Pan-Lepidosauria J. A. Gauthier and K. de Queiroz, new clade name

Registration Number: 118 2009; Evans and Borsuk-Bialylicka, 2009; Evans and Jones, 2010; but see Müller, 2004; De!nition: !e total clade of the crown clade Jones et al., 2013). More recently, kuehneosaurs Lepidosauria. !is is a crown-based total-clade have been inferred to be stem archosaurs close de"nition. Abbreviated de"nition: total ∇ of to Trilophosaurus buettneri by Pritchard and Lepidosauria. Nesbitt (2017), although Simões et al. (2018) placed them as either stem saurians or stem Etymology: Derived from pan (Greek), here archosaurs depending on the analysis, but in referring to “pan-monophylum,” another term either case only distantly related to T. buettneri. for “total clade,” and Lepidosauria, the name !e Early Triassic Sophineta cracoviensis and the of the corresponding crown clade; hence, “the Middle Jurassic Marmoretta oxoniensis have been total clade of Lepidosauria.” more consistently regarded as stem lepidosaurs, although that inference depends upon cor- Reference Phylogeny: Gauthier et al. (1988) rect association among disarticulated remains Figure 13, where the clade in question is named (Evans, 1991; Waldman and Evans, 1994; Lepidosauromorpha and is hypothesized to Evans, 2009; Evans and Borsuk-Bialylicka, include Younginiformes, which is no longer con- 2009; Evans and Jones, 2010; Jones et al., 2013; sidered part of the clade (see Composition). Renesto and Bernardi, 2014). Depending on the analysis, Simões et al. (2018) inferred S. Composition: Pan-Lepidosauria is composed cracoviensis to be either a stem lepidosaurian of Lepidosauria (Pan-Sphenodon plus Pan- or a stem squamatan, but they consistently Squamata, see entry in this volume) and all inferred M. oxoniensis as sister to Megachirella extinct species (stem lepidosaurs) that share a wachtleri on the squamate stem. Renesto and more recent common ancestor with the species Bernardi (2014) previously placed the Middle of Lepidosauria than they do with any other Triassic Megachirella wachtleri either inside or extant amniotes (Aves, Testudines, Crocodylia, outside of the lepidosaurian crown. !e Early and Mammalia). !e Late Permian fossils or Middle Jurassic Tamaulipasaurus morenoi Saurosternon bainii and Lanthanolania ivakh- may be a stem or a crown lepidosaur, but its nenkoi, as well as the Permo-Triassic fossils relationships are di#cult to assess because of Paliguana whitei and Palaeagama vielhaueri, its high degree of modi"cation for head-"rst have sometimes been inferred to be stem lepido- burrowing (Clark and Hernandez, 1994). !e saurs (e.g., Gauthier et al., 1988; Modesto and highly modi"ed Late Triassic drepanosaurs Reisz, 2002; Evans and Jones, 2010) but other have sometimes been inferred to be stem archo- times to be stem saurians (e.g., Modesto and saurs (Laurin, 1991; Simões et al., 2018) or stem Reisz, 2002; Müller, 2004). !e Upper Triassic lepidosaurs (Evans, 2009), but most authors Kuehneosauridae and its inferred Early Triassic have inferred drepanosaurs to be stem sauri- relative Pamelina polonica (but see Simões et ans that must accordingly have diverged much al., 2018) have often been inferred to be stem earlier than the fossil record currently indicates lepidosaurs (Gauthier et al., 1988; Evans, 1988, (i.e., by the Mid-Permian; Guadalupian) (e.g., Pan-Lepidosauria

Müller, 2004; Senter, 2004; Renesto et al., Synonyms: Lepidosauria Haeckel 1866 of vari- 2010; Pritchard and Nesbitt, 2017; Simões et al., ous authors (e.g., Romer, 1933, 1945, 1956, 2018). !e authors of several early phylogenetic 1966; Gardiner, 1982) and Lepidosauromorpha analyses inferred that Younginiformes are stem Gauthier in Benton 1983 of Benton (1983, lepidosaurs (e.g., Benton, 1985; Evans, 1988; 1985) and Evans (1984) are approximate syn- Gauthier et al., 1988); however, more recent onyms. Eosuchia (e.g., of Romer, 1933, 1945, analyses have placed Younginiformes, either as a 1956, 1966), characterized as being composed clade or a paraphylum, on the stem of a more of “primitive lepidosaurians” and “direct ances- inclusive crown clade composed of (in addi- tors of the lizards and snakes”, and Holapsida tion to lepidosaurs) archosaurs and sometimes Underwood 1957, composed of “Eosuchia and also turtles (e.g., Laurin, 1991; Gauthier, 1994; Rhynchocephalia”, are partial (and approximate) Caldwell, 1996; deBraga and Rieppel, 1997; synonyms. Eolacertilia Romer 1966 (often Müller, 2004; Evans and Borsuk-Bialylicka, incorrectly attributed to Robinson, 1967) sensu 2009; Renesto and Bernardi, 2014; Bever et al., Carroll (1975, 1977) and Estes (1983) is a par- 2015; Pritchard and Nesbitt, 2017; Simões et al., tial (and approximate) synonym in the context 2018). See Evans and Jones (2010) for other taxa of phylogenies that place all of the included taxa that are no longer considered stem lepidosaurs. in the lepidosaur stem group (see Composition). Lepidosauromorpha Gauthier in Benton 1983 Diagnostic Apomorphies: As a total clade of Gauthier et al. (1988) is an unambiguous (and therefore also a maximum clade), Pan- synonym. Lepidosauria is not necessarily expected to have diagnostic apomorphies (characters that Comments: Although the name Lepidosauro- arose simultaneously with the split of lepido- morpha has previously been de"ned phylogeneti- saur stem lineage from that of archosaurs; see cally as applying to the total clade of Lepidosauria de Queiroz, 2007); however, any of the apo- (Gauthier et al., 1988), we have chosen instead morphies that originated along the lepidosaur to apply the name Pan-Lepidosauria to that clade stem lineage (see Diagnostic Apomorphies for in the interest of promoting a standardized form Lepidosauria, this volume), as well as those diag- for the names of total clades (e.g., de Queiroz and nosing various side branches, would allow refer- Gauthier, 1992; Gauthier and de Queiroz, 2001; ral of a specimen or species to Pan-Lepidosauria. de Queiroz, 2007). If the name Lepidosauromor- Uncertainty in placement of several incomplete, pha is to be retained, it could be applied to a deep and often highly modi"ed, fossils further com- node within Pan-Lepidosauria, although that plicates the question. If, as we suspect, the Early would require careful formulation of the (non- Triassic Sophineta cracoviensis is a stem lepido- standard) de"nition, given the uncertain status of saur, it shares the following apomorphies with many of its potential early members (see Compo- the crown: (1) maxilla participates broadly in sition). Alternatively, the name could be applied ventral orbital margin; (2) postfrontal wraps to a clade coinciding with the origin of one of the around fronto-parietal suture; (3) lacrimal early-evolving lepidosaur apomorphies. reduced, con"ned largely to orbital rim; (4) In the context of phylogenies in which marginal teeth attached super"cially to lingual Sphenodon punctatus is more closely related to margin of jaw (pleurodont dentition); and (5), turtles, crocodilians, and birds than to Squamata a weak zygosphene-zygantrum accessory inter- (e.g., Hedges and Poling, 1999; Zardoya and vertebral articulation. Meyer, 2000), the names Lepidosauria (this

1076 Pan-Lepidosauria volume) and Reptilia (this volume) are syn- Clark, J. M., and R. Hernandez. 1994. A new bur- onyms, and Reptilia is to be granted precedence rowing diapsid from the Jurassic La Boca (see Lepidosauria in this volume). In the context Formation of Tamaulipas, Mexico. J. Vertebr. of such phylogenies, the name Pan-Lepidosauria Paleontol. 14:180–195. Estes, R. 1983. Sauria terrestria, Amphisbaenia. should not be applied to any clade (ICPN Art. Handbuch der Paläoherpetologie. Teil 10A. 14.5; Cantino and de Queiroz, 2020). Gustav Fischer, Stuttgart. Sophineta cracoviensis and possibly Pamelina Evans, S. E. 1984. !e classi"cation of the polonica (see Composition) are the earliest known Lepidosauria. Zool. J. Linn. Soc. 82:87–100. pan-lepidosaurs (~245Ma, late Olenekian, Early Evans, S. E. 1988. !e early history and relation- Triassic; Evans, 2009; Evans and Borsuk− ships of the Diapsida. Pp. 221–260 in !e Białynicka, 2009). But the initial divergence Phylogeny and Classi"cation of the Tetrapods. of Pan-Lepidosauria from Pan-Archosauria (this Vol. 1: Amphibians, Reptiles, Birds (M. J. Benton, ed.). Systematics Association Special volume) is likely older ( 265 Ma), based on the ~ Volume No. 35A. Clarendon Press, Oxford. age of the earliest-known putative pan-archo- Evans, S. E. 1991. A new lizard-like reptile (Diapsida: saur, Aenigmastropheus parringtoni (Capitanian, Lepidosauromorpha) from the Middle Jurassic mid-Late Permian; Ezcurra et al., 2014). of England. Zool. J. Linn. Soc. 103:391–412. Evans, S. E. 2009. An early kuehneosaurid reptile (Reptilia: Diapsida) from the Early Triassic of Literature Cited Poland. Palaeontol. Pol. 65:145–178. Evans, S. E., and M. Borsuk−Białynicka. 2009. A Benton, M. 1983. !e Triassic reptile Hyperodapedon small lepidosauromorph reptile from the Early from Elgin: functional morphology and rela- Triassic of Poland. Palaeontol. Pol. 65:179–202. tionships. Philos. Trans. R. Soc. Lond. B Biol. Evans, S. E., and M. E. H. Jones. 2010. !e origin, Sci. 302:605–717. early history and diversi"cation of lepidosau- Benton, M. 1985. Classi"cation and phylogeny of the romorph reptiles. Pp. 27–44 in New Aspects of diapsid reptiles. Zool. J. Linn. Soc. 84:97–164. Mesozoic Biodiversity (S. Bandyopadhyay, ed.). Bever, G. S., T. R. Lyson, D. J. Field, and B.-A. S. Lecture Notes in Earth Sciences 132. Bhullar. 2015. Evolutionary origin of the turtle Ezcurra, M. D., T. M. Scheyer, and R. J. Butler. skull. Nature 525:239–242. 2014. !e origin and early evolution of Sauria: deBraga, M., and O. Rieppel. 1997. Reptile phylog- reassessing the Permian saurian fossil record eny and the interrelationships of turtles. Zool. and the timing of the crocodile-lizard diver- J. Linn. Soc. 120:281–354. gence. PLOS ONE 9(2):e89165. Caldwell, M. 1996. Ichthyosauria: a preliminary phy- Gardiner, B. G. 1982. Tetrapod classi"cation. Zool. logenetic analysis of diapsid a#nities. Neues J. Linn. Soc. 74:207–232. Jahrb. Geol. Paläontol. Abh. 200:361–386. Gauthier, J. A. 1994. !e diversi"cation of the amni- Cantino, P. D., and K. de Queiroz. 2020. otes. Pp. 129–159 in Major Features of Vertebrate International Code of Phylogenetic Nomenclature Evolution: Short Courses in Paleontology (D. (PhyloCode), Version 6. CRC Press, Boca Prothero, ed.). Paleontological Society. Raton, FL. Gauthier, J. A., and K. de Queiroz. 2001. Feathered Carroll, R. L. 1975. Permo-Triassic “lizards” from dinosaurs, &ying dinosaurs, crown dino- the Karroo. Palaeontol. Afr. 18:17–87. saurs, and the name “Aves”. Pp. 7–41 in New Carroll, R. L. 1977. !e origin of lizards. Pp. 359– Perspectives on the Origin and Early Evolution 396 in Problems in Vertebrate Evolution (S. of Birds: Proceedings of the International Andrews, R. Miles and A. Walker, eds.). Linn. Symposium in Honor of John H. Ostrom (J. Soc. London, Symp. Ser. 4. Academic Press, Gauthier and L. F. Gall, eds.). Peabody Mus. London. Nat. Hist., Yale Univ., New Haven, CT.

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