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
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was superseded by “Yanliao Fauna” after it became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 to ‘Yabeinosaurus’ 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: firstname.lastname@example.org
© 2016 Geological Society of China
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ACTA GEOLOGICA SINICA (English Edition)
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
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ACTA GEOLOGICA SINICA (English Edition)
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
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.
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ACTA GEOLOGICA SINICA (English Edition)
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
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ACTA GEOLOGICA SINICA (English Edition)
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
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
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ACTA GEOLOGICA SINICA (English Edition)
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.
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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.
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ACTA GEOLOGICA SINICA (English Edition)
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.
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ACTA GEOLOGICA SINICA (English Edition)
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
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ACTA GEOLOGICA SINICA (English Edition)
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-
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
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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.