Clade of tetrapods including reptiles, birds and mammals
Amniotes are tetrapod vertebrate animals belonging to the clade Amniota , a large group that comprises the vast majority of living terrestrial and semiaquatic vertebrates. Amniotes evolved from amphibious stem tetrapod ancestors during the Carboniferous period . Those of Amniota are defined as the smallest crown clade containing humans , the Greek tortoise , and the Nile crocodile .[ 4] [ 5]
Amniotes are distinguished from the other living tetrapod clade — the non-amniote lissamphibians (frogs /toads , salamanders , newts and caecilians ) — by the development of three extraembryonic membranes (amnion for embryonic protection, chorion for gas exchange , and allantois for metabolic waste disposal or storage), thicker and keratinized skin , costal respiration (breathing by expanding/constricting the rib cage ), the presence of adrenocortical and chromaffin tissues as a discrete pair of glands near their kidneys , more complex kidneys , the presence of an astragalus for better extremity range of motion , the diminished role of skin breathing , and the complete loss of metamorphosis , gills , and lateral lines .[ 6] [ 7] [ 8] [ 9] [ 10] : 600 [ 10] : 552 [ 11] [ 10] : 694
The presence of an amniotic buffer, of a water-impermeable skin , and of a robust, air-breathing, respiratory system , allow amniotes to live on land as true terrestrial animals . Amniotes have the ability to procreate without water bodies . Because the amnion and the fluid it secretes shields the embryo from environmental fluctuations, amniotes can reproduce on dry land by either laying shelled eggs (reptiles, birds and monotremes ) or nurturing fertilized eggs within the mother (marsupial and placental mammals ). This distinguishes amniotes from anamniotes (fish and amphibians) that have to spawn in aquatic environments . Most amniotes still require regular access to drinking water for rehydration, like the semiaquatic amphibians do.
They have better homeostasis in drier environments, and more efficient non-aquatic gas exchange to power terrestrial locomotion , which is facilitated by their astragalus.
Basal amniotes resembled small lizards and evolved from semiaquatic reptiliomorphs during the Carboniferous period.[ 12] After the Carboniferous rainforest collapse , amniotes spread around Earth's land and became the dominant land vertebrates.[ 12]
They almost immediately diverged into two groups, namely the sauropsids (including all reptiles and birds ) and synapsids (including mammals and extinct ancestors like "pelycosaurs " and therapsids ). Among the earliest known crown group amniotes, the oldest known sauropsid is Hylonomus and the oldest known synapsid is Asaphestera , both of which are from Nova Scotia during the Bashkirian age of the Late Carboniferous around 318 million years ago .[ 1] [ 13]
This basal divergence within Amniota has also been dated by molecular studies at 310–329 Ma,[ 14] or 312–330 Ma,[ 15] and by a fossilized birth–death process study at 322–340 Ma.[ 16]
^ a b 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 .
^ Paton, R. L.; Smithson, T. R.; Clack, J. A. (8 April 1999). "An amniote-like skeleton from the Early Carboniferous of Scotland". Nature . 398 (6727): 508–513. Bibcode :1999Natur.398..508P . doi :10.1038/19071 . ISSN 0028-0836 . S2CID 204992355 .
^ Irmis, R. B.; Parker, W. G. (2005). "Unusual tetrapod teeth from the Upper Triassic Chinle Formation, Arizona, USA". Canadian Journal of Earth Sciences . 42 (7): 1339–1345. Bibcode :2005CaJES..42.1339I . doi :10.1139/e05-031 . S2CID 46418796 .
^ Laurin, Michel; Reisz, Robert R. "Amniota" . RegNum . Retrieved 19 July 2024 .
^ Queiroz, Kevin de; Cantino, Philip D.; Gauthier, Jacques A. (30 April 2020). Phylonyms: A Companion to the PhyloCode (1 ed.). CRC Press. doi :10.1201/9780429446276 . ISBN 978-0-429-44627-6 .
^ Benton, Michael J. (1997). Vertebrate Palaeontology . London: Chapman & Hall. pp. 105–109. ISBN 978-0-412-73810-4 .
^ Cieri, R.L., Hatch, S.T., Capano, J.G. et al. (2020). Locomotor rib kinematics in two species of lizards and a new hypothesis for the evolution of aspiration breathing in amniotes. Sci Rep 10 . 7739. https://doi.org/10.1038/s41598-020-64140-y
^ Janis, C. M., Napoli, J. G., & Warren, D. E. (2020). Palaeophysiology of pH regulation in tetrapods. Philosophical Transactions of the Royal Society B: Biological Sciences , 375 (1793), 20190131. https://doi.org/10.1098/rstb.2019.0131
^ Hickman, Cleveland P. Jr (17 October 2016). Integrated principles of zoology (Seventeenth ed.). McGraw-Hill . pp. 563–567. ISBN 978-1-259-56231-0 .
^ a b c Kardong, Kenneth V. (16 February 2011). Vertebrates: Comparative Anatomy, Function, Evolution . McGraw-Hill. ISBN 978-0-07-352423-8 .
^ Clack, Jennifer A. (27 August 2023). Gaining Ground: The Origin and Evolution of Tetrapods . Indiana University Press. p. 370. ISBN 978-0-253-35675-8 .
^ a b Benton, M.J.; Donoghue, P.C.J. (2006). "Palaeontological evidence to date the tree of life" . Molecular Biology and Evolution . 24 (1): 26–53. doi :10.1093/molbev/msl150 . PMID 17047029 .
^ Mann, Arjan; Gee, Bryan M.; Pardo, Jason D.; Marjanović, David; Adams, Gabrielle R.; Calthorpe, Ami S.; Maddin, Hillary C.; Anderson, Jason S. (5 May 2020). Sansom, Robert (ed.). "Reassessment of historic 'microsaurs' from Joggins, Nova Scotia, reveals hidden diversity in the earliest amniote ecosystem". Papers in Palaeontology . 6 (4). Wiley: 605–625. Bibcode :2020PPal....6..605M . doi :10.1002/spp2.1316 . ISSN 2056-2802 .
^ Delsuc, Frédéric; Philippe, Hervé; Tsagkogeorga, Georgia; Simion, Paul; Tilak, Marie-Ka; Turon, Xavier; López-Legentil, Susanna; Piette, Jacques; Lemaire, Patrick; Douzery, Emmanuel J. P. (13 April 2018). "A phylogenomic framework and timescale for comparative studies of tunicates" . BMC Biology . 16 (1): 39. doi :10.1186/s12915-018-0499-2 . ISSN 1741-7007 . PMC 5899321 . PMID 29653534 .
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^ Didier, Gilles; Laurin, Michel (1 November 2020). "Exact Distribution of Divergence Times from Fossil Ages and Tree Topologies" . Systematic Biology . 69 (6): 1068–1087. doi :10.1093/sysbio/syaa021 . PMID 32191326 .