BARBATTEIIDAE, Codrea & Venczel & Solomon, 2017

BARBATTEIIDAE FAM. NOV.

Type genus: Barbatteius Venczel & Codrea, 2016

Distribution: Maastrichtian of Romania.

Diagnosis: Differs from all the other lizards by the following unique combination of characters: extensive osteodermal crust covers and strongly fuses to the skull roofing bones; outer surface of osteodermal sculpture bears the impressions of cephalic scales; parietal foramen and parietal ventral lappets are present; temporal muscles insert dorsally onto the lateral (i.e. descending) flanges of the parietal and supratemporal processes; frontals fused and constricted between the orbits; upper temporal fenestra is not occluded by either the postfrontal or postorbital; squamosal temporal ramus widens posteriorly possessing a dorsal process; the prearticular lacks a prearticular crest, whereas the pterygoideus process (=angular process of Oelrich, 1956) is well-defined.

Remarks: Presence of an extensive osteodermal crust on the skull roofing bones bearing impressions of the cephalic scales ( Smith, 2009) was considered a synapomorphy of Autarchoglossa ( Estes et al., 1988; Conrad, 2008; Gauthier et al., 2012; Longrich, Bhullar & Gauthier, 2012). However, the monophyletic status of Autarchoglossa clustering Anguimorpha and Scincomorpha is supported neither by molecular phylogenies (e.g. Vidal & Hedges, 2009) nor by combined morphological and molecular phylogenies ( Müller et al., 2011; Reeder et al., 2015). Within Laterata, except for some rhineurids, amphisbaenians lack a dermal sculpturing on the skull roofing bones ( Estes et al., 1988). Nevertheless, Cryptolacerta hassiaca Müller et al., 2011 emerges as transitional between lacertids and amphisbaenians possessing an extensive dermal sculpturing on the skull roof ( Müller et al., 2011). The dorsum of Meyasaurus , a possible teiioid known from the Early Cretaceous of Spain ( Evans & Barbadillo, 1997), is also ornamented by a vermiculate sculpture with deep grooves marking the original positions of the overlying scale impressions ( Richter, 1994; Borsuk-Białynicka, Lubka & Böhme, 1999) ( Fig. 2H). In some Recent teiids (e.g. Ameiva , Cnemidophorus and Kentropyx ) the dermal sculpturing on the skull roofing bones is present ( Presch, 1974; Estes, 1983) ( Fig. 2E, F), although less extensive when compared to Barbatteidae ( Fig. 2A–D), else it is completely lost as in Callopistes , Dracaena and Tupinambis ( Venczel & Codrea, 2016) ( Fig. 2G). The osteodermal crust in lacertids consists of several large scales ( Friederich, 1978; Borsuk-Białynicka et al., 1999; Čerňanský & Augé, 2013) ( Fig. 2I), whereas in Recent teiids there is a trend of strong fragmentation of the parietal scales ( Cope, 1869; Harvey et al., 2012). The osteodermal crust in Barbatteiidae appears intermediary between lacertids and teiids (i.e. more fragmented than in lacertids, but less fragmented than in teiids), and distinctively it covers the postfrontal–postorbital and the posterodorsal part of the squamosal ( Venczel & Codrea, 2016).

In Recent teiids the parietal foramen is lacking, or infrequently present, as noticed by Estes (1983: 75) in a Holocene specimen of Teius teyou . In Barbatteiidae the parietal foramen is uniformly present in all the specimens assigned to the new barbatteiid taxon from ODA (n = 3, see below), located within the interparietal scute. However, the presence of a parietal foramen could not be unambiguously documented in the holotype of Barbatteius vremiri because the interparietal scute of that specimen is partially damaged ( Venczel & Codrea, 2016).

The ventral lappets of the parietal (=parietal tabs) are constantly present in members of lacertids, teiids and gymnophthalmids, but are lacking in amphisbaenians ( Conrad, 2008), presumably due to their highly modified skulls. The presence of this structure in some iguanids, agamids, chamaeleontids, cordylids and scincids has been considered as of independent origin ( Estes et al., 1988).

In Barbatteiidae the parietal temporal muscles are inserted dorsally on the parietal table and on the parietal supratemporal processes, thus retaining the plesiomorphic squamate condition similar to other teiioids, amphisbaenians and Meyasaurus . In lacertids the temporal muscles originate ventrally on the parietal table and on the parietal supratemporal processes, whereas some gymnophthalmids retain an intermediate condition (e.g. Pholidobolus ), in which the temporal muscles originate ventrally on the parietal table and dorsally on the supratemporal processes ( Gauthier et al., 2012).

In Barbatteiidae the prearticular lacks a prearticular crest, whereas in Meyasaurus it is faintly developed. The pterygoideus process (=angular process of Oelrich, 1956) in both taxa is well-developed. In lacertids the prearticular is devoid of a prearticular crest and pterygoideus process, whereas in teiids and some gymnophthalmids (e.g. Pholidobolus View in CoL ) there is a well-developed prearticular crest and the pterygoideus process is imbedded in the prearticular crest ( Conrad, 2008).