Sauropsid

Sauropsida ( Greek for "lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia, though typically used in a broader sense to also include extinct stem-group relatives of modern reptiles and birds (which, as theropod dinosaurs, are nested within reptiles as more closely related to crocodilians than to lizards or turtles).Gauthier J.A. (1994): ''The diversification of the amniotes''. In: D.R. Prothero and R.M. Schoch (ed.) Major Features of Vertebrate Evolution: 129–159. Knoxville, Tennessee: The Paleontological Society. The most popular definition states that Sauropsida is the sibling taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early synapsids have historically been referred to as "mammal-like reptiles", all synapsids are more closely related to mammals than to any modern reptile. Sauropsids, on the other hand, include all amniotes more closely related to modern reptiles than to mammals. This includes Aves (birds), which are recognized as a subgroup of archosaurian reptiles despite originally being named as a separate class in Linnaean taxonomy.

The base of Sauropsida is traditionally divided into main groups of "reptiles": Eureptilia ("true reptiles") and Parareptilia ("next to reptiles"). Eureptilia encompasses all living reptiles (including birds), as well as various extinct groups. Parareptilia is typically considered to be an entirely extinct group, though a few hypotheses for the origin of turtles have suggested that they belong to the parareptiles. The clades Recumbirostra and Varanopidae, traditionally thought to be lepospondyls and synapsids respectively, may also be basal sauropsids. The term "Sauropsida" originated in 1864 with Thomas Henry Huxley, who grouped birds with reptiles based on fossil evidence. The divisions of "Eureptilia" and "Parareptilia" have been challenged in a number of recent studies, who find that they do not represent monophyletic groups.

History of classification

Huxley and the fossil gaps

The term ''Sauropsida'' ("lizard faces") has a long history, and hails back to Thomas Henry Huxley, and his opinion that birds had risen from the dinosaurs. He based this chiefly on the fossils of '' Hesperornis'' and '' Archaeopteryx'', that were starting to become known at the time. In the Hunterian lectures delivered at the Royal College of Surgeons in 1863, Huxley grouped the vertebrate classes informally into mammals, sauroids, and ichthyoids (the latter containing the anamniotes), based on the gaps in physiological traits and lack of transitional fossils that seemed to exist between the three groups. Early in the following year he proposed the names Sauropsida and Ichthyopsida for the two latter. Huxley did however include groups on the mammalian line (synapsids) like '' Dicynodon'' among the sauropsids. Thus, under the original definition, Sauropsida contained not only the groups usually associated with it today, but also several groups that today are known to be in the mammalian side of the tree.

Sauropsids redefined (Goodrich, 1916)

By the early 20th century, the fossils of Permian synapsids from South Africa had become well known, allowing palaeontologists to trace synapsid evolution in much greater detail. The term Sauropsida was taken up by E. S. Goodrich in 1916 much like Huxley's, to include lizards, birds and their relatives. He distinguished them from mammals and their extinct relatives, which he included in the sister group Theropsida (now usually replaced with the name Synapsida). Goodrich's classification thus differs somewhat from Huxley's, in which the non-mammalian synapsids (or at least the dicynodontians) fell under the sauropsids. Goodrich supported this division by the nature of the hearts and blood vessels in each group, and other features such as the structure of the forebrain. According to Goodrich, both lineages evolved from an earlier stem group, the Protosauria ("first lizards"), which included some Paleozoic amphibians as well as early reptiles predating the sauropsid/synapsid split (and thus not true sauropsids). His concept differed from modern classifications in that he considered a modified fifth metatarsal to be an apomorphy of the group, leading him to place Sauropterygia, Mesosauria and possibly Ichthyosauria and Araeoscelida in the Theropsida.

Detailing the reptile family tree

In 1956, D. M. S. Watson observed that sauropsids and synapsids diverged very early in the reptilian evolutionary history, and so he divided Goodrich's Protosauria between the two groups. He also reinterpreted the Sauropsida and Theropsida to exclude birds and mammals respectively, making them paraphyletic, unlike Goodrich's definition. Thus his Sauropsida included Procolophonia, Eosuchia, Protorosauria, Millerosauria, Chelonia (turtles), Squamata (lizards and snakes), Rhynchocephalia, Rhynchosauria, Choristodera, Thalattosauria, Crocodilia, " thecodonts" (paraphyletic basal Archosauria), non- avian dinosaurs, pterosaurs and sauropyterygians. However, his concept differed from the modern one in that he placed reptiles without an otic notch, such as araeoscelids and captorhinids, with the theropsids.

This classification supplemented, but was never as popular as, the classification of the reptiles (according to Romer's classic ''Vertebrate Paleontology'', 3rd ed., 1966.) into four subclasses according to the positioning of '' temporal fenestrae'', openings in the sides of the skull behind the eyes. Since the advent of phylogenetic nomenclature, the term Reptilia has fallen out of favor with many taxonomists, who have used Sauropsida in its place to include a monophyletic group containing the traditional reptiles and the birds.

Cladistic definitions

The class Reptilia has been known to be an evolutionary grade rather than a clade for as long as evolution has been recognised. Reclassifying reptiles has been among the key aims of phylogenetic nomenclature. The term Sauropsida had from the mid 20th century been used to denote a branch-based clade containing all amniote species which are not on the synapsid side of the split between reptiles and mammals. This group encompasses all now-living reptiles as well as birds, and as such is comparable to Goodrich's classification. The main difference is that better resolution of the early amniote tree has split up most of Goodrich's "Protosauria", though definitions of Sauropsida essentially identical to Huxley's (i.e. including the mammal-like reptiles) are also forwarded. Some later cladistic work has used Sauropsida more restrictively, to signify the crown group, i.e. all descendants of the last common ancestor of extant reptiles and birds. A number of phylogenetic stem, node and crown definitions have been published, anchored in a variety of fossil and extant organisms, thus there is currently no consensus of the actual definition (and thus content) of Sauropsida as a phylogenetic unit.

Some taxonomists, such as Benton (2004), have co-opted the term to fit into traditional rank-based classifications, making Sauropsida and Synapsida class-level taxa to replace the traditional Class Reptilia, while Modesto and Anderson (2004), using the PhyloCode standard, have suggested replacing the name Sauropsida with their redefinition of Reptilia, arguing that the latter is by far better known and should have priority.

Cladistic definitions of Sauropsida include:

* Sauropsida as the total group of reptiles: "Reptiles plus all other amniotes more closely related to them than they are to mammals" (Gauthier, 1994). This is a branch-based total group definition. Gauthier (1994) considered turtles to be descended from parareptiles, thus defining Reptilia as a more restricted crown group encompassing diapsids and parareptiles (apart from mesosaurs, which he considered to be the most basal branch of sauropsids).
* Sauropsida as a total group, synonymous with Reptilia sensu lato: "The most inclusive clade containing '' Lacerta agilis'' and '' Crocodylus niloticus'', but not ''Homo sapiens''" (Modesto & Anderson, 2004). This total group definition leaves the question of turtle ancestry unresolved.
* Sauropsida as a broad node-based group: "The last common ancestor of mesosaurs, testudines and diapsids, and all its descendants" (Laurin & Reisz, 1995). Though formulated differently, this grouping was similar in scope and intention to the definition provided by Gauthier (1994).

Subdivisions

Eureptilia ("true reptiles") is one of the two major subgroups of the clade Sauropsida, the other one being Parareptilia. Eureptilia includes Diapsida (the clade containing all modern reptiles and birds), as well as a number of primitive Permo-Carboniferous forms previously classified under Anapsida, in the old (no longer recognised) order " Cotylosauria".

Eureptilia is characterized by the skull having greatly reduced supraoccipital, tabular, and supratemporal bones that are no longer in contact with the postorbital. Aside from Diapsida, the group notably contains Captorhinidae, a diverse and long lived (Late Carboniferous-Late Permian) clade of initially small carnivores that later evolved into large herbivores. Other primitive eureptiles such as the " protorothyrids" were all small, superficially lizard-like forms, that were probably insectivorous. One primitive eureptile, the Late Carboniferous "protorothyrid" ''Anthracodromeus,'' is the oldest known climbing tetrapod. Diapsids were the only eureptilian clade to continue beyond the end of the Permian.

The traditional classification of eureptilians has been challenged in recent studies, with several studies in the early 2020s finding that captorhinids and '' Protorothyris'' are not even sauropsids, but stem-amniotes.

Evolutionary history

Sauropsids evolved from basal amniotes approximately 320 million years ago, in the Carboniferous Period of the Paleozoic Era. In the Mesozoic Era (from about 250 million years ago to about 66 million years ago), sauropsids were the largest animals on land, in the water, and in the air. The Mesozoic is sometimes called the Age of Reptiles. In the Cretaceous–Paleogene extinction event, the large-bodied sauropsids died out in the global extinction event at the end of the Mesozoic era. With the exception of a few species of birds, the entire dinosaur lineage became extinct; in the following era, the Cenozoic, the remaining birds diversified so extensively that, today, nearly one out of every three species of land vertebrate is a bird species.

Phylogeny

The cladogram presented here illustrates the "family tree" of sauropsids, and follows a simplified version of the relationships found by M.S. Lee, in 2013. All genetic studies have supported the hypothesis that turtles (formerly categorized together with ancient '' anapsids'') are diapsid reptiles, despite lacking any skull openings behind their eye sockets; some studies have even placed turtles among the archosaurs, though a few have recovered turtles as lepidosauromorphs instead. The cladogram below used a combination of genetic (molecular) and fossil (morphological) data to obtain its results.

Laurin & Piñeiro (2017) and Modesto (2019) proposed an alternate phylogeny of basal sauropsids. In this tree, parareptiles include turtles and are closely related to non-araeoscelidian diapsids. The family Varanopidae, otherwise included in Synapsida, is considered by Modesto a sauropsid group.

In a number of recent studies, the " microsaur" clade Recumbirostra, historically considered lepospondyl reptiliomorphs, have been recovered as early sauropsids, though this assertion has been disputed by a number of authors, who contend that microsaurs are still reptillomorph stem-amniotes.

Simoes et al (2022) found Captorhinidae, '' Protorothyris'' and Araeoscelidia to form a clade that was the sister group to crown Amniota (containing true sauropsids and synapsids). The same study also considered parareptiles to be polyphyletic, with some groups being closer to the crown group of reptiles than others.

Cladogram after Simoes et al (2022):

Structure difference with synapsids

The last common ancestor of synapsids and Sauropsida lived at around 320mya during Carboniferous, known as Reptiliomorpha.

Thermal and secretion

The early synapsids inherited abundant glands on their skins from their amphibian ancestors. Those glands evolved into sweat glands in synapsids, which granted them the ability to maintain constant body temperature but made them unable to save water from evaporation. Moreover, the way synapsids discharge nitrogenous waste is through urea, which is toxic and must be dissolved in water to be secreted. Unfortunately, the upcoming Permian and Triassic periods were arid periods. As a result, only a small percent of early synapsids survived in the land from South Africa to Antarctica in today's geography. Unlike synapsids, sauropsids do not have those glands on the skin; their way of nitrogenous waste emission is through uric acid which does not require water and can be excreted with feces. As a result, sauropsids were able to expand to all environments and reach their pinnacle. Even today, most vertebrates that live in arid environments are sauropsids, snakes and desert lizards for example.

Brain structure

Different from how synapsids have their cortex in six different layers of neurons which is called neocortex, the cerebrum of Sauropsida has a completely different structure. For the corresponding structure of the cerebrum in the classic view, the neocortex of synapsids is homology with only the archicortex of the avian brain. However, in the modern view appeared since the 1960s, behavioral studies suggested that avian neostriatum and hyperstriatum can receive signals of vision, hearing, and body sensations, which means they act just like the neocortex. Comparing an avian brain to that to a mammal, nuclear-to-layered hypothesis proposed by Karten (1969), suggested that the cells which form layers in synapsids' neocortex, gather individually by type and form several nuclei. For synapsids, when one new function is adapted in evolution it will be assigned to a separate area of cortex, so for each function, synapsids will have to develop a separate area of cortex, and damage to that specific cortex may cause disability. However, for Sauropsida functions are disassembled and assigned to all nuclei. In this case, brain function is highly flexible for Sauropsida, even with a small brain, many Sauropsida can still have a relatively high intelligence compared to mammals, for example, birds in the family Corvidae. So, it is possible that some non-avian dinosaurs, like '' Tyrannosaurus,'' which had tiny brains compared to their enormous body size, were more intelligent than previously thought.Jarvis, Güntürkün, O., Bruce, L., Csillag, A., Karten, H., Kuenzel, W., Medina, L., Paxinos, G., Perkel, D. J., Shimizu, T., Striedter, G., Wild, J. M., Ball, G. F., Dugas-Ford, J., Durand, S. E., Hough, G. E., Husband, S., Kubikova, L., Lee, D. W., ... Butler, A. B. (2005). "Avian brains and a new understanding of vertebrate brain evolution". ''Nature Reviews''. Neuroscience, 6(2), 151–159. .

References

{{Taxonbar, from=Q329457

Amniotes
Extant Pennsylvanian first appearances
Fossil taxa described in 1956
Taxa named by D. M. S. Watson