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Crown Group Lizards (Reptilia, Squamata) from the Middle Jurassic of the British Isles

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Abstract

An assemblage of crown-group lizards is described from a middle Jurassic (Bathonian) horizon in the British Isles (Kirlington Quarry, Oxfordshire). These are amongst the earliest true lizards on record. At least five distinct genera (four of them new) are represented with three scincomorphs, an anguimorph (probably varanoid) and possible gekkotans. The assemblage resembles that of later Jurassic and early Cretaceous localities in being dominated by scincomorphs. These include one or more paramacellodids and a highly derived form which was apparently long-bodied with reduced limbs. The taxonomic diversity of the assemblage supports the view that the radiation of true squamates had begun by at least the early Jurassic, despite the absence of any reliable Triassic or early Jurassic record. Taxonomic and palaeogeographic implications are discussed.
... The Toxicofera [34] are split in this tree as a the Iguania are placed with other unidentatan squamates in a sister group including Scincomorpha, Lacertoidea and †Polyglyphanodontia with Elgaria as an anomaly (it is usually positioned as an anguimorph). The large clade of unidentatan squamates lies crownwards of †Scandensia ciervensis [38], †Hongshanxi xeie [39] and the Gekkonomorpha (including †Eichstaettisaurus schroederi [40]), while †Bellairsia [41], †Oculudentavis khaungraae [10] and †Huehuecuetzpalli mixtecus [42] are more basal. We treated the †Cryptovaranoides premaxillae as unfused in this analysis to encompass the juvenile holotype only. ...
... The Gekkonomorpha, including †Eichstaettisaurus schroederi [40], lie more basally, in a polytomy with †Hongshanxi xeie [39] and †Scandensia ciervensis [38]. This grouping is at the base of the Squamata but is crownwards of the pan-squamates †Oculudentavis khaungraae [10], †Bellairsia [41] and †Huehuequetzpalli mixtecus [42]. ...
... When the same matrix is processed using the molecular constraints and methodology of [10], and removing †Vellbergia and †Scandensia as rogue taxa, †Cryptovaranoides is in a poorly resolved polytomy of unidentatan and gekkonomorph crown squamates (figure 7b). This includes †Hongshanxi xiei [39], †Ardeosaurus brevipes [37] and †Eichstaettisaurus schroederi [40] but all are crownward of †Oculudentavis khaungraae [10], †Bellairsia [41] and †Huehuequetzpalli mixtecus [42]. In these analyses based on morphology only and morphology with molecular constraints, †Cryptovaranoides is therefore certainly the oldest described crown squamate, as we have previously stated [1] and when better resolved is positioned as a unidentatan, crownwards of the Gekkonomorpha. ...
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†Cryptovaranoides microlanius from the latest Triassic of England was described in 2022 as a crown-clade squamate, of importance as the oldest such modern-type lizard, extending their temporal range downward by 35 Myr. This view was challenged in 2023, and †Cryptovaranoides was reinterpreted as an archosauromorph. These decisions matter because the original view has an impact on our understanding of the early stages of squamate evolution; the revised view removes the species from such a role. The revisers emphasized the need to make careful observations of the fossils and to interpret the morphological data appropriately in terms of relationships; here, we find many errors of observation and interpretation in the work of the revisers, and we correct these with reference to the fossils, both in the rock and in the computed tomography scans we had made for the original description. We show that when the observational errors are corrected and the taxa recoded, every phylogenetic analysis confirms our original conclusion that †Cryptovaranoides is not an archosauromorph, but a lepidosauromorph, a lepidosaur, a pan-squamate and a crown squamate.
... This is a focus for the "tree of life" program because correctly dated phylogenetic trees provide insights into origins of biodiversity, innovations in evolution, and the impact of environmental drivers on macroevolution. Dating squamate origins, a clade with more than 11,000 extant species (1), has proved problematic, with estimates as latest Early Jurassic [~175 million years (Ma)] from fossils (2)(3)(4)(5), Late Triassic (~205 Ma) from molecular data (6), and Triassic-Jurassic (172 to 213 Ma) from combined data (7,8). The oldest examples of modern squamates are incomplete Middle Jurassic fossils [~168 Ma; (2)(3)(4)(5)], and the first articulated skeletons occur in the Late Jurassic Solnhofen Formation [146 to 151 Ma; (9)]. ...
... Dating squamate origins, a clade with more than 11,000 extant species (1), has proved problematic, with estimates as latest Early Jurassic [~175 million years (Ma)] from fossils (2)(3)(4)(5), Late Triassic (~205 Ma) from molecular data (6), and Triassic-Jurassic (172 to 213 Ma) from combined data (7,8). The oldest examples of modern squamates are incomplete Middle Jurassic fossils [~168 Ma; (2)(3)(4)(5)], and the first articulated skeletons occur in the Late Jurassic Solnhofen Formation [146 to 151 Ma; (9)]. ...
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Mammals, birds, and squamates (lizards, snakes, and relatives) are key living vertebrates, and thus understanding their evolution underpins important questions in biodiversity science. Whereas the origins of mammals and birds are relatively well understood, the roots of squamates have been obscure. Here, we report a modern-type lizard from the Late Triassic of England [202 million years (Ma)], comprising a partial skeleton, skull, and mandibles. It displays at least 15 unique squamate traits and further shares unidentatan and anguimorph apomorphies. The new discovery fixes the origin of crown Squamata as much older than had been thought, and the revised dating shows substantial diversification of modern-type squamates following the Carnian Pluvial Episode, 232 Ma ago.
... Extant squamates have a near global distribution except for polar areas (Vitt & Caldwell 2014), but the early Mesozoic record of squamates is sparse, with the earliest stem squamate Megachirella wachtleri † Renesto & Posenato, 2003 found in the mid-Triassic of Europe ). The Jurassic is better represented, with most specimens located in Europe (Hoffstetter 1964(Hoffstetter , 1967Waldman & Savage 1972;Seiffert 1975;Estes 1983;Evans 1994aEvans , 1998Caldwell et al. 2015;Conrad 2018), as well as North America (Prothero & Estes 1980;Evans & Chure 1998b;Evans 1996;Brownstein et al. 2022) and Asia (Hecht & Hecht 1984;Federov & Nessov 1992;Nydam et al. 2013;Dong et al. 2019;Meyer et al. 2023). There is also some fragmentary Jurassic evidence of squamates in Africa (Broschinski 1999). ...
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A pressing issue in historical biogeography for clades with long evolutionary histories and intercontinental distributions is the ever-changing position of tectonic plates. Over time, this shifting alters the proximity between areas, a factor that very few models can easily account. In addition, the absence of fossils from most molecular phylogenies often erases biogeographic signals not retained in trees of extant taxa. Here, we adapt the paleogeographic model from Landis (2017), a potentially powerful tool for ancestral-range estimation at global scales through time. This model creates “communicating classes” of areas by assigning discrete geographic units to adjacency matrices that change across multiple time slices in the Phanerozoic to model continental drift. We apply this algorithm to three existing total-evidence datasets incorporating extinct and extant tips from the order Squamata. Our results corroborate the origin of Squamata on the Eurasian continent – specifically Europe and northeastern Asia – although phylogenies sampling more Jurassic squamate lineages showed higher support for a purely European origin. Eurasia continued to be a major source of diversification throughout the Mesozoic, with dispersals into North America in the Late Jurassic and South America in the mid-Cretaceous. For Serpentes, the ancestral ranges were unclear and inconsistent across the phylogenetic hypotheses, likely influenced by the disparate and incomplete sampling in the three phylogenies.
... Anurans (frogs) and lizards are among the most widespread ectothermic vertebrates today. They were also important components of ancient terrestrial ecosystems, with the fossil record of frogs extending back to the Upper Triassic (Rage and Roček, 1989;Stocker et al., 2019) and the record of lizards extending to the Middle Jurassic (Evans, 1998;Talanda et al., 2022). Yet their fossil record is poor, and their relative abundance is only known from locales where isolated bones have been recovered by intensive screen-washing of weakly lithified strata (Eaton, 2004), or in Lagerstätten where articulated or soft-tissuebearing specimens occur (e.g., Trueb et al., 2005;Čerňanský et al., 2022;Du et al., 2024). ...
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Vertebrate tracks are described from the upper Campanian Kaiparowits Formation of Grand Staircase-Escalante National Monument, Utah, U.S.A. The traces are hypothesized to have been deposited along the periphery of a shallow alkaline lake or pond. One type of track consists of a quadripartite radiating array of small tapering digit imprints which may represent the impression of a manus. A second type of tracks consists of a pentameric radiating array of small tapering digit imprints that is associated with an elongate non-tapering impression that extends in the opposite direction from the fan-shaped digit array. The latter resembles a pes imprint. An elongate impression that is associated with one of the pes imprints, may represent a drag mark. One set of manus and pes tracks appears to be associated and shares similarities with anuran tracks. An alternative interpretation is that these fossils represent partially preserved lizard tracks. These tracks differ substantially from dinosaur tracks from the Kaiparowits Formation, which are illustrated for comparison.
... We report a near-complete (about 70%) skeleton ( Fig. 1 and Extended Data Fig. 1 21 . However, the disassociated nature of these specimens has limited their usefulness for understanding early squamate evolution. ...
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Squamates (lizards and snakes) include more than 10,000 living species, descended from an ancestor that diverged more than 240 million years ago from that of their closest living relative, Sphenodon. However, a deficiency of fossil evidence1–7, combined with serious conflicts between molecular and morphological accounts of squamate phylogeny8–13 (but see ref. ¹⁴), has caused uncertainty about the origins and evolutionary assembly of squamate anatomy. Here we report the near-complete skeleton of a stem squamate, Bellairsia gracilis, from the Middle Jurassic epoch of Scotland, documented using high-resolution synchrotron phase-contrast tomography. Bellairsia shares numerous features of the crown group, including traits related to cranial kinesis (an important functional feature of many extant squamates) and those of the braincase and shoulder girdle. Alongside these derived traits, Bellairsia also retains inferred ancestral features including a pterygoid–vomer contact and the presence of both cervical and dorsal intercentra. Phylogenetic analyses return strong support for Bellairsia as a stem squamate, suggesting that several features that it shares with extant gekkotans are plesiomorphies, consistent with the molecular phylogenetic hypothesis that gekkotans are early-diverging squamates. We also provide confident support of stem squamate affinities for the enigmatic Oculudentavis. Our findings indicate that squamate-like functional features of the suspensorium, braincase and shoulder girdle preceded the origin of their palatal and vertebral traits and indicate the presence of advanced stem squamates as persistent components of terrestrial assemblages up to at least the middle of the Cretaceous period.
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