Content uploaded by Xiaolin Wang
Author content
All content in this area was uploaded by Xiaolin Wang on Jan 19, 2017
Content may be subject to copyright.
A new toothed pteranodontoid (Pterosauria, Pterodactyloidea) from the Jiufotang
Formation (Lower Cretaceous, Aptian) of China and comments on Liaoningopterus gui
Wang and Zhou, 2003
Taissa Rodrigues
a
, Shunxing Jiang
b
, Xin Cheng
b,c
, Xiaolin Wang
b
*and Alexander W.A. Kellner
c
a
Department of Biology, Agrarian Sciences Center, Universidade Federal do Espı
´rito Santo, Alto Universita
´rio s/n, Caixa Postal 16,
Guararema CEP 29500-000, Alegre, ES, Brazil;
b
Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate
Paleontology and Paleoanthropology, Chinese Academy of Sciences, P.O. Box 643, Beijing 100044, P.R. China;
c
Laboratory of
Systematics and Taphonomy of Fossil Vertebrates, Department of Geology and Paleontology, Museu Nacional/Universidade Federal do
Rio de Janeiro, Quinta da Boa Vista s/n, Sa
˜o Cristo
´va
˜o CEP 20940-040, Rio de Janeiro, RJ, Brazil
(Received 17 February 2015; accepted 18 February 2015)
Pteranodontoids consist of a diverse and cosmopolitan clade of Cretaceous pterodactyloid pterosaurs. In the Jiufotang
Formation (Lower Cretaceous, Aptian) of northeastern China, pterodactyloids are represented by azhdarchoids and
pteranodontoids, including istiodactylids and anhanguerians. Here, we describe a new pterosaur from this unit that
represents a new species of toothed pteranodontoid. Its overall morphology is consistent with other toothed pteranodontoids
but shows some interesting features such as the orbit being more ventrally positioned than in all other species from this
clade. It differs markedly from all other pterosaurs from this unit, including Liaoningopterus,Guidraco and Ikrandraco,
with which the new taxon is possibly related. In addition to the description of the new taxon, we also describe the
anhanguerid Liaoningopterus gui in more detail. The new species, Liaoningopterus gui,Guidraco venator and Ikrandraco
avatar, are large pterosaurs with very distinct tooth morphologies, suggesting that they had different prey preferences,
partially explaining how the Jiufotang Formation could bear such a high diversity of pterosaur species.
Keywords: Lower Cretaceous; Jiufotang Formation; China; Pterosauria; Linlongopterus;Liaoningopterus
1. Introduction
Pteranodontoids are derived pterosaurs that lived during
the Cretaceous period, being found on all continents
(Kellner 2003). The clade was defined as ‘the most recent
common ancestor of Anhanguera and Pteranodon and all
their descendants’ (Kellner 2003) and has been recovered
by posterior, independent phylogenetic studies (e.g.
Andres and Ji 2008; Andres and Myers 2013; Andres
et al. 2014). Pteranodontoids are diagnosed by some
postcranial features (both axial and appendicular), such as
tall and spike-shaped neural spines of the midcervical
vertebrae; scapula shorter than the coracoid and with a
suboval proximal surface; and humerus with a warped
deltopectoral crest, posteriorly directed medial crest and
subtriangular distal end (Kellner 2003).
Among pteranodontoids, the recently erected clade
Anhangueria has been defined as ‘all pteranodontoids
more closely related to Anhanguera blittersdorffi than to
Istiodactylus latidens and Cimoliopterus cuvieri’ (Rodri-
gues and Kellner 2013). These pterosaurs are cosmopo-
litan, being known from Cretaceous strata of all continents
(except Antarctica), and are outstandingly rich in the
Romualdo Formation of Brazil (Kellner et al. 2013) and
the Cambridge Greensand of England (Rodrigues and
Kellner 2013).
The Jiufotang Formation (Aptian; He et al. 2004)of
northeastern China presents a rich and diverse pterosaur
assemblage, which to current knowledge is represented
only by dsungaripteroids. From these, there is one basal
species (Wang, Kellner, et al. 2008), exceptionally well-
represented azhdarchoids – chaoyangopterids and tape-
jarines (Dong et al. 2003; Wang and Zhou 2002,2003;
Kellner and Campos 2007;Lu
¨, Unwin, et al. 2008) and,
among pteranodontoids, some species of istiodactylids
(Wang, Campos, et al. 2008;Lu
¨, Xu, et al. 2008), the
recently described Ikrandraco avatar (Wang et al. 2014),
and two anhanguerians: Liaoningopterus gui and Gui-
draco venator (Wang and Zhou 2003; Wang et al. 2012).
In this paper, we describe a new toothed pteranodon-
toid pterosaur from the Jiufotang Formation, possibly
related to anhanguerians, and provide new osteological
information of Liaoningopterus gui.
1.1 Institutional abbreviations
IVPP, Institute of Vertebrate Paleontology and Paleoan-
thropology, Chinese Academy of Sciences, Beijing,
China; SNSB-BSPG, Staatliche Naturwissenschaftliche
Sammlungen Bayerns/Bayerische Staatssammlung fu
¨r
Pala
¨ontologie und Geologie, Munich, Germany.
q2015 Taylor & Francis
*Corresponding author. Email: wangxiaolin@ivpp.ac.cn
Historical Biology, 2015
Vol. 27, No. 6, 782–795, http://dx.doi.org/10.1080/08912963.2015.1033417
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
2. Material and methods
Specimens IVPP V15549 and IVPP V13291 are deposited at
the IVPP in Beijing, China, where they were first-hand
analysed. In order to assess the phylogenetic position of the
species herein described, an analysis was performed based on
a modified version of a previously published matrix (Wang
et al. 2014), which was selected among other pterosaur
phylogenies for the focus on toothed pteranodontoids instead
of the interrelationships of more inclusive groups. Analysis
was performed on Tree Analysis Using New Technology
(TNT), version 1.1 for Windows with a menu interface
(Goloboff et al. 2008). All characters were unordered and
given the same weight. Ornithosuchus longidens was
considered the primary outgroup. Analysis was performed
using the traditional search, with the tree bisection-
reconnection (TBR) algorithm (although the software uses
both TBR and subtree pruning-regrafting [SPR]), with 5000
random addition sequence replicates. Branches were
collapsed when their minimum length was equal to zero.
3. Systematic palaeontology
Pterosauria Kaup, 1834
Pterodactyloidea Plieninger, 1901
Pteranodontoidea Marsh, 1876,sensu Kellner, 2003
3.1 Linlongopterus jennyae gen. et sp. nov.
3.1.1 Linlongopterus gen. nov.
Type species.Linlongopterus jennyae, type by monotypy.
Etymology. From the Chinese words lin, meaning forest, and
long, dragon; and pteros, from the Greek meaning wing.
Diagnosis. The same as for the type species.
3.1.2 Linlongopterus jennyae sp. nov.
Etymology. Species name in honour of the late Elfriede
Kellner, also known as Jenny, a great supporter of
palaeontological studies.
Holotype. IVPP V15549, partial associated skull and
mandible, deposited at the IVPP in Beijing, China.
Locality and horizon. Jianchang, Jianchang County,
western Liaoning Province, China. Jiufotang Formation,
Aptian (120 Ma) (He et al. 2004).
Diagnosis. Pteranodontoid pterosaur with the orbit ventrally
positioned, with the ventral margin almost level with the
nasoantorbital fenestra. The following combination of
characters also distinguishes it from other members of this
clade: lacrimal process of the jugal thin; lacrimal process of
the jugal oriented vertically; lower temporal fenestra with a
broad ventral margin; slight lateral bend on the anterior end
of the rostral process of the pterygoid.
Description and comparison
Generalities: Specimen IVPP V15549 (Figures 1 and 2)
is preserved on a slab that was broken into two piec es. These
slabs were brought to the preparation laboratory of the IVPP
in Beijing and were glued together and then prepared.
On the smaller slab there are the tips of the rostrum and
mandible, and the larger slab includes most of the preserved
skull and the mandibular rami. Because some little
fragments are missing in the processing of the collection,
it makes the two pieces seem not contiguous. Actually, the
boundaries of them match very well, and there is no missing
part between the two pieces, and they belong to the same
individual. The external surface of the cortical bone in some
portions of the skull is also missing.
Ontogeny: Although representing a large individual,
the holotype of Linlongopterus jennyae shows several
bones displaced from their original anatomical position,
clearly showing that they were not fused, such as the jugal,
quadrate and pterygoid. The holotype of Anhanguera
santanae (Wellnhofer 1985) shows the same displacement
of cranial and lower jaw elements, and also includes
unfused carpals, indicating osteological immaturity.
Furthermore, the bone surface in this new pterosaur is
rough and consistent with Bennett’s (1993) immature
grains, suggesting that it is a sub adult.
Skull: As mentioned above, the skull is partially
preserved and exposed mostly in right lateral view. Due to
compression, part of the palate is displaced ventrally to the
skull. Some bones are dislocated from their original
anatomical position, such as the left jugal, quadratojugal
and quadrate and the right pterygoid. Similar to other
toothed pteranodontoids, the nasoantorbital fenestra is
large and elongate, with ventral and dorsal margins that
gradually diverge posteriorly.
Premaxilla: The right premaxilla comprises the tip of the
rostrum and extends posteriorly, forming the dorsal part of
the skull, including the dorsal margin of the nasoantorbital
fenestra (Figure 1). The preserved portions do not possess a
median crest, and if one was present, it would be
uncommonly short and restricted to the middle part of the
rostrum. Based on the premaxillary crest morphology and
modest length of the rostrum of known anhanguerians (about
50% of the skull length; e.g. Kellner and Tomida 2000; Wang
and Zhou 2003), we find this unlikely.
Five alveoli are preserved on the anterior part of the
right side of the rostrum, the first and the last showing the
tooth preserved in situ, although broken (Figure 3).
It cannot be confidently determined if alveoli are present
on the very tip of the skull, as seen in anhanguerids and
related taxa (e.g. Rodrigues and Kellner 2013). On the
dorsal margin of the premaxillary tip, the cortex of the
right side is broken and some of the teeth from the left side
can be observed.
Maxilla: The maxilla occupies great part of the lateral
surface of the skull and forms the ventral margin of the
nasoantorbital fenestra. From the breakage until the level
of the anterior margin of the nasoantorbital fenestra, two
teeth can be seen, still in their sockets (Figure 1), but due to
Historical Biology 783
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
Figure 1. (Colour online) General view of specimen IVPP V15549, holotype of Linlongopterus jennyae gen. et sp. nov. (A) Photograph;
(B) respective line drawing. afo, adductor fossa; art, articular; d, dentary; j, jugal; l, left; m, maxilla; naof, nasoantorbital fenestra; pl,
palatine; pm, premaxilla; pty, pterygoid; q, quadrate; qj, quadratojugal; r, right; te, teeth. Scale bar: 50mm.
Figure 2. Line drawing reconstruction of the skull and mandible of Linlongopterus jennyae based on holotype IVPP V15549. Preserved
bones in white. Scale bar: 50 mm.
784 T. Rodrigues et al.
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
poor preservation of the rest of the alveolar margin, it is
not possible to determine the total number of alveoli in this
bone, nor determine its distribution. Immediately ante-
riorly and ventrally to the nasoantorbital fenestra, the bone
is lower than in the adjacent portion and it is likely that the
right bone was lost and that the medial part of the left
maxilla is observable. Ventrally to the nasoantorbital
fenestra, the ventral-most margin of the maxilla was lost.
Jugal: The left jugal is seen in medial view and is
closely associated but disarticulated with the quadrate.
It lies still in articulation with most of the quadratojugal
(Figure 4). The articular surface for the condyloid process
is comparatively large and has an oval shape, similar to
Pteranodon (Bennett 2001).
Only the base of the maxillary process of the jugal is
preserved, but it can be observed that it forms a straight
angle with the lacrimal process, which is similar to
Liaoningopterus gui and other anhanguerids. However, the
lacrimal process is vertically oriented and thin, thus
differing from the broad condition seen in Liaoningopterus
gui and Guidraco venator (Wang and Zhou 2003; Wang
et al. 2012).
The base of the postorbital process of the jugal is
inclined posteriorly and overlain by the quadrate. Based on
this configuration, it is clear that the orbit is more ventrally
positioned than in other pteranodontoids, with the ventral
margin almost level with the ventral margin of the
nasoantorbital fenestra.
Figure 4. (Colour online) Posterior part of the skull of specimen IVPP V15549, holotype of Linlongopterus jennyae gen. et sp. nov.,
showing some dislocated bones. art, articular; j, jugal; l, left; m, maxilla; pty, pterygoid; q, quadrate; qj, quadratojugal; r, right. 1,
postorbital process of the jugal; 2, lacrimal process of the jugal; 3, maxillary process of the jugal. Scale bar: 50 mm.
Figure 3. (Colour online) Tip of the skull of specimen IVPP V15549, holotype of Linlongopterus jennyae gen. et sp. nov., in right lateral
view. Teeth and alveoli in the lower half are from the right side; teeth in the upper half are from the left side. te, teeth. Scale bar: 50 mm.
Historical Biology 785
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
Quadratojugal: Due to the dislocation of the quadrate,
the left quadratojugal is clearly visible in medial view
(Figure 4). Its articular head is detached from the rest of
the bone and is still attached to the quadrate, participating
in the condyloid process of the jaw articulation. The main
part of the bone lays in anatomical position, just posterior
to the jugal, and consists of an elongated bar. The
anterodorsal margin of the quadratojugal and the
postorbital process of the jugal are divergent, and form a
wide angle, thus resulting in a comparatively large lower
temporal fenestra with a broad ventral margin, differing in
this respect from anhanguerids, Guidraco venator and
Ikrandraco avatar (Campos and Kellner 1985; Wellnhofer
1985,1991; Kellner and Tomida 2000; Veldmeijer 2003;
Wang and Zhou 2003; Wang et al. 2012,2014).
Quadrate: The left quadrate is seen in anterior view
(Figure 4). The medial (pterygoid) process of the articular
head is tongue shaped, long but not as seen in the
istiodactylid Hongshanopterus lacustris (Wang, Campos,
et al. 2008)orAnhanguera santanae (Wellnhofer 1985).
On the opposite side, the quadratojugal is still connected
with the quadrate, with both forming the condyloid
process. There is a helical jaw joint, a common feature
among dsungaripteroids (Kellner 2003). The quadrate
shaft is rod-like and curved medially. The quadrate flange
(termed medial lamella by Wellnhofer [1985]) shows a
rugose surface with several small openings and
depressions, but no pneumatic foramen could be visually
assessed.
Palatine: The compression of the skull resulted in
some distortion, and part of the palatal region, most
probably the palatine (sensu Wellnhofer 1978; Bennett
2001; Kellner 2013; but see O
˝si et al. 2010; Pinheiro et al.
2012 for different interpretation) is exposed ventrally to
the maxillary teeth row (Figure 1), as well as sending a
process posteriorly, partially overlying the jugal and
quadrate. The bone is crushed and no detailed anatomical
features can be observed. The main exposed part is located
anteriorly to the nasoantorbital fenestra and the palatal
fenestrae cannot be observed.
Pterygoid: One pterygoid, probably the right, can be
observed isolated from other palatal elements (Figure 4).
The marginal borders are elevated in relation to the centre
of the bone, suggesting that it is exposed in dorsal view.
A small process is located anteriorly in the medial margin
and, when in articulation, contacted the vomer. The rostral
process is an elongate lateral ramus that extends anteriorly,
forming a concavity that comprises the posterior margin of
the right choana. This process is slightly curved anteriorly
and a little divergent in relation to the rostromedial
process. There is no lateral process into the subtemporal
fenestra, as present in Anhanguera araripensis (Well-
nhofer 1985; Kellner and Tomida 2000). If a pterygoid
fenestra is present, it is not visible in dorsal view.
Mandible: The mandible is seen mostly in left lateral
view and lacks part of the middle portion (Figure 1). The
anterior part of the symphysis and both mandibular rami
can be observed. The left ramus is seen in left lateral view
and some patches of bone along its length were lost,
allowing the visualisation of the right ramus in medial
view underneath it. The posterior part of the left ramus is
preserved near some of the skull elements (jugal, quadrate)
and seen in lateral view. The posterior part of the right
ramus is displaced and shows a low and elongated
adductor fossa, which is located farther from the articular
than in anhanguerids and Pteranodon (Wellnhofer 1985,
1987,1991; Bennett 2001; Veldmeijer 2003).
Dentary: The dentaries constitute most of the lateral
surface of the mandible. The mandibular symphysis has a
straight dorsal margin and bears no crest, at least not on the
tip (Figure 5), as seen in crested pteranodontoids (e.g.
Wellnhofer 1987; Kellner and Tomida 2000; Veldmeijer
Figure 5. (Colour online) Tip of the mandible of specimen IVPP V15549, holotype of Linlongopterus jennyae gen. et sp. nov., in left
lateral view, showing both right and left side teeth. d, dentary; l, left; te, teeth. Scale bar: 50 mm.
786 T. Rodrigues et al.
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
2003; Wang and Zhou 2003; Kellner et al. 2013; Wang
et al. 2014). A portion of the dorsal part of the symphysis is
visible but no sulcus can be identified because of the poor
preservation.
Some teeth are preserved next to the mandibular
symphysis, suggesting the presence of seven pairs in the
preserved distal portion. Because of the absence of the
cortical bone on the alveolar margin, only four alveoli can
be seen on the left side, two of them with replacement teeth.
There is one tooth preserved between the mandibular rami,
but it is unclear if it was positioned inside its alveolus.
Surangular, angular and articular: Both surangulars
are present, located anteriorly to the articular and
comprising the dorsal margin of the adductor fossae,
with a thin and elongate anterior process (Figure 6). The
right angular is a thin and elongate bone and comprises the
ventral margin of the mandibular ramus in medial view
(Figure 6). Its sutures with the surrounding bones are
unclear. Only the left articular is preserved and the
articular surface has two paired depressions (Figure 4).
The retroarticular process is elongate but it cannot be
determined if it has a foramen, as seen in anhanguerids and
in Pteranodon (Wellnhofer 1985,1987,1991; Bennett
2001; Veldmeijer 2003).
Splenial and prearticular: The right splenial and
prearticular are present and comprise the ventral margin of
the adductor fossae (Figure 6). Sutures between these
bones cannot be discerned.
Dentition: The holotype of Linlongopterus jennyae has
several preserved teeth, most of them isolated but close to
their sockets. The first two pairs of lower teeth have gently
curved crowns and roots, but posteriorly the crowns are
slightly smaller and straighter (Figure 5). Crowns are
either smaller or about the same size as the roots; some of
the latter are a bit expanded. The enamel is dark-brown and
presents faint longitudinal ridges. There is limited
information on alveolar pattern, but the ones preserved
(on the tips of the rostrum and mandible) are about the
same size and equally spaced.
Pterosauria Kaup, 1834
Pterodactyloidea Plieninger, 1901
Pteranodontoidea Marsh, 1876,sensu Kellner, 2003
Anhangueria Rodrigues and Kellner, 2013
Anhangueridae Campos and Kellner, 1985
3.2 Liaoningopterus gui Wang and Zhou, 2003
Holotype. IVPP V13291, partial skull and mandible
preserved mostly in left lateral view, atlas and axis, and
one midcervical vertebra, deposited at the IVPP in Beijing,
China. The holotype is also the only known specimen.
Horizon and locality. Xiaoyugou, Lianhe, Chaoyang,
western Liaoning Province, China (Wang and Zhou 2003).
Jiufotang Formation, Aptian (He et al. 2004).
Revised comparative diagnosis. Anhanguerid pterosaur
diagnosed by the following combination of characters: low
and symmetric premaxillary crest, thus differing from
Anhanguera blittersdorffi,Anhanguera araripensis and
Anhanguera spielbergi; premaxillary crest begins near but
not at the tip of the skull, thus differing from
Tropeognathus mesembrinus; premaxillary crest ends far
from the anterior margin of the nasoantorbital fenestra,
differing from Anhanguera blittersdorffi,Anhanguera
araripensis,Anhanguera piscator and Anhanguera spiel-
bergi; dorsal margin of the rostrum posterior to the
premaxillary crest and anterior to the nasoantorbital
fenestra straight, differing from Anhanguera blittersdorffi,
Anhanguera araripensis,Anhanguera piscator,Anhan-
guera santanae and Anhanguera spielbergi.
Expanded description and comparison
Generalities: The specimen is flattened and incom-
plete. From the skull, there is the rostrum seen in left
lateral view, preserved until the first third or half of the
nasoantorbital fenestra, and both left and right jugals,
quadratojugals and quadrates are exposed in their lateral
views. The mandible is partially preserved, showing a
breakage that separates most of the symphysis from the
mandibular rami. The first three cervical vertebrae are also
Figure 6. (Colour online) Posterior part of the mandible of specimen IVPP V15549, holotype of Linlongopterus jennyae gen. et sp. nov.,
in left lateral view. afo, adductor fossa; ang, angular; par, prearticular; r, right; san, surangular; spl, splenial. Scale bar: 50 mm.
Historical Biology 787
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
present, besides a very flattened and eroded long bone,
which seems to be a humerus (Figure 7).
Premaxilla: Only the left premaxilla is visible, from
the tip of the skull until the nasoantorbital fenestra. The
preserved part shows that the dorsal margin of the rostral
portion of the skull was straight (Figure 8). Due to
compression, the palatal region is exposed, including part
of the ventral portion of the premaxilla. This portion
possesses a few small foramina (Figure 9(A)), which may
indicate innervation or vascularisation.
The tip of the rostrum has its ventral margin gently
curving upward. This feature is common among
pteranodontoids such as anhanguerians and Cimoliopterus
cuvieri (Bowerbank 1851), but is present in different
degrees in different taxa. In Liaoningopterus gui, this
gentle curvature is similar to that present in Anhanguera
Figure 8. (Colour online) General view of the anterior portion of the skull of specimen IVPP V13291, holotype of Liaoningopterus gui,in
left lateral view. naof, nasoantorbital fenestra; pm, premaxilla; pmcr, premaxillary crest; prf, prefrontal; prid, palatal ridge; te, teeth. Scale
bar: 50 mm.
Figure 7. (Colour online) General view of specimen IVPP V13291, holotype of Liaoningopterus gui. (A) Photograph; (B) respective
line drawing. cv, cervical vertebra; d, dentary; hu?, possible humerus; j, jugal; l, left; la, lacrimal; naof, nasoantorbital fenestra; pm,
premaxilla; prf, prefrontal; qj, quadratojugal; r, right; sq, squamosal. Scale bar: 50 mm.
788 T. Rodrigues et al.
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
and Caulkicephalus (Campos and Kellner 1985; Well-
nhofer 1991; Kellner and Tomida 2000; Steel et al. 2005),
and is very distinct from the steeper inclination seen in
Coloborhynchus,Siroccopteryx and Uktenadactylus
(Rodrigues and Kellner 2008).
There is a very low and symmetric premaxillary crest
(Figure 8), 120 mm long, and 17 mm maximum height
(Wang and Zhou 2003). It begins near, but not at, the tip of
the rostrum, at the level of the 5th tooth, and ends well
before the nasoantorbital fenestra, at the 12th tooth (Wang
and Zhou 2003). It contrasts with Tropeognathus, in which
the crest is symmetric but higher and begins at the tip of
the rostrum, and Anhanguera blittersdorffi and Anhan-
guera spielbergi, which have strongly asymmetric crests
(Campos and Kellner 1985; Veldmeijer 2003). Anhan-
guera piscator also has a low and symmetric crest, but it
ends nearer the nasoantorbital fenestra (Kellner and
Tomida 2000). The crest ends well before the nasoantor-
bital fenestra in Anhanguera santanae (based on the
holotype, SNSB-BSPG 1982 I 90; Wellnhofer 1985;
Kellner and Tomida 2000), but in this species its shape
cannot be assessed. In Caulkicephalus trimicrodon this
structure also begins near but not at the tip of the rostrum
and ends well before the nasoantorbital fenestra (Steel
et al. 2005), but its shape cannot be determined. In any
case, the premaxillae of Caulkicephalus trimicrodon show
a marked dorsal concavity that is absent in Liaoningop-
terus gui.
Being an anhanguerid, Lianingopterus gui probably
had a distal expansion of the skull, a structure that, as
exemplified by Anhanguera blittersdorffi and all other
species of the family, is located near the anterior end of the
rostrum, although it is not terminal. As the specimen is
laterally flat, an expansion is hard to be confirmed, but two
features make a strong case for its presence. One is that the
premaxillary alveolar margin is not straight, but rather
makes a gentle upward curve from alveoli 2 to 4, and then
descends until alveoli 5 and 6; the latter alveoli would thus
be located in a constriction that follows the expansion
(Figure 9(A)). Another feature is that, as typical of
anhanguerids, the alveoli 2 to 4 are quite large, which is
normally associated with the distal expansion, alveoli 3
being the largest and usually positioned in the broader part
of the rostral tip. Alveoli 5 and 6 are smaller, and, as
observed in non-flattened specimens, are associated with
the constriction that follows the expansion, after which the
Figure 9. (Colour online) Detail of several bones of specimen IVPP V13291, holotype of Liaoningopterus gui. (A) tip of the skull in left
lateral view. Scale bar: 10mm; (B) left jugal and associated bones in left lateral view. Scale bar: 50mm; (C) right jugal and associated
bones in right lateral view. Scale bar: 50mm. j, jugal; l, left; la, lacrimal; ltf, lower temporal fenestra; pmcr, premaxillary crest; q,
quadrate; qj, quadratojugal; r, right; sq, squamosal; te, teeth. Arrow points to a large substitution tooth.
Historical Biology 789
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
rostrum usually broadens again (see Campos and Kellner
1985; Steel et al. 2005).
Maxilla: The suture between premaxilla and maxilla is
largely obliterated, and therefore their limits are not
visible. Some cracks on the surface of the bone also
prevent the assessment if the putative suture line between
the premaxilla and maxilla in front of the nasoantorbital
fenestra is present, as observed in other anhanguerid skulls
(Campos and Kellner 1985; Wellnhofer 1985,1991).
Most the left maxilla can be seen, but parts of the right
one are also visible below the ventral margin of the skull.
The maxilla comprises most of the rostrum in lateral view
and its jugal process extends until under the nasoantorbital
fenestra, as showed by the presence of a toothed margin.
Lacrimal: Only part of the left lacrimal is visible, in
articulation with the lacrimal process of the left jugal. The
suture between those bones is clearly visible, and the
lacrimal overlies the jugal. It presents a small posterior
process directed into the orbit and shows the indication of
a lacrimal foramen.
Jugal: As remarked above, both jugals are preserved
but disarticulated from the skull (Figure 9(B),(C)). They
conform to the general morphology present in other
anhanguerids (Campos and Kellner 1985; Wellnhofer
1985,1991; Kellner and Tomida 2000; Veldmeijer 2003),
such as the presence of a well-marked ridge at the base of
the lacrimal process, which is robust and vertically
oriented (Wang and Zhou 2003).
The left jugal lacksthe maxillary process, but shows both
the lacrimal and postorbital processes (Figure 9(B)). The
postorbital process is robust and, together with the lacrimal
process, forms a rounded ventral margin of the orbit. Because
both processes are divergent, the orbit had the dorsal part
broader thanthe ventral, a common feature of anhanguerids.
The right jugal shows a very long and slender
maxillary process that gets thinner gradually, ending in a
sharp point (Figure 9(C)). The lacrimal process is broad,
and its dorsal part is rugose, marking the area where it was
overlain by the lacrimal. The external surface of the
postorbital process is damaged and the interior part of the
bone is exposed.
Quadratojugal: Only the left quadratojugal can be
observed (Figure 9(B)). It is exposed in lateral view and
fused to adjacent bones. It is elongate and participates in
the ventral border of the lower temporal fenestra forming a
slightly concave margin.
Quadrate: The right quadrate is very fragmented while
the left quadrate is well preserved but lacks the articulation
with the lower jaw (Figure 9(B),(C)). The ascending
process is elongate and, with the quadratojugal and
postorbital process of the jugal, forms the inferior margin
of the lower temporal fenestra.
Squamosal: The left squamosal is partially preserved
and is connected with the quadrate (Figure 9(B)). Its
posterior margin marks the rear end of the skull in lateral
view. The anterior process is partially preserved and forms
part of the intertemporal bar, which is longer than the
quadratojugal and postorbital process of the jugal, showing
that the lower temporal fenestra was broader dorsally than
ventrally. The ventrolateral process of the squamosal is
also elongated and oriented ventrally.
Prefrontal: An isolated bone located at the dorsopos-
terior portion of the preserved skull (Figure 7) is consistent
with the prefrontal, but the outer surface is not well
preserved.
Mandible: The mandible is broken in two pieces
(Figure 10). On the slab, the symphyseal portion is located
ventrally to the rami, which is unusual. The symphysis and
left ramus are exposed in left lateral view, while the right
ramus is shown in medial view. The curvature of the left
ramus, as seen in its dorsal and ventral margins, indicates
that they were bowed, as seen in tridimensionally
preserved anhanguerids (Wellnhofer 1985,1987; Kellner
and Tomida 2000; Veldmeijer 2003).
Dentary: The mandibular symphysis is not well
preserved. The tip of the left dentary has lost the external
surface exposing the internal part of the bone. Although
some teeth are present, no alveoli can be seen.
The preservation of this specimen hinders direct
observation of the tip of the dentaries, but it can be
confidently assumed that only a minor portion was lost, as
the preserved teeth are rather large, as typical from the
anterior-most portion of anhanguerid mandibles (see
Kellner and Tomida 2000).
The high tip of the mandible (45.4 mm at the highest
point), in comparison to the height of the rami (36.3 mm at the
highest point of the left ramus) and to the preserved caudal
part of the symphysis, shows that a median dentary crest was
present (Figure 10), although its exact shape cannot be
determined. In all known anhanguerid mandibles, a dentary
crest is present and is located at the anterior-most tip of the
symphysis, with variable shapes and heights (Wellnhofer
1987; Kellner and Tomida 2000; Veldmeijer 2003)
Surangular, angular and articular: In both the left and
right mandibular rami, the sutures between the surangular
and adjacent bones are not recognisable, but this bone can
be identified by its position, forming the dorsal margin of
the adductor fossa (Bennett 2001) which, in Liaoningop-
terus gui as in other anhanguerids, is long and narrow
(Wellnhofer 1985,1987; Veldmeijer 2003). The angular
can be observed in medial view, in the right ramus where it
forms part of the ventral margin of the mandible. It is
elongated and relatively thin. The suture with the splenial
is well marked. The right articular shows the articulation
surface with the upper jaw as paired fossae, which receive
the two condyles of the helical jaw joint. The retroarticular
process is fragmentary but was an elongated element.
Splenial and prearticular: The right splenial is a
laminar bone, located in the medial portion of the ramus.
It overlies dorsally the dentary and ventrally the angular,
790 T. Rodrigues et al.
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
forming the ventral margin of the adductor fossa in its
entire length.
Dentition:Liaoningopterus gui has a remarkable
dentition, with extremely long and robust upper anterior
teeth (Wang and Zhou 2003), much large in absolute size
than in any other anhanguerid. The third upper tooth is the
largest, just over 80 mm long (excluding the base, which
was not preserved), but its size seems to be nearly followed
by the second upper tooth, which was about to be shed (a
large replacement tooth can be seen). These teeth are also
quite thick in comparison to the others.
In the upper jaw, the anterior-most pairs of teeth are
located at the tip of the rostrum, which, combined with its
gently upward curvature, makes them procumbent. The
first pair of upper teeth is located slightly dorsally to the
second pair, with the space between them less than the size
of the alveolus. Upper alveoli 1 – 4 are larger, 6 is smaller
and 7– 10 relatively large, but not as much as the anterior-
most ones. Due to the poor preservation of the fifth
alveolus, it is uncertain if it was smaller than alveoli 4 and
7, a feature normally considered distinctive of the genus
Anhanguera (Kellner 2003). From the 10th onwards, the
alveoli are smaller and extend until under the nasoantor-
bital fenestra. The anterior eight pairs of teeth are more
densely placed, while the separation of the posterior teeth
is larger than the size of their alveoli.
The crowns are long and gently curved in the first 10
pairs of teeth, while the posterior ones are shorter and
somewhat conical. All teeth show enamel with longitudi-
nal ridges, as observed before in other specimens referred
to the Anhangueridae and related taxa (Rodrigues and
Kellner 2010).
The lower jaw is not well preserved and the alveoli
cannot be seen, so their location and separation cannot be
measured. Apparently only the left teeth are preserved.
Nonetheless, the overall pattern is also similar to the upper
jaw, with larger anterior teeth and smaller posterior ones.
Atlas and axis: Just next to the preserved third cervical
vertebra, a round articulation surface can be seen. This
articulation is interpreted as the anterior cotyle of the
intercentrum of the atlas (Figure 11). The atlas is fused to
the centrum of the axis, which is partially visible, the main
part crushed underneath the third cervical vertebra. A bone
fragment preserved on the other side of the third cervical,
opposed to the atlas, may be the neural spine of the axis.
Third cervical vertebra: One cervical vertebra from
the mid series, possibly cervical 3, is exposed in ventral
view (Figure 11). The centrum is short and a lateral
pneumatic foramen on the left side can be observed. The
prezygapophyses were not preserved, but the left
postzygapophysis and both postexapophyses are present.
The postzygapophysis is short and faces posteriorly. The
postexapophyses are horn-like and very divergent, likely
partially due to taphonomy.
Possible humerus: Lying dorsally to the main part of the
skull is a flattened bone, in which most of the external surface
was lost. It is elongate but broad. The margin preserved next
to the skull is concave, but the other margin was broken.
Figure 10. (Colour online) Mandible of specimen IVPP V13291, holotype of Liaoningopterus gui. The mandibular rami are preserved
above the symphysis. afo, adductor fossa; ang, angular; art, articular; d, dentary; dcr, dentary crest; r, right; rapr, retroarticular process; spl,
splenial; te, teeth. Scale bar: 50 mm.
Historical Biology 791
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
It also presents an articulation surface with apparently two
condyles. It is tentatively identified a humerus.
4. Discussion and conclusions
In order to assess the phylogenetic relationships of
Linlongopterus jennyae, a cladistic analysis was performed
and resulted in 90 equally parsimonious trees with 213 steps
each. From these, a strict consensus tree was obtained
(Figure 12). The species was recovered within the
Dsungaripteroidea (sensu Kellner 2003) based on an
unambiguous synapomorphy: the presence of a helical jaw
joint. However, the position within this group is less clear
because the inclusion of this taxon collapsed several nodes,
as the holotype and only known specimen of Linlongop-
terus jennyae is quite incomplete. It can be excluded from
the Dsungaripteridae, which possesses a very particular
dentition, and from the edentulous Azhdarchoidea, there-
fore, being referred to the Pteranodontoidea. The latter is
further divided into three lineages. It can be confidently
excluded from the edentulous Pteranodontidae and from the
Istiodactylidae, which shows very characteristic laterally
compressed and triangular tooth crowns (Kellner 2003).
The remaining clade includes the Anhangueria and its sister
groups (Rodrigues and Kellner 2013). The new specimen
has features consistent with this clade, being excluded from
the Anhangueridae due to the lack of median crests on the
upper and lower jaws, and by the possession of anterior
teeth that are similar in size, thus lacking the typical teeth
size variation seen in Anhanguera, for instance.
Linlongopterus jennyae can be easily distinguished
from the two known anhanguerians from the Jiufotang
Formation, Liaoningopterus gui and Guidraco venator,
and from the related form Ikrandraco avatar. To cite only
the most outstanding features, Liaoningopterus gui has
very enlarged upper anterior teeth, a premaxillary crest
that begins near the tip of the rostrum, and a dentary crest
(Wang and Zhou 2003), all features absent in Linlongop-
terus.Guidraco venator has very particular thin and
elongate upper anterior teeth (Wang et al. 2012) which are
also absent in the new species. Ikrandraco avatar has an
elongate skull, a large, blade-like mandibular crest, and
short teeth (Wang et al. 2014), features not seen in the new
taxon.
Regarding morphologically similar pterosaurs from the
older Yixian Formation, Haopterus gracilis has teeth that
are overall similar (Wang and Lu
¨2001), but have a
constriction between crown and root (Figure 13), thus
differing from the present species. Comparisons with
Yixianopterus jingangshanensis, regarded as a ‘loncho-
dectid’ (Lu
¨et al. 2006), are complicated because the
holotype and only known specimen of this taxon was heavily
artificially modified, with several fake bones added (as
shown in its description) and it cannot be determined if the
few original teeth are from the upper or the lower jaw.
Concerning Liaoningopterus gui, different phyloge-
nies have indicated slightly different relationships with the
Brazilian anhanguerids, e.g. Anhanguera and Tropeog-
nathus (Lu
¨and Ji 2006; Andres et al. 2014). In the case of
the analysis by Lu
¨and Ji (2006), Liaoningopterus was
Figure 11. (Colour online) Cervical vertebral elements of specimen IVPP V13291, holotype of Liaoningopterus gui: ax, axis; fopn,
foramen pneumaticum; ic, intercentrum; poex, postexapophysis; poz, postzygapophysis. Scale bar: 50 mm.
792 T. Rodrigues et al.
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
recovered as the sister-group of Anhanguera and
Tropeognathus, and thus actually outside the Anhanguer-
idae as defined by Kellner (2003). However, as noted
before (Rodrigues and Kellner 2013), the coding
responsible for this phylogenetic position is incorrect,
and instead should be identical to the coding for the
species of Anhanguera. A more recent phylogenetic
proposal for pterosaurs (Andres et al. 2014) recovered
Liaoningopterus as the sister-group of Anhanguera,with
Tropeognathus being closer to other taxa such as
Ornithocheirus and Coloborhynchus. Although a review
of this phylogenetic analysis is beyond the scope of this
paper, we note that there are problems in the coding too.
For instance, the authors consider the absence or presence
of a dentary crest (character 133) as a missing data, with
which we disagree. Furthermore, they have coded one
character (55; orbit, dorsal position) as multistate in
Liaoningopterus, even though the taxon is only known by
its holotype (a fact acknowledged in their list of species
and specimens used in the phylogenetic analysis, provided
as supplementary material). A recent phylogenetic
proposal focusing on anhanguerians and related taxa
(Rodrigues and Kellner 2013) recovered Liaoningopterus
as one of the several wildcard taxa that brought instability
to the analysis; this was mainly due to the amount of
missing data in characters key to the group. In the present
analysis, Liaoningopterus gui was recovered in a polytomy
with Cearadactylus atrox,Caulkicephalus trimicrodon,
Tropeognathus mesembrinus and Anhanguera.
Liaoningopterus gui is referred to the Anhangueridae
by the possession of blade-like premaxillary and dentary
crests which are confined to the anterior portions of the
Figure 12. Phylogenetic relationships of Linlongopterus jennyae gen. et sp. nov. and Liaoningopterus gui (see Supplementary Material).
Historical Biology 793
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
skull and mandible (Wang and Zhou 2003). It is
remarkably similar to the species of Anhanguera and to
Caulkicephalus trimicrodon. Although it cannot be
assessed if Liaoningopterus gui hadanenlarged
frontoparietal crest, it can be distinguished from Caulk-
icephalus trimicrodon because it lacks the characteristic
small 5th, 6th and 7th alveoli of the latter (Steel et al.
2005). It also differs from most species of Anhanguera by
the combination of a low and symmetric crest located
comparatively far from the nasoantorbital fenestra
(Kellner and Tomida 2000). It can be distinguished from
Anhanguera santanae by having a straight dorsal margin
of the skull, which in the latter is concave (Wellnhofer
1985). Besides the anatomical features, Liaoningopterus
and Anhanguera come from different deposits, which were
quite separated spatially in the Lower Cretaceous.
To date, the Jiufotang Formation has shown a variety
of pterosaurs, with Liaoningopterus gui,Linlongopterus
jennyae and Guidraco venator representing large species.
They were collected in distinct geographic areas, but in the
same stratigraphic unit. However, it is difficult to be sure if
these taxa were indeed contemporaneous and sympatric
species. If they were, those pterosaurs show very distinct
tooth morphologies. Although only in Guidraco, there is
some indirect evidence suggesting that it preyed on fish
(Wang et al. 2012), all three are regarded to be piscivorous,
likely a general condition among anhanguerians and
closely related taxa. This variation in dentition points to
possible different prey item preferences and/or perhaps
also diverse feeding habits among pterosaurs from this
deposit.
Acknowledgements
We thank Yutong Li for the preparation, and Wei Gao for the
photographs of the specimen IVPP V15549. TNT is provided by the
Willi Hennig Society. We also thank Oliver Rauhut for access to
specimens under his care, and Felipe Pinheiro and an anonymous
reviewer for comments that greatly improved the manuscript.
Disclosure statement
No potential conflict of interest was reported by the authors.
Funding
This study was supported by the Hundred Talents Project of the
Chinese Academy of Sciences, the National Key Basic Research
Program of China [grant number 2012CB821900], and the
National Science Fund for Distinguished Young Scholars [grant
number 40825005]. T. Rodrigues received funding from the
Conselho Nacional de Desenvolvimento Cientı
´fico e Tecnolo
´-
gico/Fundaca
˜o de Amparo a
`Pesquisa e Inovaca
˜o do Espı
´rito
Santo [grant number 52986870/2011] and Conselho Nacional de
Desenvolvimento Cientı
´fico e Tecnolo
´gico [grant number
460784/2014-5], and A.W.A. Kellner acknowledges funding
from the Fundaca
˜o Carlos Chagas Filho de Amparo a
`Pesquisa do
Rio de Janeiro [grant number E-26/102.737/2012] and the
Conselho Nacional de Desenvolvimento Cientı
´fico e Tecnolo
´-
gico [grant number 304780/2013-8].
References
Andres B, Myers TS. 2013. Lone Star pterosaurs. Earth Environ Sci Trans
R Soc Edinb. 103:383–398.
Andres B, Clark J, Xu X. 2014. The earliest pterodactyloid and the origin
of the group. Curr Biol. 24(9):1011 – 1016. doi:10.1016/j.cub.2014.
03.030.
Figure 13. (Colour online) Teeth of specimen IVPP V11726, holotype of Haopterus gracilis: d, dentary; l, left; pm, premaxilla; r, right.
Arrows point to the constriction between crown and root of the teeth. Scale bar: 10 mm.
794 T. Rodrigues et al.
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
Andres B, Ji Q. 2008. A new pterosaur from the Liaoning province of
China, the phylogeny of the Pterodactyloidea, and convergence in
their cervical vertebrae. Palaeontology. 51(2):453–469. doi:10.
1111/j.1475-4983.2008.00761.x.
Bennett SC. 1993. The ontogeny of Pteranodon and other pterosaurs.
Paleobiology. 19:92–106.
Bennett SC. 2001. The osteology and functional morphology of the Late
Cretaceous pterosaur Pteranodon. Palaeontogr Abt A. 260:1– 112.
Bowerbank JS. 1851. On the pterodactyles of the Chalk Formation. Proc
Zool Soc Lond. 19(1):14–20. doi:10.1111/j.1096-3642.1851.
tb01125.x.
Campos DA, Kellner AWA. 1985. Panorama of the flying reptiles study
in Brazil and South America. An Acad Bras Cienc. 57(4):453 – 466.
Dong Z-M, Sun Y-W, Wu S-Y. 2003. On a new pterosaur from the Lower
Cretaceous of Chaoyang Basin, Western Liaoning, China. Global
Geol. 22:1–7.
Goloboff PA, Farris JS, Nixon KC. 2008. TNT, a free program for
phylogenetic analysis. Cladistics. 24(5):774–786. doi:10.1111/j.
1096-0031.2008.00217.x.
He HY, Wang XL, Zhou ZH, Wang F, Boven A, Shi GH, Zhu RX. 2004.
Timing of the Jiufotang Formation (Jehol Group) in Liaoning,
northeastern China, and its implications. Geophys Res Lett. 31(12):
L12605. doi:10.1029/2004GL019790.
Kaup JJ. 1834. Versuch einer Eintheilung der Saugethiere in 6 Sta
¨mme
und der Amphibien in 6 Ordnungen [An attempt to arrange mammals
in 6 lineages and amphibians in 6 orders]. Isis. 3:311– 315.
Kellner AWA. 2003. Pterosaur phylogeny and comments on the
evolutionary history of the group. In: Buffetaut E, Mazin J-M,
editors. Evolution and palaeobiology of pterosaurs. London:
Geological Society of London; p. 105 – 137.
Kellner AWA. 2013. A new unusual tapejarid (Pterosauria, Pterodacty-
loidea) from the Early Cretaceous Romualdo Formation, Araripe
Basin, Brazil. Earth Environ Sci Trans R Soc Edinb. 103:1 – 13.
Kellner AWA, Campos DA. 2007. Short note on the ingroup relationships
of the Tapejaridae (Pterosauria, Pterodactyloidea). Bol Mus Nac N S
Geol. 75:1–14.
Kellner AWA, Campos DA, Saya
˜o JM, Saraiva AAF, Rodrigues T,
Oliveira G, Cruz LA, Costa FR, Silva HP, Ferreira JS. 2013. The
largest flying reptile from Gondwana: a new specimen of
Tropeognathus cf. T. mesembrinus Wellnhofer, 1987 (Pterodacty-
loidea, Anhangueridae) and other large pterosaurs from the
Romualdo Formation, Lower Cretaceous, Brazil. An Acad Bras
Cienc. 85(1):113– 135. doi:10.1590/S0001-37652013000100009.
Kellner AWA, Tomida Y. 2000. Description of a new species of
Anhangueridae (Pterodactyloidea) with comments on the pterosaur
fauna from the Santana Formation (Aptian-Albian), northeastern
Brazil. Natl Sci Mus Monogr. 17:1 – 135.
Lu
¨J, Ji Q. 2006. Preliminary results of a phylogenetic analysis of the
pterosaurs from western Liaoning and surrounding areas. J Paleontol.
Soc Korea. 22(1):239–261.
Lu
¨J, Ji S, Yuan C, Gao Y, Sun Z, Ji Q. 2006. New pterodactyloid
pterosaur from the Lower Cretaceous Yixian Formation of western
Liaoning. In: Lu
¨J, Kobayashi Y, Huang D, Lee Y-N, editors. Papers
from the 2005 Heyuan International Dinosaur Symposium. Beijing:
Geological Publishing House; p. 195 – 203.
Lu
¨J, Unwin DM, Xu L, Zhang X. 2008. A new azhdarchoid pterosaur
from the Lower Cretaceous of China and its implications for
pterosaur phylogeny and evolution. Naturwissenschaften. 95:
891–897.
Lu
¨J, Xu L, Ji Q. 2008. Restudy of Liaoxipterus (Istiodactylidae:
Pterosauria), with comments on the Chinese istiodactylid pterosaurs.
Zitteliana Reihe B. 28:229–241.
Marsh OC. 1876. Notice of a new sub-order of Pterosauria. Am J Sci Arts.
s3–11(66):507– 509. doi:10.2475/ajs.s3-11.66.507.
O
˝si A, Prondvai E, Frey E, Pohl B. 2010. New interpretation of the palate
of pterosaurs. Anat Rec. 293(2):243–258. doi:10.1002/ar.21053.
Pinheiro FL, Schultz CL. 2012. An unusual pterosaur specimen
(Pterodactyloidea, ?Azhdarchoidea) from the Early Cretaceous
Romualdo Formation of Brazil, and the evolution of the
pterodactyloid palate. PLoS ONE. 7(11):e50088. doi:10.1371/
journal.pone.0050088.
Plieninger F. 1901. Beitra
¨ge zur Kenntnis der Flugsaurier [Contributions
to the knowledge of pterosaurs]. Palaeontogr. 48:65 – 90.
Rodrigues T, Kellner AWA. 2008. Review of the pterodactyloid
pterosaur Coloborhynchus. Zitteliana Reihe B. 28:219– 228.
Rodrigues T, Kellner AWA. 2010. Note on the pterosaur material
described by Woodward from the Reco
ˆncavo Basin, Lower
Cretaceous, Brazil. Rev Bras Paleontol. 13(2):159 – 164. doi:10.
4072/rbp.2010.2.08.
Rodrigues T, Kellner AWA. 2013. Taxonomic review of the
Ornithocheirus complex (Pterosauria) from the Cretaceous of
England. ZooKeys. 308:1– 112. doi:10.3897/zookeys.308.5559.
Steel L, Martill DM, Unwin DM, Winch JD. 2005. A new pterodactyloid
pterosaur from the Wessex Formation (Lower Cretaceous) of the Isle
of Wight, England. Cretac Res. 26(4):686 – 698. doi:10.1016/j.
cretres.2005.03.005.
Veldmeijer AJ. 2003. Description of Coloborhynchus spielbergi sp. nov.
(Pterodactyloidea) from the Albian (Lower Cretaceous) of Brazil.
Scr Geol. 125:35–139.
Wang X, Campos DA, Zhou Z, Kellner AWA. 2008. A primitive
istiodactylid pterosaur (Pterodactyloidea) from the Jiufotang
Formation (Early Cretaceous), northeast China. Zootaxa. 1813:
1–18.
Wang X, Kellner AWA, Jiang S, Cheng X. 2012. New toothed flying
reptile from Asia: close similarities between early Cretaceous
pterosaur faunas from China and Brazil. Naturwissenschaften. 99(4):
249– 257. doi:10.1007/s00114-012-0889-1.
Wang X, Kellner AWA, Zhou Z, Campos DA. 2008. Discovery of a rare
arboreal forest-dwelling flying reptile (Pterosauria, Pterodactyloi-
dea) from China. Proc Natl Acad Sci USA. 105(6):1983 – 1987. doi:
10.1073/pnas.0707728105.
Wang X, Rodrigues T, Jiang S, Cheng X, Kellner AWA. 2014. An Early
Cretaceous pterosaur with an unusual mandibular crest from China
and a potential novel feeding strategy. Sci Rep. 4:6329. doi:10.1038/
srep06329.
Wang X, Zhou Z. 2002. A new pterosaur (Pterodactyloidea, Tapejaridae)
from the Early Cretaceous Jiufotang Formation of western Liaoning,
China and its implications for biostratigraphy. Chin Sci Bull. 47:
1521– 1528.
Wang X, Zhou Z. 2003. Two new pterodactyloid pterosaurs from the
Early Cretaceous Jiufotang Formation of western Liaoning, China.
Vertebrat Palasiatic. 41(1):34–41.
Wang X, Lu
¨J. 2001. Discovery of a pterodactylid pterosaur from the
Yixian Formation of western Liaoning, China. Chin Sci Bull. 46(13):
1112– 1117.
Wellnhofer P. 1978. Pterosauria. Handbuch der Pala
¨oherpetologie, Teil
19 [Pterosauria. Encyclopedia of Paleoherpetology, Part 19].
Stuttgart and New York: Gustav Fischer Verlag.
Wellnhofer P. 1985. Neue Pterosaurier aus der Santana-Formation (Apt)
der Chapada do Araripe, Brasilien [New pterosaurs from the Santana
Formation (Aptian) of the Chapada do Araripe, Brazil]. Palaeontogr
Abt A. 187:105–182.
Wellnhofer P. 1987. New crested pterosaurs from the lower Cretaceous of
Brazil. Mitt Bayer Staatsslg Pala
¨ontol Hist Geol. 27:175–186.
Wellnhofer P. 1991. Weitere Pterosaurierfunde aus der Santana-
Formation (Apt) der Chapada do Araripe, Brasilien [Additional
pterosaur remains from the Santana Formation (Aptian) of the
Chapada do Araripe, Brazil]. Palaeontogr Abt A. 215:43 – 101.
Historical Biology 795
Downloaded by [Instituto de Economia - UFRJ] at 07:58 08 July 2015
