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A new teiid lizard from the Late Cretaceous of theHaţeg Basin, Romania and its phylogenetic andpalaeobiogeographical relationshipsMárton Venczela & Vlad A. Codreab
a Department of Natural History, Ţării Crişurilor Museum, Oradea, Romaniab Department of Geology and Palaeontology, Babeş-Bolyai University, Cluj-Napoca, RomaniaPublished online: 27 Apr 2015.
To cite this article: Márton Venczel & Vlad A. Codrea (2015): A new teiid lizard from the Late Cretaceous of the HaţegBasin, 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
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A new teiid lizard from the Late Cretaceous of the Hateg Basin, Romania and itsphylogenetic and palaeobiogeographical relationships
M�arton Venczela* and Vlad A. Codreab
aDepartment of Natural History, T �arii Crisurilor Museum, Oradea, Romania; bDepartment of Geology and Palaeontology,Babes-Bolyai University, Cluj-Napoca, Romania
(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 lowerjaws from the Pui Islaz locality (Late Cretaceous, early Maastrichtian) in the Hateg Basin, western Romania. Barbatteiusvremiri gen. et sp. nov. is diagnosed by a unique combination of symplesiomorphies and synapomorphies. A nested set ofsynapomorphies support assigning Barbatteius to Teiidae as the first unambiguous Late Cretaceous record of this familyfrom Laurasia. Barbatteius differs from other teiids by having more extensive osteodermal sculpture on the skull roof andsuspensorium, and by a pentagonal occipital osteoscute exhibiting more or less parallel lateral margins. Barbatteius is alarge-bodied lizard, estimated to be up to 800 mm in total length. It has weakly heterodont dentition, but without enlargedposterior crushing teeth, suggesting that it fed on arthropods, small vertebrates and plants. The mix of taxa with affinities toEuramerica (paramacellodid and borioteiioid lizards) and Gondwana (madtsoiid snakes and the teiid Barbatteius) currentlyknown for the Maastrichtian squamate assemblage from Hateg Basin supports the growing realization that ‘Hateg 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 super-
continent 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 ter-
restrial 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; Mak�adi 2013a, b), whereassnakes 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).
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 fau-
nas is represented by the peculiar latest Cretaceous verte-
brate fauna of ‘Hateg Island’, western Romania, which
was first reported by Nopcsa (1905). Investigations of
Benton et al. (2010) have shown that at least some of the
Hateg dinosaurs (e.g. the herbivorous genera likeMagyar-
osaurus, 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 Nopcsa’s times (e.g. turtles, crocodili-
ans, dinosaurs and pterosaurs), including fishes (Grigor-
escu 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
*Corresponding author. Email: [email protected]
� The Trustees of the Natural History Museum, London 2015. All Rights Reserved.
Journal of Systematic Palaeontology, 2015
http://dx.doi.org/10.1080/14772019.2015.1025869
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(Grigorescu & Hahn 1987; R�adulescu & Samson 1996,
1997; Csiki et al. 2005; Codrea et al. 2010).
Research on squamate reptiles from the Hateg Basin
started with reports by Grigorescu et al. (1999) of isolated
bones belonging to indeterminate anguimorph and scinco-
morph 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 (Bicuspi-
don hatzegiensis, Folie & Codrea 2005) and to indetermi-
nate 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. Bicuspi-
don aff. hatzegiensis; Mak�adi 2006). Another novelty for
the Hateg 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 M�aty�as Vremir during a field
inspection at the Pui Islaz locality (Fig. 1), in uppermost
Cretaceous (lower Maastrichtian) rocks in the river-bed of
B�arbat stream in the Hateg 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) evalu-
ate the phylogenetic relationships of the new taxon; and
(4) comment on its palaeogeographical and palaeoenvir-
onmental implications.
Geological setting and age
Stable isotope studies (Melinte-Dobrinescu & Bojar 2010)
indicate that marine sedimentation in the Hateg Basin
closed around the latest Campanian and that continental
deposition started in the earliest Maastrichtian at the lat-
est. The continental vertebrate-bearing strata of the Hateg
Basin are separated into two distinct lithostratigraphical
units, the Densus-Ciula and the Sanpetru formations
(Grigorescu 1992). The Pui beds, from where the speci-
men originates, are correlated with the lower part of the
Sanpetru Formation (Nopcsa 1905; Grigorescu et al.
1985, 1999). Palaeomagnetic studies of Panaiotu & Pan-
aiotu (2010) indicate that the continental sequence along
the Sibisel valley (Sanpetru Formation) was deposited
between chron 32n.1 and the end of chron 31n (about 72
to 67.8 Ma). Although the palaeoenvironment of the
Sanpetru Formation typically is represented by hydromor-
phic (immature) palaeosols dominated by areas of
impeded drainage, the Pui beds consist of mature palaeo-
sols dominated by moderately to well-drained floodplains
(Therrien 2005). The strata of the Pui beds are subhori-
zontal, 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 docu-
mented by red coloration and the presence of calcrete nod-
ules or continuous calcrete layers (Van Itterbeeck et al.
2004; Csiki et al. 2005; Therrien 2005). Occasional slick-
ensides, 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 (Pan�a et al. 2001). The micro-
vertebrates in this horizon are represented by albanerpe-
tontids, 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 articu-
lated and slightly displaced bones in the skull roof, sus-
pensorium, 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
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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 Crisurilor 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 ‘B�arbat’ river in the Hateg
Basin, which transects the Pui beds that yielded the holo-
type 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, neurocra-
nium, posterior part of the palatal complex and associated
fragmentary lower jaws.
Diagnosis. Large Late Cretaceous teiid lizard with esti-
mated total body length up to 800 mm. It differs from all
other lizards by the following unique combination of fea-
tures: upper temporal fenestra is not occluded by the post-
orbital; 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 pres-
ent; epipterygoid�parietal contact overlaps parietal tem-
poral 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 1. Geological map of the southern part of the Hateg Basin and location of the Pui Islaz fossil locality.
New teiid lizard from the Late Cretaceous of the Hateg Basin 3
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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 den-
tary 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 bear-
ing the impressions of cephalic scales. Differs further
from the possible crown lacertids Succinilacerta (middle
Eocene, Poland and Lithuania) and Plesiolacerta (middle
Figure 2. Partial skull roofing bones and suspensorium in the holotype (UBB V.440) of Barbatteius vremiri gen. et sp. nov. Frontoparie-tal region and supratemporal arch in A, dorsal and B, ventral views. Photographs (above) and interpretive figures using different levels ofgrey to highlight particular bones (below). Abbreviations: adt: anterodorsal tuberosity; app: alar process of prootic; ccf: crista craniifrontalis; ept: epipterygoid; fr: frontal; ipvl: imprint of parietal ventral lappet; ju: jugal; os: occipital scute; pa: parietal; pffp: frontal pro-cess 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. Scalebar D 5 mm.
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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 M�aty�as 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, Hateg Basin, Transylvania, Romania,
Europe; Late Cretaceous, early Maastrichtian, Sanpetru
Formation.
Description
General observations. It is assumed that the fragmen-
tary skull and the lower jaws belonged to the same indi-
vidual, because they were preserved in contact, although
the jaws had shifted slightly posteriorly and rotated
90�100� counter-clockwise from their original anatomi-
cal 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 3. Skull roofing bones and suspensorium in the holotype (UBB V.440) of Barbatteius vremiri gen. et sp. nov., and RecentAmeiva ameiva (Teiidae). A, frontoparietal region with parietal downgrowth of Barbatteius in left lateral view. B, reconstruction of Bar-batteius skull in dorsal view (missing part is Recent Tupinambis teguixin ZFMK 53531). C, partial skull of Ameiva ameiva (ZFMK59021) in dorsal view with details of osteodermal scutes covering the frontoparietal region. D, interpretive drawings of osteodermalscutes 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.
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Frontal. Only the posterior part of the extensively sculp-
tured, 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 osteo-
scutes (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 cor-
ner of the frontal is braced ventrolaterally by the forking
postfrontal. In ventral view, several irregular cracks
extend longitudinally and transversally across the
frontal’s ventral surface (Fig. 2B). The crista cranii fronta-
lis, 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 cov-
ered partially by sediment (Fig. 2A). The entire dorsal sur-
face 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 osteo-
dermal 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 frontopar-
ietal 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 fron-
toparietal scutes have sagittal contact anterior to the inter-
parietal scute, and similar to lacertiform lizards (e.g.�Cer�nansk�y & Aug�e 2013) the frontoparietal scutes extendforward 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 pro-
cesses are wide and robust, diverging posterolaterally at
Figure 4. Partial skull roofing bones and suspensorium in the holotype (UBB V.440) of Barbatteius vremiri gen. et sp. nov. A, supra-temporal processes of parietal, supratemporals and squamosals in posterior view; B, supratemporal arch and partial frontal in right lateralview. 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 ofsquamosal; st: supratemporal. Scale bar D 5 mm.
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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 parie-
tal lappets and the overlying frontal (Estes et al. 1988) are
readily apparent on each side. Nevertheless, the right ven-
tral parietal lappet is broken off leaving on the frontal’s
posteroventral margin an imprint of a deep and roughly
triangular concavity. The crista cranii parietalis is moder-
ately high and there is a well-preserved parietal down-
growth on the left lateral side; a fragmentary bone in front
of the alar process of the prootic, embedded in sediment
and cemented to the parietal’s ventral side, probably rep-
resents the vestiges of the epipterygoid, which in the liv-
ing animal contacted the parietal downgrowths (Fig. 3A).
The supratemporal processes are relatively wide and mod-
erately long. Their posteriormost parts are turned down-
ward 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 ante-
rodorsal 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 pro-
cess 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 postorbi-
tal�squamosal joint. The posterior ramus of the postfron-
tal�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 squa-
mosal; its posterior terminus, similar to other crown liz-
ards, is situated at the medial side of the squamosal
(Gauthier et al. 2012).
Jugal. A small piece of bone partly embedded in sedi-
ment on the left side of the skull, situated lateral to the
anterodorsal tuberosity of the postfrontal�postorbital,
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 ventralview. 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: cristainterfenestralis; 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: palatineartery foramen; bs: basisphenoid; oto: otooccipital; po: prootic; poV: posterior opening of Vidian canal; ppp: posterior process ofprootic; pt: pterygoid; qu: quadrate; quf: quadrate foramen; so: supraoccipital; sot: sphenooccipital tubercle; stp: supratrigeminal pro-cess; tn: trigeminal notch. White arrows point to furrows produced presumably by plant roots. Scale bars D 5 mm.
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probably represents the distal tip of the postorbital ramus
of the jugal (Fig. 2B); the outer surface of this bony frag-
ment 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 sedi-
ment; 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 dor-
sally 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 osteo-
scutes: a larger one facing dorsally and a smaller one fac-
ing 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 jugal’s postorbital ramus is pre-
served on the left lateral side of the postorbital. The squa-
mosal 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 parietal’s supratemporal processes
(Fig. 4A). In dorsal view the supratemporal normally
would be hidden by the ascending process of the squamo-
sal; 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 parietal’s 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 lab-
yrinth (Oelrich 1956). The alar process is broken off, but
it was found embedded in sediment, adhering to the ven-
tral part of the parietal (Fig. 2B). In ventral view, this
structure has an anteriorly open V shape contacting at its
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 andright lower jaw in dorsomedial view; B, partial left lower jaw in medial view and right lower jaw in lateral view. Arrow points to furrowpresumably 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 mylohyoidforamen; san: surangular; sds: subdental shelf; sp: splenial; sur: surangular ridge. Scale bars D 5 mm.
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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 pre-
served (Fig. 5A, C). Its ventral part diverges laterally to
the posterior border of the trigeminal notch. The supratri-
geminal process appears as a bulging prominence on the
medial side of the alar process (Fig. 5C). The inferior pro-
cess 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 pro-
tractor 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 lat-
erally. 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 ventrome-
dial 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 ori-
entation. 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 ven-
tral view, the basipterygoid processes are rather large and
prominent; they are roughly as wide as long, with the ven-
tral 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 cultri-
form process is preserved. On the right side, the sphenooc-
cipital 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 sub-
divided 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 basiptery-
goid process, respectively (Fig. 5C).
Otooccipital. This is a composite bone derived in most
squamates from the fusion of the exoccipital and opis-
thotic (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 mag-
num. The fenestra vestibuli (D foramen ovale) lies in a
dorsoposterior position relative to the recessus scalae tym-
pani (D occipital recess) and sphenooccipital tubercle; its
anterior and dorsal margins are delimited by the prootic
(Norell & Gao 1997), whereas its ventral margin is demar-
cated by the prominent and roughly horizontally displayed
interfenestral crest (Fig. 5B, D). The recessus scalae tym-
pani lays just posterior to and on the same level with the
sphenooccipital tubercle, as an elongated hole, bordered
anterodorsally by the interfenestral crest and posteroven-
trally 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 medi-
ally the otooccipital wall, as seen in Recent lacertids; the
anterodorsal margin of the MARST reaches the postero-
ventral 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 reces-
sus 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 sphenooc-
cipital 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 basioccipital’s 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 basisphe-
noid processes. A breakage line is observed on the right
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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 poste-
rior 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 articu-
lation and position. The left quadrate, except for the miss-
ing 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� fromits original position (Fig. 5A, C). In lateral view, it
appears straight and rather robust. The well-developed
cephalic condyle articulates with the squamosal dorsolat-
erally; the cemented sediment still fixes the tip of the
squamosal’s left ventral ramus to the cephalic condyle.
The mesioventral lappet for sliding contact with the poste-
rior 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 anteri-
orly and is indented by a distinct notch for insertion of the
posterior process of the pterygoid. The mandibular con-
dyle 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 posi-
tioned near the mandibular condyle on the pterygoid lap-
pet (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 deter-
mine whether they had an anteroventral or a posteroven-
tral 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 dis-
plays 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 den-
tary 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 posterolat-
eral border is notched for the surangular joint. The poster-
oventral 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 hetero-
dont. 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 bicus-
pid, whereas the ninth is already tricuspid. The 10th�20th
teeth are the largest and tallest, and their bases are broad-
ened labiolingually. They are gently curved posterolin-
gually 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 replace-
ment was present in all dentary teeth. A semicircular fur-
row, similar to those seen on the basioccipital surface, is
present on the lingual side of the left lower jaw and proba-
bly also is the trace of a plant root (Fig. 6B).
Coronoid. The coronoid is triradiate, covering the poste-
rior margins of the dentary and clasping it labiolingually
(Fig. 6A). The anteromedial ramus articulates with the
splenial on the lingual side, whereas the anterolateral den-
tary 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 coro-
noid 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.
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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 pro-
cess 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 poste-
rior surangular foramen cannot be determined with cer-
tainty because of damages posterodorsally on the
mandible’s labial side. Broken parts of the surangular dor-
sal 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 medio-
ventrally. 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-devel-
oped pterygoideus process (D angular process). The pre-
articular crest is absent.
Remarks
The fragmentary skull and associated lower jaws of Bar-
batteius are notable for their three-dimensional appear-
ance 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 roof-
ing 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 squamosal’s postorbital
ramus (Fig. 4). Separable osteodermal crust occurs in
cordylids, scincids, lacertids, xenosaurs, anguids,
Lanthanotus and helodermatids, as well as in some vara-
nids and the iguanid Amblyrhynchus, which is considered
by Estes et al. (1988) a synapomorphy of both Anguimor-
pha 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 occur-
rence 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, Isodontosauri-
dae, 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, Hoplocer-
cinae, Rhineuridae, Tchingisaurus, Sineoamphisbaena)
(Nydam et al. 2010; Gauthier et al. 2012). The presence
of vermiculate sculpture on these cephalic scale impres-
sions 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 Kentro-
pyx) 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 Sakura-
saurus shokawensis, known from the Early Cretaceous,
Japan, the dorsal surface of the paired frontals (and proba-
bly 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 fea-
tures probably shared by early lacertoid lizards. In Barbat-
teius the pileus morphology (i.e. the pattern of scalation
on the skull; e.g. �Cer�nansk�y & Aug�e 2013) seems transi-
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 sag-
ittal contact, extending forward to the frontal scute
(�Cer�nansk�y & Aug�e 2013). However, in the possible
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crown lacertid Succinilacerta succinea, known from the
Middle Eocene Baltic amber of Poland and Lithuania
(Borsuk-Bia»ynicka et al.1999), and Plesiolacerta lydek-
keri, known from the Middle Eocene�Early Oligocene
(MP14�MP21) of France (�Cer�nansk�y & Aug�e 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
Late Oligocene of southern Germany (�Cer�nansk�y & Aug�e2013), in having similarly narrow occipital scute, but dif-
fers 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 frontal’s posterolateral margin
a sutural imprint of roughly triangular shape. In the teiid-
like 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, Polyglypha-
nodon and Sineoamphisbaena) the presence of this struc-
ture 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. Cher-
minsaurus, 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 primi-
tively by the otooccipital (e.g. in lacertids; see Gauthier et
al. 2012), whereas in teiioids and Barbatteius it is bor-
dered 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 over-
laps the coronoid (Evans & Manabe 1999). The weakly
developed subdental shelf that is present in Barbatteius
may represent, after Gauthier et al. (2012), a synapomor-
phy of Teiidae (shared also by Meyasaurus). An inflated,
widely open adductor fossa, resulting from the extension
of the adductor mandibulae posterior muscle into
Meckel’s 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 lacerti-
form lizards, is as follows: ventral margin of dentary and
subdental shelf are arched ventrally (Aug�e 2005; Bolet &Evans 2012); Meckel’s groove is open up to the symphy-
sis and covered by the hypertrophied splenial that almost
reaches the symphysis (Aug�e 2005; �Cer�nansk�y & Aug�e2013); mandibular symphysis is relatively small; the coro-
noid process of the dentary does not extend onto the ante-
rolateral surface of the coronoid, unlike that of most
scincoids in which the posterolabial process of the dentary
is large and extensively overlying the coronoid (Aug�e2005; Conrad 2008); the dentition is heterodont, consist-
ing 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 differ-
entiate from those of Asian Polyglyphanodontia, known
from the Campanian of Mongolia, and Euramerican Bor-
ioteioiidea, known from the lower Cenomanian�Maastrichtian of North America, Santonian of Ihark�ut,Hungary and Maastrichtian of Hateg 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 Long-
rich et al. 2012). The Chamopsiidae (i.e. Chamops,
Cnephasaurus, Gerontoseps, Glyptogenys, Haptosphenus,
Harmondontosaurus, Leptochamops, Meniscognathus,
Pelsochamops, Socognathus, Sphenosiagon, Stypodonto-
saurus) 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 U-
shaped 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 Pelsocha-
mops (Santonian, Ihark�ut, Hungary) in having its coronoid
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not fused to the dentary (Mak�adi 2013a). Barbatteius dif-fers from Haptosphenus, an aberrant taxon having its den-
tary, 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 polyglyphano-
dontian 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 Eur-
american Polyglyphanodontini (sensu Nydam et al. 2007,
see also Mak�adi 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 replace-
ment in adults (Bicuspidon, Dicothodon, Distortodon,
Paraglyphanodon, Peneteius (D Manangysaurus) and
Polyglyphanodon). The dentition of Barbatteius, consist-
ing 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 char-
acters 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 ofMeyasaurus), 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 taxo-
nomic units (OTU) and 610 multistate characters, we per-
formed 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 recon-
nection) 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
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 Gauth-
ier 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 origi-
nal 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 Bar-
batteius (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. Barbat-
teius is recovered, similar to the first analysis, as the sister
taxon of Meyasaurus, whereas the clade of Barbatteius CMeyasaurus appears within Teiidae as the sister taxon of
Teiinae and Tupinambinae. More inclusive clades within
lacertiforms are the Gymnophthalmidae (Pholidobolus CColobosaura), 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 D12), nodal support for Lacertiformes and the rest of its
less-inclusive clades was moderate (bootstrap value below
60%, decay indexD 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), con-
ceivably 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 synapo-
morphies 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 unambigu-
ous synapomorphies: 78(2) postorbital overlaps squamo-
sal dorsally, 90(0) parietal temporal muscles originate
dorsally on parietal table and parietal supratemporal pro-
cesses (shared also by Meyasaurus), 294(1) epiptery-
goid�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) squamo-
sal temporal ramus widens posteriorly (also present in
New teiid lizard from the Late Cretaceous of the Hateg Basin 13
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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 Pol-
yglyphanodon sternbergi and Gilmoreteius (D Macroce-
phalosaurus) 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 Cre-
taceous of North America (Nydam 2013). They also have
been reported from the European Late Cretaceous
(e.g. Bicuspidon, Chamops and Distortodon from the San-
tonian of Ihark�ut, Hungary (Mak�adi 2006, 2013a, b), andBicuspidon from the Maastrichtian of the Hateg 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 lat-
eratan groups in the Jurassic and split of Teiidae and Gym-
nophthalmidae in the Late Cretaceous (about 86 Ma).
Palaeobiogeographical and palaeoenviron-
mental implications
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
Hateg 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 Hateg 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). Palaeo-
geographical reconstructions (see Benton et al. 2010)
depict ‘Hateg Island’ as having been circumscribed by
deep marine basins, separated from the nearest land-
masses by about 200�300 km and lying within the sub-
tropical belt at approximately latitude 27� N (Grigorescu
2005; Panaiotu & Panaiotu 2010). Other emerged terrains
relatively close to ‘Hateg 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 estab-
lished between these emerged areas and the Ibero-Armori-
can 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 Maas-
trichtian, 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 ‘Hateg Island’ palaeoecosystems, Barbatteius
appears as a large-bodied lizard with generalized teiid
attributes. The Teiioidea shifted from the general lacerti-
form body plan, in elongation of the body and the appar-
ent thickening and lengthening of the tail (Vitt & Pianka
2004), which are considered important factors in locomo-
tion (Ballinger et al. 1979). The resulting higher size-
Figure 7. Relationships of Barbatteius vremiri gen. et sp. nov.,based on our phylogenetic analysis. A, strict consensus of fourtrees generated with TNT, based on 57 OTUs with 610 pheno-typic characters (derived from Gauthier et al. 2012),Meyasaurus(based on published data of Evans & Barbadillo 1997) and Bar-batteius (see Supplementary Material). Numbers below nodesindicate decay indices of Bremer (D steps). B, the clade of Lac-ertiformes and supporting unambiguous synapomorphies atnodes. The character numbers and character states are fromGauthier et al. (2012) and Longrich et al. (2012).
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specific mass in Teiioidea is in agreement with their
nearly exclusive restriction to terrestrial microhabitats
(Vitt & Pianka 2004). The total body length of Barbat-
teius, 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 hetero-
dont dentition without enlarged posterior crushing teeth,
as seen in many recent teiids (Kosma 2004 and references
therein), suggests that Barbatteius mainly fed on arthro-
pods (e.g. insects, millipedes and spiders), small verte-
brates (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 ‘Hateg 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 dromaeo-
saurid theropod Balaur bondoc (Csiki et al. 2010). Alloda-
posuchus 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-cor-
dylid grade lizards, approaching Paramacellodus in mor-
phology, 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 infre-
quens in the Santonian of Ihark�ut, Hungary (Mak�adi2006, 2013a, b). On the other hand, the presence of the
sebecosuchian crocodyliform Doratodon (Weishampel et
al. 2010; Rabi & Sebo��k in press) and madtsoiid snakes
(Folie & Codrea 2005; Vasile et al. 2013), together with
teiid lizards, strengthens the view that some faunal ele-
ments of the Transylvanian landmass were of Gondwanan
origin. For the basal alethinophidian snake Nidophis insu-
laris, reported recently from the Hateg Basin, Vasile et al.
(2013) advanced the idea of an early (i.e. pre-Cenoma-
nian) dispersal event of madtsoiid snakes from Africa into
Europe, followed by subsequent diversification and distri-
bution across Alpine Europe (Hateg) and cratonic south-
western Europe (Spain). A similar scenario might be
applicable to Barbatteius as well. According to Estes
(1970, 1983), the geologically oldest record of this pres-
ently American distributed group (i.e. Teiidae and Gym-
nophthalmidae) 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 lac-
ertiforms, 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 pres-
ence 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 increas-
ingly 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 fos-
sils are still lacking) and some of their descendants
extended their distribution into Europe, Africa and South
America in pre-Cenomanian times. A subsequent radia-
tion took place in the Campanian�Maastrichtian from the
Ibero-Armorican landmass or directly from Africa (as a
transoceanic drift) onto the Transylvanian landmass, fol-
lowed by insular evolution of Barbatteius. The second
scenario is that the origin and diversification of basal lac-
ertiforms took place in Gondwana (before the split of
Africa and South America) and some of their descendants
extended their distribution into cratonic Europe in pre-
Cenomanian times; a subsequent immigration into Tran-
sylvanian 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 con-
tinents of origin (i.e. South America, Africa or Asia).
Regardless of which scenario is correct, only the South
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American stock of teiioids survived the Cretaceous/Ceno-
zoic extinction event.
Conclusions
Barbatteius vremiri represents the first unambiguous Late
Cretaceous record of Teiidae in Laurasia and the first pre-
Miocene 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 sis-
ter taxon relationship is weak, because of the high number
of plesiomorphic character states in Meyasaurus and con-
siderable 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 Eurameri-
can origin paramacellodid and borioteiioid lizards of
‘Hateg Island’, as a new distinctive element representing
Teiidae, suggestive of a more complex palaeobiogeo-
graphical 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 B€ohme
and Dr Dennis R€odder, Zoologisches Forschungsmuseum
Alexander Koenig, Bonn, for their support in accessing
the Recent lizard skeletal collection. Dr Cristina F�arcaskindly helped producing the geological map of the Hateg
Basin. Jean-Claude Rage (Mus�eum National d’Histoire
Naturelle, Paris), Randall L. Nydam (Midwestern Univer-
sity, Glendale), James D. Gardner (Royal Tyrrell
Museum, Drumheller) and an anonymous reviewer pro-
vided very helpful comments and suggestions that
improved the paper. The English was improved by Dr
James D. Gardner. This work was supported by the Roma-
nian Ministry of Education and Research CNCS under
Grant [PN-II-IDPCE-2011-3-0381].
Supplemental material
Supplemental material for this article can be accessed
here: http://dx.doi.org/10.1080/14772019.2015.1025869
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