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Sauropsida

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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).[2] 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.

Sauropsids
Temporal range: Pennsylvanian (Late Carboniferous)-Present[1]
Clockwise from top left: Pareiasaurus (an extinct pareiasaurian parareptile), Mesosaurus (an extinct mesosaurian parareptile), Smaug breyeri (a lizard), Dinemellia dinemelli (the white-faced buffalo-weaver), Crocodylus niloticus (the Nile crocodile), and Labidosaurikos (an extinct captorhinid eureptile)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Superclass: Tetrapoda
Clade: Reptiliomorpha
Clade: Amniota
Clade: Sauropsida
Watson, 1956
Subclades

The base of Sauropsida forks into two 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,[3] who grouped birds with reptiles based on fossil evidence.

History of classification

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Huxley and the fossil gaps

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The Berlin specimen of Archaeopteryx lithographica, a historically important fossil which helped to establish birds as a component of the reptile family tree

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.[4] 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.[3] 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.[5]

Sauropsids redefined (Goodrich, 1916)

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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.[5]

Detailing the reptile family tree

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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 reptiles without an otic notch, such as araeoscelids and captorhinids, were believed to be theropsids.[6]

This classification supplemented, but was never as popular as, the classification of the reptiles (according to Romer's classic Vertebrate Paleontology[7]) 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

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Sauropsida and the 19th-/20th-century conception of the class Reptilia. Both are superimposed on a cladogram of tetrapods, showing the difference in coverage.

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.[8] 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.[9][10] 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.[11]

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.[11]

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).[2] 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).[11] 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).[12] Though formulated differently, this grouping was similar in scope and intention to the definition provided by Gauthier (1994).

Evolutionary history

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Main article: Evolution of reptiles
 
Mesozoic sauropsids: non-avialan dinosaurs (Europasaurus and iguanodonts) alongside the early bird Archaeopteryx perched on the foreground tree stump.

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

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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.[13] 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,[13][14][15][16][17][18] though a few have recovered turtles as lepidosauromorphs instead.[19] The cladogram below used a combination of genetic (molecular) and fossil (morphological) data to obtain its results.[13]

Amniota

Synapsida (mammals and their extinct relatives)  

Sauropsida /
†Parareptilia
Eureptilia
Reptilia (total group)

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.[20][21]

In recent studies, the "microsaur" clade Recumbirostra, historically considered lepospondyl reptiliomorphs, have been recovered as early sauropsids.[22][23]

A 2024 study defines Captorhinidae and Araeoscelidia as sister groups that split off before the formation of crown amniota (synapsids and sauropsids). The same study also considers parareptiles to be polyphyletic, with some groups being closer to the crown group of reptiles than others.[24]

Structure difference with synapsids

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

Thermal and secretion

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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

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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.[25] 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.[26]

References

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  1. ^ Marjanović, D. (2021). "The Making of Calibration Sausage Exemplified by Recalibrating the Transcriptomic Timetree of Jawed Vertebrates". Frontiers in Genetics. 12. 521693. doi:10.3389/fgene.2021.521693. PMC 8149952. PMID 34054911.
  2. ^ a b 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.
  3. ^ a b Huxley, Thomas Henry (1864). "The Structure and Classification of the Mammalia". Medical Times and Gazette. Huxley Archives. Retrieved 2023-03-16.
  4. ^ Huxley, Thomas Henry (1877). "Lectures on Evolution". Collected Essays IV. Retrieved 2023-03-16. {{cite book}}: |website= ignored (help)
  5. ^ a b Goodrich, E.S. (1916). "On the classification of the Reptilia". Proceedings of the Royal Society of London. 89B (615): 261–276. Bibcode:1916RSPSB..89..261G. doi:10.1098/rspb.1916.0012.
  6. ^ Watson, D.M.S. (1957). "On Millerosaurus and the early history of the sauropsid reptiles". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 240 (673): 325–400. Bibcode:1957RSPTB.240..325W. doi:10.1098/rstb.1957.0003.
  7. ^ Romer, A.S. (1933). Vertebrate Paleontology. University of Chicago Press., 3rd ed., 1966.
  8. ^ Gauthier, .A., Kluge, A.G & Rowe, T. (1988). The early evolution of the Amniota. Pages 103–155 in Michael J. Benton (ed.): The Phylogeny and Classification of the Tetrapods, Volume 1: Amphibians, Reptiles, Birds. Syst. Ass. Spec. Vol. 35A. Clarendon Press, Oxford.
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  10. ^ Pearse, A.S. (ed, 1947): Zoological Names: a List of Phyla, Classes, and Orders. Prepared for Section F, American Association for the Advancement of Science. Second edition. Durham, North Carolina, U.S.A., pp. 1-22
  11. ^ a b c Modesto, S.P.; Anderson, J.S. (2004). "The phylogenetic definition of Reptilia". Systematic Biology. 53 (5): 815–821. doi:10.1080/10635150490503026. PMID 15545258.
  12. ^ Laurin, Michel; Reisz, Robert R. (1995). "A reevaluation of early amniote phylogeny". Zoological Journal of the Linnean Society. 113 (2): 165–223. doi:10.1111/j.1096-3642.1995.tb00932.x.
  13. ^ a b c Lee, M.S.Y. (2013). "Turtle origins: Insights from phylogenetic retrofitting and molecular scaffolds". Journal of Evolutionary Biology. 26 (12): 2729–2738. doi:10.1111/jeb.12268. PMID 24256520. S2CID 2106400.
  14. ^ Mannen, Hideyuki; Li, Steven S.-L. (Oct 1999). "Molecular evidence for a clade of turtles". Molecular Phylogenetics and Evolution. 13 (1): 144–148. doi:10.1006/mpev.1999.0640. PMID 10508547.
  15. ^ Zardoya, R.; Meyer, A. (1998). "Complete mitochondrial genome suggests diapsid affinities of turtles". Proc Natl Acad Sci U S A. 95 (24): 14226–14231. Bibcode:1998PNAS...9514226Z. doi:10.1073/pnas.95.24.14226. ISSN 0027-8424. PMC 24355. PMID 9826682.
  16. ^ Iwabe, N.; Hara, Y.; Kumazawa, Y.; Shibamoto, K.; Saito, Y.; Miyata, T.; Katoh, K. (2004-12-29). "Sister group relationship of turtles to the bird-crocodilian clade revealed by nuclear DNA-coded proteins". Molecular Biology and Evolution. 22 (4): 810–813. doi:10.1093/molbev/msi075. PMID 15625185.
  17. ^ Roos, Jonas; Aggarwal, Ramesh K.; Janke, Axel (Nov 2007). "Extended mitogenomic phylogenetic analyses yield new insight into crocodylian evolution and their survival of the Cretaceous–Tertiary boundary". Molecular Phylogenetics and Evolution. 45 (2): 663–673. doi:10.1016/j.ympev.2007.06.018. PMID 17719245.
  18. ^ Katsu, Y.; Braun, E.L.; Guillette, L.J. Jr.; Iguchi, T. (2010-03-17). "From reptilian phylogenomics to reptilian genomes: Analyses of c-Jun and DJ-1 proto-oncogenes". Cytogenetic and Genome Research. 127 (2–4): 79–93. doi:10.1159/000297715. PMID 20234127. S2CID 12116018.
  19. ^ Lyson, Tyler R.; Sperling, Erik A.; Heimberg, Alysha M.; Gauthier, Jacques A.; King, Benjamin L.; Peterson, Kevin J. (2012-02-23). "MicroRNAs support a turtle + lizard clade". Biology Letters. 8 (1): 104–107. doi:10.1098/rsbl.2011.0477. PMC 3259949. PMID 21775315.
  20. ^ Laurin, Michel; Piñeiro, Graciela H. (2017). "A reassessment of the taxonomic position of mesosaurs, and a surprising phylogeny of early amniotes". Frontiers in Earth Science. 5: 88. Bibcode:2017FrEaS...5...88L. doi:10.3389/feart.2017.00088. hdl:20.500.12008/33548. ISSN 2296-6463.
  21. ^ Modesto, Sean P. (January 2020). "Rooting about reptile relationships". Nature Ecology & Evolution. 4 (1): 10–11. doi:10.1038/s41559-019-1074-0. ISSN 2397-334X. PMID 31900449. S2CID 209672518. Retrieved 29 December 2020.
  22. ^ Pardo, Jason D.; Szostakiwskyj, Matt; Ahlberg, Per E.; Anderson, Jason S. (June 2017). "Hidden morphological diversity among early tetrapods". Nature. 546 (7660): 642–645. Bibcode:2017Natur.546..642P. doi:10.1038/nature22966. hdl:1880/113382. ISSN 1476-4687. PMID 28636600. S2CID 2478132.
  23. ^ Mann, Arjan; Pardo, Jason D.; Maddin, Hillary C. (2019-09-30). "Infernovenator steenae, a new serpentine recumbirostran from the 'Mazon Creek' Lagerstätte further clarifies lysorophian origins". Zoological Journal of the Linnean Society. 187 (2): 506–517. doi:10.1093/zoolinnean/zlz026. ISSN 0024-4082.
  24. ^ Successive climate crises in the deep past drove the early evolution and radiation of reptiles
  25. ^ Karten, H. J. in Comparative and Evolutionary Aspects of the Vertebrate Central Nervous System (ed. Pertras, J.) 164–179 (1969).
  26. ^ 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. doi:10.1038/nrn1606.
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