The Tritosauria – An Overlooked Third Clade of Lizards

Traditionally there have been just two lizard clades in the Squamata. The Iguania included Iguana, Draco, Phrynosoma and other similar lizards. The Scleroglossa included Tupinambis, Chalcides, Varanus, Heloderma and all the snakes and amphisbaenids. Squamate outgroups within the Lepidosauria included members of the Rhynchocephalia (such as Sphenodon) and the basal lepidosaur, Homoeosaurus, which probably appeared in the Permian, but is only known from the Late Jurassic.

Traditional Nesting
Wikipedia reports the following about the Squamata, “Squamates are a monophyletic  group that is a sister group to the tuatara. The squamates and tuatara together are a sister group to crocodiles and birds, the extant archosaurs.” This is the traditional concept, but testing this in a larger study finds that lizards and archosaurs are not closely related. Not by a long shot.

The Tritosauria, a new lizard clade that was previously overlooked.

Figure 1. Click to enlarge. The Tritosauria, a new lizard clade that was previously overlooked.

The New Heretical Tritosauria
The large study (Peters 2007) recovered a third clade of squamates just outside of the Squamata (Iguania + Scleroglossa), but inside the Lepidosauria (which includes Sphenodon and the other Rhynchocephalia). At the base of this third clade, called the Tritosauria (“third lizards”), are three very lizardy forms, none of which had fused proximal ankle bones, a trait shared by most squamates (at least those that have legs!). Lacertulus, Meyasaurus and Huehuecuetzpalli are known from crushed but articulated fossils. Lacertulus was considered a possible biped (Carroll and Thompson 1982) based on its long hind legs. It is likely that Huehuecuetzpalli (Reynoso 1998) was also a biped. All three were considered close to the base of the lepidosauria, not closely related to any living lizards.

The Tritosauria
A Clade of Misplaced and Enigmatic “Weird-Ohs”

Phylogenetically following Huehuecuetzpalli three distinct clades emerge within the Tritosauria. Some of these were formerly considered “prolacertiforms” (Peters 2000), but now we know that none are related to ProlacertaAll three subclades have some pretty weird members.

The Tanystropheidae
This clade was named by Dilkes (1998) to include “the most recent common ancestor of MacrocnemusTanystropheus and Langobardisaurus and all of its descendants.” Clade members include several long-necked taxa, some of which, like Dinocephalosaurus, preferred swimming to walking. Tanystropheus was the largest, attaining 4.5 meters in length.

The Jesairosauridae
This clade includes Jesairosaurus (Jalil 1991) and the drepanosaurs, from Hypuronector to Drepanosaurus.  This clade included several arboreal, hook-tailed taxa with short-toed feet that were able to grasp slender branches in their slow-motion quest for insects. All were rather small.

The Fenestrasauria
This clade was named by Peters (2000) to include “Cosesaurus, Preondactylus, their common ancestor and all of its descendants.” This clade started off with bipeds that flapped their arms, probably for display during mating rituals because some members, like Longisquama were exotically decorated with extradermal membranes and plumes. Powered gliding (as in Sharovipteryx) was followed by flapping flight in pterosaurs, the first flying vertebrates. Several pterosaurs secondarily developed a quadrupedal pace. Quetzalcoatlus was the largest tritosaur, attaining a wingspan of 10 meters.

Summary
Due to the wide gamut and large inclusion list of the present phylogenetic analysis, many former enigmas, mismatches and leftovers came together in a new clade of lepidosaurs that was previously overlooked. Together, the Tritosauria include some of the strangest and, at times largest, of all lizards. Hyper-elongated necks and hyper-elongated fingers, together with experiments in both a sedentary marine lifestyle (Dinocephalosaurus) and a homeothermic aerial lifestyle (Dimorphodon, for example) make this a truly dynamic and diverse clade. Some of these out-of-the-ordinary morphologies seem to have been kick-started by early experiments with bipedalism. While the arboreal niches of drepanosaurs and pterosaurs are relatively easy to identify, the long-necked tanystropheids may also have used bipedalism and a long neck to reach into tree boughs to snatch prey, creating their own arboreal niche.

Unfortunately, only pterosaurs and Huehuecuetzpalli survived the end of the Triassic and they did not survive the end of the Cretaceous. So tritosaurs are the only major clade of lizards that is extinct today.

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

References
Carroll and Thompson 1982. A bipedal lizardlike reptile fro the Karroo. Journal of Palaeontology 56:1-10.
Peters D 2000. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.
Peters D 2007. The origin and radiation of the Pterosauria. In D. Hone ed. Flugsaurier. The Wellnhofer pterosaur meeting, 2007, Munich, Germany. p. 27.
Reynoso V-H 1998. Huehuecuetzpalli mixtecus gen. et sp. nov: a basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodríguez, Central México. Philosophical Transactions of the Royal Society, London B 353:477-500.

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