, including the Yaolugou and
Bawanggou localities, have been suggested to also
represent the Lanqi/Tiaojishan Formation (Xu et al.,
2015). Volcanic rocks overlying the Tianyulong-bearing
bed at the Yaolugou Locality have been radiometrically
dated to 158.5 ±1.6 Ma (Liu et al., 2012).
The most common elements of the Yanliao fossil
assemblages include the plants Yanliaoia, Yimaia and
Schmeissneria, the conchostracans Euestheria
luanpingensis and Qaidamestheria, the bivalves
Ferganoconcha sibirica and Arguniella, the insect
Yanliaocorixa chinensis, the fish Liaosteus, the
salamander Chunerpeton, the pterosaur Darwinopterus,
and the dinosaur Anchiornis (Huang, 2015). Among these
fossils, some are common in the lower Haifanggou/
Jiulongshan Formation (e.g., Euestheria luanpingensis,
Ferganoconcha sibirica, and Yanliaocorixa chinensis),
some common in the upper Lanqi/Tiaojishan (e.g.,
Arguniella, Darwinopterus and Anchiornis), and some in
both (e.g., Chunerpeton and Yanliaoia). Only a small
number of species are known in both the Haifanggou/
Jiulongshan Formation and the Lanqi/Tiaojishan
Formation, and only a few small clades are unique to the
Yanliao Biota. Among the vertebrate fossils, the theropod
dinosaur groups Scansoriopterygidae and Anchiornithinae
(here defined as the most inclusive clade including
Anchiornis but not Archaeopteryx, Gallus, Troodon,
Dromaeosaurus, Unenlagia, or Epidexipteryx) are
probably restricted to the Yanliao Biota, although the
juvenile holotype of Zhongornis haoae from the Jehol
Biota has been tentatively interpreted as a Cretaceous
scansoriopterygid (O'Connor and Sullivan, 2014).The
scansoriopterygid and anchiornithine clades occur in both
the Haifanggou/Jiulongshan Formation and the Lanqi/
Tiaojishan Formation. However, scansoriopterygids come
mostly from the Haifanggou/Jiulongshan Formation,
whereas most anchiornithines (Pedopenna is likely to be
an anchiornithine, based on its pedal morphology) come
from the Lanqi/Tiaojishan Formation.
Because of the compositional differences between the
fossil assemblages of the older Haifanggou/Jiulongshan
Formation and the younger Lanqi/Tiaojishan Formation,
the Yanliao Biota has been divided into the geologically
older Daohugou Biota and the younger Linglongta Biota
(Huang, 2015). The term ‘Daohugou Biota’, although
sometimes used in reference to the entirety of the Yanliao
Biota (e.g. Sullivan et al., 2014), was originally introduced
to refer to fossils from the exposed strata at the Daohugou
Locality (Ji and Yuan, 2002; Zhang, 2002). Various ages
have been proposed for the Daohugou exposures, ranging
from Middle Jurassic to Early Cretaceous (Wang et al.,
2000; Wang, 2000; Ji and Yuan, 2002; Ren and Yin, 2002;
Zhang, 2002; Gao and Shubin, 2003; Wang et al., 2005; Xu
and Zhang, 2005; Ji et al., 2006), but their assignment to
the Bathonian-Callovian Haifanggou/Jiulongshan
Formation is now widely accepted (Huang, 2015). The
term ‘Linglongta Biota’ was introduced because the fossil
assemblages from the Oxfordian Lanqi/Tiaojishan
Formation are best known from several localities, including
the Daxishan Locality, that lie in Linglongta Town,
Jianchang County, Liaoning Province (Huang, 2015).
3 Taphonomy, Paleoenvironment, and
Paleoclimate of the Yanliao Biota
The Yanliao Biota bears a close taphonomic
resemblance to the Jehol Biota. For example, the majority
of the fossils recovered from the Yanliao Biota are
complete and articulated; many fine-scale structures are
preserved, including wing veins and mouthparts of insects
(Zhang and Kluge, 2007; Ren et al., 2010; Wang and
Zhang, 2010) and tiny branchiopod eggs (Shen and
Huang, 2008); specimens often retain preserved soft
tissue, including setae and hairs in insects (Zhang et al.,
2008c; Ren et al., 2010; Gao et al., 2012), skin
impressions in salamanders (Wang and Evans, 2006;
Wang et al., 2010d) and lizards (Evans and Wang, 2007;
Wang et al., 2010d), filamentous structures and
membranes in pterosaurs (Ji and Yuan, 2002; Wang et al.,
2002), feathers and membranes in dinosaurs (Zhang et al.,
2008a; Hu et al., 2009; Xu et al., 2015), and fur in
mammaliaforms (Meng et al., 2006; Zhou et al., 2013);
stomach contents are preserved in some fossils (Dong et
al., 2012); and some rarely preserved organisms, such as
lichens, are plausibly represented in the biota (Wang et al.,
2010a). A few Yanliao fossils are even preserved in ways
that provide behavioral information. For example, some
male-female pairs of the pachymeridiid Peregrinpa-
chymeridium comitcola are preserved in a mating position,
with the head of the male near the tail of the female and
vice versa (Lu et al., 2011).
Recurrent volcanism probably explains the apparently
sudden deaths, rapid burial, and excellent preservation of
the Yanliao organisms (Chang et al., 2009; Sullivan et al.,
2014). Other possible causal factors include severe
environmental and climatic changes, and even periodic
collapse of the aquatic ecosystem as argued for the Jehol
Biota (Pan et al., 2011). Possible mechanisms of
exceptional fossil preservation for the Yanliao Biota
include microbial processes (O'Brien et al., 2002; Jiang et
Dec. 2016 Vol. 90 No. 6 2233
al., 2011) and pyritization (Wang et al., 2008), although
the mechanism may also have been an unknown one
operating within an unusual fossilization microenvironment
produced by clay minerals (Briggs, 2003; Martínez-
Delclòs et al., 2004).
In general, the Yanliao paleoenvironment and climate
are poorly known, though some tentative inferences are
possible. Study of some insect fossils suggested that the
Daohugou area might have been a near–shore shallow
lacustrine basin with a warm, humid climate, diverse and
abundant vegetation, and highly aquiferous soil (Tan and
Ren, 2002; Ren et al., 2010; Selden et al., 2011), though
several different microenvironments may have coexisted
in the Daohugou paleolakes (Wang et al., 2008). In
general, this inference is consistent with paleobotanical
data from the Haifanggou Formation of western Liaoning
Province, which suggest warm but seasonal and/or moist
conditions (Sun et al., 2008). Some fossil evidence
supports a high paleoaltitude for the Daohugou area,
including the presence of mountain insects (Ren, 2003;
Engel and Ren, 2008) and mountain salamanders (Wang,
2004). Most paleobotanical data suggest a hot, dry climate
in the Yanliao Area during the period when the Lanqi
Formation was being deposited (Zheng, 2013), though
some silicified coniferous wood fossils suggest a
subtropical, humid and seasonal climate in western
Liaoning Province (Jiang et al., 2008). Consequently,
there seem to have been some temporal and spatial
variations in the Yanliao paleoenvironment and climate.
4 Biodiversity and Paleoecology of the
Yanliao Biota
To date, at least 216 plant species (Jiang et al., 2008;
Wang, 2010a; Zheng, 2013), more than 500 insect and
other invertebrate species, and 36 vertebrate species have
been reported from the Yanliao Biota (Xu et al., 2016).
Known plant fossils include spores, leaves, and wood
(Zheng, 2013), from lichens, horsetails, lycopsids, filicins,
bennettitaleans, cycads, ginkgos, czekanowskialeans,
conifers, osmundaceans, dipterids, dicksonians, and
possibly angiosperms (Fig. 3) (Pan, 1983; Wang, 2010a;
Wang and Wang, 2010; Zheng, 2013). Most of these
fossils are from Liaoning, with 124 species from the
Haifanggou Formation and 92 species from the Lanqi
Formation (Zheng, 2013).
Insect fossils are represented by at least 500 species
from 24 major insect groups, among which coleopterans,
dipterans, homopterans, hemipterans and hymenopterans
are especially high in specific diversity (Fig. 4). Other
invertebrate groups (Fig. 4) include branchiopods (Huang
and Cai, in press), gastropods (Chang and Sun, 1997;
Jiang, 2006), bivalves (Wang et al., 1989; Chang and Sun,
1997; Jiang, 2006; Duan et al., 2009), conchostracans
(Shen et al., 2003), ostracods (Duan et al., 2009) and
arachnids (Selden et al., 2008; Huang et al., 2009; Selden
and Huang, 2010; Selden et al., 2011).
Among vertebrates (Fig. 5), fish from the Yanliao Biota
include acipenseriforms (Lu, 1995; Jin, 1999) and some
undescribed ptycholepids (Duan et al., 2009). Caudates of
uncertain systematic position are present (Wang, 2000;
Wang and Rose, 2005; Wang and Evans, 2006), in
addition to a cryptobranchid (Gao and Shubin, 2003), a
hynobiid (Wang, 2004) and salamandroids (Gao and
Shubin, 2012). Lizards are represented by a species of
uncertain systematic position (Evans and Wang, 2012) and
two possible scleroglossans (Evans and Wang, 2007;
Evans and Wang, 2009). Pterosaurs are diverse, and
include anurognathids (Wang et al., 2002; Lü and Hone,
2012), a rhamphorhynchine (Lü et al., 2012),
scaphognathines (Czerkas and Ji, 2002; Lü et al., 2010b;
Lü and Bo, 2011; Cheng et al., 2012), wukongopterids
(Lü, 2009; Wang et al., 2009; Lü et al., 2010a; Wang et
al., 2010c; Lü et al., 2011), and a possible istiodactylid
(Lü and Fucha, 2010; Martill and Etches, 2012).
Dinosaurs include several maniraptoran theropods
(Czerkas and Yuan, 2002; Zhang et al., 2002; Xu and
Zhang, 2005; Zhang et al., 2008a; Xu et al., 2009; Xu et
al., 2011; Godefroit et al., 2013a; Godefroit et al., 2013b)
and the heterodontosaurid ornithischian Tianyulong (the
holotype of Tianyulong confuciusi was originally claimed
to be from the Lower Cretaceous Yixian Formation
(Zheng et al., 2009), but later studies have considered the
taxon to belong to the Yanliao Biota (Liu et al., 2012)).
The mammaliaform assemblage is relatively diverse, and
includes the gliding mammal Volaticotherium (Meng et
al., 2006), triconodonts (Yabe and Shikama, 1938; Zhou et
al., 1991), a yinotherian (Luo et al., 2007), docodonts (Ji
et al., 2006; Luo et al., 2015b; Meng et al., 2015),
haramiyidans (Zheng et al., 2013; Zhou et al., 2013; Bi et
al., 2014), a multituberculate (Yuan et al., 2013), and a
eutherian (Luo et al., 2011).
Yanliao insect herbivores include various cicadomorphs
(Huang and Cai, in press) and the prophalangopsid
Bacharaboilus lii, whose mouthparts appear suitable for
feeding on vegetation (Gu et al., 2011). The biota also
includes scorpionflies that may have fed on pollination
drops secreted by gymnosperms (Ren, 1998; Ren et al.,
2009; Shih et al., 2011). The inferred mutualistic
relationship between the scorpionflies and the
gymnosperms considerably preceded “the similar and
independent coevolution of nectar-feeding flies, moths, and
beetles on angiosperms” (Ren et al., 2009). Direct evidence
for insect herbivory comes from stomach contents
2234 Vol. 90 No. 6 Dec. 2016
preserved in some permopsocid fossils, which probably
represent sporangia (Huang et al., 2016). Vertebrate
herbivores include the haramiyidan mammal Megaconus
mammaliaformis (Zhou et al., 2013), the multituberculate
mammal Rugosodon eurasiaticus (Yuan et al., 2013), and
the heterodontosaurid dinosaur Tianyulong confuciusi
(Zheng et al., 2009).
Yanliao predatory insects include various odonatans
(Huang and Cai, in press) such as the damsel-dragonfly
Zygokaratawia reni (Nel et al., 2008), above-ground
diurnal geinitziids (Cui et al., 2012), the raphidiomimid
cockroach Graciliblatta bella (Liang et al., 2012), and the
flower bug Pumilanthocoris (Hou et al., 2012), all of
which fed on other insects. Predatory spiders include the
insect-eating nephilid Nephila jurassica (Selden et al.,
2011), among others. Vertebrate predators are diverse.
Stomach contents show that the salamander Jeholotriton
paradoxus fed on the conchostracan Euestheria
luanpingensis (Dong et al., 2012), whereas the salamander
Chunerpeton tianyiensis fed on the corixid insect
Yanliaocorixa chinensis (Dong et al., 2012). Several
Yanliao pterosaurs were probably piscivorous (Sullivan et
Fig. 3. Selected Yanliao plant fossils.
(a), the possible angiosperm Schmeisseria; (b), the bennettitalean Anomozamites sp.; (c), the filicalean Coniopteris sp.; (d), the cycad Nilssonia sp.;
(e), the conifer Yanliaoa sp.; (f), the ginkgoalean Yimaia capituliformis. Not to scale.
Dec. 2016 Vol. 90 No. 6 2235
al., 2014), but the anurognathid pterosaur Jeholopterus
ningchengensis was probably an aerial insectivore
(Bennett, 2007; Sullivan et al., 2014) and the pterosaur
Darwinopterus robustodens has been suggested to have
fed on hard-carapaced coleopterans (Lü et al., 2011). The
theropod Anchiornis was probably carnivorous and/or
insectivorous, and most Yanliao mammaliaforms were
probably insectivorous (Luo, 2007).
The most common aquatic insects include
Yanliaocorixa chinensis and ephemeropteran nymphs
(Huang and Cai, in press). Other aquatic insects include
the chresmodid Jurachresmoda gaskelli, which might have
been able to skate across the surfaces of bodies of fresh
water (Zhang et al., 2008c). Aquatic vertebrates include a
few fish, the salamanders Jeholotriton and Chunerpeton
(Dong et al., 2012), and the mammaliaform Castorocauda
Fig. 4. Selected Yanliao invertebrate fossils.
(a), the orthopteran Allaboilus gigantus; (b), the palaeontinid Daohugoucossus shii; (c), the ephemeropteran Jurassonuru samoenus; (d), the sipho-
napteran Pseudopulex jurassicus; (e), the corixid Yanliaocorixa chinensis; (f), the dipteran Strashila daohugouensis; (g), the harvestman Mesobunus
martensi; (h), the bivalve Ferganoconcha sibirica; (i), the conchostracan Euestheria luanpingensis. Not to scale.
2236 Vol. 90 No. 6 Dec. 2016
Fig. 5. Selected Yanliao vertebrate fossils.
(a), the ptycholepid fish PMOL-AF00743; (b), the cryptobranchid salamander Chunerpeton tianyiensis; (c), a juvenile lizard of uncertain sys-
tematic position; (d), the wukongopterid pterosaur Darwinopterus robustodens; (e), the anurognathid pterosaur Jeholopterus ningchengensis; (f),
the deinonychosaurian theropod Anchiornis huxleyi; the scansoriopterygid theropods Epidexipteryx hui (g) and Yi qi (h); (i), the haramiyidan
mammaliaform Shenshou lui; (j), the basal mammal Volaticotherium antiquus; (k), the eutherian mammal Juramaia sinensis. Not to scale.
Dec. 2016 Vol. 90 No. 6 2237
lutrasimilis (Ji et al., 2006).
The majority of Yanliao mammaliaforms were probably
terrestrial (Yuan et al., 2013; Zhou et al., 2013). However,
some may have been fossorial (Luo et al., 2007), or
perhaps even subterranean (Luo et al., 2015b). Similarly,
several Yanliao mammaliaforms were scansorial (Luo et
al., 2011) or even arboreal (Meng et al., 2006; Zheng et
al., 2013; Meng et al., 2015). Arboreal vertebrate species
probably also include several theropod dinosaurs (Zhang
et al., 2008a; Hu et al., 2009; Xu et al., 2009) and at least
one of the known lizards (Evans and Wang, 2009).
Capable flyers include numerous insects and various
pterosaurs (Sullivan et al., 2014); vertebrates that probably
possessed a more limited capacity for aerial locomotion
include the mammaliaform Volaticotherium (Meng et al.,
2006), the theropod Anchiornis huxleyi and its close
relatives (Xu et al., 2011), and the bizarre
scansoriopterygid Yi qi, whose wings were membranous
rather than feathered (Xu et al., 2015).
A few studies have identified evidence for specific
paleoecological interactions within the Yanliao Biota.
Several ectoparasites have been identified, including stem-
fleas which probably fed on dinosaurs, pterosaurs and
mammals (Gao et al., 2012; Huang et al., 2012). Several
species, including alloioscarabaeid insects, have been
suggested to have fed on decaying organic materials (Bai
et al., 2012), and Yanliao silphids are likely to have fed on
carcasses of small vertebrates such as mammals (Cai et al.,
2014). The exceptionally well-preserved stridulatory
structures present in a particular basal katydid suggest that
low-frequency insect song may have been a normal part of
the auditory background in the Yanliao Biota ecosystem
(Gu et al., 2012).
Yanliao Biota specimens provide evidence for
interactions between insects and plants, beyond the
presence of the probable insect herbivores mentioned
above. The bennettitalean Anomozamites villosus
frequently shows signs of herbivorous insect damage in
the distal parts of the leaves, but the proximal parts bear
protective ‘hairs’ and generally remain intact (Pott et al.,
2011). Some Yanliao insects appear to have mimicked
parts of gymnosperms, just as many modern insects mimic
parts of angiosperms. For example, two lacewings appear
to have mimicked pinnate cycadophyte leaves (Wang et
al., 2010e), and a hangingfly seemingly mimicked a type
of ginkgo leaf (Wang et al., 2012).
The Yanliao Biota displays several unusual
paleoecological features (Sullivan et al., 2014; Xu et al.,
2016): 1) a total absence of sauropodomorphs, and other
vertebrates larger than a few kg; 2) relative rarity of
herbivorous vertebrates; 3) absence of definitively aquatic
or semi-aquatic reptiles; 4) relative rarity of fish; 5)
numerical dominance of salamanders among aquatic
vertebrates; and 6) numerical dominance of theropods, as
opposed to ornithischians, among dinosaurs. The
extremely unusual characteristics of this ecosystem, as
preserved in the fossil record, are unquestionably due at
least in part totaphonomic filtering and collection bias
(Sullivan et al., 2014; Xu et al., 2016).
5 Significance of the Yanliao Biota
The Yanliao fossils have significant implications for our
understanding of many Mesozoic terrestrial and aquatic
groups. The earliest known definitive angiosperms are
from the Early Cretaceous (Sun et al., 2002; Friis et al.,
2005), considerably later than the predicted date of
origination for angiosperms based on molecular clock
studies (Sanderson et al., 2004; Smith et al., 2010).
Several plants from the Yanliao Biota might be helpful in
understanding the origin of flowering plants (Wang et al.,
2007; Wang, 2010a; b; Wang and Wang, 2010) and/or the
evolution of key angiosperm features (Wang et al., 2010b),
though this suggestion has received little attention. The
Yanliao Biota contains the earliest known members of
many insect groups, including fleas (Gao et al., 2012;
Huang et al., 2012), webspinners (Huang and Nel, 2009),
and mantophasmatods (Huang et al., 2008b), among
others (Huang, 2014). These fossils provide significant
information on the origins of the groups they represent,
and on patterns of early character evolution within those
groups. For example, several stem-fleas from the Yanliao
Biota support the hypothesis that fleas, which rank among
the major ectoparasitic insect groups, are nested within the
scorpionflies, and these fossils additionally shed new light
on the evolution of parasite-host relationships in fleas
(Gao et al., 2012; Huang et al., 2012). Yanliao fossils also
reveal a high degree of evolutionary stasis in many insect
groups (Huang et al., 2009; Ren et al., 2010), and in some
spider lineages (Selden et al., 2008; Selden and Huang,
2010). Yanliao strashilid fossils deserve special note
because they have helped to demonstrate that strashilids
are highly specialized flies (Diptera) rather than an
enigmatic group of ectoparasites, and that paedomorphism
is present among endopterygote insects (Huang et al.,
2013). In addition, there is evidence for parental care in
carrion beetles from the Yanliao Biota (Cai et al., 2014).
The Yanliao caudate fossils include the earliest known
members of three extant clades: cryptobranchids,
hynobiids, and salamandroids (Gao and Shubin, 2003;
Wang, 2004; Wang and Evans, 2006; Gao and Shubin,
2012; Jia and Gao, 2016). The Yanliao pterosaur fossils
represent an important transitional stage in pterosaur
evolution and demonstrate that modularity can play a key
2238 Vol. 90 No. 6 Dec. 2016
role in major evolutionary transformations (Lü et al.,
2010a). The Yanliao theropods strongly suggest that
several salient features, such as pennaceous feathers, the
four-winged condition, and even aerial locomotion, are
primitive for Paraves and date back to approximately the
Middle Jurassic (Hu et al., 2009; Xu et al., 2015). Among
mammaliaforms, Yanliao fossils extend the known record
of aquatic taxa (Ji et al., 2006), gliding taxa (Meng et al.,
2006) and eutherians (Luo et al., 2011) significantly
backward in time. The holotype of the Yanliao
haramiyidan Megaconus is preserved with typical
mammalian integument (Zhou et al., 2013), but the
evolutionary significance of this feature is uncertain
because of continuing controversy over the phylogenetic
position of haramiyidans. Several other Yanliao
haramiyidan fossils were reported to confirm that
multituberculates are phylogenetically nested within
Haramiyida and that haramiyidans lie within the
mammalian crown group, a conclusion that would imply
Triassic haramiyidans represent the oldest known crown-
mammals (Zheng et al., 2013; Bi et al., 2014). However, a
recent study of Triassic haramiyidan fossils suggested
instead that haramiyidans are not closely related to
multituberculates and fall outside the mammalian crown
(Luo et al., 2015a), which would imply that crown-group
mammals probably originated in the Jurassic as
traditionally believed. If haramiyidans indeed fall outside
the crown-group, however, then Megaconus can be taken
to demonstrate that a full pelage first appeared in non-
crown mammaliaforms.
The Yanliao Biota shares some common faunal
elements (e.g., haramiyid and docodont mammaliaforms,
and transitional pterosaurs) with contemporaneous
continental fossil assemblages such as the Shishugou
Biota of northwestern China (Clark et al., 2006; Hu et al.,
2007; Martin et al., 2010; He et al., 2013; Andres et al.,
2014) and the Shaximiao Biota of southwestern China
(Peng et al., 2005), but is compositionally distinct from
these more typical Jurassic assemblages in several
important respects. Relative to the Shishugou and
Shaximiao biotas, the Yanliao Biota contains much greater
taxonomic diversity within some clades, such as the
Pterosauria and Mammaliaformes, but much less diversity
in other clades such as the Testudines, Crocodylomorpha,
and Dinosauria; contains more crownward taxa within
various clades such as the Theropoda and Mammalia; and
contains many fewer herbivorous vertebrates, partly
reflecting a lack of tritylodontids, stegosaurian
ornithischians, and sauropods. These significant
differences may partially represent a preservational artifact
(lacustrine deposition for the Yanliao Biota vs fluvial
deposition for the Shishugou and Shaximiao biotas) or a
consequence of environmental differences (a more closed
forest habitat for the Yanliao Biota vs a more open habitat
for the Shishugou and Shaximiao biotas). However, they
may also suggest that some important clades genuinely
originated and underwent their early diversification in the
Yanliao Area, whereas others were genuinely rare or
absent in northeast China during Yanliao times. In any
case, the Yanliao Biota provides a unique window into the
poorly known terrestrial ecosystems of the Middle-Late
Jurassic transitional period and advances our
understanding of the early evolution of many clades.
Although significant advances have been made in
research on the Yanliao Biota over the last decade, our
understanding of this important biota is still limited. In the
future, the following areas will be particularly worthy of
attention and investigation: 1) further analysis of the
anatomy, phylogenetic positions and functional
morphology of key Yanliao taxa in order to achieve a
better understanding of important evolutionary events; 2)
scientific as opposed to commercial excavation of Yanliao
fossils, to make possible relatively unbiased
reconstructions of the Yanliao ecosystem; 3) establishing a
high-precision temporal framework for the Yanliao Biota,
and accurately determining the geographical and
stratigraphic distributions of various Yanliao fossils; 4)
reconstruction of the Yanliao paleoenvironment; and 5)
analysis of the evolution of the Yaoliao ecosystem in a
global context, in order to understand the evolution of the
wider earth-life system around the critical point in time
represented by the Middle-Late Jurassic boundary.
The authors thank Diying Huang, Zhexi Luo and Paul
Barrett for constructive reviews of the manuscript, and Ge
Sun, Shaolin Zheng, Diying Huang, Yongqing Liu, Keqin
Gao, Dongyu Hu, Changfu Zhou, Lijun Zhang, Xin Wang,
Haichun Zhang, David Hone, and Zhiyan Zhou for
discussion and for providing references and photographs.
This project was supported by the National Natural
Science Foundation of China (41120124002) and the
National Basic Research Program of China
Manuscript received July 1, 2016
accepted Aug. 10, 2016
edited by Liu Lian
Andres, B., Clark, J., and Xu, X., 2014. The earliest
pterodactyloid and the origin of the group. Current Biology,
24(9): 1011–1016.
Dec. 2016 Vol. 90 No. 6 2239
Bai, M., Ahrens, D., Yang, X., and Ren, D., 2012. new fossil
evidence of the early diversifiction of scarabs:
Alloioscarabaeus cheni (Coleoptera: Scarabaeoidea) from the
Middle Jurassic of Inner Mongolia, China. Insect Science, 19:
Bennett, S.C., 2007. A second specimen of the pterosaur
Anurognathus ammoni. Paläontologische Zeitschrift, 81: 376–
Bi, S., Wang, Y., Guan, J., Sheng, X., and Meng, J., 2014. Three
new Jurassic euharamiyidan species reinforce early
divergence of mammals. Nature, 514: 579–584.
Briggs, D.E.G., 2003. The role of decay and mineralization in the
preservation of soft-bodied fossils. Annual Review of Earth
and Planetary Sciences, 31: 275301.
Cai, C.Y., Thayer, M.K., Engel, M.S., Newton, A.F., and Ortega-
Blanco, J., 2014. Early origin of parental care in Mesozoic
carrion beetles. Proceedings of the National Academy of
Sciences, USA, 111: 14170–14174.
Chang, J.P., and Sun, Y.W., 1997. Aquatic biocoenose of the
Haifanggou Formation during the Middle Jurassic in Beipiao
Basin, Liaoning Province. Journal of Changchun University
of Earth Sciences, 27(3): 241–245.
Chang, S.C., Zhang, H., Hemming, S.R., Mesko, G.T., and Fang,
Y., 2013. 40Ar/39Ar age constraints on the Haifanggou and
Lanqi formations: when did the first flowers bloom. Geological
Society London Special Publications, 378(1): 277–284.
Chang, S.C., Zhang, H.C., Renne, P.R., and Fang, Y., 2009.
High-precision 40Ar/39Ar age constraints on the basal Lanqi
Formation and its implications for the origin of angiosperm
plants. Earth and Planetary Science Letters, 279: 212–221.
Chen, W., Ji, Q., Liu, D., Zhang, Y., Song, B., and Liu, X., 2004.
Isotope gechronology of the fossil-bearing beds in the
Daohugou area, Ningcheng, Inner Mongolia. Geological
Bulletin of China, 23(12): 1165–1169.
Chen, Y., Chen, W., Zhou, X., Li, Z., Liang, H., and Li, Q., 1997.
Liaoxi and Adjacent Mesozoic Volcanic Rocks: Chronology,
Geochemistry and Tectonic Setting. Beijing: The
Seismological Press.
Cheng, X., Wang, X., Jiang, S., and Kellner, A.W.A., 2012. A
new scaphognathid pterosaur from western Liaoning, China.
Historical Biology, 24: 101–111.
Clark, J.M., Xu, X., Eberth, D., Forster, C.A., Malkus, M.,
Hemming, S., and Hernandez, R., 2006. The Mid-Late
Jurassic terrestrial transition: new discoveries form the
Shishugou Formation, China. In: Barrett, P.M., Batten, D.J.,
Evans, S., (eds.), Ninth International Symposium on Mesozoic
Terrestrial Ecosystems and Biota. Manchester, England:
Department of Palaeontology, Natural History Museum of
London, 26–28.
Cui, Y., Storozhenko, Y.S., and Ren, D., 2012. New and little-
known species of Geinitziidae (Insecta: Grylloblattida) from
the Middle Jurassic of China, with notes on taxonomy, habitus
and habitat of these insects. Alcheringa: An Australasian
Journal of Palaeontology, 36(2): 251–261.
Czerkas, S.A., and Ji, Q., 2002. A new rhamphorhynchoid with a
head crest and complex integumentary structures. In: Czerkas,
S.J. (ed.), Feathered dinosaurs and the origin of flight.
Blanding: The Dinosaur Museum,15–41.
Czerkas, S.A., and Yuan, C.X., 2002. An arboreal maniraptoran
from Northeast China In: Czerkas, S.J. (ed.), Feathered
dinosaurs and the origin of flight. Blanding: The Dinosaur
Museum, 63–95.
Dong, L.P., Huang, D.Y., and Wang, Y., 2012. Two Jurassic
salamanders with stomach contents from Inner Mongolia,
China. Chinese Science Bulletin, 57(1): 72–76.
Duan, Y., Zheng, S.L., Hu, D.Y., Zhang, L.J., and Wang, W.L.,
2009. Preliminary report on Middle Jurassic strata and fossils
from Linglongta area of Jianchang, Liaoning. Global Geology,
28(2): 143–147.
Engel, M.S., and Ren, D., 2008. New snakeflies from the
Jiulongshan Formation of Inner Mongolia, China
(Raphidioptera). Journal of the Kansas Entomological
Society, 81: 188–193.
Evans, S.E., and Wang, Y., 2007. A juvenile lizard specimen with
well-preserved skin impressions from the Upper Jurassic/
Lower Cretaceous of Daohugou, Inner Mongolia, China.
Naturwissenschaften, 94: 431–439.
Evans, S.E., and Wang, Y., 2009. A long-limbed lizard from the
Upper Jurassic/Lower Cretaceous of Daohugou, Ningcheng,
Nei Mongol, China. Vertebrata PalAsiatica, 47(1): 21–34.
Evans, S.E., and Wang, Y., 2012. New material of the Early
Cretaceous lizard Yabeinosaurus from China. Cretaceous
Research, 34: 48–60.
Friis, E.M., Pedersen, K.R., and Crane, P.R., 2005. When Earth
started blooming: insights from the fossil record. Current
Opinion of Plant Biology, 8: 5–12.
Gao, K.Q., and Shubin, N., 2012. Late Jurassic salamandroid
from western Liaoning, China. Proceedings of the National
Academy of Science of the America of United States, 109(15):
Gao, K.Q., and Shubin, N.H., 2003. Earliest known crown-group
salamanders. Nature, 422: 424–428.
Gao, T.P., Shih, C.K., Xu, X., Wang, S., and Ren, D., 2012. Mid-
Mesozoic flea-like ectoparasites of feathered or haired
vertebrates. Current Biology, 22: 732–735.
Godefroit, P., Cau, A., Hu, D.Y., Escuillié, F., Wu, W.H., and
Dyke, G., 2013a. A Jurassic avialan dinosaur from China
resolves the early phylogenetic history of birds. Nature, 498:
Godefroit, P., Demuynck, H., Dyke, G., Hu, D.Y., Escuillié, F.,
and Claeys, P., 2013b. Reduced plumage and flight ability of a
new Jurassic paravian theropod from China. Nature
Communications, 4: 1394.
Gu, J.J., Montealegre-Z, F., Robert, D., Engel, M.S., Qiao, G.X.,
and Ren, D., 2012. Wing stridulation in a Jurassic katydid
(Insecta, Orthoptera) produced low-pitched musical calls to
attract females. Proceedings of the National Academy of
Science of the America of United States, 109: 3868–3873.
Gu, J.J., Qiao, G.X., and Ren, D., 2011. A exceptionally-
preserved new species of Barchaboilus (Orthoptera:
Prophalangopsidae) from the Middle Jurassic of Daohugou,
China. Zootaxa, 2909: 64–68.
Guo, X.Q., Han, J.G., and Ji, S.A., 2012. Advances in the study
of vertebrate fossils of the Middle Jurassic Yanliao biota in
western Liaoing Province and adjacent areas. Geological
Bulletin of China, 31(6): 928–935.
He, H.Y., Wang, X.L., Zhou, Z.H., Zhu, R.X., Jin, F., Wang, F.,
Ding, X., and Boven, A., 2004. 40Ar/39Ar dating of ignimbrite
from Inner Mongolia, northeastern China, indicates a post-
Middle Jurassic age for the overlying Daohugou Bed.
Geophyical Research Letters, 31: 1–4.
He, Y., Clark, J., and Xu, X., 2013. A large theropod metatarsal
from the upper part of Jurassic Shishugou Formation in
2240 Vol. 90 No. 6 Dec. 2016
Junggar Basin, Xinjiang, China. Vertebrata PalAsiatica, 51
(1): 29–42.
Hoffstetter, R., 1964. Les Sauria du Jurassique supérieur et
specialement les Gekkota de Baviére et de Mandchourie.
Senckenbergiana Biologie, 45: 281–324.
Hong, Y.C., 1983. Middle Jurassic fossil insects in North China.
Beijing: Geological Publishing House.
Hou, W.J., Yao, Y.Z., Zhang, W.T., and Ren, D., 2012. The
earliest fossil flower bugs (Heteroptera: Cimicomorpha:
Cimicoidea: Vetanthocoridae) from the Middle Jurassic of
Inner Mongolia, China. European Journal of Entomology,
109: 281–288.
Hu, D.Y., Hou, L.H., Zhang, L.J., and Xu, X., 2009. A pre-
Archaeopteryx troodontid from China with long feathers on
the metatarsus. Nature, 461: 640–643.
Hu, Y.M., Meng, J., and Clark, J.M., 2007. A new Late Jurassic
docodont (Mammalia) from northeastern Xinjiang, China.
Vertebrata PalAsiatica, 45: 173–194.
Huang, D.Y., 2014. Trace back the origin of recent insect orders
evidence from the Middle Jurassic Daohugou Biota.
Science Foundation in China, 22(1): 34–42.
Huang, D.Y., Engel, M.S., Cai, C.Y., Wu, H., and Nel, A., 2012.
Diverse transitional giant fleas from the Mesozoic era of
China. Nature, 483: 201–204.
Huang, D.Y., Nel, A., and Petrulevičius, J.F., 2008a. New
evolutionary evidences of Grylloblattida from the Middle
Jurassic of Inner Mongolia, north–east China (Insecta,
Polyneoptera). Zoological Journal of Linnean Society, 152:
Huang, D., Bechly, G., Nel, P., Engel, M.S., and Prokop, J., 2016.
New fossil insect order Permopsocida elucidates major
radiation and evolution of suction feeding in hemimetabolous
insects (Hexapoda: Acercaria). Scientific Reports, 6: 23004.
Huang, D., Nel, A., Cai, C.Y., and Engel, M.S., 2013.
Amphibious flies and paedomorphism in the Jurassic period.
Nature, 495: 94–97.
Huang, D., Nel, A., Zompro, O., and Waller, A., 2008b.
Mantophasmatodea now in the Jurassic. Naturwissenschaften,
95(10): 947–952.
Huang, D.Y., 2015. Yanliao Biota and Yanshan Movement. Acta
Palaeontologica Sinica, 54(4): 501–546.
Huang, D.Y., and Cai, C., (in press). The Arthropoda. In: Huang,
D.Y. (eds), The Daohugou Biota. Shanghai: Shanghai Science
and Technology Press.
Huang, D.Y., and Nel, A., 2009. Oldest webspinners from the
Middle Jurassic of Inner Mongolia, China (Insecta:
Embiodea). Zoological Journal of the Linnean Society, 156:
Huang, D.Y., Selden, P.A., and Dunlop, J.A., 2009. Harvestmen
(Arachnida: Opiliones) from the Middle Jurassic of China.
Naturwissenschaften, 96: 955–962.
Ji, Q., Luo, Z.X., Yuan, C.X., and Tabrum, A.R., 2006. A
swimming mammaliaform from the Middle Jurassic and
ecomorphological diversification of early mammals. Science,
311: 1123–1127.
Ji, Q., and Yuan, C.X., 2002. Discovery of two kinds of
protofeathered pterosaurs in the Mesozoic Daohugou Biota in
the Ningcheng region and its stratigraphic and biologic
significances. Geological Review (China), 48: 221–224.
Jia, J., and Gao, K.Q., 2016. A new basal Salamandroid
(Amphibia, Urodela) from the Late Jurassic of Qinglong,
Hebei Province, China. PLoS ONE, 11(5): e0153834.
Jiang, B.Y., 2006. Non-marine Ferganoconcha (Bivalvia) from
the Middle Jurassic in Daohugou Area, Ningcheng County,
Inner Mongolia, China. Acta Palaeontologica Sinica, 45(2):
Jiang, B.Y., Fursich, F.T., Shang, J.G., Wang, B.X., and Niu, Y.,
2011. Early Cretaceous volcanism and its impact on fossil
preservation in Western Liaoning NE China.
Palaeogeography Palaeoclimatology Palaeoecology, 302:
Jiang, H.E., Ferguson, D.K., Li, C.S., and Cheng, Y.M., 2008.
Fossil coniferous wood from the Middle Jurassic of Liaoning
Province, China. Palaeobotany & Palynology, 150: 37–47.
Jin, F., 1999. Middle and Late Mesozoic acipenseriforms from
northern Hebei and western Liaoning, China. Palaeoworld,
11: 188–280.
Lü, J., and Bo, X., 2011. A new rhamphorhynchid pterosaur
(Pterosauria) from the Middle Jurassic Tiaojishan Formation
of Western Liaoning, China. Acta Geologica Sinica (English
Edition), 85(5): 977–983.
Lü, J., and Fucha, X., 2010. A new pterosaur (Pterosauria) from
Middle Jurassic Tiaojishan Formation of western Liaoning,
China. Global Geology, 13 113–118.
Lü, J., and Hone, D.W.E., 2012. A new Chinese anurognathid
pterosaur and the evolution of pterosaurian tail lengths. Acta
Geologica Sinica (English Edition), 86(6): 1317–1325.
Lü, J., Unwin, D.M., Jin, X., Liu, Y., and Ji, Q., 2010a. Evidence
for modular evolution in a long-tailed pterosaur with a
pterodactyloid skull. Proceedings of the Royal Society B:
Biological Series 277(1680): 383–389.
Lü, J., Unwin, D.M., Zhao, B., Gao, C., and Shen, C., 2012. A
new rhamphorhynchid (Pterosauria: Rhamphorhynchidae)
from the Middle/Upper Jurassic of Qinglong, Hebei Province,
China. Zootaxa, 3158: 1–19.
Lü, J., Xu, L., Chang, H., and Zhang, X., 2011. A new
darwinopterid pterosaur from the Middle Jurassic of western
Liaoning, northeastern China and its ecological implications.
Acta Geologica Sinica (English Edition), 85(3): 507–514.
Lü, J.C., 2009. A new non-pterodactyloid pterosaur from
Qinglong County, Hebei Province of China. Acta Geologica
Sinica (English Edition), 83: 189–199.
Lü, J.C., Fucha, X.H., and Chen, J.M., 2010b. A new
scaphognathine pterosaur from the Middle Jurassic of western
Liaoning, China. Acta Geoscientica Sinica, 31(2): 263–266.
Liang, J.H., Huang, W., and Ren, D., 2012. Garciliblatt bella
gen. et sp. n. –a rare carnivorous cockroach (Insecta, Balttida,
Raphidiomimidae) from the Middle Jurassic sediments of
Daohugou in Inner Mongolia, China. Zootaxa, 3449: 62–68.
Liu, J., Zhao, Y., and Liu, X., 2006a. Age of the Tiaojishan
Formation volcanics in the Chengde Basin, northern Hebei
province. Acta Petrologica Sinica, 22(11): 2617–2630.
Liu, P.J., Huang, J.D., and Ren, D., 2010. Palaeocology of the
Middle Jurassic Yanliao Entomofauna. Acta Zootaxonomica
Sinica, 35(3): 568–584.
Liu, Y.Q., Kuang, H., Jiang, X., Peng, N., Xu, H., and Sun, H.,
2012. Timing of the earliest known feathered dinosaurs and
transitional pterosaurs older than the Jehol Biota.
Palaeogeography Palaeoclimatology Palaeoecology, 323–
325: 1–12.
Liu, Y.X., Liu, Y.Q., and Zhong, H., 2006b. LA-ICPMS zircon
U-Pb dating in the Jurassic Daohugou beds and correlative
strata in Ningcheng of Inner Mongolia. Acta Geologica Sinica
(English Edition), 80: 733–742.
Dec. 2016 Vol. 90 No. 6 2241
Lu, L.W., (1995). Fish fossils. In: D. Ren, L. W. Lu and Z. G.
Guo Jurassic-Cretaceous faunas and stratigraphy of Beijing
and neighboring areas. Beijing: Seismological Press: 121–
Lu, Y., Yao, Y.Z., and Ren, D., 2011. Two new genera and
species of fossil true bugs (Hemiptera: Heteroptera:
Pachymeridiidae) from northeastern China. Zootaxa, 2835:
Luo, Z.X., Gatesy, S.M., Jenkins, F.A., and Amaral, A.A., 2015a.
Mandibular and dental characteristics of Late Triassic
mammaliaform Haramiyavia and their ramifications for basal
mammal evolution. Proceedings of National Academy of
Sciences USA, 112(51): E7101–E7109.
Luo, Z.X., Ji, Q., and Yuan, C.X., 2007. Convergent dental
evolution in pseudotribosphenic and tribosphenic mammals.
Nature, 450: 93–97.
Luo, Z.X., Meng, Q.J., Ji, Q., Liu, D., Zhang, Y.G., and Neander,
A.I., 2015b. Evolutionary development in basal
mammaliaforms as revealed by a docodontan. Science, 347:
Luo, Z.X., 2007. Transformation and diversification in early
mammal evolution. Nature, 450: 1011–1019.
Luo, Z.X., Yuan, C.X., Meng, Q.J., and Ji, Q., 2011. A Jurassic
eutherian mammal and divergence of marsupials and
placentals. Nature, 476: 442–445.
Martínez-Delclòs, X., Briggs, D.E.G., and Peñalver, E., 2004.
Taphonomy of insects in carbonates and amber.
Palaeogeography Palaeoclimatology Palaeoecology, 203:
Martill, D.M., and Etches, S., 2012. A new monofenestratan
pterosaur from the Kimmeridge Clay Formation (Upper
Jurassic, Kimmeridgian) of Dorset, England. Acta
Palaeontologica Polonica, 58(2): 285–294.
Martin, T., Averianov, A.O., and Pfretzschner, H.U., 2010.
Mammals from the Late Jurassic Qigu Formation in the
southern Junggar Basin, Xinjiang, Northwest China.
Palaeodiversity and Palaeoenvironment, 90: 295–319.
Meng, J., Hu, Y.M., Wang, Y.Q., Wang, X.L., and Li, C.K., 2006.
A Mesozoic gliding mammal from northeastern China.
Nature, 444: 889–893.
Meng, Q.J., Ji, Q., Zhang, Y.G., Liu, D., Grossnickle, D.M., and
Luo, Z.X., 2015. An arboreal docodont from the Jurassic and
mammaliaform ecological diversification. Science, 347: 764–
Nel, A., Huang, D.Y., and Lin, Q., 2008. A new genus of
isophlebioid damsel-dragonflies with "calopterygid"-like wing
shape from the Middle Jurassic of China (Odonata:
Isophlebioidea: Campterophlebiidae). European Journal of
Entomology, 105: 783–787.
O'Brien, N.R., Meyer, H.W., Reilly, K., Ross, A.M., and
Maguire, S., 2002. Microbial taphonomic processes in the
fossilization of insects and plants in the late Eocene Florissant
Formation, Colorado. Rocky Mountain Geology, 37(1): 1–11.
O'Connor, J., and Sullivan, C., 2014. Reinterpretation of the
Early Cretaceous maniraptoran (Dinosauria: Theropoda)
Zhongornis haonae as a scansoriopterygid-like non-avian, and
morphological resemblances between scansoriopterygidfs and
basal oviraptorosaurs. Vertebrata Pal Asiatica, 52: 3–30.
Pan, G., 1983. The Jurassic precursors of angiosperms from
Yanliao region of North China and the origin of angiosperms.
Chinese Science Bulletin, 28: 1520.
Pan, Y., Sha, J., Fürsich, F.T., Wang, Y., Zhang, X., and Yao, X.,
2011. Dynamics of the lacustrine fauna from the Early
Cretaceous Yixian Formation, China: implications of volcanic
and climatic factors. Lethaia, 45: 299–314.
Peng, G.Z., Ye, Y., Gao, Y.H., Shu, C.K., and Jiang, S., 2005.
Jurassic dinosaur faunas in Zigong. Chengdu: Sichuan
People's Publishing House.
Peng, N., Liu, Y., Kuang, H., Jiang, X., and Xu, H., 2012.
Stratigraphy and geochronology of vertebrate fossil-bearing
Jurassic strata from Linglongta, Jianchang County, western
Liaoning, northeastern China. Acta Geologica Sinica (English
Edition), 86(6): 1326–1339.
Pott, C., McLoughlin, S., Wu, S.Q., and Friss, E.M., 2011.
Trichomes on the leaves of Anomozamites villosus sp. nov.
(Bennettitales) from the Daohugou beds (Middle Jurassic),
Inner Mongolia, China: Mechanical defence against
herbivorous arthropods. Review of Palaeobotany and
Palynology, 169: 48–60.
Ren, D., 1998. Flower-associated Brachycera flies as fossil
evidence for Jurassic angiosperm origins. Science, 280: 85–
Ren, D., 2003. Study on the Jurassic Fossil Raphidioptera and
Neuroptera from Northeast China. Ph. D., Beijing Forestry
Ren, D., Labandeira, C.C., Santiago-Blay, J.A., Rasnitsyn, A.,
Shih, C., Bashkuev, A., Logan, M.A.V., Hotton, C.L., and
Dilcher, D., 2009. A probable pollination mode before
angiosperms: Eurasian, long-proboscid scorpionflies. Science,
326: 840–847.
Ren, D., Lu, L.W., Guo, Z.G., and Ji, S.A., 1995. Faunas and
Stratigraphy of Jurassic-Cretaceous in Beijing and the
adjacent areas. Beijing: Seismic Press.
Ren, D., Shih, C.K., Gao, T.P., Yao, Y.Z., and Zhao, Y.Y., 2010.
Silent stories-Insect fossil treasures from Dinosaur Era of the
northeastern China. Beijing: Science Press.
Ren, D., and Yin, J.C., 2002. A new Middle Jurassic species of
Epiosmylus (Neuroptera: Osmylidae) from Inner Mongolia,
China. Acta Zootaxonomica Sinica, 27(2): 274–277.
Sanderson, M.J., Thorne, J.L., Wilk, N., and Bremer, K.G., 2004.
Molecular evidence on plant divergence times. American
Journal of Botany, 91: 1656–1665.
Selden, P.A., and Huang, D.Y., 2010. The oldest haplogyne
spider (Araneae: Plectreuridae), from the Middle Jurassic of
China. Naturwissenschaften, 97: 449–459.
Selden, P.A., Huang, D.Y., and Ren, D., 2008. Palpimanoid
spiders from the Jurassic of China. The Journal of
Arachnology, 36: 306–321.
Selden, P.A., Shih, C.K., and Ren, D., 2011. A golden orb-
weaver spider (Araneae: Nephilidae: Nephila) from the
Middle Jurassic of China. Biology Letter, 7: 775–778.
Shen, Y.B., Chen, P.J., and Huang, D.Y., 2003. Age of the fossil
conchostracans from Daohugou of Ningcheng, Inner
Mongolia. Journal of Stratigraphy, 4: 311–313.
Shen, Y.B., and Huang, D.Y., 2008. Extant clam shrimp egg
morphology: taxonomy and comparison with other fossil
branchiopod eggs. Journal of Crustacean Biology, 28(2):
Shih, C.K., Yang, X.G., Labandeira, C.C., and Ren, D., 2011. A
new long-proboscid genus of Pseudopolycentropodidae
(Mecoptera) from the Middle Jurassic of China and its plant-
host specializations. Zookeys, 130: 281–297.
2242 Vol. 90 No. 6 Dec. 2016
Smith, S.A., Beaulieu, J.M., and Donoghue, M.J., 2010. An
uncorrelated relaxed-clock analysis suggests an earlier origin
for flowering plants. Proceedings of the National Academy of
Science of the America of United States, 107(13): 5897–5902.
Sullivan, C., Wang, Y., Hone, D.W.E., Wang, Y., Xu, X., and
Zhang, F., 2014. The vertebrates of the Jurassic Daohugou
Biota of northeastern China. Journal of Vertebrate
Paleontology, 34(2): 243–280.
Sun, C.L., Dilcher, D.L., Wang, H.S., Sun, G., and Ge, Y.H.,
2008. A study of Ginkgo leaves from the Middle Jurassic of
Inner Mongolia, China. International Journal of Plant
Science, 169(8): 1128–1139.
Sun, G., Ji, Q., Dilcher, D.L., Zheng, S.L., and Nixon, K.C.,
2002. Archaefructaceae, a new basal angiosperm family.
Science, 296: 899–904.
Sun, G., Zhang, L., Zhou, C., Duan, Y., and Hu, D., 2009. A
general outline for the Liaoning Paleontological Museum.
Shenyang: Liaoning Paleontological Museum.
Sun, G., Zhang, L., Zhou, C., Duan, Y., and Hu, D., 2011. The
fossils over last 300 My in Liaoning Province. Shanghai:
Shanghai Science & Technology Press.
Tan, J.J., and Ren, D., 2002. Palaeoecology of insect community
from Middle Jurassic Jiulongshan Formation in Ningcheng
County, Inner Mongolia, China. Acta Zootaxonomica Sinica,
27(3): 428–434.
Wang, B., and Zhang, H.C., 2010. Earliest evidence of fishflies
(Megaloptera: Corydalidae): an exquisitely preserved larva
from the Middle Jurassic of China. Journal of Paleontology,
84(4): 774–780.
Wang, B.X., Li, J.F., Fang, Y., and Zhang, H.C., 2008.
Preliminary elemental analysis of fossil insects from the
Middle Jurassic of Daohugou, Inner Mongolia and its
taphonomic implications. Chinese Science Bulletin, 54(5):
Wang, L., Hu, D., Zhang, L., Zheng, S., He, H.Y., Deng, C.,
Wang, X.L., Zhou, Z.H., and Zhu, R.X., 2013. SIMS U-Pb
zircon age of Jurassic sediments in Linglongta, Jianchang,
western Liaoning: Constraint on the age of oldest feathered
dinosaurs. Chinese Science Bulletin (Chinese version), 58(1):
Wang, W.L., Zheng, S.L., Zhang, L.J., Pu, R.G., Zhang, W., Wu,
H.Z., Ju, R.H., Dong, G.Y., and Yuan, H., 1989. Mesozoic
stratigraphy and palaeontology of western Liaoning. Beijing:
Geological Publishing House.
Wang, X.F., Duan, S., Geng, B., Cui, J., and Yang, Y., 2007.
Schmeissneria: a missing link to angiosterms? BMC
Evolutionary Biology, 7(14): 1–13.
Wang, X., 2010a. Flower-related fossils from the Jurassic.
Lecture Notes in Earth Sciences, 121: 91–153.
Wang, X., 2010b. Schmeissneria: an angiosperm from the Early
Jurassic. Journal of Systematics and Evolution, 48(5): 326–
Wang, X., Krings, M., and Taylor, T.N., 2010a. A thalloid
organism with possible lichen affinity from the Jurassic of
northeastern China. Review of Palaeobotany and Palynology,
162: 591–598.
Wang, X., and Wang, S.J., 2010. Xingxueanthus: An enigmatic
Jurassic seed plant and its implications for the origin of
angiospermy. Acta Geologica Sinica (English Edition), 84(1):
Wang, X., Zheng, S.L., and Jin, J.H., 2010b. Structure and
relationships of Problematospermum, an enigmatic seed from
the Jurassic of China. International Journal of Plant Science,
171(4): 447–456.
Wang, X.L., Kellner, A.W.A., Jiang, S., Cheng, X., Meng, X.,
and Rodrigues, T., 2010c. New long-tailed pterosaurs
(Wukongopteridae) from western Liaoning, China. Anais da
Academia Brasileira de Ciências, 82(4): 1045–1062.
Wang, X.L., Kellner, A.W.A., Jiang, S.X., and Meng, X., 2009.
An unusual long-tailed pterosaur with elongated neck from
western Liaoning of China. Anais da Academia Brasileira de
Ciências, 81: 793–812.
Wang, X.L., Wang, Y.Q., Zhang, F.C., Zhang, J.Y., Zhou, Z.H.,
Jin, F., Hu, Y.M., Gu, G., and Zhang, H.C., 2000. Vertebrate
biostratigraphy of the Lower Cretaceous Yixian Formation in
Linghuan, western Liaoning and its neighboring southern Nei
Mongol (Inner Mongolia), China. Vertebrata PalAsiatica, 38
(2): 81–99.
Wang, X.L., Zhou, Z.H., He, H.Y., Jin, F., Wang, Y.Q., Zhang,
J.Y., Wang, Y., Xu, X., and Zhang, F.C., 2005. Stratigraphy
and age of the Daohugou Bed in Ningcheng, Inner Mongolia.
Chinese Science Bulletin (Chinese version), 20: 2369–2376.
Wang, X.L., Zhou, Z.H., Zhang, F.C., and Xu, X., 2002. A nearly
completely articulated rhamphorhynchoid pterosaur with
exceptionally well-preserved wing membranes and “hairs”
from Inner Mongolia, northeast China. Chinese Science
Bulletin, 47(3): 226–230.
Wang, Y., 2000. A new salamander (Amphibia: Caudata) from
the Early Cretaceous Jehol Biota. Vertebrata PalAsiatica, 38
(2): 100–103.
Wang, Y., 2004. A new Mesozoic caudate (Liaoxitriton
daohugouensis sp. nov.) from Inner Mongolia, China. Chinese
Science Bulletin, 49(8): 858–860.
Wang, Y., Dong, L.P., and Evans, S.E., 2010d. Herpetofaunas
from the Jehol associated strata in NE China: evolutionary and
ecological implications. Bulletin of the Chinese Academy of
Sciences, 24(2): 76–79.
Wang, Y., and Evans, S.E., 2006. A new short-bodied salamander
from the Upper Jurassic/Lower Cretaceous of China. Acta
Palaeontologica Polonica, 51: 127–130.
Wang, Y., and Rose, C., 2005. Jeholtriton paradoxus (Amphibia:
Caudata) from the Lower Cretaceous of southeastern Inner
Mongolia, China. Journal of Vertebrate Paleontology, 25:
Wang, Y.J., Labandeira, C.C., Shih, C.K., Ding, Q.L., Wang, C.,
Zhao, Y.Y., and Ren, D., 2012. Jurassic mimicry between a
hangingfly and a ginkgo from China. Proceedings of the
National Academy of Science of the America of United States,
109(50): 20514–20519.
Wang, Y.J., Liu, Z.Q., Wang, X., Shih, C.K., Zhao, Y.Y., Engel,
M.S., and Ren, D., 2010e. Ancient pinnate leaf mimesis
among lacewings. Proceedings of the National Academy of
Science of the America of United States, 107(37): 16212–
Wu, G., Li, C., and Wang, W., 2004. Geochemical features and its
geological implication of Middle Mesozoic Haifanggou
Formation in West Liaoning. Acta Petrol Mineral, 23: 97–105.
Xu, K., Yang, J., Tao, M., Liang, H., Zhao, C., Li, R., Kong, H.,
Li, Y., Wan, C., and Peng, W., 2003. Jurassic system in the
North of China (VII): the stratigraphic region of Northeast
China. Beijing: Petroleum Industry Press.
Xu, X., You, H., Du, K., and F., H., 2011. An Archaeopteryx-like
theropod from China and the origin of Avialae. Nature, 475:
Dec. 2016 Vol. 90 No. 6 2243
Xu, X., and Zhang, F.C., 2005. A new maniraptoran with long
metatarsalian feathers from China. Naturwissenschaften, 92:
Xu, X., Zhao, Q., Norell, M.A., Sullivan, C., Hone, D., Erickson,
G.M., Wang, X.L., Han, F.L., and Guo, Y., 2009. A new
feathered maniraptoran dinosaur fossil that fills a
morphological gap in avian origin. Chinese Science Bulletin
54: 430–435.
Xu, X., Zheng, X., Sullivan, C., Wang, X., Xing, L., Wang, Y.,
Zhang, X., Connor, J.K.O., Zhang, F., and Pan, Y., 2015. A
bizarre Jurassic maniraptoran theropod with preserved
evidence of membranous wings. Nature, 521: 70–73.
Xu, X., Zhou, Z.H., Sullivan, C., and Wang, Y., (2016). The
Yanliao Biota: a trove of exceptionally preserved Middle-Late
Jurassic terrestrial life forms. In: Fraser, N.C., Sues, H.D.,
(eds), Terrestrial Conservation Lagerstätten: Windows into
the Evolution of Life on Land. Edinburgh: Dunedin Academic
Press Ltd, 301–330.
Yabe, H., and Shikama, T., 1938. A new Jurassic Mammalia from
South Manchuria. Proceedings of the Imperial Academy (of
Japan), 14(9): 353–357.
Yang, W., and Li, S.G., 2004. The chronological framework of
the Mesozoic volcanic rocks of Western Liaoning and its
implications for the Mesozoic lithosphere thinning in Eastern
China. Abstracts of Symposium on National Petrology and
Continental Geodynamics. Haikou, China, Petroleum
Committee of Chinese Society for Geololgy, Haikou, China.
Yang, W., and Li, S.G., 2008. Geochronology and geochemistry
of the Mesozoic volcanic rocks in Western Liaoning:
implications for lithospheric thinning of the North China
Craton. Lithos, 102: 88–117.
Yuan, C.X., Ji, Q., Meng, Q.-J., Tabrum, A.R., and Luo, Z.X.,
2013. Earliest evolution of multituberculate mammals
revealed by a new Jurassic fossil. Science, 341(6147): 779–
Zhang, F., Zhou, Z., Xu, X., and Wang, X., 2002. A juvenile
coelurosaurian theropod from China indicates arboreal habits.
Naturwissenschaften, 89: 394–398.
Zhang, F.C., Zhou, Z.H., Xu, X., Wang, X.L., and Sullivan, C.,
2008a. A bizarre Jurassic maniraptoran from China with
elongate ribbon-like feathers. Nature, 455: 1105–1108.
Zhang, H., Wang, M.X., and Liu, X., 2008b. Constraint on the
upper boundary age of the Tiaojishan Formation volcanic
rocks in western Liaoning- northern Hebei by LA-ICP-MS.
Chinese Science Bulletin, 53(22): 3574–3584.
Zhang, J.F., 2002. Discovery of Daohugou Biota (Pre-Jehol
Biota) with a discussion on its geological age. Journal of
Stratigraphy, 26(3): 173–177.
Zhang, J.F., and Kluge, N.J., 2007. Jurassic larvae of mayflies
(Ephemeroptera) from the Daohugou Foramtion in Inner
Mongolia, China. Oriental Insects, 41: 351–366.
Zhang, L.J., and Wang, L.X., 2004. Mesozoic salamander fossil-
bearing strata in Reshuitang area near Lingyuan, western
Liaoning. Geology and Resources (China), 13(4): 202–206
Zhang, X.W., Ren, D., Pang, H., and Shih, C.K., 2008c. A water-
skiiing chresmodid from the Middle Jurassic in Daohugou,
Inner Mongolia, China (Polyneoptera: Ornthopterida).
Zootaxa, 1762: 53–62.
Zheng, S.L., 2013. Yanliao Flora. Shenyang.
Zheng, X., Bi, S., Wang, X., and Meng, J., 2013. A new arboreal
haramiyid shows the diversity of crown mammals in the
Jurassic period. Nature, 500: 199–202.
Zheng, X.T., You, H.L., Xu, X., and Dong, Z.M., 2009. A
heterodontosaurid dinosaur from the Early Cretaceous of
China with filamentous integumentary structures. Nature, 458:
Zhou, C.F., Wu, S., Martin, T., and Luo, Z.X., 2013. A Jurassic
mammaliaform and the earliest mammalian evolutionary
adaptations. Nature, 500: 163–167.
Zhou, M.Z., Cheng, Z.W., and Wang, Y.Q., 1991. A mammalian
lower jaw from the Jurassic of Lingyuan, Liaoning. Vertebrata
PalAsiatica, 29(3): 165–175.
Zhou, Z.H., Jin, F., and Wang, Y., 2010. Vertebrate assemblages
from the Middle-Late Jurassic Yanliao Biota in northeast
China. Earth Science Frontiers, 17: 252–254.
About the first author
XU Xing, Male; born in 1969 in Xinyuan, Xinjiang. He studies
dinosaurian taxonomy and systematics, focusing on bird
origins and related issues, and is also involved in studies
of other Mesozoic tetrapods.
... Holotype of Novaboilus ovatus sp. nov., CNU-ORT-NN2006070PC.Jiulongshan Formation, Bathonian-Callovian boundary interval(Xu et al. 2016), Middle Jurassic. ...
Prophalangopsidae was a diverse family during the Late Mesozoic, but the variation and sexual dimorphism in their forewing morphologies are rarely discussed. Based on 43 specimens – of both sexes – from eight species, an investigation into wing venation variation among/within species of Prophalangopsidae was performed using geometric morphometrics and morphological comparisons. The results indicate that wing characters are reliable for taxonomy in fossil Prophalangopsidae and that variation in wing shape and venation is common within species. The structures of the forewings are analogous between sexes within species, and it is possible to pair males and females for a fossil species. Due to the potential existence of synonyms arising from the lack of knowledge on wing venation variation within species and sexes, the species richness of fossil prophalangopsids is probably over-estimated. The role of wing venation characters in systematics and phylogenetic analysis needs to be further analysed. In addition, two new species of Prophalangopsidae from the Middle Jurassic are described.
... The Daohugou beds, yielding abundant and diverse fossil plants and animals, are now considered among the most important insect Lagerstätten (Rasnitsyn et al., 2006). The age of the Daohugou beds is somewhat controversial (e.g., Zhang, 2002;Shen et al., 2003;Rasnitsyn and Zhang, 2004;Wang et al., 2015;Chen and Wang, 2016;Xu et al., 2016). Recent radiometric dating results (Liu et al., 2006(Liu et al., , 2012Wang et al., 2013) and paleontological studies (Wang et al., 2018, and references therein), however, suggest that the Daohugou beds are of latest Middle Jurassic-earliest Late Jurassic age (Callovian-Oxfordian). ...
The extinct family Hylicellidae, as the ancestral group of modern cicadomorphans, had a wide distribution and a very high species-level biodiversity from the Triassic to Early Cretaceous. We herein report 11 new hylicellid specimens from the Jurassic Daohugou beds of Inner Mongolia, NE China, and execute geometric morphometric analysis (GMA) to elucidate their systematic position. Our GMA and subsequent morphometric statistics indicate that 10 of our new specimens can be compared to the holotype of Cycloscytina gobiensis , and one is new to science. Cycloscytina incompleta new species is erected based on this specimen, with the following discriminatory tegminal traits: C3 almost as long as and slightly narrower than C2, and the forking position of stem M distinctly migrates towards wing apex and much apicad of the stem CuA bifurcating. Additionally, Cycloscytina plachutai is herein transferred to the procercopid genus Procercopina , resulting in P . plachutai new combination. To date, just a few body structures of Hylicellidae have been revealed, and the new whole-bodied hylicellids reported herein provide some novel insights on the evolution of basal Clypeata. This study also emphasizes the use of morphometric analysis in the systematics of wing-bearing insects such as hylicellids. UUID: .
... Locality and Horizon-The holotype was recovered from Mutoudeng, Qinglong County in Hebei Province, China, from the Middle-Upper Jurassic Tiaojishan Formation (Xu et al., 2016). ...
... The Yanliao Biota was proposed to include all fossils from the Middle Jurassic Haifanggou Formation and the Upper Jurassic Tiaojishan Formation [24]. Although the palaeoenvironment and climate of this biota are poorly known, the majority of fossils are complete and articulated in fine-grained lacustrine deposits [25,26]. These two specimens both belong to the Upper Jurassic Tiaojishan Formation of the Yanliao Biota, and both pterosaur skeletons are nearly complete and articulated [27,28], indicating limited transportation of the carcasses. ...
Knowledge about the pterosaur diet and digestive system is limited, and there is little direct evidence in the fossil record. Here, we report two specimens of the wukongopterid Kunpengopterus sinensis , a juvenile and an adult, from the Late Jurassic Yanliao Biota of China with associated bromalites. Both of these concentrations are identified as emetolites, fossilized gastric pellets. These pellets contain scales of an unnamed palaeonisciform fish, confirming the pterosaur was a piscivore. It probably vomited the pellets, indicating the presence of two-part stomachs and efficient antiperistalsis in both juveniles and adults. Comparing the ganoid scales found in the pellets with those of complete fishes, it was possible to determine that the prey of the smaller pellet is an average-sized individual, while the prey of the larger pellet represents a large specimen. Kunpengopterus sinensis might have preyed on the same fish during ontogeny, with adults being able to feed on larger individuals. This article is part of the theme issue ‘The impact of Chinese palaeontology on evolutionary research’.
... Ji and Yuan 2002;Zhang 2002;Sullivan et al. 2014), since abundant and diverse fossils were reported from the latest Middle Jurassic Daohugou outcrops in Ningcheng, Inner Mongolia. Yanliao Biota was divided into the Bathonian-Callovian Daohugou phase and the Oxfordian Linglongta phase in Xu et al. (2016). Up to now, 79 cicadomorphan species have been documented in this biota, but only one fulgoromorphan fulgoridiid has been reported (Appendix A). ...
True hoppers, consisting of Fulgoromorpha and Cicadomorpha, are plant feeders with very high species-level diversity. A large amount of true hopper fossils have been reported from eastern Asia, especially from the Middle to Late Jurassic Yanliao Biota, the Early Cretaceous Jehol Biota and mid-Cretaceous Kachin amber in the last two decades. Herein, true hoppers from the Jurassic and Cretaceous of eastern Asia are reviewed, and combining palaeontological data from other regions of the world and recent advances of molecular studies, the evolutionary history of true hoppers in the mid-late Mesozoic is discussed. Permocicada beipiaoensis Wang, 1987 and Archijassus plurinervis Zhang, 1985 are here excluded from Prosboloidea and Archijassidae respectively. To the end of 2020, a total of 203 species with definite systematic position have been documented in the Jurassic and Cretaceous of eastern Asia (China, Myanmar, Siberia, Mongolia, Japan and Korea), and were attributed to 116 genera in 22 families and 7 superfamilies. Available fossil data suggest that true hopper components strongly changed in the Cretaceous: primitive groups reduced and went extinct successively, and the origin and/or early diversification of most lineages (family or subfamily level) occurred, likely owing to the displacement of host-plants in the angiosperm floristic revolution.
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The question why non-avian dinosaurs went extinct 66 million years ago (Ma) remains unresolved because of the coarseness of the fossil record. A sudden extinction caused by an asteroid is the most accepted hypothesis but it is debated whether dinosaurs were in decline or not before the impact. We analyse the speciation-extinction dynamics for six key dinosaur families, and find a decline across dinosaurs, where diversification shifted to a declining-diversity pattern ~76 Ma. We investigate the influence of ecological and physical factors, and find that the decline of dinosaurs was likely driven by global climate cooling and herbivorous diversity drop. The latter is likely due to hadrosaurs outcompeting other herbivores. We also estimate that extinction risk is related to species age during the decline, suggesting a lack of evolutionary novelty or adaptation to changing environments. These results support an environmentally driven decline of non-avian dinosaurs well before the asteroid impact.
Well-sampled dinosaur communities from the Jurassic through the early Late Cretaceous show greater taxonomic diversity among larger (>50 kg) theropod taxa than communities of the Campano-Maastrichtian, particularly to those of eastern/central Asia and Laramidia. The large carnivore guilds in Asiamerican assemblages are monopolized by tyrannosaurids, with adult medium-sized (50–500 kg) predators rare or absent. In contrast, various clades of theropods are found to occupy these body sizes in earlier faunas, including early tyrannosauroids. Assemblages with “missing middle-sized” predators are not found to have correspondingly sparser diversity of potential prey species recorded in these same faunas. The “missing middle-sized” niches in the theropod guilds of Late Cretaceous Laramidia and Asia may have been assimilated by juvenile and subadults of tyrannosaurid species, functionally distinct from their adult ecomorphologies. It is speculated that if tyrannosaurids assimilated the niches previously occupied by mid-sized theropod predators, we would expect the evolution of distinct transitions in morphology and possibly the delay of the achievement of somatic maturity in species of this taxon.
The late Early Jurassic (Toarcian) terrestrial succession exposed in the Tianshifu Basin, eastern Liaoning Province, northeastern China, was examined for palynomorphs. This sequence is part of the upper Changliangzi Formation and is mainly composed of shale and siltstone intercalated with several sandstone layers. The palynoflora in the upper part of the Changliangzi Formation is dominated by pteridophyte spores (average of 64.01%) and gymnosperm pollen grains (average of 34.99%), while the bryophyte spores are less than 1%. The dominant component of this palynoflora alternates between pteridophyte spores and gymnosperm pollen. The vegetation reconstructed from the palynoflora mainly comprises lowland ferns and cycads/ginkgophytes accompanied by upland conifers in low amounts, which is similar to the macroflora components of the Changliangzi Formation. The composition and evolution of this palynoflora indicates that eastern Liaoning was dominated by hot and subhumid climatic conditions with several warm and moderately humid intervals during the late Early Jurassic (Toarcian). Compared to the lower part of the Changliangzi Formation with coal-bearing strata, which is characterised by a rather moist climate, the upper part of this formation might represent a drier climate, although the inferred difference in temperature is small. Although there are several records of warming and drought in some inland regions in northern China during that time, which might be associated with the famous global geological event called the Toarcian oceanic anoxic event, most areas in northern China were dominated by a hot/warm and moist climate, especially in the eastern coastal region. The warm and wet flows from the Proto-Pacific Ocean may be responsible for this hot and humid climate.
Checklists of all described organisms from the Daohugou biota, and insects from the Haifanggou Formation at Haifeng Village (Beipiao City, Liaoning Province) and the ‘Jiulongshan Formation’ at Zhouyingzi Village (Luanping County, Chengde City, Hebei Province), are provided. Fossil insects from the Daohugou biota are summarized, including a total of 760 valid species reported in 396 research papers from 2001 to April, 2021. The heyday of exploration of Daohugou insects has been lasted for a decade from 2006 to 2016 according to the number of published papers.
Pterosaurs, which lived during the Mesozoic, were the first known vertebrates to evolve powered flight.¹,² Arboreal locomotion has been proposed for some taxa,³,⁴ and even considered to have played a role in the origin of pterosaur flight.⁵,⁶ Even so, there is still need for comprehensive quantitative ecomorphological analyses.³,⁴ Furthermore, skeletal adaptations correlated to specialized lifestyles are often difficult to recognize and interpret in fossils. Here we report on a new darwinopteran pterosaur that inhabited a unique forest ecosystem from the Jurassic of China. The new species exhibits the oldest record of palmar (or true) opposition of the pollex, which is unprecedented for pterosaurs and represents a sophisticated adaptation related to arboreal locomotion. Principal-coordinate analyses suggest an arboreal lifestyle for the new species but not for other closely related species from the same locality, implying a possible case of ecological niche partitioning. The discovery adds to the known array of pterosaur adaptations and the history of arborealism in vertebrates. It also adds to the impressive early bloom of arboreal communities in the Jurassic of China, shedding light on the history of forest environments.
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Abstract A new genus and species of docodontid (Docodonta, Mammalia), Acuodulodon sunae, represented by a partial left lower jaw and dentition, is described. It is from the upper part of the Shishugou Formation in the Wucaiwan area of the Junggar Basin in northeastern Xinjiang, China, with an estimated age of 159~161 Ma (Oxfordian, early Late Jurassic). The new mammal is typical of docodonts in having a cusp b in front of cusp a, a cusp c distolingual to cusp a and a cusp g mesiolingual to cusp a on lower molariforms. Differing from other docodonts, it has no cusp e or crest b–e developed on lower molariforms. Unique among docodonts, cusps a and c of the new animal maintained their sharpness while cusp g and crest b–g wore away fast, indicating that both shearing and crushing/grinding occurred in the chewing cycle and probably last for most of the life span of the animal. Phylogenetic analysis of a data matrix with 24 lower molariform characters across 15 taxa recovers a monophyletic Docodonta, which has distinct diagnostic characters in lower molariforms. Within docodonts, Docodon and Borealestes are successively basal to other docodonts; Acuodulodon and Itatodon + (Simpsonodon, Castorocauda + (Tegotherium + Sibirotherium)) form a monophyletic clade. Tegotheriid genera are nested within Docodontidae, but a monophyletic tegotheriid clade composed of Tegotherium, Sibirotherium, Itatodon, and Tashkumyrodon is not recovered. The dentary of Acuodulodon is typical of docodonts. It has a shallow postdentary trough and a wide and sharp medial ridge, both of which do not extend onto the medial side of the condylar peduncle, indicating looser contact between postdentary bones and the dentary than in morganucodontids, a more derived condition in the evolution toward the definitive mammalian middle ear. Key words Xinjiang, Late Jurassic, Docodonta, Acuodulodon, phylogeny
A new genus with a new species (Jurachresmoda gaskelli gen. et sp. nov.) of Chresmodidae (Insecta: Polyneoptera: Orthopterida) is described and illustrated. They were collected from the Middle Jurassic in Daohugou, Jiulongshan Formation, Inner Mongolia (Nei Mongol Autonomous Region), China. J. gaskelli have fringing hairs on tarsi and a part of tibiae of mid legs. These hairs and leg structures were probably associated with its water-skiing locomotion on freshwater. A high number of tarsomeres, e. g. ultra-articulated tarsi as reported in other species of Chresmodidae, are also found for this new species. But, these true tarsi are mixed with some wrinkles resulted from dehydration and compression during fossilization process. Nymphs of the new species are also described. This is the first time that long and segmented cerci of the nymph are described in Chresmodidae.
Northeastern China contains widely distributed Jurassic terrestrial strata that have yielded many spectacular mammal and pterosaur fossils, in addition to feathered dinosaur fossils and more recent discoveries from Jianchang, particularly from western Liaoning. However, the fossil‐bearing stratigraphic succession, regional correlation, and age estimates of the fossils found in Jianchang County and nearby areas have been contentious. Here, we report on the vertebrate fossil‐bearing Jurassic stratigraphy from Linglongta, Jianchang County, western Liaoning, including a SHRIMP U‐Pb zircon date unambiguously associated with the fossil horizons. The primary goal was to determine the vertebrate fossil‐bearing succession. A further aim was to provide age estimations for the fossil‐bearing horizon as well as the earliest appearance of feathered dinosaurs, the eutherian–placental clade, and transitional pterosaurs. Field investigations showed that the vertebrate fossil‐bearing stratigraphic succession in Jianchang County mainly consists of basal andesites overlain by rhythmic tuffs and tuffaceous lacustrine sediments, with the upper intermediate or acidic lavas interbedded with laminated more or less tuffaceous lacustrine deposits. This sequence correlates well with the Middle Jurassic Lanqi/Tiaojishan Formation in northeastern China. Detailed and accurate field observations showed that the well‐preserved vertebrate fossils were buried in either the middle or the upper fine‐grained laminated lacustrine deposits. Previous and current SHRIMP U‐Pb zircon dates provide an age estimation of 161–159 Myr for the fossil‐bearing horizon and vertebrates. This indicates that the earliest appearance of feathered dinosaurs here was more than 159 Myr ago and unquestionably older than Archaeopteryx from Germany, making these the earliest known feathered dinosaurs in the world. Furthermore, the eutherian–placental clade and the known transitional pterosaurs first emerged no later than 161 Myr. The vertebrate assemblage unearthed recently from Linglongta and neighboring areas in Jianchang County belongs to the Daohugou Biota. In addition to feathered dinosaurs, this biota was characterized by mammals, primitive pterosaurs, insects, and plants and was present in Inner Mongolia, western Liaoning, and northern Hebei in northeastern China during the Middle–Late Jurassic.
FIGURE 5. Photographs of J. gaskelli n. g. n. sp., Paratypes, male, nymph: A. CNU-CH-DHG 2007011, scale 10 mm; B. CNU-CH-DHG 2007012, scale 10 mm.