A New Teiid Lizard From The Late Cretaceous of The Haţeg Basin, Romania and Its Phylogenetic and Palaeobiogeographical Relationships

This article was downloaded by: [New York University]
On: 27 April 2015, At: 12:58

Publisher: Taylor & Francis
Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House,
37-41 Mortimer Street, London W1T 3JH, UK
Journal of Systematic Palaeontology

Publication details, including instructions for authors and subscription information:
http://www.tandfonline.com/loi/tjsp20
A new teiid lizard from the Late Cretaceous of the

Haeg Basin, Romania and its phylogenetic and
palaeobiogeographical relationships
a
Mrton Venczel & Vlad A. Codrea

a
Department of Natural History, rii Criurilor Museum, Oradea, Romania
Department of Geology and Palaeontology, Babe-Bolyai University, Cluj-Napoca, Romania

Published online: 27 Apr 2015.
Click for updates

To cite this article: Mrton Venczel & Vlad A. Codrea (2015): A new teiid lizard from the Late Cretaceous of the Haeg
Basin, Romania and its phylogenetic and palaeobiogeographical relationships, Journal of Systematic Palaeontology, DOI:
10.1080/14772019.2015.1025869
To link to this article: http://dx.doi.org/10.1080/14772019.2015.1025869
PLEASE SCROLL DOWN FOR ARTICLE

Taylor & Francis makes every effort to ensure the accuracy of all the information (the Content) contained
in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no
representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the
Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and
are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and
should be independently verified with primary sources of information. Taylor and Francis shall not be liable for
any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever
or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of
the Content.
This article may be used for research, teaching, and private study purposes. Any substantial or systematic
reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any
form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http://
www.tandfonline.com/page/terms-and-conditions
Journal of Systematic Palaeontology, 2015

http://dx.doi.org/10.1080/14772019.2015.1025869
A new teiid lizard from the Late Cretaceous of the Hat eg Basin, Romania and its
phylogenetic and palaeobiogeographical relationships
Marton Venczela* and Vlad A. Codreab
Department of Natural History, T
arii Crisurilor Museum, Oradea, Romania; bDepartment of Geology and Palaeontology,
Babes-Bolyai University, Cluj-Napoca, Romania
Downloaded by [New York University] at 12:58 27 April 2015
(Received 2 March 2014; accepted 23 January 2015)

A new lizard genus and species is described based on a three-dimensionally preserved partial skull and associated lower
jaws from the Pui Islaz locality (Late Cretaceous, early Maastrichtian) in the Hat eg Basin, western Romania. Barbatteius
vremiri gen. et sp. nov. is diagnosed by a unique combination of symplesiomorphies and synapomorphies. A nested set of
synapomorphies support assigning Barbatteius to Teiidae as the first unambiguous Late Cretaceous record of this family
from Laurasia. Barbatteius differs from other teiids by having more extensive osteodermal sculpture on the skull roof and
suspensorium, and by a pentagonal occipital osteoscute exhibiting more or less parallel lateral margins. Barbatteius is a
large-bodied lizard, estimated to be up to 800 mm in total length. It has weakly heterodont dentition, but without enlarged
posterior crushing teeth, suggesting that it fed on arthropods, small vertebrates and plants. The mix of taxa with affinities to
Euramerica (paramacellodid and borioteiioid lizards) and Gondwana (madtsoiid snakes and the teiid Barbatteius) currently
known for the Maastrichtian squamate assemblage from Hat eg Basin supports the growing realization that Hat eg Island
has a complex palaeobiogeographical history.
http://zoobank.org/urn:lsid:zoobank.org:pub:75C2D80F-8DDB-4FB2-9844-1552D626F63D
Keywords: Teiidae; taxonomy; Squamata; Gondwana; Maastrichtian; Europe
Introduction
Squamata are a monophyletic group of reptiles containing
more than 9800 living taxa of lizards, amphisbaenians and
snakes (Uetz 2013). Calibrated molecular dating analyses
(see e.g. Jones et al. 2013 and references cited therein)
place the origin of crown-group Squamata in the Early
Jurassic (213.2 176 Ma), around the breakup of the supercontinent Pangaea. Unequivocal squamate fossils are
known from the Early Middle Jurassic onwards (Evans et
al. 2002; Evans 2003). The origin and emergence of most
major squamate crown-groups may be placed later in the
Cretaceous (Jones et al. 2013), perhaps as an adaptive
response to circumstances of warm global climate, major
continental fragmentation and considerable alteration of terrestrial biota. The available Upper Cretaceous record shows
a disproportionate global distribution of squamates: lizards
are distributed mostly on northern (i.e. Laurasia) continents
(Gao & Norell 2000; Eaton & Kirkland 2003; Nydam &
Voci 2007; Nydam et al. 2010; Makadi 2013a, b), whereas
snakes were more common on southern (i.e. Gondwanan)
landmasses (Rage & Werner 1999; Rage et al. 2004; de la
Fuente et al. 2007; Cavin et al. 2010).
*Corresponding author. Email: mvenczel@gmail.com

The Trustees of the Natural History Museum, London 2015. All Rights Reserved.
The Late Cretaceous tetrapod assemblages of Europe

evolved throughout the Cenomanian early Campanian in
conditions of high eustatic levels (Golonka & Kiessling
2002), which transformed the European continent into an
archipelago. A good example of the resultant island faunas is represented by the peculiar latest Cretaceous vertebrate fauna of Hat eg Island, western Romania, which
was first reported by Nopcsa (1905). Investigations of
Benton et al. (2010) have shown that at least some of the
Hat eg dinosaurs (e.g. the herbivorous genera like Magyarosaurus, Telmatosaurus and possibly Zalmoxes) were of
much smaller size than their close relatives from Asia and
North America, thus demonstrating the effects of insular
dwarfing. Recent reports also have revealed a much higher
diversity of vertebrates by adding new groups to those
already known in Nopcsas times (e.g. turtles, crocodilians, dinosaurs and pterosaurs), including fishes (Grigorescu et al. 1999), lissamphibians (albanerpetontids and
frogs) (Grigorescu et al. 1999; Duffaud 2000; Venczel &
Csiki 2003; Folie & Codrea 2005; Codrea et al. 2010),
squamate reptiles (Grigorescu et al. 1999; Folie & Codrea
2005; Codrea et al. 2010; Vasile et al. 2013), birds (Wang
et al. 2011a, b) and multituberculate mammals
Marton Venczel and Vlad A. Codrea
(Grigorescu & Hahn 1987; Radulescu & Samson 1996,

1997; Csiki et al. 2005; Codrea et al. 2010).
Research on squamate reptiles from the Hat eg Basin
started with reports by Grigorescu et al. (1999) of isolated
bones belonging to indeterminate anguimorph and scincomorph lizards. Folie & Codrea (2005) documented from
the Pui Islaz locality isolated bones of lizards assigned to
Paramacellodidae (Becklesius nopcsai, Folie & Codrea
2005, B. cf. hoffstetteri), Polyglyphanodontidae (Bicuspidon hatzegiensis, Folie & Codrea 2005) and to indeterminate lizards. These initial reports already revealed an
intriguing mix of Laurasian and Gondwanan squamate
groups. Paramacellodids, apparently related to cordylids
(Hoffstetter 1967; Estes 1983), ranged from the Jurassic
to Late Cretaceous of Europe and North America (Evans
1996, 2003; Evans & Chure 1998a, b, 1999), and their
date of origin probably preceded the fragmentation of
Pangaea (Evans 2003). Similarly, the polyglyphanodontid
Bicuspidon demonstrates close palaeobiogeographical
relationships to taxa previously described from the Late
Cretaceous of North America (Nydam 1999, 2002, 2013;
Nydam & Cifelli 2002, 2005; Nydam et al. 2007, 2010)
and from the Santonian of Ihark
ut, Hungary (e.g. Bicuspidon aff. hatzegiensis; Makadi 2006). Another novelty for
the Hat eg fauna is the occurrence of Madtsoiidae, a basal
clade of alethinophidian snakes of Gondwanan origin
(Rage 1981; Werner & Rage 1994; Rage & Werner 1999;
LaDuke et al. 2010), which expands the known European
distribution of this group from the Campanian of Spain
(Rage 1996, 1999) to the Maastrichtian of Romania (Folie
& Codrea 2005; Vasile et al. 2013).
Here, we describe a new lizard taxon on the basis of a
three-dimensionally preserved partial skull and associated
lower jaws that display diagnostic characters of Teiidae.
The specimen was originally enclosed in a compact piece
of sedimentary rock of about 6 7 cm diameter and was
discovered by the geologist Matyas Vremir during a field
inspection at the Pui Islaz locality (Fig. 1), in uppermost
Cretaceous (lower Maastrichtian) rocks in the river-bed of
Barbat stream in the Hat eg Basin, Romania. Discernible
elements exposed on the outer surface of the block were
the labial side of the left lower jaw with its posterior part
broken off, the posteroventral part of the neurocranium
and part of the palatal complex, and the ventral side of the
posterior portion of the fused frontals. Unfortunately, the
anterior part of the skull and the postcoronoid part of the
left lower jaw were missing. In the present paper we: (1)
diagnose the teiid lizard from the Pui Islaz locality and
assign it to a new genus and species of Teiidae; (2)
describe and compare the known elements of this new
teiid with those of other relevant lizard groups; (3) evaluate the phylogenetic relationships of the new taxon; and
(4) comment on its palaeogeographical and palaeoenvironmental implications.
Geological setting and age

Stable isotope studies (Melinte-Dobrinescu & Bojar 2010)
indicate that marine sedimentation in the Hat eg Basin
closed around the latest Campanian and that continental
deposition started in the earliest Maastrichtian at the latest. The continental vertebrate-bearing strata of the Hat eg
Basin are separated into two distinct lithostratigraphical
units, the Densus -Ciula and the Sânpetru formations
(Grigorescu 1992). The Pui beds, from where the specimen originates, are correlated with the lower part of the
Sânpetru Formation (Nopcsa 1905; Grigorescu et al.
1985, 1999). Palaeomagnetic studies of Panaiotu & Panaiotu (2010) indicate that the continental sequence along
the Sibis el valley (Sânpetru Formation) was deposited
between chron 32n.1 and the end of chron 31n (about 72
to 67.8 Ma). Although the palaeoenvironment of the
Sânpetru Formation typically is represented by hydromorphic (immature) palaeosols dominated by areas of
impeded drainage, the Pui beds consist of mature palaeosols dominated by moderately to well-drained floodplains
(Therrien 2005). The strata of the Pui beds are subhorizontal, composed of coarse-grained channel deposits and
red, fine-grained overbank deposits; the latter were
formed during flood events (Van Itterbeeck et al. 2004).
Between inundations, palaeosol development is documented by red coloration and the presence of calcrete nodules or continuous calcrete layers (Van Itterbeeck et al.
2004; Csiki et al. 2005; Therrien 2005). Occasional slickensides, red mottles, and more frequent drab-haloed trace
roots and burrows, were reported from these calcareous
palaeosols by Therrien (2005). Below the calcrete horizon
a dinosaur bone accumulation has been found by Van
Itterbeeck et al. (2004), consisting of a titanosaurid
humerus and about 10 connected vertebrae. Above the
calcrete horizon, the red silts contain abundant operculae
of cyclophorid gastropods (Pana et al. 2001). The microvertebrates in this horizon are represented by albanerpetontids, anurans, lizards, madtsoiid snakes (Folie &
Codrea 2005) and multituberculate mammals (Grigorescu
et al. 1985; Smith & Codrea 2003; Van Itterbeeck et al.
2004; Csiki et al. 2005). The new lizard specimen
described below was also collected from this level.
Material and methods

The fossil material reported here consists of an anteriorly
incomplete, three-dimensional skull consisting of articulated and slightly displaced bones in the skull roof, suspensorium, neurocranium, palatal complex and lower
jaws. The piece of sedimentary rock that enclosed the
specimen was prepared in the Laboratory of Vertebrate
Palaeontology of the Babes -Bolyai University. Following
New teiid lizard from the Late Cretaceous of the Hat eg Basin
Figure 1. Geological map of the southern part of the Hat eg Basin and location of the Pui Islaz fossil locality.
sediment removal, the skull and lower jaws were detached

in three parts (see Figs 2, 5, 6) for detailed morphological
examination. The photographs in Figures 2 6 were taken
at the T
arii Cris urilor Museum, Oradea, Romania, using a
Canon EOS 5D Mark III digital camera equipped with a
Carl Zeiss 100 mm f/2 macro lens. Parsimony analyses
were conducted with the phylogenetic software package
TNT version 1.1 (Goloboff et al. 2008). Common English
terms and the standard anatomical orientation system are
used throughout this paper; the anatomical nomenclature
of lizards follows Rieppel (1985) and Gauthier et al.
(2012).
The fossil skull and lower jaws described herein belong
to the collections of Babes -Bolyai University (UBB),
Cluj-Napoca, Romania. Comparative materials of Recent
tegu lizards used in this study belong to Zoologisches
Forschungsmuseum Alexander Koenig (ZFMK), Bonn,
Germany.
Systematic palaeontology
Order Squamata Oppel, 1811
Suborder Lacertiformes Estes, de Queiroz &
Gauthier, 1988
Superfamily Teiioidea Estes, de Queiroz &
Gauthier, 1988
Family Teiidae Gray, 1827
Genus Barbatteius gen. nov.
Type species. Barbatteius vremiri gen. et sp. nov.

Diagnosis. As for the type and only known species.
Derivation of name. After Barbat river in the Hat eg
Basin, which transects the Pui beds that yielded the holotype specimen, and the suffix teius, a genus name of
tegu lizard, suggesting the close relationships to Teiidae.
Barbatteius vremiri sp. nov.
(Figs 2 6)
Holotype. UBB V.440, a three-dimensionally preserved,
partial skull consisting of skull roofing bones, neurocranium, posterior part of the palatal complex and associated
fragmentary lower jaws.
Diagnosis. Large Late Cretaceous teiid lizard with estimated total body length up to 800 mm. It differs from all
other lizards by the following unique combination of features: upper temporal fenestra is not occluded by the postorbital; extensive osteodermal sculpture covers the skull
roof and suspensorium; frontals fused with well-marked
interorbital constriction; parietal ventral lappet forms a
prominent V-shaped, flat process; postorbital overlaps
squamosal dorsally; squamosal ascending process is present; epipterygoid parietal contact overlaps parietal temporal muscle origin; prootic forms part of medial aperture
of the recessus scalae tympani; the dentary has a weakly
developed subdental shelf (D subdental lamina); dentary
Figure 2. Partial skull roofing bones and suspensorium in the holotype (UBB V.440) of Barbatteius vremiri gen. et sp. nov. Frontoparietal region and supratemporal arch in A, dorsal and B, ventral views. Photographs (above) and interpretive figures using different levels of
grey to highlight particular bones (below). Abbreviations: adt: anterodorsal tuberosity; app: alar process of prootic; ccf: crista cranii
frontalis; ept: epipterygoid; fr: frontal; ipvl: imprint of parietal ventral lappet; ju: jugal; os: occipital scute; pa: parietal; pffp: frontal process of postfrontal; pfpo: postfrontal postorbital; pfpp: parietal process of postfrontal; por: posterior ramus of postfrontal postorbital;
ps: parietal scute; psp: supratemporal process of parietal; sq: squamosal; sqap: ascending process of squamosal; st: supratemporal. Scale
bar D 5 mm.
teeth are heterodont, part of them with bi- or tricuspid

crowns and tooth replacement being present in all dentary
teeth; anterolateral dentary process on coronoid overlaps
dentary past level of tooth row; angular process on dentary terminates anterior to coronoid apex. Differs from all
other teiids and the possible teiid Meyasaurus (Early
Cretaceous, Spain) in having extensive osteodermal crust

that strongly fuses to the skull roof and suspensorium,
and the outer surface of osteodermal sculpture also bearing the impressions of cephalic scales. Differs further
from the possible crown lacertids Succinilacerta (middle
Eocene, Poland and Lithuania) and Plesiolacerta (middle
Figure 3. Skull roofing bones and suspensorium in the holotype (UBB V.440) of Barbatteius vremiri gen. et sp. nov., and Recent
Ameiva ameiva (Teiidae). A, frontoparietal region with parietal downgrowth of Barbatteius in left lateral view. B, reconstruction of Barbatteius skull in dorsal view (missing part is Recent Tupinambis teguixin ZFMK 53531). C, partial skull of Ameiva ameiva (ZFMK
59021) in dorsal view with details of osteodermal scutes covering the frontoparietal region. D, interpretive drawings of osteodermal
scutes of the frontoparietal region of Barbatteius and supratemporal arch in dorsal view. Abbreviations: app: alar process of prootic; fr:
frontal; fs: frontal scute; fps: frontoparietal scute; ept: epipterygoid; ips: interparietal scute; ju: jugal; os: occipital scute; pa: parietal; pd:
parietal downgrowths; popf: postorbital postfrontal; ps: parietal scute; pvl: parietal ventral lappet; sq: squamosal; st: supratemporal.
Not to scale.
Eocene late Oligocene, France and Germany) in having a

narrow and pentagonal occipital scute with more or less
parallel lateral margins.
Derivation of name. To acknowledge Matyas Vremir,
geologist from Cluj-Napoca, Romania, who collected the
holotype specimen at the Pui Islaz locality.
Occurrence. Pui Islaz locality, approximately 500 m
south of Pui village, Hat eg Basin, Transylvania, Romania,
Europe; Late Cretaceous, early Maastrichtian, Sânpetru
Formation.
Description
General observations. It is assumed that the fragmentary skull and the lower jaws belonged to the same individual, because they were preserved in contact, although
the jaws had shifted slightly posteriorly and rotated
90 100 counter-clockwise from their original anatomical position. Also anatomical structures of the skull and
lower jaws fit in the same dimensional range and represent
the same taxonomic group. Considering the robustness
and ontogenic fusion of bones, especially those in the
occipital region, the skull corresponds to an adult and
rather old individual.
Figure 4. Partial skull roofing bones and suspensorium in the holotype (UBB V.440) of Barbatteius vremiri gen. et sp. nov. A, supratemporal processes of parietal, supratemporals and squamosals in posterior view; B, supratemporal arch and partial frontal in right lateral
view. Photographs (above) and interpretive figures using different levels of grey to highlight particular bones (below). Abbreviations: fr:
frontal; pa: parietal; popf: postorbital postfrontal; psp: supratemporal process of parietal; sq: squamosal; sqap: ascending process of
squamosal; st: supratemporal. Scale bar D 5 mm.
Frontal. Only the posterior part of the extensively sculptured, azygous frontal is preserved. The preserved part is
strongly constricted between the orbits and its interorbital
width is approximately half the frontoparietal suture
length. In dorsal view, there are several fracture lines and
parts of the sculpted surface are missing. The remaining
dorsal surface is covered by a sculpted osteodermal crust
left by impressions of the epidermal scutes (Smith 2009).
This surface consists of pits and anastomosing grooves,
with deep furrows marking the limits between the osteoscutes (Fig. 2A). The frontal scute is single and situated
anteriorly. The paired frontoparietal scutes are large and
situated posterolaterally; they also extend posteriorly
across the frontoparietal joint and cover the anterolateral
portions of the parietal (Fig. 3D). The posteroventral corner of the frontal is braced ventrolaterally by the forking
postfrontal. In ventral view, several irregular cracks
extend longitudinally and transversally across the
frontals ventral surface (Fig. 2B). The crista cranii frontalis, preserved on the right ventral side only, emerges as a
low and arched anteroposterior ridge extending almost
parallel to the lateral margins. Posteromedial to the crista
cranii frontalis there are clear indications (i.e. in form of
deep, triangular sutural imprints) of the ventral parietal
lappets (Fig. 2B); the frontal parietal abutment on its
medial part is simple.
Parietal. In dorsal view, the parietal plate is moderately
damaged and its left anterolateral dorsal surface is covered partially by sediment (Fig. 2A). The entire dorsal surface of the parietal plate is elongate and somewhat
constricted in the middle, whereas the anterior part is
broadened and reaches its widest expansure at the
frontoparietal suture. At the anterolateral corner of the

parietal, there is a wedge-like area for articulation with
the underlying postfrontal. Similar to the azygous frontal,
the dorsal surface of the parietal is covered by an osteodermal crust bearing irregularly distributed pits and the
limits between the scutes are marked by deep furrows
(Fig. 2A). The occipital scute is pentagonal with more or
less parallel lateral margins; its anterior part is tapered
and has some damage at its anteriormost limit. The paired
parietal scutes are elongate; their anterolateral margins
apparently do not extend beyond the posterior quarter of
the interparietal scute, and, due to damage on both sides,
it is unclear whether the parietal and frontoparietal scutes
contacted each other (Fig. 3D). The interparietal scute,
which filled the space between the parietal and frontoparietal scutes, also is damaged by cracks and dislocations.
Anteriorly it extends slightly beyond the frontoparietal
joint, whereas the tapering posteriormost part meets the
anteriormost limit of the occipital scute; its posteromedial
third is damaged and it is hard to recognize whether it is
perforated by a parietal foramen (Figs 2A, 3D). The frontoparietal scutes have sagittal contact anterior to the interparietal scute, and similar to lacertiform lizards (e.g.
nansky & Auge 2013) the frontoparietal scutes extend
Cer
forward to the azygous frontal scute. Posterolaterally the
frontoparietal scutes extend beyond the frontoparietal
joint to a point approximately level with the presumed
parietal foramen. The posterolateral borders of the parietal
are steeply inclined ventrolaterally, and expose elongate
and posteriorly deepening supratemporal fossae, which in
the living animal were invaded up to the sculpted surface
by the jaw adductor musculature. The supratemporal processes are wide and robust, diverging posterolaterally at
Figure 5. Partial neurocranium, suspensorium and palatal complex in the holotype (UBB V.440) of Barbatteius vremiri gen. et sp. nov.
Photographs in A, dorsal; B, ventral; and C, anterior views. D, details of the sphenooccipital region, with bone limits depicted, in ventral
view. E, details of the sphenooccipital region in slightly oblique posteroventral view. Abbreviations: anf: abducens nerve foramen; bo:
basioccipital; bot: basioccipital tubercle; bpp: basipterygoid process; cap: crista alaris of prootic; ccf: cerebral carotid foramen; ci: crista
interfenestralis; cp: crista prootica; ct: crista tuberalis; cu: cultriform process; ds: dorsum sellae; ept: epipterygoid; ff: facial foramen; fv:
fenestra vestibuli; juf: jugular foramen; marst: medial aperture of recessus scalae tympani; rst: recessus scalae tympani; paf: palatine
artery foramen; bs: basisphenoid; oto: otooccipital; po: prootic; poV: posterior opening of Vidian canal; ppp: posterior process of
prootic; pt: pterygoid; qu: quadrate; quf: quadrate foramen; so: supraoccipital; sot: sphenooccipital tubercle; stp: supratrigeminal process; tn: trigeminal notch. White arrows point to furrows produced presumably by plant roots. Scale bars D 5 mm.
about 100 . The posterolateral side of the supratemporal

processes are covered by the supratemporals and by the
ascending processes of the squamosal.
In ventral view, the sutural contacts of the ventral parietal lappets and the overlying frontal (Estes et al. 1988) are
readily apparent on each side. Nevertheless, the right ventral parietal lappet is broken off leaving on the frontals
posteroventral margin an imprint of a deep and roughly
triangular concavity. The crista cranii parietalis is moderately high and there is a well-preserved parietal downgrowth on the left lateral side; a fragmentary bone in front
of the alar process of the prootic, embedded in sediment
and cemented to the parietals ventral side, probably represents the vestiges of the epipterygoid, which in the living animal contacted the parietal downgrowths (Fig. 3A).
The supratemporal processes are relatively wide and moderately long. Their posteriormost parts are turned downward and have some thickening of their distal end.
Postfrontal postorbital. These bones are almost
completely preserved and apparently fused forming a
single structural unit. The exception is the jugal ramus on
the postorbital, which is broken off. On the right side the
bone is roughly in its original position, whereas on the left
side it experienced a counter-clockwise rotation. The anterodorsal tuberosity is preserved only on the left side
(Fig. 2B). The forking and medially curved postfrontal
part is robust and relatively short; the frontal process is
somewhat longer than the parietal process. The dorsal and
lateral surfaces of the postfrontal postorbital are covered,
similar to the parietal and frontal, by a strong vermiculate
dermal sculpture; the osteoscutes are separated by deep
grooves. The first osteoscute, situated above the jugal process and facing dorsolaterally, comes in contact with two
small lateral scutes. More posteriorly are three scutes,
also facing dorsolaterally, that cover the postorbital and
the squamosal and thereby also extend over the postorbital squamosal joint. The posterior ramus of the postfrontal postorbital is elongate and tapers posteriorly; it
extends more than three-quarters the length of the upper
temporal fenestra and also extends dorsally onto the squamosal; its posterior terminus, similar to other crown lizards, is situated at the medial side of the squamosal
(Gauthier et al. 2012).
Jugal. A small piece of bone partly embedded in sediment on the left side of the skull, situated lateral to the
anterodorsal tuberosity of the postfrontal postorbital,
Figure 6. Lower jaws in the holotype (UBB V.440) of Barbatteius vremiri gen. et sp. nov. A, partial left lower jaw in lateral view and
right lower jaw in dorsomedial view; B, partial left lower jaw in medial view and right lower jaw in lateral view. Arrow points to furrow
presumably produced by plant roots; C, partial left lower jaw in ventrolateral view and right lower jaw in medial view. Photographs
(left) and interpretive figures using different levels of grey to highlight particular bones (right). Abbreviations: ac: adductor crest; aiaf:
anterior inferior alveolar foramen; amf: anterior mylohyoid foramen; an: angular; art: articular; asf: anterior surangular foramen; cor:
coronoid; den: dentary; pvd: posteroventral process of dentary; maf: mandibular fossa; mf: mental foramen; pmf: posterior mylohyoid
foramen; san: surangular; sds: subdental shelf; sp: splenial; sur: surangular ridge. Scale bars D 5 mm.
probably represents the distal tip of the postorbital ramus

of the jugal (Fig. 2B); the outer surface of this bony fragment is faintly ornamented. The impression left by that
ramus on the lateral side of the postfrontal postorbital
may indicate that a mobile joint existed here.
Squamosal. In dorsal view, the squamosal is a hooked
bone covered with prominent dermal sculpture. Its
ascending process is rather large and dorsolaterally covers
the supratemporal process of the parietal. A deep diagonal
fossa on the upper temporal bar is partly filled with sediment; this marks the posterior limit of contact between the
postorbital ramus of the squamosal and the postorbital.
Anterior to this contact line, the squamosal is covered dorsally by the postorbital. In lateral view, the squamosal is
crescentric in outline and is covered by prominent dermal
sculpture similar to that on other roofing bones (see
above). The ascending process is covered by two osteoscutes: a larger one facing dorsally and a smaller one facing posteriorly. The anterior terminus of the postorbital
ramus of the squamosal was in contact with or close to the
jugal (Fig. 2B), based on the observation that the
impression left by the jugals postorbital ramus is preserved on the left lateral side of the postorbital. The squamosal ventral ramus is relatively short and robust, with a
somewhat thickening ventral terminus that articulates
with the quadrate.
Supratemporal. In posterior view, the supratemporal
appears as a faintly sinuous lamellar bone adhering to the
lateral margins of the parietals supratemporal processes
(Fig. 4A). In dorsal view the supratemporal normally
would be hidden by the ascending process of the squamosal; yet in the specimen it became exposed thanks to the
squamosal having been slightly detached from its original
position postmortem. Ventrally, the supratemporal and
the parietals supratemporal processes form part of the
articular surface for the quadrate.
Prootic. A triradiate bone forming the anterolateral wall
of the braincase and housing part of the membranous labyrinth (Oelrich 1956). The alar process is broken off, but
it was found embedded in sediment, adhering to the ventral part of the parietal (Fig. 2B). In ventral view, this
structure has an anteriorly open V shape contacting at its
dorsolateral corner (i.e. apex) the parietal table and also
the dorsal head of the epipterygoid. Part of the crista alaris
forming the anterior border of alar process is still preserved (Fig. 5A, C). Its ventral part diverges laterally to
the posterior border of the trigeminal notch. The supratrigeminal process appears as a bulging prominence on the
medial side of the alar process (Fig. 5C). The inferior process of the prootic is relatively short and firmly fused
anteroventrally to the basisphenoid; it delimits the ventral
margin of the relatively large and rounded trigeminal
notch. The crista prootica, mostly embedded in sediments
and having a partly damaged posterior part (Fig. 5C),
extends between the posterior and the anterior inferior
processes and serves as an attachment surface for the protractor pterygoideus muscle (Oelrich 1956). Fracture lines
on each side, along the base of the posterior part of the
crista prootica, indicate that the crests were shifted laterally. The small-sized facial foramen is positioned on the
same level as the crista interfenestralis and near to the
posterior border of the prootic (Fig. 5D). The ventromedial part of the prootic forms part of the medial aperture
of the recessus scalae tympani (MARST), similar to
crown Teiioidea (Gauthier et al. 2012); this character is
best observed on the left side of the neurocranium where
it is free of sediment (Fig. 5E). The posterior process of
the prootic is relatively long and has a posterolateral orientation. It overlaps and sutures to the dorsal surface of
the paroccipital process of the otooccipital.
Basisphenoid. This rather robust median bone forms part

of the floor of the cranial cavity (Oelrich 1956); it is
strongly sutured with the basioccipital posteriorly and
with the prootic dorsolaterally. The left contact zone with
the basioccipital is strongly eroded (Fig. 5B, D, E). In ventral view, the basipterygoid processes are rather large and
prominent; they are roughly as wide as long, with the ventral carina slightly damaged. Laterally they are still in
contact with the posterior rami of the pterygoids. Between
the basipterygoid processes, a small fragment of the cultriform process is preserved. On the right side, the sphenooccipital tubercle is prominent and has a medially concave
surface. The posterior opening of the Vidian canal, which
in the living animal houses the internal carotid artery and
the platine branch of the facial nerve, is visible on each
side. In anterior view, the dorsum sellae is high and, as
seen on the left side, pierced about mid-height by the
abducens nerve foramina. The lateral sides of the dorsum
sellae display two distinct inferior (clinoid) processes
serving as an attachment surfaces for the pilae antoticae
(Rieppel & Zaher 2000). The internal carotid artery is subdivided within the Vidian canal into a dorsal (cerebral
carotid) and a ventral (palatine artery) branches (Rieppel
& Zaher 2000). The anterior openings of the cerebral
carotid and the palatine artery are seen at the anterior side
of dorsum sellae and at the anterior base of the basipterygoid process, respectively (Fig. 5C).
Otooccipital. This is a composite bone derived in most
squamates from the fusion of the exoccipital and opisthotic (Oelrich 1956; Conrad 2004; Bever et al. 2005;
Head et al. 2009). It also forms the lateral part of the
occipital tubercle and laterally delimits the foramen magnum. The fenestra vestibuli (D foramen ovale) lies in a
dorsoposterior position relative to the recessus scalae tympani (D occipital recess) and sphenooccipital tubercle; its
anterior and dorsal margins are delimited by the prootic
(Norell & Gao 1997), whereas its ventral margin is demarcated by the prominent and roughly horizontally displayed
interfenestral crest (Fig. 5B, D). The recessus scalae tympani lays just posterior to and on the same level with the
sphenooccipital tubercle, as an elongated hole, bordered
anterodorsally by the interfenestral crest and posteroventrally by the crista tuberalis. The anteromedial duct within
the recessus scalae tympani represents the passage of the
glossopharyngeal nerve in non-ophidian squamates
(Rieppel & Zaher 2000). The canal passes anteromedially
and slightly dorsally (Fig. 5E), instead of piercing medially the otooccipital wall, as seen in Recent lacertids; the
anterodorsal margin of the MARST reaches the posteroventral border of the prootic; the ventral side of the
MARST is bordered by the basioccipital. The jugular or
vagus foramen (Rieppel 1985), delimited anterodorsally
by the crista tuberalis, is positioned posterior to the recessus scalae tympani within a deep, slit-like fossa; it is
accompanied posteroventrally by two smaller foramina
for the hypoglossal nerve.
Supraoccipital. This nearly rectangular bone is strongly
fused laterally to the prootic (Fig. 5A); the anterior border
and the processus ascendens are broken off, but these
parts were found within the matrix adhering to the ventral
side of the parietal.
Basioccipital. This is robustly fused to the basisphenoid
anteriorly and to the otooccipitals laterally. The right side
displays a well-preserved and rather prominent sphenooccipital tubercle, whereas the left side is so strongly eroded
that the sphenooccipital tubercle and part of the occipital
tubercle have broken off (Fig. 5B, D, E). On the remaining
surface, at the level where the occipital tubercle would
have been, a semicircular furrow about 1 mm wide
extends anteromedially; this peculiar furrow may be an
impression left by plant roots (Jean-Claude Rage pers.
comm.). A second furrow is observed on the right side of
the basioccipitals ventral surface, in line with the crista
tuberalis.
Pterygoid. Only the posterior (i.e. quadrate) processes
are preserved (Fig. 5A C). They are shifted dorsally
from their original articulating points with the basisphenoid processes. A breakage line is observed on the right
10
side and the posterior part of the right quadrate process

has shifted medially. Posteriorly these processes contacted
the quadrate ventromedial surface (Oelrich 1956). The
fossa columellae, representing the articulation point with
the epipterygoid, is seen on the dorsal sides of the posterior processes where small remnants of the epipterygoids
are also preserved.
Quadrate. From the right quadrate only a small fragment
is preserved, probably close to its original point of articulation and position. The left quadrate, except for the missing margins of the pterygoid lappet and of the mandibular
condyle (see below), is almost completely preserved, near
its point of articulation with the pterygoid posterior ramus.
However, it has been rotated clockwise by about 45 from
its original position (Fig. 5A, C). In lateral view, it
appears straight and rather robust. The well-developed
cephalic condyle articulates with the squamosal dorsolaterally; the cemented sediment still fixes the tip of the
squamosals left ventral ramus to the cephalic condyle.
The mesioventral lappet for sliding contact with the posterior ramus of the pterygoid appears relatively small, but a
breakage surface indicates that part of its anterior margin
was broken off and, thus, was originally larger. In anterior
view, the quadrate has a vertically oval shape, only
slightly higher (11 mm) than wide (9 mm), and bears a
broad and posteriorly curved tympanic crest on its lateral
edge. The medial crest (D pterygoid lappet) is bent anteriorly and is indented by a distinct notch for insertion of the
posterior process of the pterygoid. The mandibular condyle is relatively small, but this condition resulted from a
postmortem breakage that affected both the lateral and
medial margins of the condyle. The quadrate foramen,
filled completely by sediment, is relatively small and positioned near the mandibular condyle on the pterygoid lappet (Fig. 5C). The posterior face is completely embedded
in sediment that hides the posterior crest. Postmortem
rotation of both quadrates means we were unable to determine whether they had an anteroventral or a posteroventral slope, which is unfortunate because this is a useful
character in cladistic analyses (see Gauthier et al. 2012).
Dentary. Both dentaries are complete. The left one displays a postmortem break at the level of the 15th tooth
position producing an inward curve from its roughly
straight outline (Fig. 6A C). The mandible symphyseal
region is relatively small, with lingually rounded margins.
The Meckelian canal is apparently open anteriorly to the
symphysis and closed posteriorly by the hypertrophied
splenial. Unfortunately, the anterior terminal part of the
splenial is not preserved. The ventral margin of the dentary is more or less arched ventrally and the bone becomes
slightly deeper posteriorly. The labial surface is
completely smooth and is perforated by at least seven
small mental foramina (foramina pro rami nervorum
alveolarium inferiorum); the spacing between the
foramina increases posteriorly (Fig. 6C). The posterolateral border is notched for the surangular joint. The posteroventral process, which is enclosed by the bifurcate
angular, terminates slightly anterior to the coronoid apex.
The subdental shelf is relatively small, being both shallow
and lingually narrow. The teeth have at their bases some
cementum depositions and are placed relatively close to
the subdental shelf margin. The alveolar shelf supports 26
tooth positions; the teeth are closely spaced and heterodont. The anteriormost five teeth are the smallest and are
medially curved; unfortunately, all have their tooth
crowns broken off. The 6th 10th teeth are larger in size
and at least appear to mark a posteriorward transition
from bicuspid to tricuspid crowns: the sixth tooth is bicuspid, whereas the ninth is already tricuspid. The 10th 20th
teeth are the largest and tallest, and their bases are broadened labiolingually. They are gently curved posterolingually and provided with tricuspid tooth crowns,
composed of a main central cusp and two secondary
cusps; the mesial secondary cusp is always larger than the
distal one. The tooth crown surfaces are without any trace
of striations; however, a single tooth (the 11th in the left
dentary) preserves some striae. The teeth project about
one-third of their total height above the dental parapet.
The posterior five or six teeth in each dentary are strongly
worn or broken off and partly embedded in sediment;
apparently they were of smaller size and of diminished
height. Large circular resorption pits are observed along
the dentary tooth row, indicating that the tooth replacement was present in all dentary teeth. A semicircular furrow, similar to those seen on the basioccipital surface, is
present on the lingual side of the left lower jaw and probably also is the trace of a plant root (Fig. 6B).
Coronoid. The coronoid is triradiate, covering the posterior margins of the dentary and clasping it labiolingually
(Fig. 6A). The anteromedial ramus articulates with the
splenial on the lingual side, whereas the anterolateral dentary process extends anteroventrally and articulates with
the surangular on the labial side; anteriorly it overlaps
past the level of the tooth row (e.g. Gauthier et al. 2012),
in a manner resembling some teiids (e.g. Tupinambis) and
lacertids (e.g. Lacerta). The posteromedial process
contacts the surangular laterally and the prearticular
medially.
Splenial. This is a well-developed bone that contacts the
anteromedial ramus of the coronoid dorsally and overlaps
the prearticular and the angular medially (Fig. 6B, C).
Posteriorly it does not extend beyond the apex of the coronoid process. The anterior inferior alveolar foramen and
the anterior mylohyoid foramen are situated relatively
close to each other and are each fully enclosed by the
splenial.
Angular. The angular extends on the ventral margin of

the lower jaw and turns anteroposteriorly from the lingual
to the labial side of the mandible, filling the space
between the prearticular and the surangular (Fig. 6B).
Anteriorly it bifurcates and clasps the posteroventral process of the dentary. Posteriorly it reaches the level of the
articular condyle (Fig. 6B, C). The posterior mylohyoid
foramen (D angular foramen) is located entirely within
the angular and situated near the posteroventral limits of
the splenial.
Articular complex. The articular complex is composed
of a fused surangular, prearticular and articular (Fig. 6A,
B). The surangular occupies most of the lateral surface of
the mandible. The lateral surface also bears an adductor
crest for the attachment of jaw muscles (Gao & Norell
2000). The anterior surangular foramen lies laterally, at
the level of the coronoid apex. The location of the posterior surangular foramen cannot be determined with certainty because of damages posterodorsally on the
mandibles labial side. Broken parts of the surangular dorsal margin have been turned medially and shifted into a
ventromedial position, occluding somewhat the originally
widely open adductor fossa (Fig. 6A). The sigmoid fossa
and the articular fossa are wide. However, a deep fracture
line on the lateral side of the articular condyle suggests
that it also has been turned medially and shifted medioventrally. The retroarticular process narrows posteriorly,
but its tip is broken off. Medial to the sigmoid fossa, and
covered by patches of sediments, there is a well-developed pterygoideus process (D angular process). The prearticular crest is absent.
Remarks
The fragmentary skull and associated lower jaws of Barbatteius are notable for their three-dimensional appearance and the fact that many of the bones are preserved in
or close to their in-life positions. However, osteological
and taxonomic interpretations are somewhat encumbered
by strong fusion of neurocranial bones, by weathering of
bone surfaces and by patches of sediment firmly attached
to some surfaces. Unfortunately, referable remains of the
postcranial skeleton also are lacking.
One of the most intriguing features of Barbatteius is the
presence of osteodermal crust that fuses to the skull roofing bones and suspensorium. The outer surfaces of the
osteoderms also bear the impressions of cephalic scales,
thus providing information on the pileus pattern. The
fusion line between the cephalic osteoderms and skull
bones of Barbatteius is exposed, amongst others, along
the ventrolateral margin of the squamosals postorbital
ramus (Fig. 4). Separable osteodermal crust occurs in
cordylids, scincids, lacertids, xenosaurs, anguids,
11
Lanthanotus and helodermatids, as well as in some varanids and the iguanid Amblyrhynchus, which is considered
by Estes et al. (1988) a synapomorphy of both Anguimorpha and Scincoidea. In lacertids only a few osteoderms on
the periphery of the skull table are separable, whereas
non-separable dermal sculpture occurs in some iguanians,
gymnophthalmids, teiids, xantusiids and amphisbaenians
(Estes et al. 1988). The condition seen in Barbatteius
(cephalic osteoderms fused to the skull roofing bones and
suspensorium with the fusion lines exposed in some parts
of the dorsum) apparently indicates the possible occurrence of separable osteoderms that fuse ontogenetically.
The presence of non-separable dermal sculpturing on the
parietal and frontal may represent a synapomorphy of
Autarchoglossa (see Estes et al. 1988: character 129(1);
Conrad 2008: character 10(1); Gauthier et al. 2012:
character 572(2)). In several lizard groups the dermal
skull bones are smooth (e.g. Leiolepidinae, Isodontosauridae, Mosasauria, Eischstaettisaurus, Gekkota, Krypteia,
Adamisaurus and the scincomorphan Kleskunsaurus),
whereas in others the ornamentation is weakly defined
about the frontoparietal suture (e.g. Acontias, Feylinia,
Varanidae, Gobinatus, Gilmoreteius, Polyglyphanodon)
or extends over the dorsum (e.g. Leiosaurinae, Hoplocercinae, Rhineuridae, Tchingisaurus, Sineoamphisbaena)
(Nydam et al. 2010; Gauthier et al. 2012). The presence
of vermiculate sculpture on these cephalic scale impressions was tentatively considered by Estes et al. (1988) to
be a synapomorphy of Scincomorpha with reversals in
gymnophthalmids, teiids, xantusiids and scincids. In some
Recent teiids (e.g. Ameiva, Cnemidophorus and Kentropyx) the osteodermal crust on the skull is present (Presch
1974; Estes 1983; see also Fig. 3C), whereas in others it is
reduced (e.g. Callopistes, Dracaena, Tupinambis). In the
possible teiid Meyasaurus diazromerali, known from the
Early Cretaceous, Spain, the dorsum is ornamented by a
vermiculate sculpture with deep grooves marking the
original positions of the overlying scale impressions
(Evans & Barbadillo 1997). In the scincomorphan Sakurasaurus shokawensis, known from the Early Cretaceous,
Japan, the dorsal surface of the paired frontals (and probably that of the parietal) bears shallow pustulate sculpture
without overlying scale marks (Evans & Manabe 1999).
In Barbatteius the presence of cephalic osteoderms that
fuse to the skull roofing bones and suspensorium, covered
by a vermiculate sculpture and by impressions of the
cephalic scales, appears as a unique combination of features probably shared by early lacertoid lizards. In Barbatteius the pileus morphology (i.e. the pattern of scalation
nansky & Auge 2013) seems transion the skull; e.g. Cer
tional between lacertids and teiids. Similar to lacertids
there is a single occipital scute separating the parietal
scutes posteriorly and the frontoparietal scutes are in sagittal contact, extending forward to the frontal scute
nansky & Auge 2013). However, in the possible
(Cer
12
crown lacertid Succinilacerta succinea, known from the

Middle Eocene Baltic amber of Poland and Lithuania
(Borsuk-Biaynicka et al.1999), and Plesiolacerta lydekkeri, known from the Middle Eocene Early Oligocene
nansk
(MP14 MP21) of France (Cer
y & Auge 2013), the
occipital scute is strongly widened posteriorly, whereas in
Barbatteius it remains narrow and pentagonal with more
or less parallel lateral margins. In contrast, Barbatteius
resembles Plesiolacerta eratosthenesi, known from the
nansk
Late Oligocene of southern Germany (Cer
y & Auge
2013), in having similarly narrow occipital scute, but differs from the latter in having a pointed anterior terminus
of the occipital scute.
In Barbatteius the lateral ventral lappets (D parietal
tabs) of the parietal are evident on both sides, similar to
Lacertiformes; on the right side, however, the lappet is
broken off, leaving on the frontals posterolateral margin
a sutural imprint of roughly triangular shape. In the teiidlike Polyglyphanodontia, known from the Late Cretaceous
of Asia and Euramerica, the ventral lappet of the parietal
was found in Tchingisaurus multivagus, whereas in others
(e.g. Adamisaurus, Gilmoreteius, Gobinatus, Polyglyphanodon and Sineoamphisbaena) the presence of this structure remains uncertain (Longrich et al. 2012,
supplemental material). In Meyasaurus the presence of
ventral lappets was reported by Evans & Barbadillo
(1997, see appendix 1, character 22). In Barbatteius the
frontal is fused similar to Gekkota, Teiioidea, Carusiidae,
Lygosominae, Xenosauridae and Bipes (see Gauthier et
al. 2012). Paired frontals are typical for lacertoids and
borioteiioids (Nydam et al. 2007), but fused frontals with
a strong interorbital constriction like in Barbatteius also
occur in lacertids (Bolet & Evans 2012).
In Barbatteius the supratemporal and squamosal remain
separate similar to some polyglyphanodontids (e.g. Cherminsaurus, Erdenetesaurus and Gobinatus), whereas in
others they are fused (e.g. Darchansaurus, Gilmoreteius
(D Macrocephalosaurus) and possibly Polyglyphanodon)
(Estes 1983); the condition in Adamisaurus remains
unknown.
In Barbatteius the prootic forms part of MARST that is
considered a unique and unreversed character of Teiioidea
(Gauthier et al. 2012). The MARST is bordered primitively by the otooccipital (e.g. in lacertids; see Gauthier et
al. 2012), whereas in teiioids and Barbatteius it is bordered at least partially by the posteroventral part of the
prootic.
The anterior extent of the anterolateral dentary process
of the coronoid in Barbatteius is similar to Recent teiids
and lacertids, whereas in Sakurasaurus (Early Cretaceous,
Japan) the dentary bears a posterodorsal process that overlaps the coronoid (Evans & Manabe 1999). The weakly
developed subdental shelf that is present in Barbatteius
may represent, after Gauthier et al. (2012), a synapomorphy of Teiidae (shared also by Meyasaurus). An inflated,
widely open adductor fossa, resulting from the extension

of the adductor mandibulae posterior muscle into
Meckels canal, occurs in the mandible of Barbatteius and
is considered a synapomorphy of Lacertiformes (Estes et
al. 1988). An additional combination of characters in the
lower jaws of Barbatteius that are reminiscent of lacertiform lizards, is as follows: ventral margin of dentary and
subdental shelf are arched ventrally (Auge 2005; Bolet &
Evans 2012); Meckels groove is open up to the symphysis and covered by the hypertrophied splenial that almost
nansk
reaches the symphysis (Auge 2005; Cer
y & Auge
2013); mandibular symphysis is relatively small; the coronoid process of the dentary does not extend onto the anterolateral surface of the coronoid, unlike that of most
scincoids in which the posterolabial process of the dentary
is large and extensively overlying the coronoid (Auge
2005; Conrad 2008); the dentition is heterodont, consisting of unicuspid teeth anteriorly and bi- and tricuspid
teeth posteriorly (Gauthier 1984). Based on the above
listed unique combination of features, the lower jaws and
marginal teeth of Barbatteius are relatively easy to differentiate from those of Asian Polyglyphanodontia, known
from the Campanian of Mongolia, and Euramerican Borioteioiidea, known from the lower Cenomanian
Maastrichtian of North America, Santonian of Ihark
ut,
Hungary and Maastrichtian of Hat eg Basin, Romania.
The Asian polyglyphanodontids clusters an array of taxa
with at least three different tooth patterns: leaf-shaped,
polycuspate teeth (e.g. Darchansaurus, Erdenetesaurus,
Gilmoreteius (D Macrocephalosaurus); large, bulbous,
conical teeth (e.g. Adamisaurus) and obliquely orientated,
chisel-like, policuspate teeth (e.g. Cherminsaurus). None
of the above morphologies approach the condition seen in
Barbatteius in having a slightly heterodont dentition and
less modified tooth morphology. After Nydam (2013), the
Euramerican distributed Borioteioiidea consists of
Chamopsiidae and Polyglyphanodontini (see also Longrich et al. 2012). The Chamopsiidae (i.e. Chamops,
Cnephasaurus, Gerontoseps, Glyptogenys, Haptosphenus,
Harmondontosaurus, Leptochamops, Meniscognathus,
Pelsochamops, Socognathus, Sphenosiagon, Stypodontosaurus) is diagnosed by Nydam et al. (2010) as follows: a
long, massive, U- or V-shaped mandibular symphysis
extending posteriorly to the level of the 4th 5th tooth
positions onto the superior and inferior margins of the
Meckelian groove; teeth that tend to be massive with a
mid-shaft swelling (barrel-shaped), tooth crowns tend to
have mesial and distal accessory ridges and cusps; the
teeth are widely spread along the tooth row. Barbatteius
differs from the above genera by lack of massive Ushaped mandibular symphysis on the dentary, by lack of
tooth crowns with conical apex and bordered by mesial
and distal accessory ridges and by mid-shaft swelling of
mandibular teeth. Barbatteius also differs from Pelsochamops (Santonian, Iharkut, Hungary) in having its coronoid
not fused to the dentary (Makadi 2013a). Barbatteius differs from Haptosphenus, an aberrant taxon having its dentary, splenial, coronoid and surangular fused (although the
limits of bones are still visible), sulcus dentalis closed and
possessing a subacrodont tooth attachment (Estes 1983).
Barbatteius resembles the incertae sedis polyglyphanodontian Obamodon (referred earlier to Leptochamops)
and Prototeius in having the mandibular symphysis
weakly developed, but differs from the latter two and also
from the chamopsid Tripennaculus in lack of tooth crowns
with a tall central cusp separated from accessory cusps by
deep lingual grooves (Longrich et al. 2012). The Euramerican Polyglyphanodontini (sensu Nydam et al. 2007,
see also Makadi 2013a, b) includes lizard taxa that share
transversely orientated, interdigitating, mammal-like teeth
in the posterior portion of the tooth row, probably used for
oral food processing, and suppression of tooth replacement in adults (Bicuspidon, Dicothodon, Distortodon,
Paraglyphanodon, Peneteius (D Manangysaurus) and
Polyglyphanodon). The dentition of Barbatteius, consisting of unicuspid and slightly recurved teeth anteriorly and
bi- and tricuspid teeth posteriorly, strongly differs from
the above depicted multicuspid tooth morphology. In
addition, tooth replacement in Barbatteius was present in
all dentary teeth.
Phylogenetic relationships
To assess the phylogenetic relationships of Barbatteius
within Squamata, we added all scorable osteological characters for the holotype specimen to the character taxon
matrix (CTM) of Gauthier et al. (2012) (196 characters,
representing 32% from the character list). Additionally,
based on published data of Evans & Barbadillo (1997),
we reviewed the characters of Meyasaurus, a presumed
early teiioid from the Early Cretaceous of Spain (Evans &
Barbadillo 1999; Evans 2003; but see Conrad 2008 for a
different placement of Meyasaurus), and added these (214
characters, representing 35% from the character list) to
the CTM of Gauthier et al. (2012). Corrections applied to
the CTM of Gauthier et al. (2012) follow the revisions of
Longrich et al. (2012, supplemental material: characters
89, 117, 360, 388, 413, 468 and 572). Using our modified
Gauthier et al. (2012) dataset of 194 operational taxonomic units (OTU) and 610 multistate characters, we performed parsimony analyses with the phylogenetic
software package TNT version 1.1 (Goloboff et al. 2008).
In New Technology search the CTM was first analysed
using sectorial search, ratchet, drift, and tree fuse options
with default parameters, and then the generated trees were
analysed under traditional TBR (i.e. tree bisection reconnection) branch swapping. Bootstrap support values using
1000 replicates in traditional search and Bremer (1994)
decay indices up to 20 steps longer than the minimum tree
13
length were also calculated. The traditional search with

the TBR branch swapping algorithm found 50 trees with a
length of 4882 steps (consistency index (CI) D 0.199;
retention index (RI) D 0.770). Barbatteius is recovered,
within the clade of Lacertiformes (Lacertoidea of Gauthier et al. 2012) as the sister taxon of Meyasaurus, whereas
the clade of Barbatteius C Meyasaurus appears within
Teiidae as the sister taxon of Teiinae and Tupinambinae.
In a second analysis, we restricted the dataset of the original 194 OTUs for easier handling to a total of 59 OTUs.
In this subset all important squamate groups, considered
most relevant to assess phylogenetic relationships of Barbatteius (e.g. most snakes were eliminated), were
included. We selected, where available, the OTUs with
more complete CTMs (see Supplementary Material). The
TBR search in TNT returned four equally parsimonious
trees of 2448 steps length (CI D 0.357; RI D 0.641). The
strict consensus tree is represented in Figure 7A. Barbatteius is recovered, similar to the first analysis, as the sister
taxon of Meyasaurus, whereas the clade of Barbatteius C
Meyasaurus appears within Teiidae as the sister taxon of
Teiinae and Tupinambinae. More inclusive clades within
lacertiforms are the Gymnophthalmidae (Pholidobolus C
Colobosaura), clustering with Teiidae in Teiioidea and
the clade of Lacertidae (Takydromus C Lacerta), which
appears as the sister taxon of Teiioidea. Except for the
clades of lacertids (bootstrap D 90%, decay index D 8)
and gymnophthalmids (bootstrap D 99%, decay index D
12), nodal support for Lacertiformes and the rest of its
less-inclusive clades was moderate (bootstrap value below
60%, decay index D 2 to 4). Nodal support for the clade of
Barbatteius C Meyasaurus is extremely low (bootstrap
support value below 50% and Bremer support D 1), conceivably because of the high percentage of unscorable
characters for both genera.
Based on the scored characters of Barbatteius and
Meyasaurus, below we detail the unambiguous synapomorphies mapped by TNT (Fig. 7B). Barbatteius shares
with Lacertiformes two unambiguous synapomorphies: 89
(1) parietal ventral lappet forms a prominent V-shaped,
flat process (present also in Meyasaurus), and 394(2)
coronoid anterolateral dentary process overlaps dentary
past level of tooth row; Barbatteius shares with Teiioidea
two unambiguous synapomorphies: 36(1) frontals fused
(also in Meyasaurus), and 314(1) prootic forms part of
MARST; Barbatteius shares with Teiidae four unambiguous synapomorphies: 78(2) postorbital overlaps squamosal dorsally, 90(0) parietal temporal muscles originate
dorsally on parietal table and parietal supratemporal processes (shared also by Meyasaurus), 294(1) epipterygoid parietal contact overlaps parietal temporal muscle
origin and 360(1) weakly developed subdental shelf
(shared also by Meyasaurus). Finally, Barbatteius and
Meyasaurus share two synapomorphies: 163(1) squamosal temporal ramus widens posteriorly (also present in
14

been reported from the European Late Cretaceous
(e.g. Bicuspidon, Chamops and Distortodon from the Santonian of Iharkut, Hungary (Makadi 2006, 2013a, b), and
Bicuspidon from the Maastrichtian of the Hat eg Basin,
Romania (Folie & Codrea 2005)). If the assignment of
Barbatteius within Teiidae is correct then it represents the
first occurrence of this group from the Late Cretaceous
(early Maastrichtian). This record is roughly in agreement
with the time tree of squamates of Hedges & Vidal (2009),
who estimated the divergence of Teiioidea from other lateratan groups in the Jurassic and split of Teiidae and Gymnophthalmidae in the Late Cretaceous (about 86 Ma).
Palaeobiogeographical and palaeoenvironmental implications
Figure 7. Relationships of Barbatteius vremiri gen. et sp. nov.,

based on our phylogenetic analysis. A, strict consensus of four
trees generated with TNT, based on 57 OTUs with 610 phenotypic characters (derived from Gauthier et al. 2012), Meyasaurus
(based on published data of Evans & Barbadillo 1997) and Barbatteius (see Supplementary Material). Numbers below nodes
indicate decay indices of Bremer (D steps). B, the clade of Lacertiformes and supporting unambiguous synapomorphies at
nodes. The character numbers and character states are from
Gauthier et al. (2012) and Longrich et al. (2012).
Xenosaurus) and 411(1) prearticular pterygoideus process

(D angular process of Oelrich 1956) is present (also
shared with Iguania).
Despite missing a substantial portion of the skeleton,
Barbatteius is placed as an early member of Teiidae based
on four unambiguous cranial synapomorphies shared with
this family. On the other hand Barbatteius is not part of
Borioteiioidea, a group of lizards distantly related to
Teiioidea (Nydam et al. 2007). From that group only Polyglyphanodon sternbergi and Gilmoreteius (D Macrocephalosaurus) were included in our analysis (see Fig. 7A),
because the remaining borioteiioidean taxa are commonly
established on fragmentary jaws. Borioteiioideans are one
of the most common groups of lizards from the Late Cretaceous of North America (Nydam 2013). They also have
In the Late Cretaceous, one of the European archipelago

islands was formed by emerged segments of continental
crust belonging to the tectonic block Tisia-Dacia. This
Hat eg Island had an estimated surface of at least 7500
km2 (Weishampel et al. 1991) or even, as suggested by
outcrops and fossils from a significantly larger area than
that of the Hat eg Basin (Nopcsa 1905; Codrea & Dica
2005; Codrea et al. 2010), had a much larger landmass up
to 80,000 km2 (Csiki 2005; Benton et al. 2010). Palaeogeographical reconstructions (see Benton et al. 2010)
depict Hat eg Island as having been circumscribed by
deep marine basins, separated from the nearest landmasses by about 200 300 km and lying within the subtropical belt at approximately latitude 27 N (Grigorescu
2005; Panaiotu & Panaiotu 2010). Other emerged terrains
relatively close to Hat eg Island were the emergent
sequences of the ALCAPA block to the west and the
Adriatic Dinaric Carbonate Platform to the south
(Benton et al. 2010). From at least the second half of the
Campanian, intermittent land routes probably were established between these emerged areas and the Ibero-Armorican landmass (Csiki & Grigorescu 1998; Benton et al.
2010; but see also Jianu & Boekschoten 1999; Csiki-Sava
et al. 2015). The calcrete horizons in the Pui beds indicate
a climate where the annual rainfall was less than 760 mm
(Royer 1999, 2000; Khadkikar et al. 2000). Applying the
thermal gradient for the late Campanian middle Maastrichtian, based on the methodology of Amiot et al.
(2004), the estimated mean annual palaeotemperature of
the Pui area might have been around 22 C.
In the Hat eg Island palaeoecosystems, Barbatteius
appears as a large-bodied lizard with generalized teiid
attributes. The Teiioidea shifted from the general lacertiform body plan, in elongation of the body and the apparent thickening and lengthening of the tail (Vitt & Pianka
2004), which are considered important factors in locomotion (Ballinger et al. 1979). The resulting higher size-
specific mass in Teiioidea is in agreement with their
nearly exclusive restriction to terrestrial microhabitats
(Vitt & Pianka 2004). The total body length of Barbatteius, based on published data on Recent teiids (Harvey et
al. 2012; Arias et al. 2013) and extrapolating from the
incomplete holotype skull, came close to 800 mm (skull
length about 65 mm, snout vent length about 260 mm
and tail length up to 520 540 mm). The weakly heterodont dentition without enlarged posterior crushing teeth,
as seen in many recent teiids (Kosma 2004 and references
therein), suggests that Barbatteius mainly fed on arthropods (e.g. insects, millipedes and spiders), small vertebrates (e.g. fish, amphibians, turtle hatchlings, other
lizards and perhaps multituberculates) and plants.
Although the only specimen of Barbatteius bears no signs
of carnivore attack, in the food chain of Hat eg Island
palaeoecosystems, Barbatteius might have been included
as prey for other carnivores, even if the latter were not
numerous or, perhaps more accurately, have rarely been
found as fossils. Amongst the top predator candidates are
the crocodilian Allodaposuchus, various small theropods
(Csiki & Grigorescu 1998), and the aberrant dromaeosaurid theropod Balaur bondoc (Csiki et al. 2010). Allodaposuchus probably controlled the fluvial ecosystems
including riparian zones, whereas the smaller theropods
and Balaur with their highly modified raptorial hind limbs
(Csiki et al. 2010) probably foraged across much larger
areas in search of food.
The presence of paramacellodid lizards (Becklesius
nopcsai and B. cf. hoffstetteri) in the early Maastrichtian
at the Pui Islaz locality (Folie & Codrea 2005) points to
the relictual nature of this assemblage. It is reminiscent of
Early Cretaceous Euramerican faunas (Weishampel et al.
2010), but with a number of endemic forms. Although no
paramacellodid species have been described from the
Late Cretaceous of North America, paramacellodid-cordylid grade lizards, approaching Paramacellodus in morphology, are known in multiple horizons on that continent
(Nydam 2013). The presence of borioteiioid lizards in the
Late Cretaceous of North America (e.g. Nydam 2013) and
Eastern Europe also suggests a faunal connection between
these continents (Csiki-Sava et al. 2015). The European
borioteiioid record consists of Bicuspidon hatzegiensis in
the early Maastrichtian of Pui, Romania (Folie & Codrea
2005), and Bicuspidon aff. hatzegiensis, Distortodon
rhomboideus and the chamopsid Pelsochamops infrequens in the Santonian of Ihark
ut, Hungary (Makadi
2006, 2013a, b). On the other hand, the presence of the
sebecosuchian crocodyliform Doratodon (Weishampel et
k in press) and madtsoiid snakes
al. 2010; Rabi & Sebo
(Folie & Codrea 2005; Vasile et al. 2013), together with
teiid lizards, strengthens the view that some faunal elements of the Transylvanian landmass were of Gondwanan
origin. For the basal alethinophidian snake Nidophis insularis, reported recently from the Hat eg Basin, Vasile et al.
15
(2013) advanced the idea of an early (i.e. pre-Cenomanian) dispersal event of madtsoiid snakes from Africa into
Europe, followed by subsequent diversification and distribution across Alpine Europe (Hat eg) and cratonic southwestern Europe (Spain). A similar scenario might be
applicable to Barbatteius as well. According to Estes
(1970, 1983), the geologically oldest record of this presently American distributed group (i.e. Teiidae and Gymnophthalmidae) comes from the late Paleocene of
Itaborai, Brazil (as Teiinae and Tupinambinae indet.). In
Europe the only putative member in this evolutionary line
is the Berriasian late Barremian Meyasaurus (a more
primitive form than Barbatteius), whose phylogenetic
position within Lacertiformes is strongly supported
(Evans & Barbadillo 1997; Evans 2003; this study). This
may concord with the divergence dates proposed for lacertiforms, as extending from the Early Jurassic (e.g. Evans
& Barbadillo 1997; Vidal & Hedges 2005) to the Middle
or Late Jurassic (Evans 2003; Wiens et al. 2006), or even
to the Early Cretaceous (Mulcahy et al. 2012). The presence of Meyasaurus in south-western Europe suggests
that diversification and radiation of these early lacertiform
stocks took place at least in pre-Cenomanian times, even
if their palaeobiogeographical origins remain difficult to
estimate. Nevertheless, a constraint in the distribution of
these terrestrial vertebrates starts around the Early/Late
Cretaceous (approximately 112 Ma), due to the opening
of the South Atlantic Ocean (Torsvik et al. 2009;
Chaboureau et al. 2012). Subsequently Africa and South
America became isolated, while the discontinuous route
of the so-called Mediterranean Sill (Rage 2002) probably
became predominantly impracticable, due to the increasingly high global sea levels (Golonka & Kiessling 2002).
In these circumstances we suggest two possible scenarios.
The first scenario is that the origin and diversification of
basal lacertiforms took place somewhere in Asia (key fossils are still lacking) and some of their descendants
extended their distribution into Europe, Africa and South
America in pre-Cenomanian times. A subsequent radiation took place in the Campanian Maastrichtian from the
Ibero-Armorican landmass or directly from Africa (as a
transoceanic drift) onto the Transylvanian landmass, followed by insular evolution of Barbatteius. The second
scenario is that the origin and diversification of basal lacertiforms took place in Gondwana (before the split of
Africa and South America) and some of their descendants
extended their distribution into cratonic Europe in preCenomanian times; a subsequent immigration into Transylvanian landmass took place from the south-western
European landmass or directly from Africa, followed by
insular evolution of Barbatteius. Deciding which of these
two scenarios may be correct is hampered by the absence
of Cretaceous teiioid fossils from any of the potential continents of origin (i.e. South America, Africa or Asia).
Regardless of which scenario is correct, only the South
16
American stock of teiioids survived the Cretaceous/Cenozoic extinction event.
Supplemental material
Supplemental material for this article can be accessed
here: http://dx.doi.org/10.1080/14772019.2015.1025869
Conclusions
Barbatteius vremiri represents the first unambiguous Late
Cretaceous record of Teiidae in Laurasia and the first preMiocene fossil evidence of this group outside South
America. Barbatteius, consisting of a three-dimensional
partial skull and associated lower jaws, preserves a unique
combination of features, which allows its taxonomic
assignment to teiid lizards. An expanded cladistic analysis
recovers Barbatteius and Meyasaurus in a sister taxon
relationship within a more inclusive clade of Teiidae
(Teiinae C Tupinambinae); however, support for this sister taxon relationship is weak, because of the high number
of plesiomorphic character states in Meyasaurus and considerable missing morphological data for both taxa.
Barbatteius vremiri is a relatively large-sized lizard
provided with extensive osteodermal sculpture on its skull
roofing bones and suspensorium, and the outer surface of
this osteodermal crust also bearing the impressions of
cephalic scales. The weakly heterodont dentition without
enlarged posterior crushing teeth suggests that it fed on
arthropods, small vertebrates and plants.
Barbatteius adds to the previously reported Euramerican origin paramacellodid and borioteiioid lizards of
Hat eg Island, as a new distinctive element representing
Teiidae, suggestive of a more complex palaeobiogeographical history for taxa on the Transylvanian Landmass.
For both the madtsoiid snake Nidophis (Vasile et al.
2013) and the teiid lizard Barbatteius Gondwanan origins
may be presumed. However, given the lack of Cretaceous
teiioid fossils from Gondwanan territories, the above
assumption needs confirmation by future research.
Acknowledgements
The authors are deeply indebted to Prof. Wolfgang Bohme
and Dr Dennis R
odder, Zoologisches Forschungsmuseum
Alexander Koenig, Bonn, for their support in accessing
the Recent lizard skeletal collection. Dr Cristina Farcas
kindly helped producing the geological map of the Hat eg
Basin. Jean-Claude Rage (Museum National dHistoire
Naturelle, Paris), Randall L. Nydam (Midwestern University, Glendale), James D. Gardner (Royal Tyrrell
Museum, Drumheller) and an anonymous reviewer provided very helpful comments and suggestions that
improved the paper. The English was improved by Dr
James D. Gardner. This work was supported by the Romanian Ministry of Education and Research CNCS under
Grant [PN-II-IDPCE-2011-3-0381].
References
Amiot, R., Lecuyer, C., Buffetaut, E., Fluteau, F., Legendre, S.
& Martineau, F. 2004. Latitudinal temperature gradient during the Cretaceous Upper Campanian Middle Maastrichtian:
d18O record of continental vertebrates. Earth and Planetary
Science Letters, 226, 255 272.
Arias, F. J., Barrios, F. & Palavecino, A. 2013. Range extension and geographic distribution of the poorly known species, Contomastix leachei Peracca, 1897 (Squamata:
Teiidae). Check List, 9, 844 846.
Auge, M. 2005. Evolution des lezards du Paleogene en Europe.
M
emoires du Mus
eum National dHistoire Naturelle, 192,
1 369.
Ballinger, R. E., Nietfeldt, J. W. & Krupa, J. J. 1979. An
experimental analysis of the role of the tail in attaining high
running speed in Cnemidophorus sexlineatus (Reptilia:
Squamata: Lacertilia). Herpetologica, 35, 114 116.
Benton, M. J., Csiki, Z., Grigorescu, D., Redelstorff, R.,
Sander, P. M., Stein, K. & Weishampel, D. B. 2010. Dinosaurs and the island rule: the dwarfed dinosaurs from Hat eg
Island. Palaeogeography, Palaeoclimatology, Palaeoecology, 293, 438 454.
Bever, G. S., Bell, C. J. & Maisano, J. A. 2005. The ossified
braincase and cephalic ostedoderms of Shinisaurus crocodilurus (Squamata, Shinisauridae). Palaeontologia Electronica, 8.1.14, 1 36.
Bolet, A. & Evans, S. E. 2012. Lizards and amphisbaenians
(Reptilia, Squamata) from the late Eocene of Sosss (Catalonia, Spain). Palaeontologia Electronica, 16.1.8A, 1 23.
Borsuk-Bialynicka, M., Lubka, M. & B
ohme, W. 1999. A lizard from Baltic amber (Eocene) and the ancestry of the
crown group lacertids. Acta Palaeontologica Polonica, 44,
349 382.
Bremer, K. 1994. Branch support and tree stability. Cladistics,
10, 295 304.
Cavin, L., Tong, H., Boudad, L., Meister, C., Piuz, A.,
Tabouelle, J., Aarab, M., Amiot, M., Buffetaut, E., Dyke,
G. J., Hua, S. & Le Loeuff, J. 2010. Vertebrate assemblages from the early Late Cretaceous of southeastern
Morocco: an overview. Journal of African Earth Sciences,
57, 391 412.
nansk
Cer
y, A. & Auge, M. L. 2013. New species of the genus
Plesiolacerta (Squamata: Lacertidae) from the Upper Oligocene (MP28) of southern Germany and a revision of the type
species Plesiolacerta lydekkeri. Palaeontology, 56, 79 94.
Chaboureau, A. C., Donnadieu, Y., Sepulchre, P., Robin, C.,
Guillocheau, F. & Rohais, S. 2012. The Aptian evaporites
of the South Atlantic: a climatic paradox? Climate of the
Past, 177, 1315 1333.
Codrea, V. A. & Dica, E. P. 2005. Upper Cretaceous lowermost Miocene lithostratigraphic units exposed in Alba Iulia
Sebes Vint u de Jos area (SW Transylvanian basin). Studia Universitatis Babes-Bolyai, Geologia, 50, 19 26.
Codrea, V., Vremir M., Jipa C., Godefroit P., Csiki Z., Smith
T. & F
arcas , C. 2010. More than just Nopcsas Transylvanian dinosaurs: A look outside the Hat eg Basin.
Palaeogeography, Palaeoclimatology, Palaeoecology, 293,
391 405.
Conrad, J. L. 2004. Skull, mandible, and hyoid of Shinisaurus
crocodilurus Ahl (Squamata, Anguimorpha). Zoological
Journal of the Linnean Society, 141, 399 434.
Conrad, J. L. 2008. Phylogeny and systematics of Squamata
(Reptilia) based on morphology. Bulletin of the American
Museum of Natural History, 310, 1 182.
Csiki, Z. 2005. Sistematica, tafonomia si paleoecologia microvertebratelor si dinosaurilor saurischieni din Maastrichtianul Bazinului Hateg. Unpublished PhD thesis, University of
Bucharest, 527 pp. [in Romanian].
Csiki, Z. & Grigorescu, D. 1998. Small theropods of the Late
Cretaceous of the Hat eg Basin (Western Romania)
an
unexpected diversity at the top of the food chain. Oryctos, 1,
87 104.
Csiki, Z., Grigorescu, D. & R
uklin, M. 2005. A new multituberculate specimen from the Maastrichtian of Pui, Romania
and reassessment of affinities of Barbatodon. Acta Palaeontologica Romaniae, 5, 73 86.
Csiki, Z., Vremir, M., Brusatte, S. L. & Norell, M. A. 2010.
An aberrant island-dwelling theropod dinosaur from the
Late Cretaceous of Romania. Proceedings of the National
Academy of Sciences USA, 107, 15357 15361.
si, A., Pereda-Suberbiola, X.
Csiki-Sava, Z., Buffetaut, E., O
& Brusatte, S. L. 2015. Island life in the Cretaceous - faunal
composition, biogeography, evolution, and extinction of
landliving vertebrates on the Late Cretaceous European
archipelago. ZooKeys, 469, 1 161.
Duffaud, S. 2000. Les faunes damphibiens du Cr
etac
e sup
erieur
a lOligoc
ene inf
erieur en Europe: pal
eobiodiversit
e,
evolution, mise en place. Unpublished PhD thesis, Museum
national dHistoire Naturelle, Paris, 221 pp.
Eaton, J. G. & Kirkland, J. I. 2003. Diversity patterns of nonmarine Cretaceous vertebrates of the Western Interior Basin.
Pp. 263 313 in P. J. Harries (ed.) High-resolution
approaches in stratigraphic paleontology. Kluwer Academic Publishers, Boston.
Estes, R. 1970. Origin of the Recent North American lower vertebrate fauna: an inquiry into the fossil record. Forma et
Functio, 3, 139 163.
Estes, R. 1983. Sauria Terrestria, Amphisbaenia. Pp. 1 248 in
P. Wellnhofer (ed.) Handbuch der Pal
aoherpetologie. Volume 10A. Gustav Fischer Verlag, Stuttgart.
Estes, R., de Queiroz, K. & Gauthier, J. A. 1988. Phylogenetic
relationships within Squamata. Pp. 119 281 in R. Estes &
G. K. Pregill (eds) Phylogenetic relationships of the lizard
families. Stanford University Press, Stanford.
Evans, S. E. 1996. Parviraptor (Squamata: Anguimorpha) and
other lizards from the Morrison Formation at Fruita, Colorado. Pp. 243 248 in M. Morales (ed.) The continental
Jurassic. Bulletin of the Museum of Northern Arizona 60,
Flagstaff, Arizona.
Evans, S. E. 2003. At the feet of the dinosaurs: the early history
and radiation of lizards. Biological Reviews, 78, 513
551.
Evans, S. E. & Barbadillo, L. J. 1997. Early Cretaceous lizards
from Las Hoyas, Spain. Zoological Journal of the Linnean
Society, 119, 23 49.
Evans, S. E. & Barbadillo, L. J. 1999. A short-limbed lizard
from the Lower Cretaceous of Spain. Palaeontology, 60,
73 85.
Evans, S. E. & Chure, D. C. 1998a. Morrison lizards: structure,
relationships and biogeography. Modern Geology, 23,
35 48.
17
Evans, S. E. & Chure, D. C. 1998b. Paramacellodid lizard

skulls from the Jurassic Morrison Formation at Dinosaur
National Monument, Utah. Journal of Vertebrate Paleontology, 18, 99 114.
Evans, S. E. & Chure, D. C. 1999. Upper Jurassic lizards from
the Morrison Formation of Dinosaur National Monument,
Utah. Pp. 151 159 in D. D. Gillette (ed.) Vertebrate Paleontology in Utah. Utah Geological Survey Miscellaneous
Publication 99 1. Utah Geological Survey, Salt Lake City.
Evans, S. E. & Manabe, M. 1999. Early Cretaceous lizards
from the Okurodani Formation of Japan. Geobios, 32,
889 899.
Evans, S. E., Prasad, G. V. R. & Manhass B. K. 2002. Fossil
lizards from the Jurassic Kota formation of India. Journal of
Vertebrate Paleontology, 22, 299 312.
Folie, A. & Codrea, V. 2005. New lissamphibians and squamates from the Maastrichtian of Hat eg Basin, Romania.
Acta Palaeontologica Polonica, 50, 57 71.
Fuente, M. S. de la, Salgado, L., Albino, A., Baez, A. M.,
Bonaparte, J. F., Calvo, J. O., Codorniu, L., Coria, R. A.,
Gasparini, Z., Gonz
ales Riga, B. J., Novas, F. E. & Pol,
D. 2007. Tetrapodos continentales del Cretacico de la
Argentina: una sntesis actualizada. Ameghiniana, 11,
137 153.
Gao, K. -Q. & Norell, M. A. 2000. Taxonomic composition and
systematics of late Cretaceous lizard assemblages from
Ukhaa Tolgod and adjacent localities, Mongolian Gobi Desert. Bulletin of the American Museum of Natural History,
249, 1 118.
Gauthier, J. 1984. A cladistic analysis of the higher systematics
categories of the Diapsida. Unpublished PhD thesis, University of California, 564 pp.
Gauthier, J. A., Kearney, M., Maisano, J. A., Rieppel, O. &
Behlke, D. B. 2012. Assembling the squamate tree of life:
perspectives from the phenotype and the fossil record. Bulletin of the Peabody Museum of Natural History, 53, 3 308.
Goloboff, P. A., Farris, J. S. & Nixon, K. C. 2008. TNT, a free
program for phylogenetic analysis. Cladistics, 24, 774 786.
Golonka, J. & Kiessling, W. 2002. Phanerozoic time scale and
definition of time slices. Pp. 11 20 in W. Kiessling, E.
Fl
ugel & J. Golonka (eds) Phanerozoic reef patterns. SEPM
Special Publication 72, Tulsa, Oklahoma.
Gray, J. E. 1827. A synopsis of the genera of saurian reptiles, in
which some new genera are indicated, and the others
reviewed by actual examination. Philosophical Magazine, 2,
54 58.
Grigorescu, D. 1992. Nonmarine Cretaceous formations of
Romania. Pp. 142 164 in N. J. Mateer & P. J. Chen (eds)
Aspects of nonmarine Cretaceous geology. China Ocean
Press, Beijing.
Grigorescu, D. 2005. Rediscovery of a forgotten land: the last
three decades of research on the dinosaur-bearing deposits
from the Hat eg basin. Acta Palaeontologica Romaniae, 5,
191 204.
Grigorescu, D. & Hahn, G. 1987. The first multituberculate
teeth from the Upper Cretaceous of Europe (Romania). Geologica et Paleontologica, 21, 237 243.
Grigorescu, D., Hartenberger, J. L., R
adulescu, C., Samson,
P. M. & Sudre, J. 1985. Decouverte de mammiferes et dinosaures dans le Cretace superieur de Pui (Roumanie). Comptes rendus de lAcad
emie des Sciences, Paris, 301,
1365 1368.
Grigorescu, D., Venczel, M., Csiki, Z. & Limberea, R. 1999.
New latest Cretaceous microvertebrate fossil assemblages
18
from the Hat eg Basin (Romania). Geologie en Mijnbouw,

98, 310 314.
Harvey, M. B., Ugueto, G. B., Ronald L. & Gutberlet, R. L.
2012. Review of teiid morphology with a revised taxonomy
and phylogeny of the Teiidae (Lepidosauria: Squamata).
Zootaxa, 3459, 1 156.
Head, J. H., Paul M., Barrett, P. M. & Rayfield, E. J. 2009.
Neurocranial osteology and systematic relationships of Varanus (Megalania) prisca Owen, 1859 (Squamata: Varanidae). Zoological Journal of the Linnean Society, 155,
445 457.
Hedges, S. B. & Vidal, N. 2009. Lizards, snakes, and amphisbaenians (Squamata). Pp. 383 389 in S. B. Hedges & S.
Kumar (eds) The timetree of life. Oxford University Press,
New York.
Hoffstetter, R. 1967. Coup doeil sur les Sauriens (Lacertiliens)
des couches de Purbeck (Jurassique superieur dAngleterre).
Problemes actuels de pal
eontologie (Evolution des
vertebr
es), Centre National de la Recherche Scientifique,
163, 349 371.
Jianu, C. M. & Boekschoten, G. J. 1999. The Hat egisland or
outpost? Deinsea, 7, 195 198.
Jones, M. E. H., Anderson, C. J., Hipsley, C. A., M
uller, J.,
Evans, S. E. & Schoch, R. 2013. Integration of molecules
and new fossils supports a Triassic origin for Lepidosauria
(lizards, snakes, and tuatara). BMC Evolutionary Biology,
13, 208.
Khadkikar, A. S., Chamyal, L. S. & Ramesh, R. 2000. The
character and genesis of calcrete in Late Quaternary alluvial
deposits, Gujarat, western India, and its bearing on the interpretation of ancient climates. Palaeogeography, Palaeoclimatology, Palaeoecology, 162, 239 261.
Kosma, R. 2004. The dentition of Recent and fossil scincomorphan lizards (Lacertilia, Squamata) - systematics, functional
morphology, palecology. Unpublished PhD thesis, University of Hannover, 187 pp.
LaDuke, T. C., Krause, D.W., Scanlon, J. D. & Kley, N. J.
2010. A Late Cretaceous (Maastrichtian) snake assemblage
from the Maevarano Formation, Mahajanga Basin, Madagascar. Journal of Vertebrate Paleontology, 30, 109 139.
Longrich, N. R., Bhullar, B. A. S. & Gauthier, J. A. 2012.
Mass extinction of lizards and snakes at the Cretaceous Paleogene boundary. Proceedings of the National
Academy of Sciences USA, 109, 21396 21401.
Makadi, L. 2006. Bicuspidon aff. hatzegiensis (Squamata: Scincomorpha: Teiidae) from the Upper Cretaceous Csehbanya
Formation (Hungary, Bakony Mts). Acta Geologica Hungarica, 49, 373 385.
Makadi, L. 2013a. The first known chamopsiid lizard (Squamata) from the Upper Cretaceous of Europe (Csehbanya
Formation; Hungary, Bakony Mts). Annales de
Pal
eontologie, 99, 261 274.
Makadi, L. 2013b. A new polyglyphanodontine lizard (Squamata: Borioteiioidea) from the Late Cretaceous Ihark
ut
locality (Santonian, Hungary). Cretaceous Research, 46,
166 176.
Melinte-Dobrinescu, M. C. & Bojar, A.-V. 2010. Late Cretaceous carbon- and oxygen isotope stratigraphy, nannofossil
events and paleoclimate fluctuations in the Hat eg area (SW
Romania). Palaeogeography, Palaeoclimatology, Palaeoecology, 293, 295 305.
Mulcahy, D. G., Noonan, B. P., Moss, T., Townsend, T. M.,
Reeder, T. W., Sites, J. W. & Wiens, J. J. 2012. Estimating
divergence dates and evaluating dating methods using
phylogenomic and mitochondrial data in squamate reptiles.

Molecular Phylogenetics and Evolution, 65, 974 991.
Nopcsa, F. 1905. Zur Geologie der Gegend zwischen
Gyulafehervar, Deva, Ruszkabanya und der r
umanischen
Landesgrenze. Mitteilungen aus dem Jahrbuch der Ungarischen geologischen Anstalt, 14, 91 279.
Norell, M. A. & Gao, K. 1997. Braincase and phylogenetic relationships of Estesia mongoliensis from the Late Cretaceous
of the Gobi Desert and the recognition of a new clade of lizards. American Museum Novitates, 3211, 1 25.
Nydam, R. L. 1999. Polyglyphanodontinae (Squamata: Teiidae)
from the medial and Late Cretaceous: new taxa from Utah,
USA and Baja California del Norte, Mexico. Pp. 303 317
in D. D. Gillette (ed.) Vertebrate paleontology in Utah. Utah
Geological Survey Miscellaneous Publication 99-1. Utah
Geological Survey, Salt Lake City.
Nydam, R. L. 2002. Lizards of the Mussentuchit local fauna
(Albian Cenomanian boundary) and comments on the evolution of the Cretaceous lizard fauna of North America.
Journal of Vertebrate Paleontology, 22, 645 660.
Nydam, R. L. 2013. Squamates from the Jurassic and Cretaceous of North America. Palaeobiodiversity and Palaeoenvironments, 93, 535 565
Nydam, R. L., Caldwell, M. W. & Fanti, F. 2010. Borioteiioidean lizard skulls from Kleskun Hill (Wapiti Formation;
Upper Campanian), west-central Alberta, Canada. Journal
of Vertebrate Paleontology, 30, 1090 1099.
Nydam, R. L. & Cifelli, R. L. 2002. A new teiid lizard from the
Cedar Mountain Formation (Albian Cenomanian boundary) of Utah. Journal of Vertebrate Paleontology, 22,
276 285.
Nydam, R. L. & Cifelli, R. L. 2005. New data on the dentition
of the scincomorphan lizard Polyglyphanodon sternbergi.
Acta Palaeontologica Polonica, 50, 73 78.
Nydam, R. L., Eaton, J. & Sankey, J. 2007. New taxa of transversely toothed lizards (Squamata: Scincomorpha) and new
information on the evolutionary history of Teiids. Journal
of Paleontology, 81, 538 549.
Nydam, R. L. & Voci, G. E. 2007. Teiid-like scincomorphan
lizards from the Late Cretaceous (Campanian) of southern
Utah. Journal of Herpetology, 41, 211 219.
Oelrich, T. M. 1956. The anatomy of the head of Ctenosaura
pectinata (Iguanidae). Miscellaneous Publications, Museum
of Zoology, University of Michigan, 94, 1 122.
Oppel, M. 1811. Die Ordnungen, Familien, und Gatungen der
Reptilien. Joseph Lindauer, M
unchen, 87 pp.
Pana, I., Grigorescu, D., Csiki, Z. & Costea, C. 2001. Paleoecological significance of the continental gastropod assemblages from the Maastrichtian dinosaur beds of the Hat eg
Basin. Acta Paleontologica Romaniae, 3, 337 343.
Panaiotu, C. G. & Panaiotu, C. E. 2010. Palaeomagnetism of
the Upper Cretaceous Sânpetru Formation (Hat eg Basin,
South Carpathians). Palaeogeography, Palaeoclimatology,
Palaeoecology, 293, 343 352.
Presch, W. 1974. Evolutionary relationships and biogeography
of the macroteiid lizards (Family Teiidae, Subfamily Teiinae). Bulletin of the Southern California Academy of Sciences, 73, 23 72.
k, N. (in press). A revised Eurogondwana
Rabi, M. & Sebo
model: Late Cretaceous notosuchian crocodyliforms and
other vertebrate taxa suggest the retention of episodic faunal
links between Europe and Gondwana during most of the
Cretaceous. Gondwana Research, DOI: 10.1016/j.
gr.2014.09.015
Radulescu, C. & Samson, P.-M. 1996. The first multituberculate skull from the Late Cretaceous (Maastrichtian) of
Europe (Hat eg Basin, Romania). Anuarul Institutului Geologic al Rom^
aniei, 69, 177 178.
Radulescu, C. & Samson, P.-M. 1997. Late Cretaceous Multituberculata from the Hat eg Basin. Sargetia, 17, 247 255.
Rage, J. -C. 1981. Les continents Peri-atlantiques au Cretace
superieur: migrations des faunes continentales et problemes
paleogeographiques. Cretaceous Research, 2, 65 84.
Rage, J. -C. 1996. Les Madtsoiidae (Reptilia, Serpentes) du
Cretace superieur dEurope: temoins gondwaniens dune
dispersion transtethysienne. Comptes Rendus de lAcad
emie
de Sciences, Paris, 322, 603 608.
Rage, J. -C. 1999. Squamates (Reptilia) from the Upper Cretaceous of La~no (Basque Country, Spain). Estudios del Museo
Ciencias Naturales de Alava, 14, 121 133.
Rage, J. -C. 2002. The continental Late Cretaceous of Europe:
toward a better understanding. Comptes Rendus Palevol, 1,
257 258.
Rage, J. -C., Prasad, G. V. R. & Bajpai, S. 2004. Additional
snakes from uppermost Cretaceous (Maastrichtian) of India.
Cretaceous Research, 25, 425 434.
Rage, J. -C. & Werner, C. 1999. Mid-Cretaceous (Cenomanian) snakes from Wadi Abu Hashim, Sudan: the earliest
snake assemblage. Palaeontologia Africana, 35, 85 110.
Rieppel, O. 1985. The recessus scalae tympani and its bearing
on the classification of reptiles. Journal of Herpetology, 19,
373 384.
Rieppel, O. & Zaher, H. 2000. The braincases of mosasaurs and
Varanus, and the relationships of snakes. Zoological Journal
of the Linnean Society, 129, 489 514.
Royer, D. L. 1999. Depth to pedogenic carbonate horizon as a
paleoprecipitation indicator? Geology, 27, 1123 1126.
Royer, D. L. 2000. Depth to pedogenic carbonate horizon as
paleoprecipitation indicator? Reply. Geology, 28, 572 573.
Smith, K. T. 2009. A new lizard assemblage from the earliest
Eocene (zone Wa0) of the Bighorn Basin, Wyoming, USA:
biogeography during the warmest interval of the Cenozoic.
Journal of Systematic Palaeontology, 7, 299 358
Smith, T. & Codrea, V. 2003. New multituberculate mammals
from the Late Cretaceous of Transylvania (Romania). Pp. 51
in V. Codrea & P. Dica (eds) Abstracts volume. Fourth
National Symposium of Paleontology, Cluj-Napoca.
Therrien, F. 2005. Palaeoenvironments of the latest Cretaceous
(Maastrichtian) dinosaurs of Romania: insights from fluvial
deposits and paleosols of the Transylvanian and Hat eg
basins. Palaeogeography, Palaeoclimatology, Palaeoecology, 218, 15 56.
Torsvik, T. H., Rousse, S., Labails, C. & Smethurst, M. A.
2009. A new scheme for the opening of the South Atlantic
19
Ocean and the dissection of an Aptian salt basin. Geophysical Journal International, 177, 1315 1333.
Uetz, P. 2013. The Reptile Database. http://www.reptile-data
base.org/, accessed November, 2013.
Van Itterbeeck, J., S
as
aran, E., Codrea, V., S
as
aran, L. &
Bultynck, P. 2004. Sedimentology of the Upper Cretaceous
mammal and dinosaur bearing sites along the Râul Mare and
Barbat rivers. Hat eg Basin, Romania. Cretaceous Research,
25, 517 530.
Vasile, S ., Csiki, Z. & Venczel, M. 2013. A new madtsoiid
snake from the Upper Cretaceous of the Hat eg Basin, western Romania. Journal of Vertebrate Paleontology, 33,
1100 1119.
Venczel, M. & Csiki, Z. 2003. New frogs from the latest Cretaceous of Hat eg Basin, Romania. Acta Palaeontologica
Polonica, 48, 609 616.
Vidal, N. & Hedges, S. B. 2005. The phylogeny of squamate
reptiles (lizards, snakes, and amphisbaenians) inferred from
nine nuclear protein-coding genes. Comptes Rendus Biologies, 328, 1000 1008.
Vitt, L. J. & Pianka, E. R. 2004. Historical patterns in lizard
ecology: what teiids can tell us about lacertids. Pp. 139 157
in V. Perez-Mellado, N. Riera, & A. Perera (eds) The Biology of Lacertid lizards. Evolutionary and Ecological Perspectives. Institut Menorqu dEstudis, Recerca 8, Menorca.
si, A. & Dyke, G. J. 2011a. The first
Wang, X., Csiki, Z., O
definitive record of a fossil bird from the Upper Cretaceous
(Maastrichtian) of the Hat eg Basin, Romania. Journal of
Vertebrate Paleontology, 31, 227 230.
Wang, X., Dyke, G. J., Codrea, V., Godefroit, P. & Smith, T.
2011b. A euenantiornithine bird from the Late Cretaceous
Hat eg Basin of Romania. Acta Palaeontologica Polonica,
56, 853 857.
Weishampel, D. B., Grigorescu, D. & Norman, D. B. 1991.
The dinosaurs of Transylvania. National Geographic
Research, 7, 196 215.
Weishampel, D. B., Csiki, Z., Benton, M. J., Grigorescu, D. &
Codrea, V. 2010. Palaeobiogeographic relationships of the
Hat eg biota Between isolation and innovation. Palaeogeography, Palaeoclimatology, Palaeoecology, 293,
419 437.
Werner, C. & Rage J. -C. 1994. Mid-Cretaceous snakes from
Sudan. A preliminary report on an unexpectedly diverse
snake fauna. Comptes Rendus de lAcad
emie des Sciences,
Paris, 319, 247 252.
Wiens, J. J., Brandley, M. C. & Reeder, T. W. 2006. Why
does a trait evolve multiple times within a clade? Repeated
evolution of snakelike body form in squamate reptiles. Evolution, 60, 123 141.

A New Teiid Lizard From The Late Cretaceous of The Haţeg Basin, Romania and Its Phylogenetic and Palaeobiogeographical Relationships

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

A New Teiid Lizard From The Late Cretaceous of The Haţeg Basin, Romania and Its Phylogenetic and Palaeobiogeographical Relationships

Uploaded by

Copyright:

Available Formats

This article was downloaded by: [New York University]

On: 27 April 2015, At: 12:58

Journal of Systematic Palaeontology

A new teiid lizard from the Late Cretaceous of the

Mrton Venczel & Vlad A. Codrea

Department of Natural History, rii Criurilor Museum, Oradea, Romania

Department of Geology and Palaeontology, Babe-Bolyai University, Cluj-Napoca, Romania

Click for updates

PLEASE SCROLL DOWN FOR ARTICLE

Journal of Systematic Palaeontology, 2015

Downloaded by [New York University] at 12:58 27 April 2015

(Received 2 March 2014; accepted 23 January 2015)

*Corresponding author. Email: mvenczel@gmail.com

The Late Cretaceous tetrapod assemblages of Europe

Downloaded by [New York University] at 12:58 27 April 2015

Marton Venczel and Vlad A. Codrea

(Grigorescu & Hahn 1987; Radulescu & Samson 1996,

Geological setting and age

Material and methods

Downloaded by [New York University] at 12:58 27 April 2015

sediment removal, the skull and lower jaws were detached

Type species. Barbatteius vremiri gen. et sp. nov.

Marton Venczel and Vlad A. Codrea

Downloaded by [New York University] at 12:58 27 April 2015

teeth are heterodont, part of them with bi- or tricuspid

Cretaceous, Spain) in having extensive osteodermal crust

Downloaded by [New York University] at 12:58 27 April 2015

Eocene late Oligocene, France and Germany) in having a

Downloaded by [New York University] at 12:58 27 April 2015

Marton Venczel and Vlad A. Codrea

frontoparietal suture. At the anterolateral corner of the

Downloaded by [New York University] at 12:58 27 April 2015

about 100 . The posterolateral side of the supratemporal

Downloaded by [New York University] at 12:58 27 April 2015

Marton Venczel and Vlad A. Codrea

probably represents the distal tip of the postorbital ramus

Downloaded by [New York University] at 12:58 27 April 2015

Basisphenoid. This rather robust median bone forms part

Marton Venczel and Vlad A. Codrea

Downloaded by [New York University] at 12:58 27 April 2015

side and the posterior part of the right quadrate process

Downloaded by [New York University] at 12:58 27 April 2015

Angular. The angular extends on the ventral margin of

Downloaded by [New York University] at 12:58 27 April 2015

Marton Venczel and Vlad A. Codrea

crown lacertid Succinilacerta succinea, known from the

widely open adductor fossa, resulting from the extension

Downloaded by [New York University] at 12:58 27 April 2015

length were also calculated. The traditional search with

Marton Venczel and Vlad A. Codrea

Downloaded by [New York University] at 12:58 27 April 2015

Palaeobiogeographical and palaeoenvironmental implications

Figure 7. Relationships of Barbatteius vremiri gen. et sp. nov.,

Xenosaurus) and 411(1) prearticular pterygoideus process

In the Late Cretaceous, one of the European archipelago

Downloaded by [New York University] at 12:58 27 April 2015

Marton Venczel and Vlad A. Codrea

American stock of teiioids survived the Cretaceous/Cenozoic extinction event.

Downloaded by [New York University] at 12:58 27 April 2015

Downloaded by [New York University] at 12:58 27 April 2015

Evans, S. E. & Chure, D. C. 1998b. Paramacellodid lizard

Downloaded by [New York University] at 12:58 27 April 2015

Marton Venczel and Vlad A. Codrea

about 100 . The posterolateral side of the supratemporal