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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: mvenczel@gmail.com

� The Trustees of the Natural History Museum, London 2015. All Rights Reserved.

Journal of Systematic Palaeontology, 2015

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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.

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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

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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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