Content uploaded by Di Liu
Author content
All content in this area was uploaded by Di Liu on Oct 26, 2017
Content may be subject to copyright.
Short note on a new anurognathid pterosaur with evidence
of perching behaviour from Jianchang of
Liaoning Province, China
JUNCHANG LU
¨1,2*, QINGJIN MENG3, BAOPENG WANG3*, DI LIU3,
CAIZHI SHEN1,2 & YUGUANG ZHANG3
1
Institute of Geology, Chinese Academy of Geological Sciences,
Beijing 100037, China
2
Key Laboratory of Stratigraphy and Palaeontology, Ministry of Land and Resources
of China, Beijing 100037, China
3
Beijing Museum of Natural History, Beijing 100050, China
*Correspondence: wbppku@163.com; Lujc2008@126.com
Abstract: A new anurognathid pterosaur, Versperopterylus lamadongensis gen. et sp. nov., is
erected based on a complete skeleton with a skull preserved. It is characterized by twoshort distinct
ridges present on the ventral surface of the cervical vertebrae; coracoids slightly longer than scap-
ula; humerus, wing phalanx 3 and tibia nearly the same in length; grooves clearly present on the
posterior surface of the wing phalanges 1–3; and the first toe reversed. It is the first anurognathid
pterosaur from China with a definitively short tail, and the first pterosaur with a reversed first toe.
The reversed first toe of Versperopterylus indicates that it had arboreal habitats. The discovery of
Versperopterylus lamadongensis from the Jiufotang Formation strongly expands the geological
age range for anurognathid pterosaurs.
Supplementary material: The character list and data matrix for phylogenetic analysis are avail-
able at http://doi.org/10.6084/m9.figshare.c.3873394
The anurognathid pterosaurs were a group of small
pterosaurs known mainly in Europe and Asia dur-
ing the Jurassic period. At present, four genera are
reported: Anurognathus, from the Late Jurassic of
Germany (Do
¨derlein 1923; Bennett 2007); Batra-
chognathus, from the Late Jurassic of Kazakhstan
(Ryabinin 1948; Unwin & Bakhurina 2000; Costa
et al. 2013) and also a specimen from Mongolia
(Bakhurina & Unwin 1995); Dendrorhynchoides,
from the Middle Jurassic of China (Ji & Ji 1998;
Ji et al. 1999; Lu
¨& Hone 2012; Jiang et al.
2015); and Jeholopterus (Ji & Yuan 2002; Wang
et al. 2002; Sullivan et al. 2014) from the Middle
to Late Jurassic of China. Here, we report a new
anurognathid pterosaur: Versperopterylus lama-
dongensis gen. et sp. nov. from the Early Creta-
ceous Jiufotang Formation of Lamamdong, in
Jianchang of Liaoning Province (Fig. 1). It repre-
sents the youngest anurognathid pterosaur in geo-
logical age and this substantially expands the
temporal range of anurognathid pterosaurs. The
reversed first toe of Versperopterylus lamadongen-
sis provides direct evidence of its perching
behaviour.
Systematic palaeontology
Pterosauria Kaup, 1834
Anurognathidae Kuhn 1937
Versperopterylus lamadongensis gen. et sp. nov.
Etymology.Versper-, Latin word for ‘dusk’
implying that the new pterosaur may seek food at
dusk; -pteryl, Latin word for ‘wing’. The specific
name is referred to the fossil locality, lamadong of
Jianchang County, Liaoning Province.
Type specimen. An almost complete skeleton
with skull and jaws (BMNHC-PH-001311). The
specimen is now stored in the Beijing Museum of
Natural History.
Locality and horizon. Lamadong goumen, Jian-
chang County, Liaoning Province; Jiufotang
Formation.
Diagnosis. A anurognathid pterosaur with the
following combination of characters: the skull is rel-
atively smaller than other anurognathids; the ratio of
length to width is approximately 79.1%; the poste-
rior region of the skull is rounded (shared with Anu-
roganthus and Bactrachognathus); the anterior end
From:Hone, D. W. E., Witton,M.P.&Martill, D. M. (eds) New Perspectives on Pterosaur Palaeobiology.
Geological Society, London, Special Publications, 455, https://doi.org/10.1144/SP455.16
#2017 The Author(s). Published by The Geological Society of London. All rights reserved.
For permissions: http://www.geolsoc.org.uk/permissions. Publishing disclaimer: www.geolsoc.org.uk/pub_ethics
of the lower jaw is horizontally expanded; the cau-
dal end of the mandible is without a distinct dorsal
‘coronoid’ eminence; postexapophyses are present
on the cervical vertebrae; the number of caudal ver-
tebrae is 15 or fewer (shared with Anuroganthus);
metacarpals I– III are the same length (shared with
Batrachognahus and Dendrorhynchoids); a promi-
nent anteriorly directed tubercle on the dorsal apex
of the external trochanter of the femur; the length
of metatarsal IV is shorter than metatarsals II–III
(shared with Jeholopterus); coracoids are slightly
longer than the scapula; the humerus, wing phalange
3 and tibia are nearly the same length; the wing
metacarpal is approximately 36% of the length of
the humerus; grooves are clearly present on the pos-
terior surface of the wing phalange 1 to the wing
phalange 3; and the first toe is reversed.
Description
The specimen is almost complete, except for miss-
ing one humerus and some cervical vertebrae (Figs
2–4; Table 1). The wing span is about 1 m. The
fused scapulocoracoid, the smooth long bone sur-
faces and the fusion of extensor process to the
wing phalanx 1 indicate that it is at least a sub-adult
animal. The skull is remarkably small on this ani-
mal. The ratio of skull width to humeral length is
0.57, which is smaller than that Dendrorhynchoides
mutoudengensis (this ratio is 0.74: Lu
¨& Hone
2012). It is unlike other anurognathids and its ven-
tral surface is exposed. The skull is slightly wider
than long. The ratio of length to width of the skull
is approximately 79.1%, which is similar to that of
Dendrorhynchoides mutoudengensis (this ratio is
80%: Lu
¨& Hone 2012). It is semi-circular in ventral
view (Fig. 3a). The premaxilla and maxilla are slen-
der, rod-like. The upper jaw teeth are relatively stout
with blunt tips (Fig. 3b), unlike some longer teeth
with curved tips found in Dendrorhynchoides
mutoudengensis (Lu
¨& Hone 2012). There are
eight teeth preserved. Although the exact number
of teeth is not sure due to the poor preservation,
there are, perhaps, 12 teeth in the upper jaw accord-
ing to the distance between two teeth and the
dentition length.
There are four cervical vertebrae preserved. The
anterior two are rectangular in ventral view. There
are two short, distinct longitudinal ridges on the cen-
tral portion of the centrum, which form a groove
between them. The cervical vertebrae bear a distinct
postexapophysis. The neural spine is high and
blade-like. There are no cervical ribs associated
with the cervical vertebrae.
The anterior dorsal vertebrae bear a distinct neu-
ral spine. There are 14 dorsal vertebrae preserved
with their ventral surfaces exposed. Both the ante-
rior and posterior articular ends are flat. The ventral
surfaces are smooth. The shaft of the third dorsal rib
is the widest of the dorsal ribs. The seventh dorsal
rib is the longest and the first one is the shortest.
At least 13 caudal vertebrae can be observed.
The anterior part of the tail is covered by the
ischium, but it is inferred that no more than two cau-
dal vertebrae were covered. This means that there
are no more than 15 caudal vertebrae (Fig. 4d).
Fig. 1. A map of Chinese anurognathid pterosaur sites: ffiDendrorhynchoides;fflJeholopterus;
Dendrorhynchoides mutoudengensis;ÐVersperopterylus (BMNHC-PH-001311).
J. LU
¨ET AL.
The anterior caudal vertebrae are rectangular in lat-
eral view. The last one bears a pointed distal end.
The left scapula and coracoid are fused into
the scapulocoracoid. The shaft of the scapula is
slightly concave and it is slightly shorter than the
coracoid. The coracoid is stouter than the scapula
and it bears a distinct brachial flange. The shaft of
the coracoid is straight. The shaft of the humerus
Fig. 2. Photograph (a) and line drawings (b)ofVersperopterylus (BMNHC-PH-001311). Abbreviations: cav, caudal
vertebrae; csk, crushed skull; co, coracoids; dv, dorsal vertebrae; dr, dorsal ribs; f, femur; fot, fourth toe; fi, fibula;
ft, first toe; h,, humerus; lj, lower jaw; mc, maniual claw; mcI-IV, metacarpals I –IV; mttI-IV, metatarsals I – IV;
mttv, metatarsal V; pt, pteroid; rd, radius; sc, scapula; ti, tibia; ul, ulna.
ANUROGNATHID PTEROSAUR FROM CHINA
Fig. 3. (a). Photograph of the skull of Versperopterylus; and (b) Close-up of the dentition, showing the teeth
(arrows point). Scale bar is 0.5 cm in (b).
J. LU
¨ET AL.
Fig. 4. Photographs of the holotype Versperopterylus (BMNHC-PH-001311): (a) whole specimen; (b) close-up of
the pes; (c) close-up of wing phalanges 1 and 2; and (d) close-up of the tail. Abbreviations: cav, caudal vertebrae;
fot, the fourth toe; gr, groove; mtt I– IV, metatarsal I–IV; rft, the reversed first toe; wph1, 2, wing phalanges 1 and
2; st, second toe; tt, third toe. Scale bars are 1 cm in (b) and (d).
ANUROGNATHID PTEROSAUR FROM CHINA
is straight. There is no pneumatic opening near its
proximal end. The deltopectoral crest of the
humerus is relatively small, and is longer than
wide. It is located proximally. There is a distinct
ridge near the distal end of the humerus. The lateral
margin of the deltopectoral crest is straight.
The ulna and radius are parallel to each other.
The ulna is longer than the radius. The width ratio
of ulna to radius is 1.17. The ulna bears a distinct
olecranon process at its proximal end. The shaft of
the radius is straight with slightly expanded distal
and proximal ends.
The proximal carpal is irregular in shape with a
concave surface in ventral view. It is larger than
any other carpals. It articulates with the distal end
of the ulna. There are two unfused distal carpals.
The proximal portion of the pteroid is slightly
curved. The distal end of the pteroid is expanded,
and knob-like. The length ratio of pteroid to
humerus is 0.19. Metacarpals I – IV are the same
length. Metacarpal IV is stout. The first phalanx of
the first manual digit is long and slender, slightly
thinner than metacarpals I– III. The ungual is deep
at its proximal end and curved with a sharp tip.
The rest of the digits of the left hand are missing.
The wing phalanges decrease in length from the
first to the third. The ventral surfaces of all the right
wing phalanges are exposed. Although the phalan-
ges were heavily depressed during preservation, a
distinct groove appears posterior to the middle
shaft of wing phalanx 1 (Fig. 4c). This groove is
on the posterior surface of the wing phalanx 1.
The extensor tendon process is fused with the first
wing phalanx. The proximal end of the first wing
phalanx is expanded. The first wing phalange is
the longest. The proximal end of the second wing
phalanx is expanded, larger than its distal end.
The distal end of the wing phalanx 2 is slightly
expanded. The proximal end of the third wing pha-
lange is much more expanded than its distal end,
which becomes thinner but is slightly expanded.
There is also a distinct groove on the posterior sur-
faces of wing phalanges 2 and 3. The fourth wing
phalanx is rod-like, quite small and straight with
pointed distal end. This is different to the curvature
seen in the distal phalanges of other anurognathids
(see Hone et al. 2015), but this is also very small
and coupled with the apparent tapering of phalanx
3; thus, it might explain the apparent absence of
the fourth phanalx in the juvenile Anurogahtus.
Table 1. Measurements (in mm) of holotype of Versperopterylus lamadongensis gen. et sp. nov
Length Width
Skull 32.1 40.6 (between quadrate)
Single cervical 9.4 6.8
Dorsals +sacrals 112.9
Tail 31.0
Scapula 37.2 –
Coracoid 38.3 3.4
Humerus 71.0 6.4
Deltopectoral crest 12.5 5.2
Ulna 96.4 5.6
Radius 92.9 4.8
Metacarpals 25.8
Pteroid 13.7 1.7
Manus digit I: 1; 2 15.8; 13.6 (claw) 1.1; 5.4 (proximal end)
Width of phalange 1 121.3 (with extensor process);
117.8 (without extensor process)
4.6
Width of phalange 2 95.4 2.9
Width of phalange 3 71.0 1.9
Width of phalange 4 14.3 1.0
Acetabulum 7.8 4.7 (height)
Pre-acetabulum process 32.0
Femur 52.6 5.7
Tibia 71.9 6.6
Fibula 47.9
Metatarsals I– IV (whole) 34.5 8.0
Metatarsals I– V 30.9; 34.0; 34.0; 33.0; 13.9 3.5; –; – ; – ; –
Pedal digit I: 1; 2 12.2; 8.1 2.1; 4.3
Pedal digit II: 1; 2; 3 6.3; 11.7; 11.2 2.5; 1.6; 4.4
Pedal digit III: 1; 2: 3; 4 5.4; 4.9; 9.8; 10.6 2.7; 2.9; 1.9; 5.1
Pedal digit IV: 1; 2; 3; 4; 5 4.6; 3.9; 2.1; 7.5; 9.2 2.6; 2.6; 2.5; 1.9; 4.1
Pedal digit V: 1; 2 19.8; – 1.8
J. LU
¨ET AL.
The pelvic girdle is not well preserved. It seems
that the pubis, ischium and ilium are fused together.
The preacetabular process of the ilium is slender and
long, covering at least four posterior dorsal verte-
brae. The acetabulum bears a distinct dorsal margin
and anterior margin. The shapes of ischium and
pubis are unclear because of being covered by the
proximal portion of the femur. The femoral head
has a distinct neck, which forms an angle of 1208
with the shaft of the femur. There is a distinct
ridge on the anterolateral surface near the proximal
end. The femoral head bears a sharp margin. The
shaft of the femur is straight. The tibia is straight
with an oblique proximal end. The proximal end
of the fibula is fused with the proximal end of the
tibia. The proximal portion of the fibula is rod-like
but below it, by about 0.7 cm, it becomes sharply
expanded, which may be a pathology. The distal
end of the tibia bears two distinct condyles. Metatar-
sals I– IV are in close contact with each other. Their
proximal ends are fused. Metatarsal II and metatar-
sal III are the same length and size. Metatarsal I is
shorter than metatarsal IV, which is, in turn, shorter
than metatarsals II and III. Metatarsal V is sepa-
rated from other metatarsals. It is stout and shorter
than other metatarsals. The first phalange of digit 5
is straight but its length cannot be determined due
to a missing distal portion. The first toe of the
right pes is clearly reversed (Fig. 4b), although
the left toe is slightly destroyed. The tips of the
pedal unguals of digits II, III and IV are pointed
in the same direction, whilst the tip of the first toe
is pointed in the opposite direction in both pedes;
thus, it is inferred that the first toe is reversed.
The naturally articulated dorsal to caudal series
and all other elements that are not missing indicate
that they are not transported far and are quickly
buried after death. Therefore, they are regarded as
genuine, not caused by preservation. The digital
formula is 2–3–4–5–2. The pedal unguals are
curved with sharp tips. They bear a large extension
process. The pedal unguals are slightly smaller than
the manual unguals.
Comparison and discussion
Versperopterylus differs from Dendrorhynchoides
(Ji & Ji 1998; Lu
¨& Hone 2012; Jiang et al. 2015)
in that the length ratio of wing phalange 2 to
Fig. 5. The living scene of Versperopterylus (drawn by Zhao Chuang).
ANUROGNATHID PTEROSAUR FROM CHINA
Fig. 6. Strict consensus of 630 most parsimonious trees (MPTs) obtained by TNT, based on analysis of 68 taxa and
124 characters, showing the phylogenetic position of Versperopterylus. gen. et sp. nov. (tree length, 503). The
numbers adjacent to each node are Bremer support values.
J. LU
¨ET AL.
phalange 1 (i.e. 0.79) is smaller than that of Dendro-
rhynchoides (i.e. 0.88). The length ratio of wing
metacarpal to humerus is also smaller than that of
Dendrorhynchoides (it is 36% in Versperopterylus:
however, it is 40.6% in Dendrorhynchoides mutou-
dengensis:Lu
¨& Hone 2012). Versperopterylus has
a tail that is shorter than Dendrorhynchoides.Den-
drorhynchoides was first reported to bear a long
tail because it was not clearly preserved, and was
once thought to have had the tail added onto the
specimen (Unwin et al. 2000). However, the discov-
ery of more material further confirms that there is a
greater likelihood that it had a long tail, as more spec-
imens indicate (Lu
¨& Hone 2012; Jiang et al. 2015).
Versperopterylus differs from Jeholopterus
(Wang et al. 2002; Sullivan et al. 2014) in that Jeho-
lopterus does not bear a tail (according to the holo-
type (IVPP V12705) and the referred specimen
(CAGS-IG-02-81), which came from the same
place, both do not have a visible tails; therefore,
we inferred that Jeholopterus does not bear a tail,
although we cannot exclude the possibility that the
tail is not preserved – even if they do have a tail,
the tail should be very short); the length ratio of
wing phalange 2 to phalange 1, and the ratio of
ulna to humerus and femur to tibia, are smaller
than those of Jeholopterus (IVPP V12705) (they
are 0.79, 1.36 and 0.73 in Versperopterylus, whilst
they are 0.88, 1.44 and 0.80 in Jeholopterus,
respectively).
Versperopterylus differs from Anurognathus
(Do
¨derlein 1923; Dalla Vecchia 2002; Bennett
2007) in that the length ratio of the wing metacarpal
to the metatarsals is greater than that of Anurogna-
thus (this ratio is 0.61 in Anurognathus (BSP
1922.I.42), whilst it is 0.76 in Vesperopterylus).
Versperopterylus differs from Batrachognathus
(Ryabinin 1948; Unwin & Bakhurina 2000) in that
its length ratio of ulna to humerus is greater than
that of Batrachognathus (the ratio is 1.0 in Batra-
chognathus, whilst it is 1.36 in Vesperopterylus).
At present, there are three kinds of anurognathid
pterosaurs from western Liaoning and its surround-
ing areas, in terms of their body bauplans: a tail-less
one, such as Jeholopterus; one with a short tail, such
as Versperopterylus; and one with a longer tail, such
as Dendrorhynchoides. The reversed first toe of Ver-
speropterylus indicates that it had arboreal habitats,
which seems like more of a gripping adaptation: ver-
tical clinging, branch climbing and/or branch cling-
ing (above or below the branch) (Fig. 5).
Phylogenetic analysis
A phylogenetic analysis was carried out which in-
cluded 88 ingroup taxa and 124 characters (based
on the modified data matrix of Lu
¨et al. 2016). We
subjected the dataset to a maximum parsimony anal-
ysis in TNT v1.1 (Goloboff et al. 2008). We first
conducted a ‘new technology’ search (with default
parameters for sectorial search, ratchet, tree drift
and tree fusion) that recovered a minimum length
tree in 10 replicates. This procedure aims to broadly
sample tree space and identify individual tree
islands. We then subjected the recovered most par-
simonious trees (MPTs) to a traditional search with
TBR branch swapping, which more fully explores
the tree islands found in the ‘new technology’
search. This process returned a total of 3 043 358
MPTs in 503 steps. Bremer values were used to
assess clade support. The strict consensus of the
630 MPTs (Fig. 6) recovers a monophyletic group
of anurognathid pterosaurs, with Versperopterylus
basal to other anurognathid pterosaurs. The clade
Anurognathidae is supported by the following syn-
apomorphies: skull broad with very short preorbital
region; palatal elements reduced to thin bars of
bone; teeth are small, peg-like and widely spaced;
cervical ribs are highly reduced or absent; the com-
bined length of the caudal vertebrae is shorter than
the dorsal series; the ulna is 133–150% of the
humerus length; the manus digit IV (wing-finger)
phalanx 1 is 1.5– 2.0 times longer compared to the
length of the tibiotarsus; the contribution of the
wing-phalanx 1 to the wing-finger length is 30 –
40%; and preacetabular process of the ilium is lon-
ger than the postacetabular process.
Conclusion
Versperopterylus is the first anurognathid pterosaur
from China preserved with a clearly reduced tail,
and it is also the youngest anurognathid pterosaur
in geological age. The reversed first toe of Versper-
opterylus may indicate that it, perhaps, has a grip-
ping adaptation.
We thank Dr Dale Winkler (Southern Methodist Univer-
sity, Dallas, TX, USA) for providing thorough comments
on the earlier version of this paper. This study was sup-
ported by the National Natural Science Foundation of
China (grant Nos 41688103 and 41672019), the Funda-
mental Research Funds for the Chinese Academy of
Geological Sciences (grant No. JB1504) and the project
from the China Geological Survey (DD 20160126) to
Junchang Lu
¨.
References
Bakhurina, N.N. & Unwin, D.M. 1995. A survey of
pterosaurs from the Jurassic and Cretaceous of the for-
mer Soviet Union and Mongolia. Historical Biology,
10, 197–245.
Bennett, S.C. 2007. A second specimen of the pterosaur
Anurognathus ammoni.Pala
¨ontologische Zeitschrift,
81, 376–398.
ANUROGNATHID PTEROSAUR FROM CHINA
Costa, F.R., Alifanov, V., Dalla Vecchia, F.M. &
Kellner, A.W.A. 2013. On the presence of an
elongated tail in an undescribed specimen of Batra-
chognathus volans (Pterosauria: Anurognathidae:
Batrachognathinae). In:Saya
´o, J.M., Costa, F.R.,
Bantim, R.A.M. & Kellner, A.W.A. (eds) Short
Communications, Rio Ptero 2013 – International Sym-
posium on Pterosaurs. Universidade Federal do Rio de
Janeiro, Rio de Janeiro, 54– 56.
Dalla Vecchia, F.M. 2002. Observations on the non-
pterodactyloid pterosaur Jeholopterus ningchengensis
from the Early Cretaceous of northeastern China.
Natura Nascosta,24, 8– 27.
Do
¨derlein, L. 1923. Anurognathus Ammoni, ein neuer
Flugsaurier. Sitzungsberichte der Mathematisch-
Naturwissenschaftlichen Abteilung der Bayerischen
Akademie der Wissenschaften zu Mu
¨nchen.
Goloboff, P.A., Farris, J.S. & Nixon, K.C. 2008. TNT.
A free program for phylogenetic analysis. Cladistics,
24, 774– 786.
Hone, D.W.E., Van Rooijen, M.K. & Habib, M.B.
2015. The wingtips of the pterosaurs: anatomy, aero-
nautical function and ecological implications.
Palaeogeography, Palaeoclimatology, Palaeoecology,
440, 431– 439.
Ji,S.A.&Ji, Q. 1998. A new fossilpterosaur (Rhamphorhyn-
choidea) from Liaoning, Dendrorhynchus curvidenta-
tus, gen. et sp. nov. Jiangsu Geology,22, 199–206.
Ji, S.A., Ji,Q.&Padian, K. 1999. Biostratigraphy of new
pterosaurs from China. Nature,398, 573– 574.
Ji,Q.&Yu a n , C.X. 2002. Discovery of two kinds of proto-
feathered pterosaurs in the Mesozoic Daohugou Biota in
the Ningcheng region and itsstrati graphic and biologic
significances. Geological Review,48, 221– 224.
Jiang, S.X., Wang,X.L., Cheng, X., Costa, F.R., Huang,
J.D. & Kellner, A.W.A. 2015. Short note on an
anurognathid pterosaur with a long tail from the
Upper Jurassic of China. Historical Biology,27,
718–722.
Lu
¨, J.C. & Hone, D.W.E. 2012. A new Chinese anurogna-
thid pterosaur and the evolution of pterosaurian tail
lengths. Acta Geologica Sinica,86, 1317– 1325.
Lu
¨, J.C., Kundra
´t,M.&Shen, C.Z. 2016. New material
of the pterosaur Gladocephaloideus Lu
¨
et al
., 2012
from the Early Cretaceous of Liaoning Province,
China, with comments on its systematic position.
PLoS ONE,11, e0154888. https://doi.org/10.1371/
journal.pone.0154888
Ryabinin, A.N. 1948. Remarks on a flying reptile from the
Jurassic of the Kara-Tau, Akademia Nauk, Paleonto-
logical Institute, Trudy,15, 86– 93.
Sullivan, C., Wang, Y., Hone, D.W., Wang, Y.Q.,
Xu,X.&Zhang, F.C. 2014. The vertebrates of
the Jurassic Daohugou biota of northeastern China.
Journal of Vertebrate Paleontology,34, 1– 37.
Unwin, D.M. & Bakhurina, N.N. 2000. Pterosaurs
from Russia, Middle Asia and Mongolia. In:Benton,
M.J., Shishkin, M.A., Unwin, D.M. & Kurochkin,
E.N. (eds) The Age of Dinosaurs in Russia and
Mongolia. Cambridge University Press, Cambridge,
420–433.
Unwin, D.M., Bkhurina, N.N. & Lu
¨, J.C. 2000. On the
systematic and stratigraphic significance of pterosaurs
from the Lower Cretaceous Yixian Formation (Jehol
Group) of Liaoning, China. Mitteilungen aus dem
Museum fu
¨r Naturkunde in Berlin, Geowissenschafli-
chen Reihe,3, 181– 206.
Wang, X.L., Zhou, Z.H., Zhang, F.C. & 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, 226– 232.
J. LU
¨ET AL.
