Cladistics Cladistics (2021) 1–38 10.1111/cla.12460 Phylogenetic value of jaw elements of lacertid lizards (Squamata: Lacertoidea): a case study with Oligocene material from France Lukardis Charlotte Marie Wencker*a , Emanuel Tschoppa,b,c , Andrea Villaa,d , Marc Louis Augee and Massimo Delfinoa,f a a degli Studi di Torino, Via Valperga Caluso, 35, Turin, 10125, Italy; bCentrum f€ Dipartimento di Scienze della Terra, Universit ur Naturkunde, at Hamburg, Martin-Luther-King-Platz 3, Hamburg, 20146, Germany; cDivision of Paleontology, American Museum of Natural History, Universit€ Central Park West & 79th Street, New York, NY, 10024, USA; dSNSB-Bayerische Staatssammlung f€ ur Pal€ aontologie und Geologie, Richard- Wagner-Straße 10, Munich, 80333, Germany; eCR2P, UMR 7207 CNRS, MNHN, Mus eum National d’Histoire naturelle, 57 rue Cuvier, Paris, 75231, France; fInstitut Catal a de Paleontologia Miquel Crusafont, Universitat Aut onoma de Barcelona, Edifici ICTA-ICP, Carrer de les Columnes s/n, Campus de la UAB, Cerdanyola del Vall es, Barcelona, 08193, Spain Accepted 5 April 2021 Abstract Several extinct species are known from the family Lacertidae, but due to poor preservation, many of them are based on single bones. Here, we compare phylogenetic signals of disarticulated premaxillae, maxillae and dentaries of lacertids from four French Oligocene localities (Coderet, La Colombiere, Roqueprune 2, Mas de Got B). We identified five morphotypes among the premaxil- lae, six among the maxillae, and ten among the dentaries. These morphotypes were scored as individual taxa per locality into three separate character matrices with the same 246 characters, one matrix for each jaw element. Subsequently, the phylogenetic position of the morphotypes was tested using maximum parsimony. The consensus trees with the dentaries and the maxillae found a large polytomy including all taxa except the outgroup taxon Gekko gecko. The consensus tree with the premaxillae showed a consider- ably more resolved topology but found all morphotype taxa outside Lacertidae. In a second step, we compared the constitution of our three datasets and the morphotype taxa. Our results suggest that a combination of convergent characters and missing data led to the outgroup position of the premaxilla morphotype taxa. The poor resolution of the maxillae strict consensus is likely a conse- quence of their fragmentary preservation. For the dentaries, a high amount of missing data due to the high number of morphotype taxa most likely caused the poor tree resolution. Indeed, tests with fewer morphotypes found tree resolutions comparable to the premaxilla data. When linking the morphotypes, five possible lacertid “species” were found. Comparison with already known French Oligocene lacertid species points to a slightly higher species richness of Lacertidae at that time than known before. Reliable species classification based on phylogeny only seems possible when combining the jaw elements or in association with other cranial and postcranial material, putting some doubt on species identifications based on single bones. © 2021 The Authors. Cladistics published by John Wiley & Sons Ltd on behalf of Willi Hennig Society. Introduction species from Eurasia and Africa (Arnold et al., 2007; Uetz et al., 2018). Lacertidae includes endemic “island The family Lacertidae constitutes the taxonomically giants” such as Gallotia stehlini and Gallotia simonyi dominant reptile group in Europe (Arnold et al., 2007; from the Canary Islands (Arnold, 1973), but also the Sillero et al., 2014; Speybroeck et al., 2016; Villa and widely distributed species Lacerta agilis, which occurs Delfino, 2019a; Speybroeck et al., 2020). These small from Western Europe all the way east to Mongolia to medium-sized lizards are represented by >300 extant (Arnold et al., 2007). The place and time of origin of Lacertidae are still being debated. Their geographical origin was suggested *Corresponding author. to be located in Europe (Estes, 1983a; Fu, 1998). Also E-mail address: lukardischarlottemarie.wencker@unito.it © 2021 The Authors. Cladistics published by John Wiley & Sons Ltd on behalf of Willi Hennig Society. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. 2 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 crown-group lacertids are thought to have originated studies of the origin of Lacertidae derive from the in Europe, based on the fact that fossils considered to state of preservation of early extinct members of the be close to or at the base of the crown group were group. found on this continent (Borsuk-Bialynicka et al., Taxonomic and systematic studies are greatly com-  nansk 1999; Cer y and Auge, 2013). Moreover, the lac- plicated because fossil squamates generally are found ertid sister clade Eolacertidae is also known from the disarticulated (Rage, 2013). Disarticulation can result Eocene of Europe (Cer  nansk y and Smith, 2018). In both from preservation and collection bias, and is terms of timing, molecular clock analyses identified a especially widespread among Palaeogene lacertid middle Cretaceous age for the most recent common lizards, so information about their early history is lim- ancestor of Lacertidae and Amphisbaenia (98 Myr  nansky and Auge, 2013). Among the few ited (Cer old; Wiens et al., 2006) but the calibration date used exceptions of articulated fossil lacertids worth men- has since been questioned (Hipsley et al., 2009). The tioning are exceptional cases of preservations in earliest putative crown-lacertid was Plesiolacerta from amber, on slabs/blocks and in dry environments. An the middle Eocene of Lissieu (Mammal Palaeogene almost complete specimen preserved in Baltic amber zone (MP) 14; Auge, 2005; Cer  nansk y and Auge, has been referred to Succinilacerta succinea by Borsuk- 2013). Additionally, a recent study found strong mor- Bialynicka et al. (1999). Thanks to a block containing phological similarities between modern Lacertidae and an almost complete and previously unpublished skull skeletal material from the early Eocene of Mutigny, with a few associated postcranial bones, the osteology France (MP 8–9; Cer nansk y et al., 2020). These studies and phylogenetic relationships of the Oligocene lacer- are further supported by molecular clock analyses tid Dracaenosaurus croizeti is now known in detail yielding estimates for the split between Lacertinae and  nansky et al., 2017). Janosikia ulmensis has been (Cer Gallotiinae during the late Palaeocene (58–56 Ma; revisited and redescribed on the basis of two partially Hipsley et al., 2009). Challenges regarding the interpre- preserved individuals represented by several cranial tation of the time of origin of Lacertidae also derive and some postcranial elements embedded in early Mio- from the difficulties in resolving higher-level phyloge- cene slabs from Germany (Cer  nansky et al., 2016). netic relationships. Two mummified and therefore articulated partial Molecular and morphological data have created skeletons of giant lacertids, coming from the Quater- contradictory relationships among major clades within nary filling of a basaltic cavity on the Canary Islands, Squamata (Losos et al., 2012). Morphological studies have been originally described by Castillo et al. (1994), recovered Iguania as sister to all other squamates and were subsequently used to extract ancient mtDNA (Scleroglossa), and Lacertidae as sister to Scincoidea by Maca-Meyer et al. (2003). However, the majority within Scincomorpha (Estes et al., 1988; Conrad, 2008; of extinct lacertid species were erected based on single Gauthier et al., 2012). However, molecular-based phy- bones only, mostly tooth-bearing bones (maxillae and logenies found Iguania nested more crown-ward within dentaries in particular) as well as parietals (for Euro- Squamata and Lacertidae as sister to Amphisbaenia pean post-Palaeogene taxa, see Villa and Delfino, (Townsend et al., 2004; Vidal and Hedges, 2005; Wiens 2019a, and references therein; for the osteology of et al., 2012; Pyron et al., 2013). Attempts with total- European lizards, see Villa and Delfino, 2019b, and evidence analyses including fossils found tree topolo- references therein). These are generally considered as gies within Squamata that were congruent with rela- being the most diagnostic elements and thus having tionships among higher-level clades recovered based the highest taxonomic value in the lacertid skull. How- on molecular data only, implying that differences ever, such an approach to taxonomy can be problem- between morphological and molecular phylogenies are atic. the result of convergence, and confirming the close Species descriptions and identifications based on relationship of Lacertidae and Amphisbaenia (Wiens highly fragmentary material can lack true diagnostic et al., 2010; M€ uller et al., 2011; Reeder et al., 2015). features. An example is the species Lacerta altenbur- Only recently, two studies found for the first time mor- gensis. This species was described by Rauscher (1992) phological phylogenies that were in agreement with based on a single maxilla with an uniquely shaped molecular and combined analyses, as they recovered zygomatic process. However, personal observation of Gekkota as the earliest squamate crown clade and the type maxilla by one of us (E. Tschopp in 2016) Iguania further up in the tree: one study was per- revealed that the proposed diagnostic shape of the dor- formed with a subset of morphological characters (ver- sal edge of the zygomatic process is most likely the tebrae) (Auge and Guevel, 2018); the second study had result of damage because the orbital margin of the a larger outgroup sampling compared to earlier mor- zygomatic process seems to have sharp edges derived phological analyses (and the whole skeleton consid- from breakage. The validity of other species is more ered) and found Lacertidae as sister to Amphisbaenia difficult to test because they are based on subtle differ- (Sim~oes et al., 2018). Additional challenges impacting ences in shapes or ratios of completely preserved bones L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 3 or teeth, without a detailed assessment of variability of Centrale, Tervuren, Belgium; NHMUK, Natural His- these traits in a fossil sample or among extant species. tory Museum, London, UK; NHMW, Naturhis- This issue obviously impacts any further analysis using torisches Museum Wien, Austria; PIMUZ, species occurrences to reconstruct evolution and/or Pal€aontologisches Institut und Museum der Univer- study biodiversity through time. sit€at Z€ urich, Switzerland; SRK, Sammlung Ralf Species identifications are crucial to understand the Kosma, Staatliches Naturhistorisches Museum Braun- systematics of a particular group, as well as its evolu- schweig, Germany; UAM, Universitad Aut onoma de tionary history. Assessments of evolutionary patterns Madrid, Spain; UCBL, Universite Claude Bernard, and rates within clades generally rely on the number of Lyon, France; YPM, Yale Peabody Museum, New reported species or genera (Barrett and Upchurch, Haven, USA; ZZSiD, Institute of Systematics and 2005; Pereira, 2015; Wiens, 2015; Tennant et al., 2016), Evolution of Animals, Polish Academy of Sciences, which can then be tested for correlations with ecology, Krakow, Poland. palaeoenvironmental or climatic changes in order to understand potential drivers of biodiversity. A great study system to analyze the impact of climatic changes Materials and methods on lacertid evolution is the Oligocene with its abruptly cold temperatures after the Palaeocene–Eocene Ther- Studied material mal Maximum (PETM) (Zachos et al., 2001, and refer- ences therein) and the warm period registered in the The studied material from four Oligocene localities in France Eocene (Prothero, 1994) leading to the so called (Coderet, La Colombiere, Roqueprune 2, Mas de Got B; Fig. 1) was “Grande Coupure Eocene/Oligocene” (Stehlin, 1909). initially assigned to Lacertidae and is stored in the collections of the The response of the lacertids to these climatic and envi- MNHN in Paris. All studied remains are associated with preliminary collection numbers which include a code indicating the collecting site ronmental changes is of high interest, especially consid- (e.g. MdGB-1, MdGB-2, . . . for remains from Mas de Got B). We ering the assumed close temporal association with the studied a selected set of samples containing 156 disarticulated jaw origin of crown-group lacertids. However, in order to elements from these localities. Although for La Colombiere, Roque- understand how these climatic changes influenced taxo- prune 2 and Mas de Got B one single sample from each locality was nomic diversity, it is of crucial importance to assess the selected, we had six samples from Coderet: Fouilles Viret, Coderet Couche 1 sup, Coderet Couche Verte sup 1-25, Coderet E1-0, validity of the species that occurred during that period Coderet H1-100 and Coderet H1-75. Each sample of Coderet corre- of time, as well as the methodology that was used to sponds to a different spot or stratigraphic layer within the locality. distinguish them at first place (see also Tschopp et al., The latter three samples were taken during excavations in the 1960s in review). Here, we test the latter by comparing the (Hugueney, 1969). The labelling of those samples corresponds to the excavation grid, which was numbered A to J from East to West, and phylogenetic signal of three tooth-bearing jaw elements 0 to 2 from South to North. Samples were taken from different of lacertid lizards that often were used to distinguish stratigraphic levels in those sectors, dividing them into increments of and erect new species from that period: premaxillae, 0.25 m from the surface level downwards (e.g. Coderet E1-0 was maxillae and dentaries. taken from sector E1 at a depth of 0–0.25 m below surface; Hugue- ney, 1969; de Bonis et al., 1999). We have no information on the Institutional abbreviations precise place and stratigraphic layer for the other samples. In order to keep the information gained from the sample sets separated, we did not create one large set for Coderet but instead retained the orig- BSPG, Bayerische Staatssammlung f€ ur Pal€ aontologie inal subsamples. und Geologie, Munich, Germany; CIPA, Osteoteca, Laboratorio Arqueociencias, Lisbon, Portugal; Geological background COMGR, Collezione Osteologica Mauro Grano, Rome, Italy; FMNH, Field Museum of Natural His- In the South of France, Roqueprune 2 is located in the Departe- tory, Chicago, USA; GMZ, Grant Museum of Zool- ment Tarn-et-Garonne and Mas de Got B in the Departement Lot; ogy and Comparative Anatomy, University College both are approximately 100 km N of Toulouse. They belong to the London, UK; HUJ-OST, Osteological Collections, karstic phosphate deposits of Quercy (Phosphorites du Quercy; Hebrew University of Jerusalem, Israel; ICP, Institut Vianey-Liaud, 1974; Vianey-Liaud and Wood, 1976) and originated from slow fissure fillings. In most cases, the deposits of Quercy did Catala de Paleontologia Miquel Crusafont, Barcelona, not retain the original order of sedimentation and younger layers are Spain; IRSNB, Institut Royal des Sciences Naturelles mixed with older ones (Vianey-Liaud and Marivaux, 2016). Therefore, de Belgique, Brussels, Belgium; MDHC, Massimo Del- the precise age of the fossils often is uncertain (de Bonis, 1974, 2011, fino Herpetological Collection in the Museum of Geol- and references therein). However, the sediments of Mas de Got B ogy and Paleontology of the Department of Earth were correlated with the MP 22, and thus represent the oldest of the fossil localities studied herein; Roqueprune 2 was correlated with the Sciences of the University of Turin, Italy; MNCN, slightly younger MP 23 (Schmidt-Kittler et al., 1987; Aguilar et al., Museo Nacional de Ciencias Naturales, Madrid, 1997). Therefore, both localities represent Rupelian deposits. Spain; MNHN, Museum National d’Histoire Natur- Coderet is located at the northern part of the Allier basin in the elle, Paris, France; MRAC, Musee Royal de l’Afrique Departement Allier, 30 km SW of Moulins at Limagne 4 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 Cod MdGB LaC Roq2 100 km Fig. 1. Map of France indicating the geographical positions of the four Oligocene localities with black stars. Cod, Coderet; LaC, La Colom- biere; Roq2, Roqueprune 2; MdGB, Mas de Got B. bourbonnaise. Limagne is a collapsed basin orientated in a N–S Fouilles Viret; CC1sup: Coderet Couche 1 sup; CCVsup: Coderet direction (Hugueney, 1969; de Bonis et al., 1999). It is backfilled Couche Verte sup 1-25; CE1-0: Coderet E1-0; CH1-100: Coderet with detrital and carbonate sediments (lacustrine and local forest H1-100; CH1-75: Coderet H1-75; LaC: La Colombiere; Roq2: resources). The quarry of Coderet is in contact with the western bor- Roqueprune 2; MdGB: Mas de Got B). For instance, the mOTU der fault of Limagne (Hugueney, 1969; de Bonis et al., 1999). It was identified as premaxilla morphotype P1 from the sample of Fouilles correlated to MP 30 (Schmidt-Kittler et al., 1987; Aguilar et al., Viret was given the name P1 FV. 1997) and is therefore of Chattian age. We then created three phylogenetic matrices with the same 246 La Colombiere is located in the Departement Herault in the sub- osteological characters (180 cranial and 66 postcranial ones), each of urbs of the city of Montpellier. It is a depositional group of narrow which included the morphotypes of a single jaw element. In this fissure fillings (Thaler, 1966). It belongs to the Coderet zone and is, way, we avoided potential problems for the algorithm resulting from therefore, of Chattian age as well (MP 30; Thaler, 1966; Schmidt- lacking anatomical overlap among the elements under study Kittler et al., 1987). (Tschopp et al., 2018a). The matrices were created with MESQUITE (v.3.51, build 898; Maddison and Maddison, 2018). They are based on the matrix of Tschopp et al. (2018b), which was, in turn, an Character matrix construction updated version of the matrix used by Villa et al. (2017). Twenty- four characters were added based on literature (including one that The three lacertid jaw elements on which we focused (premaxilla, was used by Villa et al., 2017, but excluded in Tschopp et al., 2018b) maxilla, dentary) were classified into individual morphotypes based and on personal observations (see Appendix 1 and references on morphological differences such as, for example, the complexity of therein). We excluded one character from the original matrix of their tooth crowns, which were based on the character matrix Tschopp et al. (2018b) (Character 217: Inscriptional ribs: absent (0); explained below. These morphotypes were divided into single mor- present (1)) because it had the same character state for all scored photype operational taxonomic units (mOTUs) based on their taxa and was, therefore, uninformative in our specific case. More- respective localities. We minimized the amount of missing data and over, we added seven outgroup and 11 ingroup operational taxo- kept the geographical and stratigraphic information separated by nomic units (OTUs). The matrix consists mostly of extant species, creating those bone-specific mOTUs. This procedure of treating dis- with the addition of extinct lacertid species known from the Oligo- articulated elements for phylogenetic analysis is, to our knowledge, cene: Ligerosaurus pouiti (Auge, 1993), Mediolacerta roceki Auge, performed here for the first time in squamates. To label the mOTUs, 2005, Plesiolacerta lydekkeri Hoffstetter, 1942, Pseudeumeces cadur- we used the names for the morphotypes, which contain the letter P censis (Filhol, 1877), Dracaenosaurus croitezi Gervais, 1848–1852, for premaxilla, M for maxilla and D for dentary plus a successive and “Lacerta” filholi Auge, 1988. We created two separate locality- cipher, and combined it with an abbreviation of the locality (FV: level OTUs for Dracaenosaurus croizeti, one representing the L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 5 material found in Cournon (approximately 10 km E of Clermont- ordered, as they were either quantitative characters or formed a mor- Ferrand; MP 28; Schmidt-Kittler et al., 1987; Cer  nansky et al., phocline (Brazeau, 2011). 2017), and the other including material from Coderet. The same Preliminary analyses recovered vast polytomies in the strict con- strategy was applied for “Lacerta” filholi, with one OTU represent- sensus trees, most likely due to the highly incomplete state of many ing the material found in Pech-du-Fraysse (including the holotype; of the fossil OTUs to be tested herein. Therefore, the analyses were approximately 30 km SE from Aurillac; Quercy Phosphorites; MP run with 13 constraints to ensure the generally accepted basic struc- 28; Schmidt-Kittler et al., 1987), and the other combining the infor- ture of the trees (as obtained from molecular data; Zheng and mation from material from Coderet (see the Supplementary material Wiens, 2016). The constraints defined Scincoidea (Chalcides, Broad- for a complete list of taxa and characters). leysaurus, Ablepharus), Lacertoidea (Amphisbaenia, Lacertidae, Tei- The outgroup now comprises Gekko gecko (Linnaeus, 1758) as a idae), Anguimorpha (Varanus, Anguis, Pseudopus), Amphisbaenia representative of the most basal squamatan clade Gekkota (molecu- (Blanus), Lacertidae (Gallotiinae and Lacertinae), Gallotiinae (Gallo- lar, total-evidence and morphological analyses agree on Gekkota tia and Psammodromus), Lacertinae (Eremiadini and Lacertini), being one of the most basal groups within Squamata; Gauthier Eremiadini (Acanthodactylus, Eremias, Mesalina, Ophisops), Lacertini et al., 2012; Reeder et al., 2015; Zheng and Wiens, 2015; Sim~ oes (Algyroides, Anatololacerta, Archaeolacerta, Iberolacerta, Lacerta, et al., 2018). Where specimens were available for first-hand scoring, Phoenicolacerta, Podarcis, Takydromus, Timon, Zootoca), and the the more derived outgroups were represented by more than one genera Podarcis, Algyroides, Iberolacerta and Lacerta. We excluded species, following the recommendations by Brusatte (2010) and the French mOTUs, and the extinct species Dracaenosaurus croizeti, Tschopp and Upchurch (2019). Squamatan clades represented in Ligerosaurus pouiti, Mediolacerta roceki, Pseudeumeces cadurcensis, this updated matrix are the Scincoidea, Anguimorpha, Teiidae and Plesiolacerta lydekkeri and “Lacerta” filholi from all constraints, as Amphisbaenia. Scincoidea is represented by Chalcides ocellatus no molecular data for those taxa are available. Therefore, they are (Forskal, 1775), Broadleysaurus major (Dumeril, 1851) and Able- considered to be “wild card taxa” and could be recovered at any pharus kitaibelii (Bibron and Bory de Saint-Vincent, 1833). The position in the tree by the analysis. It should be noted that the con- position of Scincoidea is unclear, with molecular data indicating a strained clades might only be recovered partly or not at all in the position near the base of the squamate tree as the sister taxon to resulting strict consensus tree as a consequence of conflicting tree Lacertoidea + Toxicofera (Pyron et al., 2013; Reeder et al., 2015; topologies in some cases. Zheng and Wiens, 2015), and morphological data recovering the group close to Lacertoidea, with which it forms the clade Scinco- Jaw element comparison morpha (Estes et al., 1988; Conrad, 2008; Gauthier et al., 2012). Anguimorpha is represented herein by Varanus exanthematicus (Bosc, 1792), Anguis veronensis Pollini, 1818, and Pseudopus apodus Character testing. In order to be able to compare the (Pallas, 1775). The relative position of this clade is unclear too, phylogenetic values of the three different jaw elements (premaxilla, being recovered in the opposite positions compared to Scincoidea. maxilla, dentary) we had to ensure the comparability of the three Molecular and morphological data find Anguimorpha within a datasets. To do so, we used both a quantitative and a qualitative clade that includes also Iguania and Serpentes (among other approach for character comparison. First, we compared the number groups), and which forms the sister taxon to Lacertoidea (Reeder of characters existing in the matrix for each jaw element et al., 2015; Zheng and Wiens, 2015; Sim~ oes et al., 2018). Within individually; characters scored for two or more elements at once Lacertoidea, Teiidae are represented by Salvator merianae Dumeril were counted multiple times (e.g. characters that code for maxillary and Bibron, 1839. Amphisbaenia is represented by Blanus rufus and dentary teeth). (Hemprich, 1820) and Blanus strauchi (Bedriaga, 1884) and is In a second step, the characters describing the three jaw elements thought to be the sister taxon of Lacertidae based on molecular were classified into “quantitative” and “qualitative” characters, with data (Zheng and Wiens, 2016). The ingroup Lacertidae is scored at a further subdivision of the quantitative characters into “meristic” species-level herein, mostly based on observations of ten or fewer and “morphometric”, and of qualitative characters into “countable/ specimens per species by one or more of the authors. In order to measurable” and “shape-describing” characters (Table 1; classifica- accurately represent intraspecific polymorphisms, we used majority tions of the individual characters can be found in the Supplementary scoring for polymorphic characters, following Wiens (1995). Where material). The distribution of the different character types then was an equal number of specimens showed different character states, we expressed as a percentage of the total number of characters for each scored the species as polymorphic. Hence, variation of character jaw element. states that occurred only in a minority of the specimens belonging to one species was not factored in, this should be born in mind Morphotype testing. Based on the number of characters during data interpretation. describing the premaxilla, maxilla and dentary obtained from the first part of the character testing, we took into account the Phylogenetic analysis preservation state or “specimen” completeness of the individual mOTUs. We calculated the actual percentages of scorable characters among the previously regarded characters describing the individual For the species-level phylogenetic analysis with a maximum parsi- jaw elements. mony approach, we used the software TNT (v.1.5; Goloboff and Catalano, 2016). Extended implied weighting with a k-value of 5, 12 The three jaw elements were represented by different numbers of and 20, and the default setting for assumed homoplasy for missing morphotypes for each sample, changing the number of total OTUs, entries was applied to reduce the influence of highly variable charac- and thus effectively the taxon sampling among the three matrices. In ters and missing data (Goloboff, 2014; Goloboff et al., 2018; see also order to avoid a potentially confounding impact on the analysis recommendations in Tschopp and Upchurch, 2019). The tree search based on the number of OTUs, we ran the phylogenetic analysis ten was performed using the New Technology search, enabling all algo- times per jaw element under the same conditions as stated above, rithms with their default settings, and stabilizing the consensus tree with subsampled datasets reduced to five randomly picked mOTUs five times with a factor of 75. The resulting best-fitting trees were for each analysis and ensuring that each mOTU was picked at least used as a starting point for a second iteration of tree bisection and once. reconnection (traditional heuristic search) to ensure a better tree In addition, we tested the stability of the mOTUs with respect to space coverage. Twenty-seven multistate characters were treated as their tree position. We followed the procedure for rogue taxa 6 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 Table 1 List of character types and subtypes used for the quantitative approach of jaw element comparison with their definitions and examples. Character type Character subtype Definition Example Quantitative Meristic character states are countable Maxilla, number of labial foramina: 6 or less (0); >6 (1) Morphometric character states are measurable Premaxilla, tooth-bearing portion, curvature, anterior- posterior length to transverse width: <0.2 (wide) (0); 0.2–0.4 (intermediate) (1); >0.4 (narrow) (2) Qualitative Countable/measurable character states are neomorphic Premaxilla, ascending nasal process, posterior end of (absent/present; Sereno, 2007) or medial ridge, shape in lateral view: bifid, with dorsal have a “countable” state and ventral spurs (0); single (1) Shape-describing character states are Dentary, subdental ridge, shape: straight (0); concave transformational (Sereno, 2007): (1) they describe shapes that are not “measurable”/ “countable” identification used in Pei et al. (2020) with minor modifications to premaxillae nor maxillae were present. The sample their TNT script (improvecombin.run). In a first step, unstable from Roqueprune 2 included 10% premaxillae and (rogue) taxa were identified heuristically (prune nelsen). We allowed a maximum of n + 1 taxa to be pruned at the same time (n = num- 90% dentaries. ber of mOTUs per jaw element). In a second step, the heuristically identified unstable taxa were checked by testing the tree resolution Premaxillae. Five different lacertid morphotypes for when adding or removing the unstable taxa in different combinations the premaxilla were identified among the sample of (prune-up-and-down). Here, combinations of up to n + 1 taxa could Roqueprune 2 and five subsamples of Coderet be reinserted in the tree and the pruning was evaluated with a factor F of 0.9. Factor F defines the penalty for the removal of a taxon, it (Table 2). The preservation of the studied elements varies between 0 and 1; the higher the number the more costly the generally comprises the tooth-bearing portion with removal. Given that our mOTUs were very fragmentary, we decided partially preserved teeth, and the anterior part of the to select a higher factor F to identify the taxa whose removal ascending nasal process, whereas the posterior end of increased the tree resolution the most. We identified unstable taxa the nasal process plus the articulation with the nasal are for all analyses of the complete datasets (k = 5, 12 and 20). As last, we included only the stable mOTUs of each jaw element dataset. never preserved. The analyses were run under the same parameters as before. Morphotype P1 (Fig. 2a) is characterized by seven As a last test, we merged the locality-level mOTUs belonging to tooth positions, of which the median tooth is enlarged. the same morphotype into a single cross-regional mOTU by follow- The preserved teeth are monocuspid and somewhat ing the majority rule. In this way, the amount of missing data could globose with narrow longitudinal striae. In one speci- be reduced in some cases. Consequently, we created, in congruence with the number of observed morphotypes, five cross-regional men, we observed some indication of bicuspidity in mOTUs for the premaxilla, six for the maxilla and ten for the den- one of the posterior-most teeth (within the sample tary dataset. The following morphotypes were present only in a sin- from Fouilles Viret). The facets for articulation with gle locality and therefore were kept as they were: P4 CE1-0, M2 the maxillae are large. The palatine processes border- CC1sup, M5 CE1-0, M6 CE1-0, D5 CE1-0, D6 CE1-0 and D10 ing the depression on the horizontal plate are pre- Roq2. We reran the analyses in two steps: first, using only one cross-regional mOTU at a time, and then using all cross-regional served in about half of the specimens. A small pit can mOTUs per jaw-element-dataset. The analyzing parameters were the be present at the transition of the tooth-bearing por- same as before. tion to the ascending nasal process. The ascending nasal process has a circular cross-section at its base. The ethmoidal foramina at the base of the ascending Results nasal process are not situated within fossae. The med- ial ridge on the ventral side of the preserved part of Morphotype descriptions the ascending nasal process is not visible and it there- fore probably does not reach far anteriorly. Among the jaw elements in our sample, the den- Morphotype P2 (Fig. 2b) is rather slender, with taries were the most abundant representing 68%, with seven tooth positions accommodating pencil-shaped maxillae and premaxillae comprising 23% and 9% of teeth, which all seem to be of the same size. The tooth the sample, respectively. In general, the sample from crowns are monocuspid with relatively less dense, nar- Coderet contained the highest number of jaw elements rowly spaced striae, compared to morphotype P1. with 91% of the studied jaw elements coming from Large maxillary facets are present, and the ethmoidal this locality. The distribution for Coderet was 66% foramina are situated within fossae, which are posteri- dentaries, 25% maxillae and 9% premaxillae. For the orly bordered by a ridge. Palatine processes were never localities La Colombiere and Mas de Got B, neither preserved, so their real absence or presence cannot be L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 7 Table 2 Occurrences of morphotypes and their abundance given in absolute numbers. P1–P5, premaxillary morphotypes; M1–M6, maxillary morpho- types; D1–D10, dentary morphotypes; Cod, Coderet; FV, Fouilles Viret; CC1sup, Coderet Couche 1 sup; CCVsup, Coderet Couche Verte sup 1 –25; CE1–0, Coderet E1–0; CH1–100, Coderet H1–100; CH1–75, Coderet H1–75; LaC, La Colombiere; Roq2, Roqueprune 2; MdGB, Mas de Got B. Locality P1 P2 P3 P4 P5 M1 M2 M3 M4 M5 M6 D1 D2 D3 D4 D5 D6 D7 D8 D9 D10 Cod FV 1 - - - 1 4 - - - - - 2 5 - - - - - - - - CC1sup 2 1 - - - 4 3 4 1 - - 21 10 6 - - - - - - - CCVsup 1 - - - - 5 - - 1 - - - 4 - 1 - - 6 3 - - CE1-0 2 2 1 1 - 7 - - 2 1 1 5 6 - 3 2 3 5 - - - CH1-100 - - - - - 1 - 1 1 - - - 3 2 - - - - 1 - - CH1-75 - - - - 1 - - - - - - - 2 2 1 - - - - - - LaC - - - - - - - - - - - - - - - - - - - 2 - Roq2 - - 1 - - - - - - - - - - - - - - - 1 7 1 MdGB - - - - - - - - - - - - - - 2 - - - - - - confirmed. The horizontal plate seems to have no and is wider than dorsoventrally high. The medial depression. The ascending nasal process is very slen- ridge is not reaching far anteriorly on its ventral side. der, dorsoventrally higher than wide, and the medial ridge on its ventral side does not reach far anteriorly. Maxillae. The preservation of the maxillae was Morphotype P3 (Fig. 2c) has an overall shape which rather fragmentary. Less than 50% of a single element is similar to morphotype P2. It has seven tooth posi- was preserved on average, mostly representing isolated tions with monocuspid, rounded and narrowly striated anterior, central and posterior parts. Anterior parts teeth. The maxillary facets on the tooth-bearing por- were generally preserved without the facial process and tion are smaller than in morphotype P2 but still large. with broken anteromedial/-lateral premaxillary The ethmoidal foramina are situated within fossae, processes. In total, six morphotypes among the which are not completely closed and delimited posteri- maxillae were identified in five subsamples of Coderet orly by a ridge. A pit at the base of the ascending (Table 2). nasal process can be present. Palatine processes might Morphotype M1 (Fig. 3a) has a maximum of 12 be absent or broken off, and the horizontal plate has tooth positions preserved in a single element, the total no depression. The preserved part of the ascending number of teeth in one element is probably between 16 nasal process has parallel lateral margins and the med- and 20. Most teeth seem to be tricuspid with smooth ial ridge on its ventral side does not seem to reach far tooth crowns. In the anterior half, large and robust anteriorly, as it is not visible on the preserved part of teeth are present, followed by abruptly smaller teeth in the ascending nasal process. the central part, which are relatively consistent in size. Morphotype P4 (Fig. 2d) differs from the other The tooth size increases gradually posteriorly, where morphotypes in having nine teeth. The teeth are slen- the most robust teeth are located. The tooth row is der, rather fang-like and curved posteriorly. Their straight in ventral view. For the articulation of the tooth crowns show no sign of striation. The tooth- palatine, a slight medial extension is present. The bearing portion is relatively wide and straight with anterolateral premaxillary process has a concave ven- steep and small maxillary facets. The ethmoidal foram- tral edge. The facial process has a uniform external ina are located lateral to the ascending nasal process, surface with a well-developed dermal ornamentation. penetrating the horizontal lamina. The ascending nasal The anterior margin of the facial process is perpendic- process is wider than dorsoventrally high and seems to ular to the dental crest for some distance and a small be straight throughout its entire length with the medial subtriangular process projects anteriorly from the ridge probably being restricted to the posterior part. anterodorsal corner of the facial process. On the med- Morphotype P5 (Fig. 2e) has seven tooth positions ial side, the anterior margin is connected with the car- with monocuspid, globose-like teeth which bear nar- ina maxillaris by a small ridge. There is no step visible rowly spaced, longitudinal striae. The maxillary facets at the orbital margin of the zygomatic process. are deep and large. The horizontal plate is depressed Morphotype M2 (Fig. 3b) was never preserved with and continues into two palatine processes. At the base a facial process. An indication of dermal ornamenta- of the ascending nasal process, two pairs of ethmoidal tion is present. The maximum number of observed foramina are present; they are not situated within fos- teeth in a single element is 12, whereas the total num- sae. The ascending nasal process has parallel margins ber is probably approximately 16–20 teeth. The 8 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 Fig. 2. Overview of premaxilla morphotypes. (a) Morphotype P1, premaxilla from Coderet Couche 1 sup (CC1sup-1) in posterior view. (b) Mor- photype P2, premaxilla from Coderet Couche 1 sup (CC1sup-3) in posterior view. (c) Morphotype P3, premaxilla from Coderet E1-0 (CE1-0-5) in anterior view. (d) Morphotype P4, premaxilla from Coderet E1-0 (CE1-0-6) in posterior view. (e) Morphotype P5, premaxilla from Fouilles Viret (FV-2) in right lateral view. The arrowheads mark important diagnostic structures: 1, enlarged median tooth; 2, ethmoidal foramen within fossa which is not completely closed and bordered posteriorly by a ridge; 3, fang-like, pointed tooth; 4, ethmoidal foramen without fossa; 5, sec- ond ethmoidal foramen without fossa. anp, ascending nasal process; et.f, ethmoidal foramen; mf, maxillary facet; pp, palatine process. Scale bar 1 mm. Individual and more detailed figures of the morphotypes can be found in the Supplementary material. majority of the tooth crowns seem to be tricuspid with Morphotype M3 (Fig. 3c) was always preserved widely spaced striae, but bicuspid teeth do occur in the without the premaxillary processes, but the nasal anterior part. The four posterior-most teeth are signifi- recess and central-posterior parts were present. The cantly smaller and less robust than the succeeding ones facial process is incomplete in all elements. Its anterior in the posterior half. Teeth in the central part decrease margin is clearly perpendicular to the dental crest and gradually in size and robustness. The tooth row is connected with the carina maxillaris on the medial side straight in ventral view, and the supradental shelf by a small ridge. The lateral surface is uniform with- extends slightly for the articulation with the palatine. out a distinct groove, but with well-developed dermal The orbital margin of the zygomatic process seems to ornamentation. The zygomatic process seems to be be smooth. slightly undulating, but without a distinct step. The L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 9 Fig. 3. Overview of maxilla morphotypes. (a) Morphotype M1, left maxilla from Coderet E1-0 (CE1-0-7) in lateral view. (b) Morphotype M2, right maxilla from Coderet Couche 1 sup (CC1sup-8) in medial view. (c) Morphotype M3, right maxilla from Coderet Couche 1 sup (CC1sup- 11) in lateral view. (d) Morphotype M4, left maxilla from Coderet Couche 1 sup (CC1sup-15) in lateral view. (e) Morphotype M5, right maxilla from Coderet E1-0 (CE1-0-16) in medial view. (f) Morphotype M6, right maxilla from Coderet E1-0 (CE1-0-17) in medial view. The arrowheads mark important diagnostic structures: 1, subtriangular process; 2, facial process which is perpendicular to dental crest; 3, dermal ornamentation; 4, concave ventral edge of anterolateral premaxillary process; 5, slight medial extension for articulation with palatine; 6, abruptly smaller and less robust posterior-most teeth; 7, larger tooth in anterior-half; 8, globose-like, monocuspid tooth; 9, fang-like, pointed tooth. alp, anterolateral pro- cess; ca.m, carina maxillaris; fp, facial process; ps, palatine shelf; zp, zygomatic process. Scale bar 1 mm. Individual and more detailed figures of the morphotypes can be found in the Supplementary material. development of tooth size and robustness throughout Morphotype M4 (Fig. 3d) is more robust and larger the tooth row is similar to morphotype M1, but the than morphotypes M1, M2 and M3, as well as M6. tri- and bicuspid teeth have widely spaced striae Only the anterior part is preserved. The lateral surface instead of being smooth. The preserved teeth in the has no groove that would separate the ventral and anterior half are more robust than the posteriorly suc- dorsal portions, but a well-developed dermal ornamen- ceeding teeth, whereas the posterior-most teeth seem tation is present. The anterior margin of the partly to be the most robust. preserved facial process is perpendicular to the dental 10 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 crest for some distance. On the medial side, the ante- the posterior part. The subdental ridge is concave; its rior margin is connected with the carina maxillaris by thickness increases anteriorly. A distinct splenial facet a small ridge. The anterolateral premaxillary process is present, which indicates a splenial extending for has a concave ventral edge in lateral view. The antero- approximately ¾ of the tooth row. The anterior-most medial premaxillary process bears a dorsomedially teeth are strongly tilted anteriorly. The Meckelian projecting spur close to the anterior end. The three to canal is triangular and reaches the symphysis, which four anterior-most teeth are smaller and less robust seems to have a distinct articular facet. The coronoid than the (approximately) two succeeding ones, which facet is distinct. are then followed by again less robust teeth; the Morphotype D2 (Fig. 4b,c) is very similar to mor- increase in size seems to be gradual. The teeth are photype D1. Its shape also is rather slender, but con- slightly fang-like with tooth apices pointing posteri- tinuously broadening distally. The majority of the orly. The tooth crowns are mostly mono- or bicuspid preserved teeth are bicuspid, a few tricuspid teeth with widely spaced striae; there is some indication of occur in the posterior half. At the tip of the dentary, tricuspidity present in the more posterior teeth. the teeth are procumbent. The tooth crowns have Morphotype M5 (Fig. 3e) is relatively robust with a widely spaced striae. The Meckelian canal reaches the uniform external surface and no dermal ornamenta- symphysis. The articulation facet of the symphysis tion. The anterior margin of the facial process is per- seems to be distinct. The subdental ridge is concave pendicular to the dental crest and does not bear a and thickens anteriorly. Two elements are preserved subtriangular process. On the medial side, the anterior with parts of the splenial (among the sample of margin is connected to the carina maxillaris by a small Fouilles Viret, Coderet E1-0), which has a bifid ante- ridge. The preserved onset of the anterolateral premax- rior end. The extent of the splenial seems to be illary process shows a concave ventral edge. The teeth approximately ¾ of the tooth row with a distinct sple- are similar to the ones of the premaxilla morphotype nial facet. The coronoid facet is distinct. P1 in being monocuspid and globose-like with nar- Morphotype D3 (Fig. 4d,e) is more robust than the rowly spaced striae on the tooth crowns. In general, other dentary morphotypes. The concave subdental the tooth size seems to increase distinctly posteriorly. ridge is very thick, with the greatest thickness anteri- Morphotype M6 (Fig. 3f) has a rather slender shape orly, and a well-developed splenial facet. Teeth at the with a uniform external surface and no dermal orna- tip of the dentary are tilted anteriorly. The most mentation. The teeth are monocuspid and fang-like robust teeth in the posterior part of the dentary show with strongly backwards curved apices. The crown sur- clear tricuspidity, whereas bicuspid teeth are present in faces have no striation. the anterior part. The tooth crowns have widely spaced longitudinal striae. The coronoid facet is dis- Dentaries. The dentaries always lack the angular tinct. The Meckelian canal is triangular and open until process, and for most of the elements, either only the the symphysis, which seems to have a distinct articula- posterior or only the anterior part is preserved. tion facet. However, one almost complete mandible and some Morphotype D4 (Fig. 4f,g) is similar to the morpho- almost complete dentaries were among the samples. In types D1 and D2. The element is continuously broad- total, ten morphotypes could be identified among the ening with the most robust teeth in the posterior part. studied bones from all four localities (Table 2). The tooth size gradually decreases anteriorly. Tricus- Morphotype D1 (Fig. 4a) is slender, but continu- pid teeth are present in the posterior part, whereas the ously broadens dorsoventrally towards its distal end. teeth become bicuspid anteriorly and might be mono- The maximum number of teeth observed in a complete cuspid close to the symphysis. The anterior-most teeth tooth row is 23. Most of the teeth seem to be bicuspid are strongly anteriorly tilted. The tooth crowns have and no striation is visible. However, tricuspid teeth are narrowly spaced striae. The subdental ridge is concave present in the posterior half. The change in tooth size and its thickness is increasing anteriorly. The splenial is gradual with the most robust teeth being located in facet is distinct and the posterior end of the splenial is Fig. 4. Overview of dentary morphotypes D1–D5. (a) Morphotype D1, right dentary from Coderet Couche 1 sup (CC1sup-16) in medial view. (b, c) Morphotype D2, right dentary from Coderet Couche Verte sup 1-25 (CCVsup-8) in medial view (b), tooth of a left dentary from Coderet Couche 1 sup (CC1sup-46) in medial view (c). (d, e) Morphotype D3, right dentary from Coderet Coche 1 sup (CC1sup-47) in medial view (d), teeth of a left dentary from Coderet H1-100 (CH1-100-7) in medial view (e). (f, g) Morphotype D4, right dentary from Coderet E1-0 (CE1-0-29) in medial view (f), tooth of a left dentary from Coderet E1-0 (CE1-0-31) in medial view (g). (h, i, j) Morphotype D5, right dentary from Coderet E1-0 (CE1-0-32) in medial (h) and ventral (i) view, tooth of a left dentary from Coderet E1-0 (CE1-0-33) in medial view (j). The arrowheads mark important diagnostic structures: 1, tricuspid tooth crown; 2, distinct articular facet on symphysis; 3, distinct splenial facet; 4, widely spaced striae; 5, narrowly spaced striae; 6, bicuspid tooth crown; 7, Meckelian canal that opened up at the tip. mk.c, Meckelian canal; spl.ft, splenial facet; sy, symphysis. Scale bars 1 mm. Individual and more detailed figures of the morphotypes can be found in the Supplementary material. L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 11 12 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 single and pointed. The articulation facet of the sym- show tricuspidity in the posterior half, whereas bicus- physis and the coronoid facet are distinct. The angular pid teeth are present in the anterior half. The coronoid process is exceeding the apex of the coronoid. facet is distinct, whereas the symphysis is relatively Morphotype D5 (Fig. 4h–j) has, on the contrary to indistinct. The teeth at the tip do not seem to be tilted all other morphotypes, a Meckelian canal which is anteriorly. closed in the anterior part for approximately ¼ of the Morphotype D9 (Fig. 5g) is similar to morphotype dental shelf, and that opens again at the very tip. It is D7. It is relatively slender, with a concave subdental situated rather ventrally. The articulation of the sym- ridge. The teeth have bicuspid tooth crowns, and no physis is fairly indistinct, and the teeth do not seem to indication of tricuspid teeth is visible. The tooth be tilted anteriorly. The tooth crowns have widely crowns appear to be smooth. The coronoid facet is spaced striae and are at least bicuspid. The tooth size very distinct. The splenial probably extended for increases posteriorly. The subdental ridge is rather approximately ⅔ of the tooth row, based on the dis- straight. The dentary seems to be rather sickle-shaped. tinct facets. The Meckelian canal is open until it No distinct splenial articulation facet is visible. How- reaches the seemingly indistinct symphysis. The most ever, only the anterior part of this morphotype is pre- robust teeth are present in the posterior part of the served. dentary, but the two posterior-most teeth seem to be a Morphotype D6 (Fig. 5a) is very slender and fragile. bit less robust than their anterior neighbours. The It appears to be sickle-shaped with a slightly concave teeth in the tip are procumbent. subdental ridge and a weakly developed splenial facet. Morphotype D10 (Fig. 5h,i) is preserved only as a The preserved teeth are bicuspid with smooth tooth single anterior tip. The teeth are upright, bulky and crowns. globose, with monocuspid tooth crowns bearing nar- Morphotype D7 (Fig. 5b–d) is slender, but its shape rowly spaced striae. The anterior-most tooth also is is continuously broadening posteriorly. The subdental the most robust one and the tooth size seems to ridge is similar to morphotype D6, and less strongly decrease abruptly posteriorly. The Meckelian canal concave than in morphotypes D1, D2, D3 and D4. seems to be triangular (but is not preserved entirely), The splenial facet is distinct, but the splenial seems to and reaches the symphysis, which has a distinct articu- extend less far anteriorly than in the other morpho- lation facet. The teeth overtop the dental crest by types (approximately ⅔ of tooth row). Therefore, the approximately ⅔, whereas in all other observed den- triangular Meckelian canal appears to be narrower taries the teeth overtop the dental crest by less than than in the aforementioned morphotypes. The open half of their height. portion of the canal reaches the symphysis, which has a distinct facet. The teeth at the tip are procumbent. Phylogenetic analysis In the posterior part, teeth with very distinct tricuspid- ity are present with widely spaced striae. The striation Premaxillae. The analysis of the complete is less distinct in the specimens found in Coderet premaxilla dataset produced by far the best strict Couche Verte sup 1–25. The tooth size gradually consensus tree resolution among the three jaw decreases anteriorly. The coronoid facet is distinct. elements. However, all mOTUs were recovered outside Morphotype D8 (Fig. 5e,f) is similar to the morpho- Lacertidae (Fig. 6). A polytomy was present at the type D3. The subtriangular element has a concave sub- base of the tree. It was composed of four OTUs (P1 dental ridge that thickens anteriorly. The splenial facet CCVsup, Ablepharus kitaibelii, Chalcides ocellatus and is distinct; it probably extended for approximately ¾ Varanus exanthematicus) and four branches: (1) a of the tooth row. The most robust teeth are present in partly recovered Anguimorpha clade with ((P4 CE1-0, the posterior part, but the two or three posterior-most Anguis veronensis), Pseudopus apodus); (2) a smaller teeth are significantly less robust than their neighbour- polytomy with ((Broadleysaurus major, P3 Roq2), (P2 ing teeth. Tooth size gradually decreases anteriorly. CC1sup, P2 CE1-0), P3 CE1-0)); (3) a second smaller The tooth crowns have narrowly spaced striae and polytomy with ((Dracaenosaurus croizeti (Cod), Fig. 5. Overview of dentary morphotypes D6–D10. (a) Morphotype D6, right dentary from Coderet E1-0 (CE1-0-34) in medial view. (b, c, d) Morphotype D7, left dentary from Coderet E1-0 (CE1-0-37) in medial view (b), left dentary from Coderet E1-0 (CE1-0-38) in medial view (c), tooth of a left dentary from Coderet Couche Verte sup 1-25 (CCVsup-14) in medial view (d). (e, f) Morphotype D8, right dentary from Coderet H1-100 (CH1-100-9) in medial view (e), tooth of a left dentary from Coderet Couche Verte sup 1-25 (CCVsup-19) in medial view (f). (g) Mor- photype D9, left dentary from La Colombiere (LaC-1) in medial view. (h, i) Morphotype D10, right dentary from Roqueprune 2 (Roq2-10) in medial view (h), tooth of the right dentary from Roqueprune 2 (Roq2-10) in lateral view (i). The arrowheads mark important diagnostic struc- tures: 1, weakly developed splenial facet; 2, very distinct tricuspid tooth crown; 3, distinct splenial facet; 4, distinct articular facet on symphysis; 5, widely spaced striae; 6, smaller and less robust posterior-most teeth; 7, narrowly spaced striae; 8, large and globose-like monocuspid tooth. Scale bars 1 mm. Individual and more detailed figures of the morphotypes can be found in the Supplementary material. L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 13 14 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 Dracaenosaurus croizeti (Cou), P1 FV, P1 CC1sup, P1 basal. Also, Ligerosaurus pouiti was only recovered at CE1-0), (P5 FV, P5 CH1-75)); and (4) an entirely a resolved position when analyzing the premaxilla resolved clade of Lacertoidea. dataset. It was found within the clade of Iberolacerta When applying a k = 5, higher taxonomic relation- (k = 12 and 20) or within Gallotiinae (k = 5). For ships were almost completely resolved, showing the Plesiolacerta lydekkeri only the analyses with general topology of (Gekkota, (Anguimorpha, (Scin- premaxillae and maxillae resulted in a resolved coidea, Lacertoidea))). The mOTUs P3 Roq2 and P4 position within Timon, as sister to Timon lepidus CE1-0 were recovered within the same clades as stated (Daudin, 1802) (k = 12 and 20) or as sister to Timon before. The other mOTUs were recovered as part of pater (Lataste, 1880) (k = 5). The two Dracaenosaurus Lacertoidea with P1 CCVsup at the base forming a croizeti OTUs (from Cournon and Coderet) were polytomy with two other branches: (1) a smaller poly- always sister to each other, and for analyses with tomy (Dracaenosaurus croizeti (Cod), Dracaenosaurus dentaries and maxillae they were recovered within croizeti (Cou), P1 FV, P1 CC1sup, P1 CE1-0, P5 FV, Anguimorpha. In the case of the premaxilla dataset, P5 CH1-75, (Blanus rufus, Blanus strauchi)); and (2) the two taxa were found outside of Lacertoidea (P2 CC1sup, P2 CE1-0, P3 CE1-0, (Salvator merianae without further resolution (k = 12 and 20) or in close + Lacertidae). relationship to Amphisbaenia (k = 5). The two OTUs of “Lacerta” filholi (from Pech-du-Fraysse and Maxillae. The strict consensus trees with the maxilla Coderet) were found, for the analysis with premaxillae, mOTUs recovered a large polytomy, which consisted nested within Gallotiinae. For the analysis with of all OTUs but the outgroup taxon Gekko gecko dentaries, only the taxon from Coderet was found (Fig. 7). No further resolved subclades including any sister to Gallotia caesaris (Lehrs, 1914). of the mOTUs were recovered, but two clades involving mOTUs within the large polytomy were Jaw element comparison found only for the analysis with k = 5: (1) a polytomy with (M3 CC1sup, M3 CH1-100, M4 CC1sup, M4 Character testing. Our character matrix contains 32 CE1-0, M4 CH1-100, “Lacerta” filholi (PdF)); and (2) characters coding for premaxillae, 28 characters for a clade with (M2 CC1sup, Ligerosaurus pouiti). maxillae and 24 characters for dentaries from the total number of 180 cranial characters. The classification of Dentaries. Similar to the recovered trees based on the characters describing the premaxilla, maxilla and the maxilla mOTUs, the strict consensus trees with the dentary into quantitative, qualitative and their dentary mOTUs included a large polytomy consisting subtypes, showed that the majority of the characters of all OTUs but the outgroup taxon (Fig. 8). Four are qualitative (84% of the premaxillary characters, resolved clades (three for k = 5) comprising mOTUs 89% of the maxillary characters and 92% of the were found within the polytomy: (1) (Iberolacerta dentary characters; Table 3). When comparing the cyreni (M€ uller and Hellmich, 1937), D5 CE1-0), distribution of characters among the subtypes Iberolacerta monticola (Boulenger, 1905)); (2) “countable/measurable”, and “shape-describing”, a (Algyroides nigropunctatus (Dumeril and Bibron, smaller number of characters is attributed to the 1839), D6 CE1-0), Algyroides fitzingeri (Wiegmann, subtype “shape-describing” (31% in the premaxillae, 1834)); (3) (Blanus strauchi, Blanus rufus), D10 Roq2); 32% in the maxillae and 41% in the dentaries). The and (4) (D9 LaC, Podarcis tiliguerta (Gmelin, 1789)) character distribution among the subtypes of (only for k = 12 and 20). quantitative characters shows that maxillary and dentary characters are exclusively meristic in the used Fossil wild card taxa. The positions of the eight dataset, whereas for the premaxillae 60% of the extinct taxa were not consistent throughout. Resolved quantitative characters are attributed to morphometric positions for Mediolacerta roceki and Pseudeumeces characters. cadurcensis were observed only for the analysis with the premaxillae. They were recovered at the base of Morphotype testing. The state of completeness was Lacertidae with Pseudeumeces cadurcensis being more calculated based on the number of applicable Fig. 6. Strict consensus tree of analysis of the complete premaxilla dataset using a value of k = 20 based on six most parsimonious trees (MPTs). Length of strict consensus: 1547 steps; CI of strict consensus: 0.180; RI of strict consensus: 0.441. Length of MPTs: 1499 steps; CI of MPTs: 0.186; RI of MPTs: 0.462. Arrowheads indicate the premaxilla mOTUs; daggers show the fossil OTUs; locks mark clades with fulfilled constrain- ing. Black branches, non-crown-lacertids; blue branches, Gallotiinae; green branches, Lacertinae. FV, Fouilles Viret; CC1sup, Coderet Couche 1 sup; CE1-0, Coderet E1-0; CH1-75, Coderet H1-75; CCVsup, Coderet Couche Verte sup 1-25; Roq2, Roqueprune 2; PdF, Pech-du-Fraysse; Cod, Coderet; Cou, Cournon. Strict consensus trees of k = 5 and 12 can be found in the Supplementary material. L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 15 Gekko gecko Varanus exanthematicus Chalcides ocellatus Ablepharus kitaibelli P1 CCVsup ◄ Pseudopus apodus Anguis veronensis P4 CE1-0 ◄ P3 CE1-0 ◄ Broadleysaurus major P3 Roq2 ◄ P2 CC1sup ◄ P2 CE1-0 ◄ Dracaenosaurus croizeti (Cod) † Dracaenosaurus croizeti (Cou) † P1 FV ◄ P1 CC1sup ◄ P1 CE1-0 ◄ P5 FV ◄ P5 CH1-75 ◄ Blanus strauchi Blanus rufus Salvator merianae Pseudeumeces cadurcensis † Mediolacerta roceki † Psammodromus algirus “Lacerta“ filholi (PdF) † Gallotia caesaris “Lacerta“ filholi (Cod) † Gallotia simonyi Gallotia stehlini Eremias velox Acanthodactylus boskianus Acanthodactylus erythrurus Mesalina guttulata Ophisops elegans Algyroides fitzingeri Algyroides nigropunctatus Zootoca vivipara Takydromus sp. Phoenicolacerta troodica Ligerosaurus pouiti † Iberolacerta cyreni Iberolacerta monticola Archaeolacerta bedriagae Anatololacerta danfordi Podarcis siculus Podarcis hispanicus Podarcis tiliguerta Podarcis muralis Podarcis waglerianus Lacerta viridis Lacerta trilineata Lacerta bilineata Lacerta strigata Lacerta agilis Lacerta media Lacerta schreiberi Timon princeps Timon kurdistanicus Timon pater Timon lepidus Plesiolacerta lydekkeri † 16 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 characters describing the premaxilla, maxilla and well within Gallotiinae (k = 20). The M4 mOTUs were dentary. However, for some dentary morphotypes we found unresolved within Lacertidae, and M4 CC1sup were also able to score characters describing the also was recovered as Gallotiinae (k = 5). Only M4 splenial and/or coronoid, given that they were found CCVsup was found in no further resolved position articulated. Nonetheless, only characters describing the within Lacertoidea. dentary were included in the calculation here. When analyzing the reduced dentary datasets, Therefore, the preservation state of the premaxillae, remarkably better resolved strict consensus trees were maxillae and dentaries allowed us to score the obtained than for the complete dataset. The resolution respective mOTUs for an average of 68% (range of the reduced dentary datasets was comparable to the 59%–75%), 39% (25%–61%) and 64% (38%–79%) of one of the premaxilla datasets, and the same general the available characters, respectively (for state of tree topology was observed. However, larger poly- completeness of the individual mOTUs and their tomies were still present in about half of the cases. individual positions during analyses, see Appendix 2). The three mOTUs D5 CE1-0, D6 CE1-0 and D10 The analysis of the reduced datasets with the pre- Roq2 were recovered at the same positions as when maxillae generally recovered the same tree topologies analyzing the complete dataset. The D1 mOTUs fluc- as seen for the complete dataset. In 80% of the cases, tuated between the clade of Gallotiinae and Lacerta, the higher systematic topologies were resolved (as seen with only D1 FV being consistently found within Lac- in complete premaxilla analysis under k = 5). The erta. Among the D2 mOTUs, D2 FV and D2 CCVsup mOTUs of P1, P4 and P5 were all recovered within were always recovered within Lacerta. mOTU D2 Anguimorpha with mOTU P4 always being sister to CE1-0 switched between a position within Gallotiinae Anguis veronensis. Under k = 5, the P5 mOTUs and (all k-values), Podarcis (k = 5), or as sister to Timon + P1 CE1-0 fluctuated between Anguimorpha and a Lacerta (k = 12 and 20), D2 CH1-100 was found unre- basal position within Lacertoidea with the P5 mOTUs solved within Lacertidae (k = 5 and 12) and Galloti- being in a clade with Amphisbaenia. mOTUs P1 FV inae (k = 20), and D2 CH1-75 was in an undetermined and P1 CC1sup were always found in a clade with position within Lacertoidea. For mOTUs D3, only D3 Amphisbaenia when applying k = 5. The mOTUs P2 CC1sup was found further resolved than Lacertoidea and P3 were recovered within the clade of Scincoidea, as part of Gallotiinae (k = 20) or Lacerta (all k- with P3 Roq2 being consistently found as the sister values). All D4 mOTUs were recovered as Lacer- taxon to Broadleysaurus major. But P2 mOTUs and toidea, with D4 CE1-0 being found within Gallotiinae P3 CE1-0 were also found as sister to or in polytomy (all k-values), Podarcis (k = 5), or as sister to Timon + with Salvator merianae + Lacertidae (k = 5). Lacerta (k = 12 and 20), and D4 MdGB being found The analyses with the reduced maxilla datasets still as part of Podarcis (k = 5) or as sister to Timon + Lac- resulted in strict consensus trees with large polytomies. erta (k = 20). The mOTU D4 CCVsup switched However, 30% of the trees had the clade of between a position within Lacertoidea or Scincoidea. Anguimorpha resolved, and in another 30%, the large mOTU D7 CE1-0 was consistently found within Lac- polytomy was restricted to Lacertidae. Within the erta, whereas D7 CCVsup was found in an unresolved polytomies, some smaller clades were found. mOTUs position within Lacertoidea. Among the D8 mOTUs, M5 CE1-0 and M6 CE1-0 were never found in a only D8 CH1-100 was found further resolved in a resolved position. The M1 mOTUs were mostly recov- clade with Gallotiinae, the others were found only as ered within Lacertidae, with M1 CCVsup (k = 5) and Lacertoidea (D8 CCVsup) or in no resolved position M1 CH1-100 found within Podarcis. Among the M1 at all (D8 Roq2). The D9 mOTUs were found either mOTUs only M1 FV (k = 12 and 20) and M1 CC1sup within Podarcis or as sister to Timon + Lacerta, and in were positioned in the clade of Gallotiinae. mOTU a few cases (under k = 20) within Gallotiinae. M2 CC1sup was always recovered as Gallotiinae. The The stability tests identified in total three of 11 pre- mOTU M3 CC1sup was found within Gallotiinae maxillae, eight of 14 maxillae and seven of 26 dentary (k = 5), and M3 CH1-100 was either recovered as sis- mOTUs as unstable (Appendix 2). After excluding the ter to “Lacerta” filholi (from Pech-du-Fraysse) within unstable mOTUs and rerunning the analyses, the tree the unresolved Lacertoidea clade (k = 5 and 12) or as resolution improved significantly compared to the Fig. 7. Strict consensus tree of analysis of the complete maxilla dataset using a value of k = 20 based on more than 100 000 MPTs (overflow). Length of strict consensus: 2269 steps; CI of strict consensus: 0.123; RI of strict consensus: 0.112. Length of MPTs: 1499 steps; CI of MPTs: 0.186; RI of MPTs: 0.455. Arrowheads indicate the maxilla mOTUs; daggers show the fossil OTUs; black locks mark clades with fulfilled con- straining, the two grey connected locks represent a single constraint which split up into two clades during analysis. Black branches, non-crown- lacertids; blue branches, Gallotiinae; green branches, Lacertinae. FV, Fouilles Viret; CC1sup, Coderet Couche 1 sup; CE1-0, Coderet E1-0; CH1- 100, Coderet H1-100; CCVsup, Coderet Couche Verte sup 1-25; PdF, Pech-du-Fraysse; Cod, Coderet; Cou, Cournon. Strict consensus trees of k = 5 and 12 can be found in the Supplementary material. L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 17 Gekko gecko Chalcides ocellatus Salvator merianae Anguis veronensis Pseudopus apodus Broadleysaurus major Ablepharus kitaibelli Blanus rufus Blanus strauchi “Lacerta“ filholi (PdF) † “Lacerta“ filholi (Cod) † M1 FV ◄ M1 CC1sup ◄ M2 CC1sup ◄ M3 CC1sup ◄ M4 CC1sup ◄ M1 CE1-0 ◄ M4 CE1-0 ◄ M5 CE1-0 ◄ M6 CE1-0 ◄ M1 CH1-100 ◄ M3 CH1-100 ◄ M4 CH1-100 ◄ M1 CCVsup ◄ M4 CCVsup ◄ Ligerosaurus pouiti † Mediolacerta roceki † Pseudeumeces cadurcensis † Varanus exanthematicus Dracaenosaurus croizeti (Cod) † Dracaenosaurus croizeti (Cou) † Gallotia simonyi Gallotia stehlini Gallotia caesaris Psammodromus algirus Eremias velox Mesalina guttulata Ophisops elegans Anatololacerta danfordi Phoenicolacerta troodica Podarcis siculus Takydromus sp. Zootoca vivipara Timon kurdistanicus Timon princeps Lacerta agilis Lacerta bilineata Lacerta media Lacerta schreiberi Lacerta strigata Lacerta trilineata Lacerta viridis Acanthodactylus boskianus Acanthodactylus erythrurus Algyroides fitzingeri Algyroides nigropunctatus Archaeolacerta bedriagae Iberolacerta cyreni Iberolacerta monticola Podarcis hispanicus Podarcis tiliguerta Podarcis muralis Podarcis waglerianus Timon pater Timon lepidus Plesiolacerta lydekkeri 18 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 Gekko gecko Chalcides ocellatus Salvator merianae Broadleysaurus major Ablepharus kitaibelli “Lacerta“ filholi (PdF) † D1 FV ◄ D2 FV ◄ D1 CC1sup ◄ D2 CC1sup ◄ D3 CC1sup ◄ D1 CE1-0 ◄ D2 CE1-0 ◄ D4 CE1-0 ◄ D7 CE1-0 ◄ D2 CH1-100 ◄ D3 CH1-100 ◄ D8 CH1-100 ◄ D2 CH1-75 ◄ D3 CH1-75 ◄ D4 CH1-75 ◄ D2 CCVsup ◄ D4 CCVsup ◄ D7 CCVsup ◄ D8 CCVsup ◄ D8 Roq2 ◄ D9 Roq2 ◄ D4 MdGB ◄ Ligerosaurus pouiti † Mediolacerta roceki † Plesiolacerta lydekkeri † Pseudeumeces cadurcensis † Varanus exanthematicus Dracaenosaurus croizeti (Cod) † Dracaenosaurus croizeti (Cou) † Anguis veronensis Pseudopus apodus D10 Roq2 ◄ Blanus rufus Blanus strauchi “Lacerta“ filholi (Cod) † Gallotia caesaris Gallotia simonyi Gallotia stehlini Psammodromus algirus Anatololacerta danfordi Podarcis hispanicus Podarcis siculus Timon pater Timon kurdistanicus Timon princeps Timon lepidus Lacerta agilis Lacerta media Lacerta schreiberi Lacerta strigata Eremias velox Acanthodactylus boskianus Acanthodactylus erythrurus Mesalina guttulata Ophisops elegans Algyroides fitzingeri Algyroides nigropunctatus D6 CE1-0 ◄ Archaeolacerta bedriagae Iberolacerta monticola Iberolacerta cyreni D5 CE1-0 ◄ Phoenicolacerta troodica Takydromus sp. Zootoca vivipara Podarcis muralis Podarcis waglerianus Podarcis tiliguerta D9 LaC ◄ Lacerta bilineata Lacerta trilineata Lacerta viridis L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 19 Fig. 8. Strict consensus tree of analysis of the complete dentary dataset using a value of k = 20 based on more than 100 000 MPTs (overflow). Length of strict consensus: 2074 steps; CI of strict consensus: 0.135; RI of strict consensus: 0.215. Length of MPTs: 1505 steps; CI of MPTs: 0.185; RI of MPTs: 0.464. Arrowheads indicate the dentary mOTUs; daggers show the fossil OTUs; locks mark clades with fulfilled constraining. Black branches, non-crown-lacertids; blue branches, Gallotiinae; green branches, Lacertinae. FV, Fouilles Viret; CC1sup, Coderet Couche 1 sup; CE1-0, Coderet E1-0; CH1-100, Coderet H1-100; CH1-75, Coderet H1-75; CCVsup, Coderet Couche Verte sup 1-25; LaC, La Colombiere; Roq2, Roqueprune 2; MdGB, Mas de Got B; PdF, Pech-du-Fraysse; Cod, Coderet; Cou, Cournon. Strict consensus trees of k = 5 and 12 can be found in the Supplementary material. analyses with the complete datasets, and all three data- ocellatus, (P1, (Blanus strauchi, Blanus rufus)), (Salva- sets resulted in trees with a similar general tree topol- tor merianae, Lacertidae)). ogy. The remaining stable premaxillary mOTUs When analyzing the single maxilla cross-regional generally were recovered in the same positions as in mOTUs, the cross-regional mOTU M1 was found in a the analyses of the reduced datasets with their respec- clade with Zootoca vivipara, Takydromus sp. and tive k-value. For the maxilla dataset, the new analyses Phoenicolacerta troodica, as the sister taxon to Zootoca recovered all stable mOTUs within crown-Lacertidae, vivipara. The mOTU M2 (no cross-regional OTU; M2 except for mOTU M5 CE1-0, which formed a grade occurs in a single sample only) was found as sister to with Salvator merianae being more basal to it, and the Ligerosaurus pouiti, within a clade composed of the lacertids Pseudeumeces cadurcensis, Ligerosaurus pouiti two Iberolacerta species and Archaeolacerta bedriagae. and Mediolacerta roceki as successively more derived Under k = 5, it was found in polytomy with Ligero- branches. Under k = 5, all stable maxilla mOTUs were saurus pouiti as part of Gallotiinae. The cross-regional either recovered within Gallotiinae or forming a sister mOTUs M3 and M4 were both found as sisters to the clade to it. With k = 12 and 20, the maxilla mOTUs clade Timon + Lacerta. The analysis including only were found in a polytomy within Lacertidae and only morphotype M5 (no cross-regional OTU; M5 was M1 CC1sup was found as Gallotiinae. The analyses found in a single sample only) recovered it in a grade with the stable dentary mOTUs found the mOTU D10 with Salvator merianae as more basal, and the lacertids Roq2, D5 CE1-0 and D6 CE1-0 in the same positions Pseudeumeces cadurcensis, Ligerosaurus pouiti and as in the previous analyses. All remaining stable den- Mediolacerta roceki in successively more derived posi- tary mOTUs were recovered within Podarcis, except tions. The strict consensus tree of the analysis with that D8 CH1-100 was found within Gallotiinae (k = 5 mOTU M6 (no cross-regional OTU; M6 is represented and 12) or as sister to Ligerosaurus pouiti, together in a single sample only) was poorly resolved with a forming the sister clade to Timon. For a tabular sum- large polytomy excluding only the outgroup taxon mary of the mOTU positions for the complete, Gekko gecko but including mOTU M6. When includ- reduced and only-stable-mOTUs analyses, see Appen- ing all cross-regional maxilla mOTUs simultaneously dix 2. (including M2, M5 and M6), the tree topology showed The analysis with the cross-regional mOTUs of the the same large polytomy as found by the analysis premaxillae found, when including only one at a time, restricted to the mOTU M6 (Fig. 10; for the trees of the cross-regional mOTUs P1, P2 and P5 within the cross-regional mOTU analyzed individually, see Anguimorpha as sister taxon to the two Dra- Supplementary material). The cross-regional mOTUs caenosaurus croizeti OTUs. Only when applying k = 5, M1 and M3 were found in a smaller polytomy within the cross-regional mOTU P1 was found as sister to the the large polytomy, including the “Lacerta” filholi two Blanus taxa. mOTU P4 (no cross-regional OTU; OTU from Pech-du-Fraysse. The cross-regional P4 occurs in a single sample only) was recovered as mOTU M4 was found at the same position as when sister to Anguis veronensis within Anguimorpha. The analyzed alone. The other cross-regional mOTUs were cross-regional mOTU P3 was found as sister to Broad- not found in a resolved position in the strict consensus leysaurus major within Scincoidea. For the analysis tree. For the analysis under k = 5, the clade of Galloti- including all premaxilla cross-regional mOTUs at the inae was resolved in the larger polytomy and included same time (including also P4), the strict consensus tree the cross-regional mOTUs M1, M2, M3 and M4. found all mOTUs recovered in the same positions as The separate analyses for the dentary cross-regional before, except for P2, which was now found within mOTUs found D1 and D7 within Gallotiinae, as sister Scincoidea, in a polytomy with cross-regional mOTU taxa to a clade including the three taxa of Gallotia and P3 and Broadleysaurus major (Fig. 9; for the trees of the two “Lacerta” filholi OTUs. The analysis with the the cross-regional mOTUs analyzed individually, see cross-regional mOTU D2 recovered it within Lacerta, Supplementary material). For the analysis of the cross- as sister to the clade Lacerta media + L. schreiberi. regional mOTUs under k = 5, the clade of Scincoidea The cross-regional mOTU D3 was recovered within collapsed into the following polytomy: (P2, P3, Able- Lacerta as well, but in a polytomy with L. media and pharus kitaibelii, Broadleysaurus major, Chalcides L. schreiberi. The analysis including the cross-regional 20 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 Table 3 Distribution of character types coding for premaxillae, maxillae, and dentaries. The distribution is given in absolute numbers and relative percen- tages in brackets (rounded to the nearest integer). morph: morphometric; c/m: countable/measurable; shape: shape describing. Quantitative Qualitative Total Meristic Morpho Total c/m Shape Premaxilla 5 (16%) 2 (6%) 3 (9%) 27 (84%) 17 (53%) 10 (31%) Maxilla 3 (11%) 3 (11%) 0 (0%) 25 (89%) 17 (61%) 8 (28%) Dentary 2 (8%) 2 (8%) 0 (0%) 22 (92%) 13 (54%) 9 (38%) mOTU D4 found it within Gallotiinae, as sister to the + Lacerta (dentary datasets: k = 20). Ligerosaurus clade Gallotia stehlini + G. simonyi. mOTU D5 (no pouiti was found at the base of Lacertidae, between cross-regional OTU; D5 occurs only in a single sam- Pseudeumeces cadurcensis and Mediolacerta roceki ple) was found as part of Iberolacerta, forming the sis- (maxilla dataset: all k-values; dentary dataset: k = 20), ter taxon to I. monticola. As before, mOTU D6 (no as part of Gallotiinae (premaxilla datasets: k = 5 and cross-regional OTU; D6 is known from a single sam- 20; maxilla datasets: k = 5; dentary dataset: all k- ple only) was found within Algyroides, forming the sis- values), within Iberolacerta (premaxilla dataset: k = 12 ter taxon to A. nigropunctatus. The cross-regional and 20; maxilla datasets: k = 12 and 20; dentary data- mOTU D8 and Ligerosaurus pouiti were found as sis- sets: k = 12) or as sister to Timon + Lacerta (dentary ter clade to Timon. Under k = 5, D8 was still close to datasets: k = 20). Plesiolacerta lydekkeri was found Ligerosaurus pouiti but those two were positioned within Timon, as sister to Timon lepidus (all datasets: within Gallotiinae. The cross-regional mOTU D9 was k = 12 and 20) or to Timon pater (premaxilla and recovered as sister to Timon + Lacerta, or in a poly- maxillae datasets: k = 5; dentary datasets: all k- tomy with Podarcis and Timon + Lacerta (k = 5). As values). The two Dracaenosaurus croizeti OTUs (Cour- already observed in the trees of the previous analysis, non and Coderet) were almost exclusively found in a mOTU D10 (no cross-regional OTU; D10 is repre- sister relationship to each other within Anguimorpha sented in a single sample only) was positioned as sister (all datasets: all k-values); in some cases, the two taxa to the clade Blanus strauchi + B. rufus (Amphisbaenia). were found in close relationship to Amphisbaenia (pre- When including all cross-regional dentary mOTUs at maxilla datasets: k = 5). The two OTUs of “Lacerta” the same time (including D5, D6 and D10), D5, D6, filholi (Pech-du-Fraysse and Coderet) were consistently D8 and D10 were recovered at the same positions as found within Gallotiinae. before (Fig. 11; for the trees of the cross-regional mOTU analyzed individually, see Supplementary mate- rial). By contrast, the other cross-regional mOTUs Discussion (D1, D2, D3, D4, D7, D9) were recovered in different positions, namely within Podarcis, where D9 was sister The recovered trees for premaxillae, maxillae and to P. tiliguerta. Under k = 5, the cross-regional mOTU dentaries from our analyses with the complete datasets D8 was found within Gallotiinae. displayed different degrees of resolution, implying that During morphotype testing, the eight other extinct the phylogenetic signal of tooth-bearing elements is taxa, which were treated as wild card taxa, were found incongruent within Lacertidae. This means that pre- mostly in the same positions as recovered by analyses maxillae, maxillae and dentaries alone are not equally with the complete datasets. Mediolacerta roceki and suited for species identification based on phylogenetic Pseudeumeces cadurcensis were always found close to analysis, and that taxa represented by single bones each other. They were positioned at the base of Lacer- only often cannot be placed in a well-resolved position tidae with Pseudeumeces cadurcensis being more basal in a phylogenetic tree. Likewise, different bones from (all datasets: all k-values), within Gallotiinae (premax- the same species, when found disarticulated, can be illa and dentary datasets: k = 20) or as sister to Timon recovered in different positions along the tree. Here, Fig. 9. Strict consensus tree of the premaxillary cross-regional analysis using a value of k = 20 based on two MPTs. Length of strict consensus and MPTs: 1494 steps; CI of strict consensus and MPTs: 0.187; RI of strict consensus and MPTs: 0.456. Length of MPTs: arrowheads indicate the premaxilla (cross-regional) mOTUs; daggers show the fossil OTUs; locks mark clades with fulfilled constraining. Grey dashed-line arrow indicates the position of the cross-regional mOTU when analyzing it individually (k = 20). Black branches, non-crown-lacertids; blue branches, Gallotiinae; green branches, Lacertinae. CE1-0, Coderet E1-0; PdF, Pech-du-Fraysse; Cod, Coderet; Cou, Cournon. Strict consensus trees of k = 5 and 12 can be found in the Supplementary material. L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 21 Gekko gecko Varanus exanthematicus P1 (cross-regional) ◄ P5 (cross-regional) ◄ Dracaenosaurus croizeti (Cod) † Dracaenosaurus croizeti (Cou) † Pseudopus apodus Anguis veronensis P4 (CE1-0) ◄ Chalcides ocellatus Ablepharus kitaibelli Broadleysaurus major P2 (cross-regional) ◄ P3 (cross-regional) ◄ Blanus strauchi Blanus rufus Salvator merianae Pseudeumeces cadurcensis † Ligerosaurus pouiti † Gallotia caesaris “Lacerta“ filholi (Cod) † Mediolacerta roceki † Psammodromus algirus “Lacerta“ filholi (PdF) † Gallotia simonyi Gallotia stehlini Eremias velox Acanthodactylus boskianus Acanthodactylus erythrurus Mesalina guttulata Ophisops elegans Zootoca vivipara Algyroides fitzingeri Algyroides nigropunctatus Archaeolacerta bedriagae Iberolacerta monticola Iberolacerta cyreni Anatololacerta danfordi Phoenicolacerta troodica Takydromus sp. Podarcis siculus Podarcis hispanicus Podarcis tiliguerta Podarcis muralis Podarcis waglerianus Lacerta viridis Lacerta trilineata Lacerta bilineata Lacerta strigata Lacerta agilis Lacerta media Lacerta schreiberi Timon princeps Timon kurdistanicus Timon pater Timon lepidus Plesiolacerta lydekkeri † 22 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 we explore the possible reasons for this issue, and morphometric characters are affected by taphonomic what this could mean for the systematics of extinct lac- deformation, given that meristic characters cannot log- ertids from the Oligocene of France. ically be altered in this way. Although differences in the relative amount of different character types in our Dataset comparison dataset are low (Table 3), they indicate that quantita- tive characters were slightly less error-prone and more The quantitative comparison of the characters reliable than qualitative characters in terms of recover- showed no significant overrepresentation of any of the ing resolved phylogenetic trees. It seems unlikely, three elements compared to the other two. Concerning though, that these were the only factors impacting tree the qualitative character comparison, our dataset dis- resolution and/or causing the sometimes conflicting played only relatively few differences in character type positions of the morphotypes. distribution among premaxillary, maxillary and den- Morphological variability differs between tooth- tary characters. On the one hand, it has been argued bearing elements, the impact of which was tested with that a high amount of “shape-describing” characters the analyses with a reduced mOTU sampling. When makes the analysis more error-prone, because state simply comparing the number of observed morpho- definitions in this kind of character often are vague types, the premaxillae and maxillae had a similar num- and could thus be interpreted differently by different ber of morphotypes, whereas the dentaries represent researchers. Consequently, qualitative, shape- the most variable jaw element (see also Table 2). The describing characters often were regarded as less objec- dentary also was the most frequent element in our tive than “countable/measurable” qualitative charac- sample at 68% of the material, whereas the maxillae ters (Catalano et al., 2010). However, qualitative and premaxillae constituted 23% and 9%, respectively. characters in general have been interpreted to be less This distribution of the three jaw elements is compara- objective than quantitative characters by some phylo- ble to other localities with squamate remains (e.g. geneticists (e.g., Poe and Wiens, 2000). On the other Monte Tuttavista, Sardinia, Italy: 65% dentaries, 32% hand, quantitative characters with discrete character maxillae, 3% premaxillae; Tschopp et al., 2018b; states, as are being used in our dataset, can be prob- Maramena, northern Greece: 73% dentaries, 25% lematic because the definition of their state boundaries maxillae, 2% premaxillae (considering only the squa- may be subjective (Archie, 1985; Rae, 1998). Indeed, if mate remains); Georgalis et al., 2019), indicating that a range of values is discretized into two distinct char- the dentaries have a higher preservation probability acter states, taxa with significantly different values can compared to the relatively thin, and thus more fragile be grouped together, whereas taxa displaying similar maxillae and premaxillae (and assuming that they are values, but on the two sides of the state boundary, will equally recognizable during picking and sorting activi- be scored differently (Farris, 1990). Therefore, several ties). It also should be kept in mind that dentaries and authors have suggested the use of continuous (Rae, maxillae are paired bones, whereas lacertid premaxillae 1998; Goloboff et al., 2006) or even morphometric are single bones making dentaries and maxillae twice characters (Catalano et al., 2010), which can both now as likely to be preserved. Hence, the observed higher also be scored and implemented directly in TNT variability might be due to preservation bias. (Goloboff et al., 2006; Goloboff and Catalano, 2016). Missing scores in the mOTUs appear to have had a However, discretizing continuous characters is a very more profound impact on tree topology than character straightforward approach, and the scoring of the number and definition, because the possibility of deter- OTUs into discrete states is easily reproducible. Dis- mination and positioning of the taxa within the phylo- cretizing continuous character states, as was done in genetic tree decreases with an increasing number of our dataset, can also partially address the problem of missing entries (Wiens, 2003). The resultant instability outliers in the dataset, which can result from abiotic of the incomplete OTU leads to a recovery of a high causes such as taphonomic deformation (Tschopp and number of conflicting most parsimonious trees Upchurch, 2019) that almost always affects fossils (MPTs), and consequently to a poorly resolved strict (Arbour and Currie, 2012; Tschopp et al., 2013). Nev- consensus tree (Gauthier, 1986; Huelsenbeck, 1991; ertheless, among continuous characters, only the Wilkinson and Benton, 1995; Pol and Escapa, 2009). Fig. 10. Strict consensus tree of the maxillary cross-regional analysis using a value of k = 20 based on 20 MPTs. Length of strict consensus: 1739 steps; CI of strict consensus: 0.160; RI of strict consensus. Length of MPTs: 1498 steps; CI of MPTs: 0.186; RI of MPTs: 0.452. Arrow- heads indicate the maxilla (cross-regional) mOTUs; daggers show the fossil OTUs; locks mark clades with fulfilled constraining. Grey dashed- line arrows indicate the position of the (cross-regional) mOTUs when analyzing them individually (k = 20). Grey asterisk marks the mOTU which was found at a resolved position when analyzed individually, but still as non-lacertid. Black branches, non-crown-lacertids; blue branches, Gallotiinae; green branches, Lacertinae. CC1sup, Coderet Couche 1 sup; CE1-0, Coderet E1-0; PdF, Pech-du-Fraysse; Cod, Coderet; Cou, Cour- non. Strict consensus trees of k = 5 and 12 can be found in the Supplementary material. L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 23 Gekko gecko Salvator merianae Anguis veronensis Pseudopus apodus Blanus rufus Blanus strauchi M2 (CC1sup) ◄ M5 (CE1-0) ◄ * M6 (CE1-0)◄◄ ◄ pouiti † Ligerosaurus Mediolacerta roceki † Pseudeumeces cadurcensis † Varanus exanthematicus Dracaenosaurus croizeti (Cod) † Dracaenosaurus croizeti (Cou) † Ablepharus kitaibelli Chalcides ocellatus Broadleysaurus major “Lacerta“ filholi (Cod) † M1 (cross-regional) ◄ M3 (cross-regional)◄◄ ◄ Psammodromus algirus “Lacerta“ filholi (PdF) † Gallotia caesaris Gallotia simonyi Gallotia stehlini Eremias velox Mesalina guttulata Ophisops elegans Acanthodactylus boskianus Acanthodactylus erythrurus Algyroides fitzingeri Algyroides nigropunctatus Zootoca vivipara Takydromus sp. Phoenicolacerta troodica Iberolacerta monticola Iberolacerta cyreni Archaeolacerta bedriagae Anatololacerta danfordi Podarcis siculus Podarcis hispanicus Podarcis tiliguerta Podarcis muralis Podarcis waglerianus M4 (cross-regional) ◄ Lacerta schreiberi ◄ Lacerta media Lacerta agilis Lacerta strigata Lacerta bilineata Lacerta trilineata Lacerta viridis Timon princeps Timon kurdistanicus Timon pater Timon lepidus Plesiolacerta lydekkeri † 24 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 Gekko gecko Varanus exanthematicus Dracaenosaurus croizeti (Cod) † Dracaenosaurus croizeti (Cou) † Anguis veronensis Pseudopus apodus Ablepharus kitaibelli Chalcides ocellatus Broadleysaurus major Blanus strauchi Blanus rufus D10 (Roq2) ◄ Salvator merianae Pseudeumeces cadurcensis † Mediolacerta roceki † Psammodromus algirus “Lacerta“ filholi (PdF) † Gallotia caesaris “Lacerta“ filholi (Cod) † Gallotia simonyi Gallotia stehlini Eremias velox Acanthodactylus boskianus Acanthodactylus erythrurus Mesalina guttulata Ophisops elegans Algyroides fitzingeri Algyroides nigropunctatus D6 (CE1-0) ◄ Zootoca vivipara Takydromus sp. Phoenicolacerta troodica D5 (CE1-0) ◄ Iberolacerta cyreni Iberolacerta monticola Archaeolacerta bedriagae Anatololacerta danfordi D1 (cross-regional) ◄ D4 (cross-regional) ◄ D7 (cross-regional) ◄ D3 (cross-regional) ◄ D2 (cross-regional) ◄ D9 (cross-regional) ◄ Podarcis tiliguerta Podarcis hispanicus Podarcis muralis Podarcis waglerianus Podarcis siculus Lacerta schreiberi Lacerta media Lacerta agilis Lacerta strigata Lacerta bilineata Lacerta trilineata Lacerta viridis Ligerosaurus pouiti † D8 (cross-regional) ◄ Timon princeps Timon kurdistanicus Timon pater Timon lepidus Plesiolacerta lydekkeri † L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 25 Fig. 11. Single MPT of the dentary cross-regional analysis using a value of k = 20. Length: 1504 steps; CI: 0.186; RI: 0.456. Arrowheads indi- cate the dentary (cross-regional) mOTUs; daggers show the fossil OTUs; locks mark clades with fulfilled constraining. Grey dashed-line arrows indicate the position of the cross-regional mOTUs when analyzing them individually (k = 20). Black branches, non-crown-lacertids; blue branches, Gallotiinae; green branches, Lacertinae. CC1sup, Coderet Couche 1 sup; CE1-0, Coderet E1-0; PdF, Pech-du-Fraysse; Cod, Coderet; Cou, Cournon. Strict consensus trees of k = 5 and 12 can be found in the Supplementary material. The state of completeness of the various morpho- appraised. If needed, more rigorous methods could be types (quantified by the relative number of characters used in a subsequent iteration. that were scorable for the individual morphotypes of the tooth-bearing elements) showed remarkable differ- Possible phylogenetic position of mOTUs ences. Whereas the premaxilla and the dentary mOTUs displayed comparable average percentages, The phylogenetic positions found for the various the portion of scorable characters for the maxilla mor- mOTUs varied considerably in the different analyses, photypes was significantly lower (most complete max- rendering taxonomic interpretations difficult for some illa mOTU: 61%; most incomplete premaxilla mOTU: morphotypes. Concerning the influence of the weight- 59%). This degree of “specimen” incompleteness in the ing strength applied to the maximum parsimony analy- case of the maxilla probably is the main reason for the ses, we generally observed poorer tree resolutions with poorly resolved strict consensus trees. Such an inter- k = 5, whereas k = 12 and 20 created better resolved pretation is also supported by the resolution of the tree topologies. Even though the latter two weighting strict consensus trees of analyses with the reduced strengths showed quite similar results, k = 20 gener- datasets. The resolution of the strict consensus trees ated the best tree resolution. The somewhat better per- with the dentaries was slightly lower than the one of formance of the milder downweighting against the premaxillae, probably owing to the somewhat homoplastic characters, represented by the higher k- lower “specimen” completeness. However, when a lar- values (12 and 20), agrees with results from analyses ger polytomy occurred, the dentary mOTUs were with simulated data (Goloboff et al., 2018). mostly recovered in the resolved part of the strict con- sensus trees. By contrast, the reduced datasets with the Premaxilla mOTUs. Even though all premaxilla maxillae still recovered larger polytomies with the mOTUs were recovered outside of Lacertidae, the majority of mOTUs involved. The stability tests outgroup position of the mOTUs belonging to revealed a strong correlation of stability and complete- morphotypes P1 (Fig. 2a), P2 (Fig. 2b), P3 (Fig. 2c) ness of the mOTUs. Generally, the higher the com- and P5 (Fig. 2e), is highly questionable and likely pleteness of an mOTU, the more stable its position inaccurate. Several morphological features that among the MPTs, regardless of the weighting strength. generally are regarded to be diagnostic for Lacertidae However, some mOTUs with low completeness values are present in these morphotypes, supporting an were nonetheless identified as stable, implying the assignment of these premaxillae to the ingroup. The importatance of not only the number of scored charac- observed number of teeth ranging from six to nine fits ters, but also the character itself (e.g. an apomorphic the range found in extant and extinct lacertids feature for a specific group in the dataset). (Barahona Quintana, 1996; Barahona and Barbadillo, By creating the cross-regional mOTUs, the amount  nansk 1998; Evans, 2008; Khosravani et al., 2011; Cer y of missing data was reduced without deleting taxa, as  and Auge, 2013; Cer  nansk nansky et al., 2016; Cer y mOTUs of the same morphotype but with different and Syromyatnikova, 2019; Villa and Delfino, 2019b). states of preservation were merged. Moreover, a possi- Moreover, the tooth arrangement and composition are ble ecomorphological signal (i.e. directed noise) was lacertid-like with closely spaced teeth and rounded weakened as data from different localities were com- tooth apices (Villa and Delfino, 2019b). However, the bined and, thereby, the real phylogenetic signal was prominent enlarged median tooth observed in P1 enhanced. The analyses did not result in completely previously was regarded as an unambiguous different positions, but especially in the case of the synapomorphy for the clade composed of maxillae, the morphotypes were recovered at much Amphisbaenia + Dibamidae (Gauthier et al., 2012), more resolved positions. and probably impacts our analyses as well. Other The different morphotype testing methods also can features found in the premaxillae morphotypes also be seen as a less laborious approach to test alternative may be unusual in lacertids generally, but can be tree hypotheses. Of course, it is less powerful than observed in some taxa. The slender ascending nasal using completely different datasets as done by Scar- process and the teeth of morphotype P2 are similar to petta (2020). But, by testing the different k-values and the extant species Lacerta strigata Eichwald, 1831 combinations of taxa, the general value of a taxon’s  nansky and Syromyatnikova, 2019, therein (e.g. Cer position in the phylogenetic tree can readily be fig. 12), and the extinct species Lacerta poncenatensis 26 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 M€ uller, 1996, from the Miocene of Poncenat, France  nansk similar to the one seen in Lacerta (e.g. Cer y and (e.g. M€ uller, 1996, therein fig. 5). The unclosed fossae Syromyatnikova, 2019, therein fig. 13). around the ethmoidal foramina present in P3 (Fig. 2c), Morphotype M2 (Fig. 3b) with the abrupt change is a feature that also occurs in Lacerta bilineata from smaller posterior-most teeth to preceding larger Daudin, 1802 (MNCN-16505) and Podarcis muralis teeth in the posterior half which also had been (MDHC 311). A second pair of ethmoidal foramina observed in specimens belonging to the large-sized lac- was present in P5. We observed this state also in the ertid genus Timon and the extinct species “Lacerta” OTUs in our dataset: Anguis veronensis (MDHC 102; siculimelitensis B€ohme and Zammit-Maempel, 1982 Anguimorpha), Blanus strauchi (MDHC 287; (B€ohme and Zammit-Maempel, 1982; Mateo Miras, Amphisbaenia), Gallotia simonyi (NHMW 849), 1988; Tschopp et al., 2018b), can also be an identified Acanthodactylus erythrurus (UAM.R. Ac-VII) and as an indeterminate lacertid but the poor preservation Iberolacerta monticola (UAM.R. Lm77; all three are renders a more precise classification difficult. lacertids). It also is known as a rare feature in Morphotype M5 (Fig. 3e) has globose teeth with Podarcis muralis (Laurentini, 1786) (Barahona deep striae, which are similar to the teeth preserved in Quintana, 1996; Villa and Delfino, 2019b). It seems premaxillary morphotype P1; thus, the two may that convergently acquired features pull the mOTUs of belong to a single species. Also, the two morphotypes the aforementioned morphotypes out of the ingroup overlap geographically (see Table 2). The phylogenetic and overrule the real phylogenetic signal. Hence, they position of morphotype M5 was never further resolved are likely representing lacertid remains. than basal most Lacertidae (or Lacertoidea), which Only the position of mOTU P4 (Fig. 2d), which was makes a more specific classification speculative. always recovered as sister to the anguimorph Anguis Morphotype M6 (Fig. 3f) is unlikely to represent a veronensis, seems to be plausible, and it was suggested lacertid because of its fang-like, slender teeth. These to indeed represent an anguid (Rage, 2016, pers. characteristics appear to be shared with varanids (e.g., comm.). It resembles the extinct anguid species Pseu- Georgalis et al., 2017, therein fig. 1; Villa et al., 2018; dopus laurillardi (Lartet, 1851) in terms of the dorsal Villa and Delfino, 2019a, therein fig. 11). Our analysis shape of the tooth-bearing portion that is almost rect- did not recover M6 at any resolved position. Hence, angular and quite robust, looking like the crossbar of based on our observations, the morphotype likely does the letter “T” (e.g. Klembara et al., 2010, therein fig. not represent a lacertid, but rather an anguimorph. 3). Given that mOTU P4 is also thought to be an angui- morph, and they both come from the same locality Maxilla mOTUs. All maxilla mOTUs generally were and sample (Coderet E1-0), those could represent the recovered within Lacertidae, except for morphotypes same (non-lacertid) species. Alternatively, they could M5 and M6. The morphotypes M1 (Fig. 3a) and M3 be palaeovaranids, which also share a similar tooth (Fig. 3c) could represent the same species owing to morphology, but their tooth crowns have longitudinal their similar appearance; they differ only in the striae (Georgalis, 2017). Their presence is not reported presence or absence of striae on their tooth crowns. from Coderet, but they are well-known from the Additionally, morphotypes M4 (Fig. 3d) and M3 also Quercy phosphorites (Georgalis, 2017, and references show similarities and differ (mostly) in their size. Also, therein). phylogenetic analyses (individual cross-regional analyses) recovered them in the same position, as sister Dentary mOTUs. Except for morphotype D10, all to Timon + Lacerta. It is possible that these two either dentary morphotypes were generally recovered as represent different species or that the size difference lacertids. The dentary morphotypes D1 (Fig. 4a), D2 originates from sexual dimorphism or ontogeny, (Fig. 4b,c) and D4 (Fig. 4f,g) are highly similar in because variation in the size of the skull between adult their overall shape, with the only differences being the males and adult females or juveniles is common presence or absence of either widely or narrowly among lacertids (e.g. Klemmer, 1957; Darevsky, 1967), spaced striae. Their phylogenetic position was mainly and it is usually male-biased (Vincent and Herrel, within Gallotiinae or at different positions within 2007, and references therein). The smaller morphotype Lacertini. Considering that the three morphotypes are M3 might represent the adult female or juvenile, and rather slender, an identification of the three the larger morphotype M4 the adult male version of a morphotypes as Lacertini rather than Gallotiinae single species. A classification of the three might be reasonable but remains speculative. morphotypes to Lacertidae (maybe Lacertini) seems The morphotypes D3 (Fig. 4d,e) and D8 (Fig. 5e,f) plausible, based on the tooth arrangement especially in are similar in being rather robust. The two are distin- the anterior part, with the small anterior teeth having guished by the distinctly smaller teeth of D8 in the slightly posteriorly pointing tooth apices. Adding to posterior-most part of the jaw, with the most robust this, the texture of the dermal ornamentation is also teeth preceding those. This morphology also was L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 27 observed in the maxillary morphotype M2, which also Dracaenosaurus croizeti was as well recovered outside was relatively robust. The peculiar tooth morphology Lacertidae (see discussion below). resembles certain specimens of Timon and the extinct Morphotype D5 (Fig. 4h–j) was present only in the species “Lacerta” siculimelitensis (B€ ohme and Zammit- sample from Coderet E1-0. It was always recovered Maempel, 1982; Mateo Miras, 1988; Tschopp et al., within the clade of Iberolacerta. However, a Meckelian 2018b). Based on the aforementioned observations, the canal which is closed in the anterior quarter and open- two morphotypes may or may not represent the same ing up at the tip was never detected by any of us in species, but their identification as lacertid is very Iberolacerta (or lacertids in general). Instead, such a likely. morphology is reported in skinks, for instance in Tra- The very slender morphotype D6 (Fig. 5a) from chylepis aurata and Chalcides ocellatus (e.g. Caputo, Coderet E1-0 always was recovered as part of the 2004; Villa and Delfino, 2019b, therein fig. 43). Based Algyroides-clade. However, its high incompleteness on these data, the position of the morphotype remains raises doubts on such a specific identification. In any uncertain. Because in lacertids, the Meckelian canal is case, a referral to Lacertidae and, given the slender known to be completely open (e.g. Estes et al., 1988), morphology and its biogeography, to Lacertini seems an identification of morphotype D5 as lacertid is to be plausible. highly unlikely. It might represent a skink or even The morphotype D7 (Fig. 5b–d) has a very dis- some other, unidentified taxon. tinctly developed tricuspidity of the tooth crowns which is an attribute that was formerly used as a Possible phylogenetic positions of the eight fossil wild diagnostic feature for the extinct species Miolacerta card taxa tenuis (Rocek, 1984). However, strong tricuspidity also has been observed in the lacertid genera Taky- Regardless of the weighting strength and number of dromus and Gallotia, in the gerrhosaurid Trachelopty- taxa (including mOTUs) in the dataset, Plesiolacerta chus madagascariensis (Barahona et al., 2000; Kosma, lydekkeri was always recovered within the genus 2004), and in specimens of “Lacerta” filholi (e.g. Timon, either as sister to Timon lepidus or Timon pater. Auge and Smith, 2009, therein fig. 2). Moreover, the  nansky and Auge, 2013; Previous studies (e.g. Cer overall appearance is “Lacerta” filholi-like with the  Cernansky and Syromyatnikova, 2019) have already rather slender subtriangular shape, the distinct coro- noticed shared features between Plesiolacerta and noid facet and the posteriorly reduced subdental shelf Timon. However, several issues render a close relation- (e.g. Auge and Smith, 2009). Hence, morphotype D7 ship of Plesiolacerta lydekkeri and Timon lepidus or might belong to “Lacerta” filholi, but currently avail- Timon pater questionable. Molecular analysis of the able data only seem to allow for a classification as six extant species of Timon have shown that they form Lacertidae indet. a monophyletic group (Pyron et al., 2013). The genus Morphotype D9 (Fig. 5g) also has strong similarities can be subdivided into two subclades that are geo- with “Lacerta” filholi in terms of shape, but tricuspid graphically separated into an eastern and a western teeth as seen in “Lacerta” filholi (Auge, 2005) are clade. Timon kurdistanicus (Suchow, 1936) and Timon absent. Morphotype D9 represents in all probability a princeps (Blanford, 1874) form the eastern clade with a lacertid and probably a member of the tribe Lacertini, distribution across Turkey, northern parts of Iraq and as it has a fairly slender appearance. Iranian areas (Zagros mountains) (Eiselt, 1968; Ander- Morphotype D10 (Fig. 5h,i) differs in its morphol- son, 1999; Ilgaz and Kumlutasß, 2008; Ahmadzadeh ogy from the other dentaries, because of the globose, et al., 2012). The western clade is composed of two very robust teeth that are distinctly different in size subclades: one is formed by Timon pater and Timon from each other and have narrowly spaced striae. This tangitanus (Boulenger, 1889), both from northwestern was also observed in morphotypes P1 and M5. A similar Africa (Paulo et al., 2008; Perera and Harris, 2010), tooth morphology has been reported for the extinct lac- whereas the second is formed by Timon lepidus and ertids Pseudeumeces cadurcensis (e.g. Auge and Hervet, Timon nevadensis (Buchholz, 1963), which are dis- 2009, therein fig. 2) and Dracaenosaurus croizeti (e.g. tributed across southwestern Europe (Miraldo et al., M€  nansk uller, 2006; Cer y et al., 2017, therein fig. 9). It 2012). Divergence time of the sister genera Lacerta should be noted that there is no geographical or strati- and Timon has been estimated to 18.6 Ma (95% high- graphic overlap of P1 and M5 with D10 (see Table 2). est posterior density (HPD) interval: 17.5–20.6 Ma) All phylogenetic analyses found the mOTU D10 form- and the split between the western and eastern clade of ing a clade within Amphisbaenia. However, only the Timon to 7.4 Ma (HPD interval: 5.9 9.0 Ma; Ahmad- very tip of the dentary was preserved, which makes an zadeh et al., 2016). Plesiolacerta lydekkeri is known identification barely possible, but a referral to lacertids from the middle Eocene (Lutetian; MP 14) until the might be plausible, also in consideration that early Oligocene (Rupelian; MP 21) (Auge, 2005; 28 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38  nansk Cer y and Auge, 2013 and references therein), one (39%). This highlights again how specimen incom- which pre-dates these estimated divergence times con- pleteness and tree resolution are correlated. siderably. Therefore, although a close relationship of Plesiolacerta and Timon seems plausible, the position Species diversity of the morphotypes of Plesiolacerta lydekkeri within Timon, as recovered by our analyses, is here regarded questionable. Given the observed morphological similarities and Estes (1983b) proposed a close relationship between the phylogenetic positions of the fossil material in the the amblyodont lacertid species Pseudeumeces cadur- several analyses, there is support for probably three censis and Dracaenosaurus croizeti, suggesting they lacertid and one anguimorphan species among the pre- may form a morphological series. Based on similarities maxillae, about four lacertid and one anguimorphan in tooth shape, a lacertid lineage from “Lacerta” filholi species among the maxillae, and about five lacertid via Mediolacerta roceki and Pseudeumeces cadurcensis and one non-lacertid species among the dentaries. to Dracaenosaurus croizeti was hypothesized (Hoffstet- The disparity between the premaxilla morphotypes ter, 1944; Rage, 1987; Auge, 2005). To our knowledge, is quite distinct, but the number of different morpho- phylogenetic analyses have been carried out only with types was rather low. By contrast, the number of den- Pseudeumeces cadurcensis and Dracaenosaurus croizeti tary morphotypes was relatively high, but the so far. Their position was consistently found within differences between those were less distinct. In the case  nansk Gallotiinae (Cer y et al., 2016, 2017; Tschopp of the maxilla, the number of morphotypes and the et al., 2018b; Garcia-Porta et al., 2019). In our analy- distinctness was in between those of the two other jaw ses, only the two “Lacerta” filholi OTUs were found elements. The variation in size observed in the dentary consistently within Gallotiinae. Pseudeumeces cadur- and maxilla morphotypes probably results from either censis and Mediolacerta roceki were almost always sexual dimorphism (Klemmer, 1957; Darevsky, 1967; found at the base of Lacertidae, with few analyses Vincent and Herrel, 2007, and references therein; Lju- recovering them within Gallotiinae or Lacertini. In an bisavljevic et al., 2010; Borczyk et al., 2014) or onto- even stronger contrast to earlier analyses, the two Dra- geny (Rocek, 1980). caenosaurus croizeti OTUs were either part of Considering possible connections between the jaw Anguimorpha or forming a clade with Amphisbaenia elements resulting from their morphological similari- (for some premaxillae analyses). Hence, our analyses ties, at least five different lacertid species (Fig. 12) and did not recover the hypothesised lacertid amblyodont two non-lacertid species are present in our sample. lineage, suggesting that amblyodonty evolved several The first lacertid species (“species 1”) probably com- times independently. Nevertheless, the position of Dra- prises the premaxilla morphotypes P1 and P5, the caenosaurus croizeti outside Lacertidae remains ques- maxilla morphotype M5, and the dentary morphotype tionable, given that other analyses found it D10, as they share the same morphology of globose  nansk consistently within Gallotiinae (Cer y et al., 2017; teeth with dense, narrowly spaced striae. A similar Garcia-Porta et al., 2019). tooth morphology occurs in the species Pseudeumeces The Miocene taxon Ligerosaurus pouiti was first cadurcensis and Dracaenosaurus croizeti (Auge, 2005); described as part of the genus Pseudeumeces (Auge, the two could thus be plausible candidates for a spe- 1993) and later assigned to its own genus despite cies referral of morphotypes P1, P5, M5 and D10. The superficial similarities in the dentition (Auge et al., second lacertid species (“species 2”) is likely repre- 2003). Our analyses found it mostly within Gallotiinae sented by the maxilla morphotype M2 and the dentary or the genus Iberolacerta. In very few cases, Ligero- morphotype D8. They have the same distribution of saurus pouiti also was recovered as sister to Timon + robust teeth with the most robust teeth being located Lacerta and at the base of Lacertidae, with in the posterior half followed by significantly less Pseudeumeces cadurcensis being more basal. The very robust and smaller teeth in the posterior-most part of inconsistently recovered closer relationship to the jaw element (maxilla, dentary) which is also known Pseudeumeces cadurcensis seems to confirm the referral in the extant genus Timon and the Pleistocene species of Ligerosaurus pouiti to a different genus. The contro- “Lacerta” siculimelitensis (B€ ohme and Zammit- versial or rather unstable positions of these wild card Maempel, 1982; Mateo Miras, 1988; Tschopp et al., taxa in our analyses were likely also caused by the 2018b). However, M2 has narrowly spaced striae, other added extinct mOTUs, which potentially render whereas D8 has widely spaced striae on their tooth their recovered positions less reliable. crowns. Given that similarities of the two mentioned Regarding the completeness of the wild card taxa, species with Plesiolacerta lydekkeri were reported Mediolacerta roceki and Ligerosaurus pouiti were the  nansky and Auge, 2013; Cer already (e.g. Cer  nansk y most incomplete taxa (9% and 6%, respectively), and Syromyatnikova, 2019), “species 2” might be clo- whereas Plesiolacerta lydekkeri was the most complete sely related with it or even belong to Plesiolacerta L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 29 lydekkeri. As a third lacertid species (“species 3”), the Colombiere, Roqueprune 2, Mas de Got B), “Lacerta” two dentary morphotypes D1 and D9 may be con- filholi is known to have been present in all of them, specific with the maxilla morphotype M1, given that Mediolacerta roceki is reported from Roqueprune 2 they all have smooth tooth crowns and a similar over- and Coderet, Dracaenosaurus croizeti occurred only in all robustness. The fourth lacertid species (“species 4”) Coderet, and no remains of Plesiolacerta lydekkeri and may be represented by the maxilla morphotypes M3 Pseudeumeces cadurcensis have been found in any of and M4, which possibly show ontogenetic variation or the four localities, although Pseudeumeces cadurcensis sexual dimorphism, with the less robust M3 potentially overlaps in age with Coderet and La Colombiere showing the adult female or juvenile constitution. (Auge, 2005). The last occurrence of Plesiolacerta Those two maxilla morphotypes might be linked to lydekkeri was in the early Oligocene in the MP 21 the dentary morphotypes D2 and D3, which also (Auge, 2005), hence, there is no chronological overlap might show a sexual dimorphism or variation in size with our four French localities, as the oldest one is due to ontogeny, with the slenderer morphotype D2 correlated to MP 22 (Mas de Got B). With respect to possibly representing the adult female or juvenile state. time, an increase in species variability is observable It is possible that the dentary morphotypes D4 and from the Rupelian to the Chattian. When reconciling D7 also are referable to this species, because the differ- our previously identified five lacertid “species” with ences among the four dentary morphotypes are quite the five already known ones, “species 5” seems to be small. However, all morphotypes assigned to this spe- unique. It is present in the sample from Coderet cies have widely spaced striae on their tooth crowns, (Fig. 12), but it does not resemble any of the five except for D4, which has narrowly spaced striae. known species due to its delicateness. However, onto- Therefore, the informative character on the constitu- geny or sexual dimorphism might also play a role here. tion of striae on tooth crowns (when present) seems to “Species 1” was found in the samples from Coderet be questionable, as also seen in the previously men- and Roqueprune 2 (Fig. 12) and resembles or is tioned “species 2”. Apart from these minor differences, related to the amblyodont species Pseudeumeces cadur- the morphotypes resemble “Lacerta” filholi, a species censis and Dracaenosaurus croizeti. The stratigraphic which is known already from Coderet, La Colombiere, range of Pseudeumeces cadurcensis and Dracaenosaurs Mas de Got B and Roqueprune 2 (Auge, 2005). The croizeti fits the age of Coderet (MP 30), but remains fifth species (“species 5”) is represented by dentary from this locality are reported only for Dracaensaurus morphotype D6, which was the most fragile element in croizeti (Auge, 2005). However, Roqueprune 2, which the sample; no comparable maxilla or premaxilla was was correlated to MP 23, pre-dates the first occur- observed for this species. It resembles Algyroides in its rences of the two amblyodont species (P. cadurcensis: delicateness. The two premaxilla morphotypes P2 and MP 25; D. croizeti: MP 28; Auge and Hervet, 2009). P3 probably represent two separate species, but those “Species 2” occurs in the samples from Coderet and are likely linked to some of the already defined five Roqueprune 2 (Fig. 12). It might resemble or be lacertid species. Both have tooth crowns with narrowly related to Plesiolacerta lydekkeri, which last occurred spaced striae, but as seen above, the condition of the in MP 21 (Auge, 2005); this pre-dates Coderet (MP striae might not be taxonomically significant at the 30) and Roqueprune 2 (MP23). “Species 4” was found species level. in the samples from Coderet and Mas de Got B One of the non-lacertid species probably consists of (Fig. 12), and probably represents “Lacerta” filholi, the premaxilla morphotype P4 and the maxilla mor- which has been reported from these localities before. photype M6, and are most plausible referred to an “Species 3” was found in Coderet, La Colombiere and anguimorphan species. The second non-lacertid species Roqueprune 2 (Fig. 12). It could represent any of the is represented by the dentary morphotype D5, which five known species from the Oligocene of France may be a skink or some other “scincomorphan” lizard. except for the two amblyodont species (Pseudeumeces However, it should be kept in mind that the dataset cadurcensis and Dracaenosaurus croizeti). Therefore, that we used was created specifically for lacertids, and the species variability of lacertid lizards in the Oligo- therefore, it has a rather low taxon sampling for the cene of France seems to be somewhat greater than outgroups. Because of this, the phylogenetic position recognised previously with possibly six different spe- of possible non-lacertid taxa is not very reliable. cies: Plesiolacerta lydekkeri, “Lacerta” filholi, Medio- During the Oligocene in France, five lacertid species lacerta roceki, Pseudeumeces cadurcensis, are certainly known to have occurred: Plesiolacerta Dracaenosaurus croizeti and “species 5”. Possibly, the lydekkeri, “Lacerta” filholi, Mediolacerta roceki, stratigraphic range of Plesiolacerta lydekkeri extends Pseudeumeces cadurcensis and Dracaenosaurus croizeti further into the Oligocene, whereas the first occurrence (Auge, 2005; Auge and Hervet, 2009). In the four of Pseudeumeces cadurcensis or Dracaenosaurus croizeti localities from where our samples are (Coderet, La might date further back than previously thought. 30 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 1 Cod 3 2 4 4 5 MdGB LaC Roq2 2 3 3 1 100 km SPECIES 1 SPECIES 4 Elements: SPECIES 3 Elements: P1+P5+M5+D10 M3+M4+D2+D3(+D4?+D7?) SPECIES 2 Elements: SPECIES 5 M1+D1+D9 Elements: Element: M2+D8 D6 Fig. 12. Distribution pattern and constitution of the possible five lacertid species among the four French localities indicated by black stars. Although the morphotypes P2 and P3 were classified as lacertids, they are not included here, because a referral to one of the five “species” is too speculative. Cod, Coderet; LaC, La Colombiere; Roq2, Roqueprune 2; MdGB, Mas de Got B. Conclusions However, based on observations of the morpho- types, possibly five lacertid “species” could be identi- Assessing the phylogenetic value of lacertid premax- fied in our samples from the four French localities. illae, maxillae and dentaries is only possible to a lim- Reconciliation with the already known lacertid species ited extent. The phylogenetic trees resulting from three in France during the Oligocene suggests a larger spe- individual datasets of jaw elements show remarkable cies diversity than thought previously, with six species differences in terms of resolution of the strict consen- in total, indicating a more rapid radiation of Lacer- sus trees and number of MPTs. Also, analyses of the tidae following the abrupt shift to colder temperatures same dataset but with different weighting strengths after the Paleocene–Eocene Thermal Maximum and methods often led to conflicting positions of the (PETM) and the warm period during the Eocene. single-bone morphotypes. However, various iterations of phylogenetic analyses L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 31 with modified datasets failed to recover clear and con- highly useful comments and corrections on our manu- sistent phylogenetic positions for most morphotypes, script. demonstrating the weakness of their phylogenetic sig- Financial support was provided for L.C.M.W. by a nal. Also, a highly resolved tree does not necessarily PhD fellowship from the Universita degli Studi di Tor- imply that there was a reliable phylogenetic signal in ino. Part of the dataset was assembled under the lead the dataset. Therefore, it is important to be aware of of E.T. when he was holding a Postdoctoral Fellow- the constitution of the data being used and to test the ship funded by the European Union’s Seventh Frame- robustness of the taxa’s positions. A precise classifica- work programme for research and innovation under tion at species-level of the individual jaw elements was the Marie Sklodowska-Curie grant agreement No. not possible based on the phylogenetic analysis. 609402 – 2020 researchers: Train to Move (T2M). Hence, species descriptions and classification based on A.V. is funded by a Humboldt Research Fellowship single tooth-bearing elements should be treated with provided by the Alexander von Humboldt Founda- caution. Although real autapomorphic features may be tion. M.D. acknowledges support from Fondi di Ate- present in some of these elements, these seem to be dif- neo dell’Universita di Torino (2017–2019), Generalitat ficult to interpret as such, and if identified correctly, de Catalunya (CERCA Programme, consolidated they do not seem to provide enough information to research group 2017 SGR 116 GRC) and Agencia consistently reveal their true phylogenetic position, Estatal de Investigacion of Spain (CGL2016-76431-P, and so phylogenetic trees should undergo a conscien- AEI/FEDER, EU). Further data collection was done tious “test of robustness”. during Synthesys grants to visit the MNHN (Paris, France) and NHMW (Vienna, Austria) financed by the European Community Research Infrastructure Acknowledgements Action under the FP7 ‘Capacities’ Programme (FR- TAF-5007 and 5839, to A.V. and E.T. respectively; We thank †Jean-Claude Rage, Salvador Bailon and AT-TAF-4591 and 5725, to A.V. and E.T., respec- Virginie Bouetel from the MNHN for kindly providing tively) and during two Erasmus+ Traineeships (Univer- the key material for this study. Moreover, we thank sity College London, UK; NHMW, AT) to L.C.M.W. the numerous people who helped during collection vis- This is the publication number 354 of the Museo di its. Especially we thank S onia Gabriel and Simon Geologia e Paleontologia collections at the Universita Davis (CIPA), Marta Calvo and Alberto Sanchez degli Studi di Torino. (MNCN), Jeff Streicher and Patrick Campbell (NHMUK), Heinz Grillitsch, Georg Gassner and Silke Conflict of interest Schweiger (NHMW), Christian Klug and Winand Brinkmann (PIMUZ), Hannah Cornish and Tannis None declared. Davidson (GMZ), and Ralf Kosma (SRK). Additional specimens could be studied thanks to loans from HUJ- OST, MRAC, UAM and ZZSiD. We would like to References specifically thank Rebecca Biton, Georgios Georgalis and Mauro Grano, who personally brought specimens Aguilar, J.-P., Agustı, J., Alexeeva, N., Antoine, P.-O., Antunes, M.T., Legendre, S. and Wolsan, M., 1997. Syntheses et tablaux from these collections to Turin for us to study, and de correlations [Overview and table of correlations]. In: Aguilar, Annelise Folie, Francisco Ortega and Zbigniew Szynd- J.-P., Legendre, S. & Michaux, J. (Eds.), Actes du Congres lar for approving and organizing loans. We further- BiochroM. Mem. Trav. Ecole Prat. Hautes Etudes, Inst. 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Revision of Varanus marathonensis (Squamata, Varanidae) based on historical and new material: morphology, systematics, 6084/m9.figshare.13277564 and paleobiogeography of the European monitor lizards. PLoS File S1. Nexus file of the premaxilla taxon-character One 13, e0207719. matrix (.nex format). Villa, A. and Delfino, M., 2019a. Fossil lizards and worm lizards (Reptilia, Squamata) from the Neogene and Quaternary of File S2. Nexus file of the maxilla taxon-character Europe: an overview. Swiss J. Palaeontol. 138, 177–211. matrix (.nex format). Villa, A. and Delfino, M., 2019b. A comparative atlas of the skull File S3. Nexus file of the dentary taxon-character osteology of European lizards (Reptilia: Squamata). Zool. J. matrix (.nex format). Linn. Soc. 187, 829–928. Villa, A., Tschopp, E., Georgalis, G.L. and Delfino, M., 2017. File S4. List of characters including their character Osteology, fossil record and palaeodiversity of the European classifications (.xlsx format). lizards. Amphibia-Reptilia 38, 79–88. File S5. List of studied specimens (.xlsx format). Vincent, S.E. and Herrel, A., 2007. Functional and ecological correlates of ecologically-based dimorphisms in squamate File S6. List of references for File S4 and File S5 reptiles. Integr. Comp. Biol. 47, 172–188. (.pdf format). Wiens, J.J., 1995. Polymorphic characters in phylogenetic File S7. List of figured mOTUs including their pre- systematics. Syst. Biol. 44, 482–500. liminary collection numbers (.xlsx format). Wiens, J.J., 2003. Missing data, incomplete taxa, and phylogenetic accuracy. Syst. Biol. 52, 528–538. File S8. TNT input files and tree files (.tre format) Wiens, J.J., 2015. Explaining large-scale patterns of vertebrate of analyses of complete premaxilla, maxilla and den- diversity. Biol. Lett. 11, 20150506. tary datasets (k = 5, 12, 20) (.zip format). Wiens, J.J., Brandley, M.C., and Reeder, T.W., 2006. Why does a File S9. TNT input files and tree files (.tre and .nex trait evolve multiple times within a clade? Repeated evolution of snakeline body form in squamate reptiles. Evolution 60, 123– format) of analyses of reduced premaxilla, maxilla and 141. dentary datasets (k = 5, 12, 20) (.zip format). Wiens, J.J., Hutter, C.R., Mulcahy, D.G., Noonan, B.P., Townsend, File S10. TNT tree files (.tre and .nex format) of sta- T.M., Sites, J.W. Jr and Reeder, T.W., 2012. Resolving the phylogeny of lizards and snakes (Squamata) with extensive bility analyses of premaxilla, maxilla and dentary data- sampling of genes and species. Biol. Lett. 8, 1043–1046. sets (k = 5, 12, 20), including the script for taxa Wiens, J.J., Kuczynski, C.A., Townsend, T., Reeder, T.W., pruning (.zip format). Mulcahy, D.G. and Sites, J.W. Jr, 2010. Combining File S11. TNT input files and tree files (.tre format) phylogenomics and fossils in higher-level squamate reptile phylogeny: molecular data change the placement of fossil taxa. of analyses of cross-regional premaxilla, maxilla, and Syst. Biol. 59, 674–688. dentary datasets (k = 5, 12, 20) (.zip format). Wilkinson, M. and Benton, M.J., 1995. Missing data and File S12. Single MPTs or strict consensus trees of rhynchosaur phylogeny. Hist. Biol. 10, 137–150. Zachos, J., Pagani, M., Sloan, L., Thomas, E. and Billups, K., 2001. the phylogenetic analyses testing the positions of the Trends, rhythms, and aberrations in global climate 65 Ma to cross-regional mOTUs individually (.pdf format). present. Science 292, 686–693. File S13. Strict consensus trees of analyses of com- Zheng, Y. and Wiens, J.J., 2015. Do missing data influence the plete premaxilla, maxilla and dentary datasets (k = 5, accuracy of divergence-time estimation with BEAST? Mol. Phylogenet. Evol. 85, 41–49. 12) (.pdf format). Zheng, Y. and Wiens, J.J., 2016. Combining phylogenomic and File S14. Individual figures of morphotypes (.zip for- supermatrix approaches, and a time-calibrated phylogeny for mat). squamate reptiles (lizards and snakes) based on 52 genes and File S15. Table of completeness of the eight fossil 4162 species. Mol. Phylogenet. Evol. 94, 537–547. wild card taxa (.pdf format). 36 L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38 Appendix 1 List of newly inserted characters, including their states and sources. No. New character Source (and original character numbers) 12 Premaxilla, tooth-bearing portion: curvature, anteroposterior length to LCMW, pers. obs., 2018 transverse width: <0.2 (wide) (0); 0.2–0.4 (intermediate) (1); >0.4 (narrow) (2). 36 Premaxilla, tooth row, shape: U-shaped (0); V-shaped (1). LCMW, pers. obs., 2018 40 Premaxilla, second pair of ethmoidal foramina: absent (0); present (1). LCMW, pers. obs., 2019 41 Premaxilla, ascending nasal process, posterodorsal end, shape in anterior Barahona & Barbadillo (1997; C3.1), modified (split in view: pointed (0); widely rounded (1). two) by Villa et al. (2017; C23) 47 Premaxilla, ascending nasal process, length of articulation with the nasals: modified after Gauthier et al. (2012; C10; exclusion of less than half nasal length (0); more than halfway to frontal between character state (2): nearly to, or articulates with nasals (1). frontals); Quadros et al. (2018; C3) 48 Premaxilla, ascending nasal process, constriction at the base: absent (0); wording modified from Quadros et al. (2018; C4); present (1). Brizuela (2010; C3) 62 Maxilla, anteromedial premaxillary process lying between vomers and Estes et al. (1988; C12); Denton and O’Neill (1995; premaxilla: absent, vomers contact premaxilla (0); present, vomers do not C4); Nydam and Cifelli (2002; C4); Nydam et al. contact premaxilla (1). (2007; C4); Conrad (2008; C25); Quadros et al. (2018; C13) 73 Nasals, anterior width: exceeds nasofrontal joint width (0); is subequal to Gauthier et al. (2012; C18); Quadro et al. (2018; nasofrontal joint width (1); less than anterior frontal width (2). C6) 74 Nasal, anterior border: nasolateral process present, forming the posterior Quadros et al. (2018; C7); modified from Conrad border and part of the labial border of the external nares (0); nasolateral (2008; C22); Gauthier et al. (2012; C22) process absent (1). 77 Dermal skull bone ornamentation, frontal/parietal: absent, dermal skull Modified after Quadros et al. (2018; C17; wording roof smooth (0); present over dorsum (1); present on jugal postorbital bar modified and exclusion of state (1): lightly rugose (2). about frontoparietal suture); modified from Estes et al. (1988; C129); Conrad (2008; C10); Gauthier et al. (2012; C572) 93 Postorbital, overlapping of squamosal: dorsomedially as slender tapering Modified from Gauthier et al. (2012; C78; exclusion of rod attached superficially (0); dorsally (1); postorbital in long V-shaped state (0): laterally into V-shaped recess in squamosal); trough dorsally and then rotating dorsolaterally posteriorly (2); squamosal Arnold (1998); Quadros et al. (2018; C35) lies in trough beneath postorbital (3). 107 Quadrate, tympanic crest shape: straight or vertically developed (0); curved, Wording modified after Quadros et al. (2018; C78); forming "C" on outer line (1). modified from Brizuela (2010; C63) 109 Squamosal, base of temporal ramus: diverges from parietal (0); base lies Quadros et al. (2018; C46); Gauthier et al. (2012; against parietal (1). C162) 110 Supratemporal position on parietal: partly ventral (0); partly ventrolateral De Queiroz (1987); Gauthier et al. (2012; C170) (1); all lateral (2); dorsolateral (3). 127 Parietal, parasagittal crest strongly developed on the dorsolateral surface Quadros et al. (2018; C22); modified from Denton and for the insertion of the adductor mandible musculature: absent (0); present O’Neill (1995; C10); Nydam and Cifelli (2002; C10); (1). Nydam et al. (2007; C10); Conrad (2008; C75); Gauthier et al. (2012; C9) 128 Epipterygoid-parietal contact: absent (0); overlaps parietal temporal muscle Gauthier et al. (2012; C294); Quadros et al. (2018; origin parietal downgrowths (1). C84) 155 Dentary, subdental ridge, shape: straight (0); concave (1). Bailon et al. (2014) 159 Dentary, Meckelian canal, shape: triangular (0), tubular (1). ET, pers. obs., 2018 160 Dentary, orientation of teeth at the anterior tip: upright (0); tilted LCMW, pers. obs., 2019 anteriorly or procumbent (1). 175 Coronoid-surangular articulation: coronoid restricted to medial aspect of Gauthier et al. (2012; C390); Quadros et al. (2018; mandible (0); coronoid extends onto dorsal surface of surangular (1); C113) coronoid arches over dorsal margin of mandible to reach lateral surface of surangular (2). 184 Teeth, premaxillary, crown morphology (if monocuspid): fang-like/pointed LCMW, pers. obs., 2018 (0); rounded (1). 185 Teeth, premaxilla, median tooth: absent (0); present (1). Wording modified from Scanlon (1996; C4); Lee (1998; C154); Gauthier et al. (2012; C413) 186 Teeth, premaxilla, median tooth: same size as neighbouring premaxillary Lee (1998; C155); modified after Gauthier et al. (2012; teeth (0); enlarged (1). C414; wording modified and exclusion of state (2) greatly enlarged median premaxillary tooth) 190 Teeth, maxilla and dentary, differences in tooth robustness: (0) absent to LCMW, pers. obs., 2019 minimal; present not more than 50% different in robustness (1); (present, 50% or more difference in robustness (2). Appendix 2 Positions of premaxilla, maxilla and dentary mOTUs in the trees with the complete and reduced datasets, and the analyses with the stable mOTUs only for each of the three k-values. Completeness scores for the individual mOTUs are given in brackets behind the taxon name. FV, Fouilles Viret; CC1sup, Coderet Couche 1 sup; CE1-0, Coderet E1-0; CH1-100, Coderet H1-100; CH1-75, Coderet H1-75; CCVsup, Coderet Couche Verte sup 1- 25; LaC, La Colombiere; Roq2, Roqueprune 2; MdGB, Mas de Got B; PdF, Pech-du-Fraysse; Cod, Coderet; T+L, Timon + Lacerta. Analysis with a k-value of 5 Analysis with a k-value of 12 Analysis with a k-value of 20 mOTU Complete Reduced Stable Complete Reduced Stable Complete Reduced Stable P1 FV (66%) Lacertoidea (clade Lacertoidea (clade Lacertoidea (clade Polytomy with D. Anguimorpha Anguimorpha Polytomy with D. Anguimorpha Anguimorpha with Amphisbaenia) with Amphisbaenia) with Amphisbaenia) croizeti croizeti P1 CC1sup (72%) Lacertoidea (clade Lacertoidea (clade Excluded Polytomy with D. Anguimorpha Excluded Polytomy with D. Anguimorpha Excluded with Amphisbaenia) with Amphisbaenia) croizeti croizeti P1 CE1-0 (75%) Lacertoidea (clade Lacertoidea (clade Lacertoidea (clade Polytomy with D. Anguimorpha Anguimorpha Polytomy with D. Anguimorpha Anguimorpha with Amphisbaenia) with Amphisbaenia) with Amphisbaenia) croizeti croizeti P1 CCVsup (59%) Lacertoidea Anguimorpha; Lacertoidea – Anguimorpha Excluded – Anguimorpha Excluded Lacertoidea P2 CC1sup (72%) Lacertoidea (sister/ Lacertoidea (sister/ Lacertoidea (sister/ Clade with B. major Scincoidea Scincoidea Clade with B. major Scincoidea Scincoidea polytomy to/with S. polytomy to/with S. polytomy to/with S. merianae + merianae + merianae + Lacertidae) Lacertidae) Lacertidae) P2 CE1-0 (75%) Lacertoidea (sister/ Lacertoidea (sister/ Lacertoidea (sister/ Clade with B. major Scincoidea Scincoidea Clade with B. major Scincoidea Scincoidea polytomy to/with S. polytomy to/with S. polytomy to/with S. merianae + merianae + merianae + Lacertidae) Lacertidae) Lacertidae) P3 CE1-0 (69%) Lacertoidea (sister to/ Lacertoidea (sister to/ Lacertoidea (sister to/ Clade with B. major Scincoidea Scincoidea Clade with B. major Scincoidea Scincoidea polytomy with S. polytomy with S. polytomy with S. merianae + merianae + merianae + Lacertidae) Lacertidae) Lacertidae) P3 Roq2 (59%) Scincoidea (sister to B. Scincoidea (sister to B. Scincoidea (sister to B. Sister to B. major Scincoidea (sister to B. Scincoidea (sister to B. Sister to B. major Scincoidea (sister to B. Scincoidea (sister to B. major) major) major) major) major) major) major) P4 CE1-0 (59%) Anguimorpha (sister Anguimorpha (sister Anguimorpha (sister Sister to A. veronensis Anguimorpha (sister Anguimorpha (sister Sister to A. veronensis Anguimorpha (sister Anguimorpha (sister to A. veronensis) to A. veronensis) to A. veronensis) to A. veronensis) to A. veronensis) to A. veronensis) to A. veronensis) P5 FV (66%) Lacertoidea (clade Anguimorpha; Excluded Clade with D. croizeti Anguimorpha Anguimorpha Clade with D. croizeti Anguimorpha Anguimorpha with Amphisbaenia) Lacertoidea (clade with Amphisbaenia) P5 CH1-75 (75%) Lacertoidea (clade Anguimorpha; Lacertoidea (clade Clade with D. croizeti Anguimorpha Anguimorpha Clade with D. croizeti Anguimorpha Anguimorpha with Amphisbaenia) Lacertoidea (clade with Amphisbaenia) with Amphisbaenia) L. C. M. Wencker et al. / Cladistics 0 (2021) 1–37 M1 FV (29%) – Lacertidae Excluded – Gallotiinae Excluded – Gallotiinae Excluded M1 CC1sup (36%) – Gallotiinae Gallotiinae – Gallotiinae Gallotiinae – Gallotiinae Gallotiinae M1 CE1-0 (61%) – Lacertidae sister to Gallotiinae – Lacertidae Lacertidae – Lacertidae Lacertidae M1 CH1-100 (25%) – Podarcis Excluded – Podarcis Excluded – Podarcis Excluded M1 CCVsup (39%) – Podarcis sister to Gallotiinae – Lacertidae Excluded – Lacertidae Excluded M2 CC1sup (36%) sister to L. pouiti Gallotiinae Gallotiinae – Gallotiinae Excluded – Gallotiinae Excluded M3 CC1sup (50%) polytomy with “L.” Gallotiinae Sister to Gallotiinae – Lacertidae Lacertidae – Lacertidae Lacertidae filholi (PdF) M3 CH1-100 (32%) polytomy with “L.” Lacertoidea (sister Sister to Gallotiinae – Lacertoidea (sister Lacertidae – Gallotiinae Lacertidae filholi (PdF) “L.” filholi (PdF)) “L.” filholi (PdF)) M4 CC1sup (32%) polytomy with “L.” Gallotiinae Sister to Gallotiinae – Lacertidae Excluded – Lacertidae xcluded filholi (PdF) M4 CE1-0 (36%) polytomy with “L.” Lacertidae Sister to Gallotiinae – Lacertidae Lacertidae – Lacertidae Lacertidae filholi (PdF) M4 CH1-100 (32%) polytomy with “L.” Lacertidae sister to Gallotiinae – Lacertidae Excluded – Lacertidae Excluded filholi (PdF) M4 CCVsup (25%) – Lacertoidea Excluded – Lacertoidea Excluded – Lacertoidea Excluded M5 CE1-0 (50%) – – Lacertoidea (sister to – – Lacertoidea (sister to – – Lacertoidea (sister to S. merianae + S. merianae + S. merianae + Lacertidae) Lacertidae) Lacertidae) M6 CE1-0 (25%) – – Excluded – – Excluded – – Excluded 37 38 Appendix 2 (Continued) Analysis with a k-value of 5 Analysis with a k-value of 12 Analysis with a k-value of 20 mOTU Complete Reduced Stable Complete Reduced Stable Complete Reduced Stable D1 FV (75%) – Lacerta Podarcis – Lacerta Podarcis – Lacerta Podarcis D1 CC1sup (75%) – Lacerta Podarcis – Gallotiinae; Lacerta Podarcis – Gallotiinae; Lacerta Podarcis D1 CE1-0 (63%) – Gallotiinae; Lacerta Podarcis – Gallotiinae Podarcis – Gallotiinae Podarcis D2 FV (71%) – Lacerta Podarcis – Lacerta Podarcis – Lacerta Podarcis D2 CC1sup (79%) – Lacerta Podarcis – Lacerta Podarcis – Lacerta Podarcis D2 CE1-0 (75%) – Gallotiinae; Podarcis Podarcis – Gallotiinae; sister to Podarcis – Gallotiinae; sister to Podarcis T+L T+L D2 CH1-100 (75%) – Lacertidae Podarcis – Lacertidae Podarcis – Gallotiinae Podarcis D2 CH1-75 (46%) – Lacertoidea Excluded – Lacertoidea Excluded – Lacertoidea Excluded D2 CCVsup (63%) – Lacerta Podarcis – Lacerta Podarcis – Lacerta Podarcis D3 CC1sup (75%) – Lacerta Podarcis – Lacerta Podarcis – Gallotiinae; Lacerta Podarcis D3 CH1-100 (75%) – Lacertoidea Podarcis – Lacertoidea Podarcis – Lacertoidea Podarcis D3 CH1-75 (42%) – Lacertoidea Excluded – Lacertoidea Excluded – Lacertoidea Excluded D4 CE1-0 (79%) – Gallotiinae; Podarcis Podarcis – Gallotiinae; sister to Podarcis – Gallotiinae; sister to Podarcis T+L T+L D4 CH1-75 (50%) – Lacertoidea Excluded – Lacertoidea Excluded – Lacertoidea Excluded D4 CCVsup (54%) – Lacertoidea; Excluded – Lacertoidea; Excluded – Lacertoidea; Excluded Scincoidea Scincoidea Scincoidea D4 MdGB (75%) – Podarcis Podarcis – Lacertoidea Podarcis – Sister to T+L Podarcis D5 CE1-0 (42%) Iberolacerta (sister to Iberolacerta (sister to Iberolacerta (sister to Iberolacerta (sister to Iberolacerta (sister to Iberolacerta (sister to Iberolacerta (sister to Iberolacerta (sister to Iberolacerta (sister to I. cyreni) I. cyreni) I. cyreni) I. cyreni) I. cyreni) I. cyreni) I. cyreni) I. cyreni) I. cyreni) D6 CE1-0 (63%) Algyroides (sister to A. Algyroides (sister to A. Algyroides (sister to A. Algyroides (sister to A. Algyroides (sister to A. Algyroides (sister to A. Algyroides (sister to A. Algyroides (sister to A. Algyroides (sister to A. nigropunctatus) nigropunctatus) nigropunctatus) nigropunctatus) nigropunctatus) nigropunctatus) nigropunctatus) nigropunctatus) nigropunctatus) D7 CE1-0 (75%) – Lacerta Podarcis – Lacerta Podarcis – Lacerta Podarcis D7 CCVsup (71%) – Lacertoidea Podarcis – Lacertoidea Podarcis – Lacertoidea Podarcis D8 CH1-100 (75%) – clade with Gallotiinae Gallotiinae – clade with Gallotiinae Gallotiinae – clade with Gallotiinae Sister to Timon D8 CCVsup (38%) – Lacertoidea Excluded – Lacertoidea Excluded – Lacertoidea Excluded D8 Roq2 (38%) – – Excluded – – Excluded – – Excluded D9 LaC (67%) – Podarcis Podarcis Sister to P. tiliguerta Podarcis; sister to T+L Podarcis sister to P. tiliguerta Gallotiinae; sister to Podarcis T+L; Podarcis; D9 Roq2 (58%) – Podarcis Excluded – Sister to T+L Excluded – Gallotiinae; sister to Excluded T+L D10 Roq2 (54%) Clade with Clade with Clade with Clade with Clade with Clade with Clade with Clade with Clade with Amphisbaenia Amphisbaenia Amphisbaenia Amphisbaenia Amphisbaenia Amphisbaenia Amphisbaenia Amphisbaenia Amphisbaenia CC1sup, Coderet Couche 1 sup; CCVsup, Coderet Couche Verte sup 1-25; CE1-0, Coderet E1-0; CH1-100, Coderet H1-100; CH1-75, Coderet H1-75; Cod, Coderet; FV, Fouilles Viret; LaC, La Colombiere; MdGB, Mas de Got B; PdF, Pech-du-Fraysse; Roq2, Roqueprune 2; T + L: Timon + Lacerta. L. C. M. Wencker et al. / Cladistics 0 (2021) 1–38