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Short note on a non-pterodactyloid pterosaur from Upper Jurassic deposits of Inner Mongolia, China

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  • Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences

Abstract and Figures

Daohugou is an important locality of the Jurassic Yanliao Biota, where only two pterosaurs have been described so far (Jeholopterus and Pterorhynchus). Here we report a new genus and species, Daohugoupterus delicatus gen. et sp. nov. (IVPP V12537), from this region, consisting of a partial skeleton with soft tissue. The skull is laterally compressed, differing from Jeholopterus. The antorbital fenestra is larger than in Pterorhynchus. The upper temporal fenestra is unusually small. The short cervical vertebrae bearing cervical ribs indicate that it is a non-pterodactyloid flying reptile. The sternal plate is triangular, being much wider than long. The deltopectoral crest of humerus is positioned proximally and does not extend further down the shaft, a typical feature of basal pterosaurs. Daohugoupterus also differs from the wukongopterids and scaphognathids from the Tiaojishan Formation at Linglongta, regarded to be about the same age as the Daohugou Bed. The new specimen increases the Jurassic non-pterodactyloid pterosaur diversity of the Yanliao Biota and is the smallest pterosaur from Daohugou area so far.
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Short note on a non-pterodactyloid pterosaur from
Upper Jurassic deposits of Inner Mongolia, China
Xin Chengab, Xiaolin Wanga, Shunxing Jiangab & Alexander W.A. Kellnerc
a Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of
Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of
Sciences, P.O. Box 643, Beijing 100044, P.R. China
b University of Chinese Academy of Sciences, Beijing 100049, P.R. China
c Laboratory of Systematics and Taphonomy of Fossil Vertebrates, Department of Geology
and Paleontology, Museu Nacional/UFRJ, Quinta da Boa Vista s/n, São Cristóvão, CEP
20940-040 Rio de Janeiro, RJ, Brazil
Published online: 17 Nov 2014.
To cite this article: Xin Cheng, Xiaolin Wang, Shunxing Jiang & Alexander W.A. Kellner (2014): Short note on a non-
pterodactyloid pterosaur from Upper Jurassic deposits of Inner Mongolia, China, Historical Biology: An International Journal
of Paleobiology, DOI: 10.1080/08912963.2014.974038
To link to this article: http://dx.doi.org/10.1080/08912963.2014.974038
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Short note on a non-pterodactyloid pterosaur from Upper Jurassic deposits of
Inner Mongolia, China
Xin Cheng
a,b
, Xiaolin Wang
a
*, Shunxing Jiang
a,b
and Alexander W.A. Kellner
c
a
Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and
Paleoanthropology, Chinese Academy of Sciences, P.O. Box 643, Beijing 100044, P.R. China;
b
University of Chinese Academy of
Sciences, Beijing 100049, P.R. China;
c
Laboratory of Systematics and Taphonomy of Fossil Vertebrates, Department of Geology and
Paleontology, Museu Nacional/UFRJ, Quinta da Boa Vista s/n, Sa
˜o Cristo
´va
˜o, CEP 20940-040 Rio de Janeiro, RJ, Brazil
(Received 15 August 2014; accepted 4 October 2014)
Daohugou is an important locality of the Jurassic Yanliao Biota, where only two pterosaurs have been described so far
(Jeholopterus and Pterorhynchus). Here we report a new genus and species, Daohugoupterus delicatus gen. et sp. nov.
(IVPP V12537), from this region, consisting of a partial skeleton with soft tissue. The skull is laterally compressed, differing
from Jeholopterus. The antorbital fenestra is larger than in Pterorhynchus. The upper temporal fenestra is unusually small.
The short cervical vertebrae bearing cervical ribs indicate that it is a non-pterodactyloid flying reptile. The sternal plate is
triangular, being much wider than long. The deltopectoral crest of humerus is positioned proximally and does not extend
further down the shaft, a typical feature of basal pterosaurs. Daohugoupterus also differs from the wukongopterids and
scaphognathids from the Tiaojishan Formation at Linglongta, regarded to be about the same age as the Daohugou Bed. The
new specimen increases the Jurassic non-pterodactyloid pterosaur diversity of the Yanliao Biota and is the smallest pterosaur
from Daohugou area so far.
Keywords: Pterosauria; non-pterodactyloid pterosaur; Daohugoupterus; Late Jurassic; China
Introduction
The research of Chinese pterosaur materials has made
great progress in the study of those extinct flying reptiles
from the beginning of twenty-first century. Many new taxa
were discovered, mostly coming from the Yixian and
Jiufotang formations of western Liaoning (e.g. Jiang and
Wang 2011; Wang et al. 2012), which belongs to the Early
Cretaceous Jehol Biota (Zhou and Wang 2010). In the last
5 years, more discoveries of flying reptiles were made in
an older deposit that belongs to the Yanliao Biota (Zhou
et al. 2010), which is almost distributed in the same area as
the layers that comprise the Jehol Biota.
Two of the most important fossil-bearing horizons of the
Yanliao Biota are found in the regions of Daohugou at the
Ningcheng County, Inner Mongolia, and Linglongta at the
Jianchang County, Liaoning. Several pterosaur specimens
have been recovered from these deposits, which so far
consists of the principal vertebrate component of this biota
(Cheng 2013;Wangetal.2014). There is still an ongoing
debate about the correct stratigraphic placement and
consequentage of those deposits. Some researchers advocate
that they represent the Daohugou Bed (or Formation), while
others regard them as part of the Tiaojishan Formation,
which has been assigned to the Middle Jurassic (Chen et al.
2004;Jietal.2005; Liu et al. 2006,2012;Lu
¨et al. 2010,
2011;Lu
¨and Bo 2011;Wangetal.2013; Sullivan et al.
2014), Late Jurassic (Zhang 2002;Heetal.2004; Zhang et al.
2008;Zhouetal.2010) or Late Jurassic– Early Cretaceous
(Wang et al. 2005,2009,2010; Cheng et al. 2012). The last
U –Pb zircon age of Linglongta has defined it as Late Jurassic
(Cheng 2013;Wangetal.2014).
The most important pterosaur materials of what can be
regarded as the Linglongta Pterosaur Fauna are undoubtedly
the wukongopterids. The members of this pterosaur group
combine primitive non-pterodactyloid features with some
derived characteristics thought previously to be restricted to
the pterodactyloids (Wang et al. 2009,2010;Lu
¨et al. 2011).
Other finds include the anurognathid Jeholopterus ning-
chengensis from the Daohugou area (Wang et al. 2002;
Kellner et al. 2010)andthescaphognathidsJianchangnathus
robu stus and Jianchangopterus zhaoianus (Lu
¨and Bo 2011;
Cheng et al. 2012; Zhou 2014). The other described taxa
from this region are Fenghuangopterus lii (Linglongta, Lu
¨
et al. 2010) and the poorly known Pterorhynchus wellnhoferi
(Daohugou, Czerkas and Ji 2002).
Here we report another pterosaur from Daohugou,
Daohugoupterus delicatus gen. et sp. nov., a new addition
to the Linglongta Pterosaur Fauna.
Systematic paleontology
Pterosauria Kaup, 1834
Daohugoupterus gen. nov.
q2014 Taylor & Francis
*Corresponding author. Email: wangxiaolin@ivpp.ac.cn
Historical Biology, 2014
http://dx.doi.org/10.1080/08912963.2014.974038
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Etymology:Daohugoupterus, from Daohugou Village,
where the specimen was collected, and pteros, from the
Greek meaning ‘wing’.
Type species: Daohugoupterus delicatus gen. et sp. nov.
Diagnosis: The same as for the species.
Daohugoupterus delicatus gen. et sp. nov.
Etymology:Delicatus, from the Latin, meaning ‘small’, in
allusion to the comparatively small and delicate body type
of this species.
Holotype: Partial skeleton with almost a complete skull
housed in the Institute of Vertebrate Paleontology and
Paleoanthropology (Chinese Academy of Sciences),
Beijing, China, under the number IVPP V12537 (Figure 1).
Locality and horizon: Daohugou, Ningcheng County, Inner
Mongolia, China. Daohugou Bed (Formation) or Tiaojishan
Formation, Late Jurassic (Cheng 2013; Wang et al. 2014).
Diagnosis: Non-pterodactyloid pterosaur with the follow-
ing (potential) autapomorphies: nasal with posterior
processes extending between the frontals, and sternal
plate with the breadth nearly 2.5 times of the length.
It can further be distinguished from other non-
pterodactyloid pterosaurs by the following combination
of characters: reduced upper temporal fenestra (smaller
than in Wukongopteridae and Scaphognathidae); slit-like
lower temporal fenestra; quadrangular deltopectoral crest
of the humerus more developed than that in the
Campylognathoididae; and elongated and straight pteroid.
Description
The holotype (IVPP V12537) of Daohugoupterus
delicatus gen. et sp. nov. is preserved in a slab and a
counterslab of grey shale (Figures 1 and 2). It shows
several conchostracans, similar to the holotype of
Jeholopterus from the same locality, indicating fresh
water, likely a lacustrine environment (Wang et al. 2002).
The new specimen is composed of a partial skeleton with
the skull and the lower jaw lacking the anterior tip. The
cervical and dorsal series are well represented but sacral
and caudal elements are missing. Parts of the forelimb and
the pectoral girdle are also exposed.
The skull of Daohugoupterus delicatus is nearly
complete, missing the rostrum anterior to the middle of the
antorbital fenestra. It was preserved dorsoventrally and
part of the right roof taken to the counterslab, which
resulted in the exposition of some ventral elements of the
Figure 1. (Colour online) Daohugoupterus delicatus gen. et sp. nov. holotype (IVPP V12537), preserved in a slab of grey shale, with
soft tissue near the neck, and many conchostracans around the skeleton. (a) Main slab. (b) Line drawing of the main slab, showing the
short cervical vertebrae bearing thin cervical ribs, the triangular sternal plate, and the deltopectoral crest of humerus positioned
proximally and not extending further down the shaft. Scale bar: 20 mm. car, carpus; con, conchostracan; cor, coracoid; cri, cervical rib; cv,
cervical vertebra; dv, dorsal vertebra; hu, humerus; l, left; ptd, pteroid; r, right; ri, rib; sca, scapula; st, sternum.
2X. Cheng et al.
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skull. Most of the cranial elements described here are from
the dorsal side on the main slab.
The bones that form the left orbit are preserved
completely, showing that it was a large rounded cranial
opening. Only the posterior part of the left antorbital
fenestra is preserved, with the bone bar that forms the
posterior margin inclined forward. Despite being incom-
plete, this opening is comparatively larger than in
Pterorhynchus (Czerkas and Ji 2002). The upper temporal
opening is rounded and very small compared with other
non-pterodactyloid taxa such as the wukongopterids (e.g.
Wang et al. 2009,2010). The lower temporal opening is
slit-like differing from most non-pterodactyloid ptero-
saurs, similar only to the condition reported for
Campylognathoides (Padian 2008a).
The posterior parts of the premaxilla extend between
almost half the length of the frontals. In the midline, the
dorsal margin of the premaxilla has a low ridge, which is
not developed into a premaxillary crest. There is no
evidence of an extended cranial crest such as the one
apparently present in Pterorhynchus (Czerkas and Ji
2002). The nasal is an elongated and flat bone positioned
laterally to the premaxilla and extends between the
frontals, differing from all other pterosaurs, including
Jianchangnathus robustus (Cheng et al. 2012). The
frontals are triangular and form the largest element of
the skull roof. The sutures with the parietal are straight.
The parietals are also flat, form the anterior margin of the
upper temporal opening, and lack any evidence of cranial
crest.
There is a plate-like bone positioned dorsal to the orbit
and anterior to the frontal that is being interpreted as the
supraorbital. Posteriorly, this bone contacts the frontal,
and anteriorly covers the dorsal margin of the lacrimal.
The lacrimal is partially overlain by the nasal and the
supraorbital dorsally, and by the jugal ventrally. The jugal
bar of this bone separates the orbit from the antorbital
fenestra and is comparatively broad. No lacrimal foramen
as the one present in the wukongopterid Kunpengopterus
sinensis is observed (Wang et al. 2010).
The jugal bears four processes, with the lacrimal
process sub-vertical. The lacrimal and postorbital
processes form the ventral margin of the orbit. The
quadrate is a broad element and contacts medially the
pterygoid.
Some palatal elements can be partially observed
through the orbits. As a result of the splitting of the slabs,
both pterygoids are exposed in different planes. The left
one is more complete and shows the dorsal surface, while
the right one lacks the dorsal surface revealing the internal
parts of this element. It forms the posterior, medial and
lateral margins of the postpalatinal fenestra, the latter
having an oval shape.
Not much of the lower jaw is observable, because it is
covered by the skull. The preserved posterior end of the
dentary is comparatively deep. Only one tooth can be
Figure 2. (Colour online) Detail of the skull of Daohugoupterus delicatus gen. et sp. nov. (a) Close up of the skull in the main slab. (b)
Line drawing of the skull, showing the large and rounded orbit, the unusually small upper temporal fenestra and the flat skull roof with no
cranial crest. Scale bar: 10 mm. anf, antorbital fenestra; art, articular; bpt, basipterygoid; bs, basisphenoid; co, cotyle; d, dentary; ec,
ectopterygoid; ex, exoccipital; f, frontal; j, jugal; l, left; la, lacrimal; ltf, lower temporal fenestra; n, nasal; op, opisthotic; or, orbit; p,
parietal; pm, premaxilla; po, postorbital; pplf, postpalatinal fenestra; pty, pterygoid; q, quadrate; r, right; soc, supraoccipitale; sor,
supraorbital; sq, squamosal; te, tooth; utf, upper temporal fenestra.
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recognised, but is not exposed sufficiently to provide any
detailed anatomical information.
Atlas and axis are very short, and no suture is visible
separating them. Another seven cervical vertebrae can be
recognised. All are short, rather robust and some bear thin
cervical ribs. As far as observation allows they are
procoelous. The pre- and postzygapophyses are strongly
build. Cervical ribs can be observed in the sixth and eighth
element.
The first dorsal vertebra is preserved in the counterslab
and covered by the sternum plate. It is much smaller than
the last cervical.
The scapula and coracoid are fused, being U-shaped.
The coracoid is shorter than the scapula and shows a well-
developed biceps tubercle.
The sternum has a developed triangular sternal plate
that is much wider than long, more than in any other non-
pterodactyloid reported so far. The width is more than
twice the length. The lateral margins form posteriorly an
obtuse angle of 1208.
Most forelimb bones are lost, and only the left
humerus, a proximal carpal and a pteroid were preserved.
The humerus is exposed lateroventrally with the caput
humeri partially covered by the sternum. No pneumatic
opening could be identified. The shaft is slightly bowed.
The deltopectoral crest is quadrangular, similar to the
condition observed in Eudimorphodon and Campylog-
nathoides, but differs from those taxa in being longer and
more extended ventrally and by not extending further
down the humerus shaft (e.g. Wellnhofer 1978). The
deltopectoral crest is also not hatched-shaped as observed
in Rhamphorhynchus (Kellner 2003) and does not display
the rounded end observed in Dorygnathus (Padian 2008b).
One fused proximal carpal series shows the pteroid
articulated and is displaced near the cervical vertebrae.
The pteroid has a relatively elongated and thin shaft that is
rather straight (Table 1).
Close to the cervical vertebrae and right coracoid,
portions of soft tissue are preserved as a dark matter
(Figure 3). Near the neck, some fibres are observed and
their general irregular aspect is consistent with the
pycnofibres that form the integumental covering in
pterosaurs (Kellner et al. 2010). In some areas, there are
parallel fibres that most likely comprise actinofibrils,
which are part of the wing membrane.
Discussion
According to the ontogenetic research done so far in
pterosaur (e.g. Bennett 1995; Kellner and Tomida 2000;
Kellner et al. 2013), the holotype (IVPP V12537) of
Daohugoupterus delicatus shows indication that it was an
adult animal when it perished, as several bones are fused:
the frontal and parietal, the elements of the proximal
carpal series, the scapula and coracoid.
The new species can be easily excluded from the
Pterodactyloidea based on the comparatively short
cervical vertebrae that bear cervical ribs. Also the shape
of the deltopectoral crest of the humerus, with a
quadrangular shape, differs from the humeri of all
known pterodactyloids (e.g. Wellnhofer 1991; Kellner
2003).
There are two species described from Dahougou,
coming from the same horizon as Daohugoupterus
delicatus:Jeholopterus ningchengensis and Pterorhynchus
Figure 3. (Colour online) Detail of the soft tissue of
Daohugoupterus delicatus gen. et sp. nov. on the counterslab. (a)
Soft tissue around the cervical vertebrae and right coracoids with
two white frames marking positions of (b) and (c). (b) Clumpy,
supple and crossed pycnofibres near the cervical vertebrae. (c) A
portion of parallel fibres similar to actinofibrils. Scale bar: 10 mm
in (a), 5 mm in (b) and (c). act, actinofibrils; car, carpus; cor,
coracoid; cv, cervical vertebra; pyc, pycnofibres; r, right.
Table 1. Measurements of Daohugoupterus delicatus gen. et
sp. nov. (mm).
Bones Length Width
cv 1 and 2 3.6 7.5
3 5.5 8.4
4 7.4 8.5
5 7.9 9.4
6 8.0 9.0
7 6.4 9.2
8 5.7 9.9
9 5.2
a
sca 31.4 left 4.5 left
cor 20.0
a
right
hu 41.8 left 5.2 left
ptd 11.0
Notes: cor, coracoid; cv, cervical vertebra; hu, humerus; ptd, pteroid; sca,
scapula.
a
Estimated length.
4X. Cheng et al.
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wellnhoferi. The most obvious difference between
Daohugoupterus delicatus and Jeholopterus ningchengen-
sis is the shape of skull. Even though the anterior tip is lost
in IVPP V12537, the skull of Daohugoupterus delicatus is
compressed laterally with the length longer than the width
differing from Jeholopterus ningchengensis and any other
anurognathid (Wang et al. 2002). Other differences can be
found in the palatal elements, which are quite reduced in
anurognathids to rod-like structures (e.g. Bennett 2007)
contrasting to the condition observed in Daohugoupterus.
Regarding Pterorhynchus wellnhoferi, a species based
on one fairly complete but poorly described specimen, the
main differences is the presence of a sagittal head crest that
extends above and across the posterior two-thirds of the
skull, and a relatively smaller antorbital fenestra (Czerkas
and Ji 2002). Even though it is uncertain if the holotype of
Pterorhynchus wellnhoferi is a mature individual, accord-
ing to the measurements (Table 2) it has a bigger size than
Daohugoupterus delicatus. Furthermore, Daohugoupterus
delicatus is the smallest pterosaur reported from the
Daohugou area so far.
There have been other pterosaurs discovered from
layers of the Tiaojishan Formation that outcrop in the
region of Linglongta and that have been correlated with
the Daohugou Bed (or Formation) (Wang et al. 2005,
2009). The most important group are the Wukongopter-
idae, which have shown to be rather diverse (e.g. Wang
et al. 2009,2010). Daohugoupterus can be excluded from
this group based mainly on the short cervical vertebrae
(e.g. Wang et al. 2009,2010).
Besides the Wukongopteridae, Scaphognathidae is
another important pterosaur clade that has been reported
from Linglongta, with three species discovered so far:
Fenghuangopterus lii,Jianchangopterus zhaoianus and
Jianchangnathus robustus (Lu
¨et al. 2010;Lu
¨and Bo
2011; Cheng et al. 2012), although it is debatable whether
Fenghuangopterus lii belongs to this group (Cheng et al.
2012). Unfortunately, Daohugoupterus delicatus lacks the
rostrum and nearly all teeth, which are the most important
diagnostic features of the Scaphognathidae. Despite all
this, Daohugoupterus delicatus differs from all scaphog-
nathids by having much smaller temporal openings,
including a slit-like lower temporal fenestra (e.g. Cheng
et al. 2012;Zhou2014; Bennett 2014). Also, the shape of
the deltopectoral crest of the humerus differs from all
scaphognathids where this bone is preserved.
In respect of Fenghuangopterus lii, if the measure-
ments provided by Lu
¨et al. (2010) are correct, the new
species has a comparatively shorter neck relative to the
length of the humerus (Table 2). Furthermore, the cervical
vertebrae of Fenghuangopterus lii appear to be longer and
narrower compared with Daohugoupterus delicatus.
Moreover, the ratio scapula/coracoid of Fenghuangopterus
lii is 0.80 according to the measurement in the published
paper (Lu
¨et al. 2010), which is an unusual feature for non-
pterodactyloids (Cheng et al. 2012), while, in the new
species, the scapula is clearly larger than the coracoid (see
Table 1).
Another important non-pterodactyloid pterosaur from
Asia is Sordes pilosus, recovered from the Karatau
Mountains in Kazakhstan (Sharov 1971). Compared with a
referred specimen of Sordes (Wellnhofer 1991), Daohu-
goupterus delicatus differs in the shape of the humerus,
that is quadrangular and lacks the rounded distal end.
Lastly, there are some features that can be regarded as
potential autapomorphies of the species, such as posterior
processes of the nasal extending between the frontals. The
most important is the particular shape of the sternum,
much wider than long, which has so far not been
documented in any other pterosaur.
Conclusions
Based on a series of combination of distinctive features, a
new genus and species, Daohugoupterus delicatus from
Daohugou (Ningcheng, Inner Mongolia), is established
and is the smallest pterosaur discovered from this area
until now. The new species differs from all other flying
reptiles recovered from the Linglongta Pterosaur Fauna
particularly due to the unusual shape of the sternum. The
holotype of Daohugoupterus also shows the preservation
of soft tissue that is consistent with the integumental
covering formed by pycnofibres, showing that such fibres
were widespread among pterosaurs.
Table 2. Comparison between Daohugoupterus delicatus gen. et sp. nov. and several pterosaur species from Jurassic of China (mm).
hu cv OS
Daohugoupterus delicatus (IVPP V12537) 41.8 49.7 (total) Adult This paper
Jeholopterus ningchengensis (IVPP V12705) 62 Adult Wang et al. (2002)
Pterorhynchus wellnhoferi (CAGS02-IG-gausa-2/DM 608) 52 63 (total) Czerkas and Ji (2002)
Fenghuangopterus lii (CYGB-0037) 63 85 (3– 7) Adult Lu
¨et al. (2010)
Jianchangnathus robustus (IVPP V16866) 51.2 (3– 5) Juvenile Cheng et al. (2012)
Wukongopterus lii (IVPP V15113) 38.7 Juvenile Wang et al. (2009)
Darwinopterus linglongtaensis (IVPP V16049) 40.4 Sub-adult Wang et al. (2010)
Kunpengopterus sinensis (IVPP V16047) 36.2 Adult Wang et al. (2010)
Notes: cv, cervical vertebra; hu, humerus; OS, ontogenetic stage.
Historical Biology 5
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Acknowledgements
The authors thank Dr Zhonghe Zhou and Yan Li for the field
work, Yutong Li for the preparation of the specimen and Jie
Zhang for the photographs.
Funding
This study was supported by the National Science Fund for
Distinguished Young Scholars [grant number 40825005], the
National Key Basic Research Program of China [grant number
2012CB821900] and the Hundred Talents Project of CAS.
Funding has also been received from the Fundaca
˜o Carlos
Chagas Filho de Amparo a
`pesquisa do Rio de Janeiro [grant
number FAPERJ # E-26/102.737/2012] and the Conselho
Nacional de Desenvolvimento Cientı
´fico e Tecnolo
´gico [grant
number CNPq # 304780/2013-8].
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... The most important localities that yield the Yanliao Biota are Daohugou in Ningcheng County of southeast Inner Mongolia (Haifanggou Fm.), Linglongta of Jianchang County of western Liaoning Province (Tiaojishan Fm.), and Mutoudeng of Qinglong County of northern Hebei Province (Tiaojishan Fm.; Lü et al., 2013;Huang, 2015Huang, , 2016. From the Haifanggou Formation at Daohugou (Liu et al., 2012), pterosaurs are relatively rare, with Jeholopterus ningchengensis (Wang et al., 2002), Pterorhynchus wellnhoferi (Czerkas & Ji, 2002) and Daohugoupterus delicatus (Cheng et al., 2015). From the slightly younger Tiaojishan Formation at the Linglongta locality, pterosaurs are abundant in number and in diversity (see Sullivan et al., 2014 for a review), with wukongopterids (Wang et al., 2009Lü et al., 2009;Cheng et al., 2017), Jianchangopterus (Lü & Bo, 2011), Jianchangnathus and Fenghuangopterus (Lü, Fucha & Chen, 2010). ...
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