{"id":2541,"date":"2017-02-06T21:24:18","date_gmt":"2017-02-06T21:24:18","guid":{"rendered":"https:\/\/courses.lumenlearning.com\/wm-biology2\/?post_type=chapter&#038;p=2541"},"modified":"2017-07-05T17:14:07","modified_gmt":"2017-07-05T17:14:07","slug":"amniotes","status":"publish","type":"chapter","link":"https:\/\/courses.lumenlearning.com\/suny-wmopen-biology2\/chapter\/amniotes\/","title":{"raw":"Amniotes","rendered":"Amniotes"},"content":{"raw":"<h2>Identify characteristics of amniotes<\/h2>\r\nAmniotes are a clade of tetrapod vertebrates comprising the reptiles, birds, and mammals.\u00a0Amniotes are characterized by having an egg equipped with an amnion, an adaptation to lay eggs on land or retain the fertilized egg within the mother.\r\n\r\nAmniote embryos, whether laid as eggs or carried by the female, are protected and aided by several extensive membranes. In eutherian mammals (such as humans), these membranes include the amniotic sac that surrounds the fetus. These embryonic membranes and the lack of a larval stage distinguish amniotes from tetrapod amphibians.\r\n<div class=\"textbox learning-objectives\">\r\n<h3>Learning Objectives<\/h3>\r\n<ul>\r\n \t<li>Identify the classes of animals that are amniotes<\/li>\r\n \t<li>Describe the main characteristics of amniotes<\/li>\r\n \t<li>Discuss the evolution of amniotes<\/li>\r\n<\/ul>\r\n<\/div>\r\n<h2>Amniotic Animals<\/h2>\r\nIn the past, the most common division of amniotes has been into the classes Mammalia, Reptilia, and Aves. Birds are directly descended, however, from dinosaurs, so this classical scheme results in groups that are not true clades. We will consider birds as a group distinct from reptiles for the purpose of this discussion with the understanding that this does not completely reflect phylogenetic history and relationships. Instead, modern phylogenetics places birds and reptiles into a larger clade together, though birds (not reptiles) are the true descendants of dinosaurs.\r\n\r\nThe amniotes\u2014reptiles, birds, and mammals\u2014are distinguished from amphibians by their terrestrially adapted egg, which is protected by amniotic membranes. The evolution of amniotic membranes meant that the embryos of amniotes were provided with their own aquatic environment, which led to less dependence on water for development and thus allowed the amniotes to branch out into drier environments. This was a significant development that distinguished them from amphibians, which were restricted to moist environments due their shell-less eggs. Although the shells of various amniotic species vary significantly, they all allow retention of water. The shells of bird eggs are composed of calcium carbonate and are hard, but fragile. The shells of reptile eggs are leathery and require a moist environment. Most mammals do not lay eggs (except for monotremes). Instead, the embryo grows within the mother\u2019s body; however, even with this internal gestation, amniotic membranes are still present.\r\n<h2>Characteristics of Amniotes<\/h2>\r\n[caption id=\"attachment_2547\" align=\"alignright\" width=\"400\"]<img class=\" wp-image-2547\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/1223\/2017\/02\/06212753\/Figure_29_04_01.png\" alt=\"The illustration shows an egg with the shell, embryo, yolk, yolk sac, and the extra-embryonic membranes\" width=\"400\" height=\"331\" \/> Figure\u00a01.\u00a0The key features of an amniotic egg are shown.[\/caption]\r\n\r\nThe amniotic egg is the key characteristic of amniotes. In amniotes that lay eggs, the shell of the egg provides protection for the developing embryo while being permeable enough to allow for the exchange of carbon dioxide and oxygen. The albumin, or egg white, provides the embryo with water and protein, whereas the fattier egg yolk is the energy supply for the embryo, as is the case with the eggs of many other animals, such as amphibians. However, the eggs of amniotes contain three additional extra-embryonic membranes: the chorion, amnion, and allantois (Figure\u00a01).\r\n\r\nExtra-embryonic membranes are membranes present in amniotic eggs that are not a part of the body of the developing embryo. While the inner amniotic membrane surrounds the embryo itself, the <b>chorion\u00a0<\/b>surrounds the embryo and yolk sac. The chorion facilitates exchange of oxygen and carbon dioxide between the embryo and the egg\u2019s external environment. The <b>amnion<\/b> protects the embryo from mechanical shock and supports hydration. The <b>allantois<\/b> stores nitrogenous wastes produced by the embryo and also facilitates respiration. In mammals, membranes that are homologous to the extra-embryonic membranes in eggs are present in the placenta.\r\n\r\nAdditional derived characteristics of amniotes include waterproof skin, due to the presence of lipids, and costal (rib) ventilation of the lungs.\r\n<div class=\"textbox exercises\">\r\n<h3>Practice Question<\/h3>\r\nWhich of the following statements about the parts of an egg are false?\r\n<ol style=\"list-style-type: lower-alpha;\">\r\n \t<li>The allantois stores nitrogenous waste and facilitates respiration.<\/li>\r\n \t<li>The chorion facilitates gas exchange.<\/li>\r\n \t<li>The yolk provides food for the growing embryo.<\/li>\r\n \t<li>The amniotic cavity is filled with albumen.<\/li>\r\n<\/ol>\r\n[reveal-answer q=\"428004\"]Show Answer[\/reveal-answer]\r\n[hidden-answer a=\"428004\"]Statement d is false.[\/hidden-answer]\r\n\r\n<\/div>\r\n<h2>Evolution of Amniotes<\/h2>\r\nThe first amniotes evolved from amphibian ancestors approximately 340 million years ago during the Carboniferous period. The early amniotes diverged into two main lines soon after the first amniotes arose. The initial split was into synapsids and sauropsids.\u00a0<b>Synapsids<\/b> include all mammals, including extinct mammalian species. Synapsids also include therapsids, which were mammal-like reptiles from which mammals evolved. <b>Sauropsids<\/b> include reptiles and birds, and can be further divided into anapsids and diapsids. The key differences between the synapsids, anapsids, and diapsids are the structures of the skull and the number of temporal fenestrae behind each eye (Figure\u00a02).\r\n\r\n[caption id=\"attachment_2549\" align=\"aligncenter\" width=\"1024\"]<img class=\"size-large wp-image-2549\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/1223\/2017\/02\/06212922\/Figure_29_04_02-1024x217.jpg\" alt=\"The illustration compares three different skull types. All three skulls are elongated and similar in shape; the only difference between them is the number of holes behind the eye. The anapsid skull (left) has no openings. The synapsid skull (middle) has one opening, and the diapsid skull (right) has two openings, one on top of the other.\" width=\"1024\" height=\"217\" \/> Figure\u00a02.\u00a0Compare the skulls and temporal fenestrae of anapsids, synapsids, and diapsids. Anapsids have no openings, synapsids have one opening, and diapsids have two openings.[\/caption]\r\n\r\n<b>Temporal fenestrae<\/b> are post-orbital openings in the skull that allow muscles to expand and lengthen. <b>Anapsids<\/b> have no temporal fenestrae, synapsids have one, and <b>diapsids<\/b> have two. Anapsids include extinct organisms and may, based on anatomy, include turtles. However, this is still controversial, and turtles are sometimes classified as diapsids based on molecular evidence. The diapsids include birds and all other living and extinct reptiles.\r\n\r\nThe diapsids diverged into two groups, the Archosauromorpha (\u201cancient lizard form\u201d) and the Lepidosauromorpha (\u201cscaly lizard form\u201d) during the Mesozoic period (Figure\u00a03). The <b>lepidosaurs<\/b> include modern lizards, snakes, and tuataras. The <b>archosaurs<\/b> include modern crocodiles and alligators, and the extinct pterosaurs (\u201cwinged lizard\u201d) and dinosaurs (\u201cterrible lizard\u201d). Clade Dinosauria includes birds, which evolved from a branch of dinosaurs.\r\n\r\n[caption id=\"attachment_2550\" align=\"aligncenter\" width=\"1024\"]<img class=\"size-large wp-image-2550\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/1223\/2017\/02\/06212953\/Figure_29_04_03-1024x636.png\" alt=\"The trunk of the amniote phylogenetic tree is the ancestral amniote. Initially, the tree branches into diapsids, anapsids, and synapsids. Synapsids give rise to mammals, which are therapsids. Anapsids are all extinct. Diapsids are subdivided into two groups, lepidosaurs and archosaurs. Lepidosauria includes plesiosaurs, ichthyosaurs, Sphenodontia and Squamata, which includes lizards and snakes. Archosauria branches into Crocodilia, pterosaurs, dinosaurs, and birds.\" width=\"1024\" height=\"636\" \/> Figure\u00a03.\u00a0This chart shows the evolution of amniotes. The placement of Testudines (turtles) is currently still debated.[\/caption]\r\n\r\n<div class=\"textbox exercises\">\r\n<h3>Practice Question<\/h3>\r\nMembers of the order Testudines have an anapsid-like skull with one opening. However, molecular studies indicate that turtles descended from a diapsid ancestor. Why might this be the case?\r\n\r\n[practice-area rows=\"2\"][\/practice-area]\r\n[reveal-answer q=\"540837\"]Show Answer[\/reveal-answer]\r\n[hidden-answer a=\"540837\"]The ancestor of modern Testudines may at one time have had a second opening in the skull, but over time this might have been lost.[\/hidden-answer]\r\n\r\n<\/div>\r\n<h2><strong>Check Your Understanding<\/strong><\/h2>\r\nAnswer the question(s) below to see how well you understand the topics covered in the previous section. This short quiz does\u00a0<strong>not<\/strong>\u00a0count toward your grade in the class, and you can retake it an unlimited number of times.\r\n\r\nUse this quiz to check your understanding and decide whether to (1) study the previous section further or (2) move on to the next section.\r\n\r\nhttps:\/\/assessments.lumenlearning.com\/assessments\/4986","rendered":"<h2>Identify characteristics of amniotes<\/h2>\n<p>Amniotes are a clade of tetrapod vertebrates comprising the reptiles, birds, and mammals.\u00a0Amniotes are characterized by having an egg equipped with an amnion, an adaptation to lay eggs on land or retain the fertilized egg within the mother.<\/p>\n<p>Amniote embryos, whether laid as eggs or carried by the female, are protected and aided by several extensive membranes. In eutherian mammals (such as humans), these membranes include the amniotic sac that surrounds the fetus. These embryonic membranes and the lack of a larval stage distinguish amniotes from tetrapod amphibians.<\/p>\n<div class=\"textbox learning-objectives\">\n<h3>Learning Objectives<\/h3>\n<ul>\n<li>Identify the classes of animals that are amniotes<\/li>\n<li>Describe the main characteristics of amniotes<\/li>\n<li>Discuss the evolution of amniotes<\/li>\n<\/ul>\n<\/div>\n<h2>Amniotic Animals<\/h2>\n<p>In the past, the most common division of amniotes has been into the classes Mammalia, Reptilia, and Aves. Birds are directly descended, however, from dinosaurs, so this classical scheme results in groups that are not true clades. We will consider birds as a group distinct from reptiles for the purpose of this discussion with the understanding that this does not completely reflect phylogenetic history and relationships. Instead, modern phylogenetics places birds and reptiles into a larger clade together, though birds (not reptiles) are the true descendants of dinosaurs.<\/p>\n<p>The amniotes\u2014reptiles, birds, and mammals\u2014are distinguished from amphibians by their terrestrially adapted egg, which is protected by amniotic membranes. The evolution of amniotic membranes meant that the embryos of amniotes were provided with their own aquatic environment, which led to less dependence on water for development and thus allowed the amniotes to branch out into drier environments. This was a significant development that distinguished them from amphibians, which were restricted to moist environments due their shell-less eggs. Although the shells of various amniotic species vary significantly, they all allow retention of water. The shells of bird eggs are composed of calcium carbonate and are hard, but fragile. The shells of reptile eggs are leathery and require a moist environment. Most mammals do not lay eggs (except for monotremes). Instead, the embryo grows within the mother\u2019s body; however, even with this internal gestation, amniotic membranes are still present.<\/p>\n<h2>Characteristics of Amniotes<\/h2>\n<div id=\"attachment_2547\" style=\"width: 410px\" class=\"wp-caption alignright\"><img loading=\"lazy\" decoding=\"async\" aria-describedby=\"caption-attachment-2547\" class=\"wp-image-2547\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/1223\/2017\/02\/06212753\/Figure_29_04_01.png\" alt=\"The illustration shows an egg with the shell, embryo, yolk, yolk sac, and the extra-embryonic membranes\" width=\"400\" height=\"331\" \/><\/p>\n<p id=\"caption-attachment-2547\" class=\"wp-caption-text\">Figure\u00a01.\u00a0The key features of an amniotic egg are shown.<\/p>\n<\/div>\n<p>The amniotic egg is the key characteristic of amniotes. In amniotes that lay eggs, the shell of the egg provides protection for the developing embryo while being permeable enough to allow for the exchange of carbon dioxide and oxygen. The albumin, or egg white, provides the embryo with water and protein, whereas the fattier egg yolk is the energy supply for the embryo, as is the case with the eggs of many other animals, such as amphibians. However, the eggs of amniotes contain three additional extra-embryonic membranes: the chorion, amnion, and allantois (Figure\u00a01).<\/p>\n<p>Extra-embryonic membranes are membranes present in amniotic eggs that are not a part of the body of the developing embryo. While the inner amniotic membrane surrounds the embryo itself, the <b>chorion\u00a0<\/b>surrounds the embryo and yolk sac. The chorion facilitates exchange of oxygen and carbon dioxide between the embryo and the egg\u2019s external environment. The <b>amnion<\/b> protects the embryo from mechanical shock and supports hydration. The <b>allantois<\/b> stores nitrogenous wastes produced by the embryo and also facilitates respiration. In mammals, membranes that are homologous to the extra-embryonic membranes in eggs are present in the placenta.<\/p>\n<p>Additional derived characteristics of amniotes include waterproof skin, due to the presence of lipids, and costal (rib) ventilation of the lungs.<\/p>\n<div class=\"textbox exercises\">\n<h3>Practice Question<\/h3>\n<p>Which of the following statements about the parts of an egg are false?<\/p>\n<ol style=\"list-style-type: lower-alpha;\">\n<li>The allantois stores nitrogenous waste and facilitates respiration.<\/li>\n<li>The chorion facilitates gas exchange.<\/li>\n<li>The yolk provides food for the growing embryo.<\/li>\n<li>The amniotic cavity is filled with albumen.<\/li>\n<\/ol>\n<div class=\"qa-wrapper\" style=\"display: block\"><span class=\"show-answer collapsed\" style=\"cursor: pointer\" data-target=\"q428004\">Show Answer<\/span><\/p>\n<div id=\"q428004\" class=\"hidden-answer\" style=\"display: none\">Statement d is false.<\/div>\n<\/div>\n<\/div>\n<h2>Evolution of Amniotes<\/h2>\n<p>The first amniotes evolved from amphibian ancestors approximately 340 million years ago during the Carboniferous period. The early amniotes diverged into two main lines soon after the first amniotes arose. The initial split was into synapsids and sauropsids.\u00a0<b>Synapsids<\/b> include all mammals, including extinct mammalian species. Synapsids also include therapsids, which were mammal-like reptiles from which mammals evolved. <b>Sauropsids<\/b> include reptiles and birds, and can be further divided into anapsids and diapsids. The key differences between the synapsids, anapsids, and diapsids are the structures of the skull and the number of temporal fenestrae behind each eye (Figure\u00a02).<\/p>\n<div id=\"attachment_2549\" style=\"width: 1034px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" aria-describedby=\"caption-attachment-2549\" class=\"size-large wp-image-2549\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/1223\/2017\/02\/06212922\/Figure_29_04_02-1024x217.jpg\" alt=\"The illustration compares three different skull types. All three skulls are elongated and similar in shape; the only difference between them is the number of holes behind the eye. The anapsid skull (left) has no openings. The synapsid skull (middle) has one opening, and the diapsid skull (right) has two openings, one on top of the other.\" width=\"1024\" height=\"217\" \/><\/p>\n<p id=\"caption-attachment-2549\" class=\"wp-caption-text\">Figure\u00a02.\u00a0Compare the skulls and temporal fenestrae of anapsids, synapsids, and diapsids. Anapsids have no openings, synapsids have one opening, and diapsids have two openings.<\/p>\n<\/div>\n<p><b>Temporal fenestrae<\/b> are post-orbital openings in the skull that allow muscles to expand and lengthen. <b>Anapsids<\/b> have no temporal fenestrae, synapsids have one, and <b>diapsids<\/b> have two. Anapsids include extinct organisms and may, based on anatomy, include turtles. However, this is still controversial, and turtles are sometimes classified as diapsids based on molecular evidence. The diapsids include birds and all other living and extinct reptiles.<\/p>\n<p>The diapsids diverged into two groups, the Archosauromorpha (\u201cancient lizard form\u201d) and the Lepidosauromorpha (\u201cscaly lizard form\u201d) during the Mesozoic period (Figure\u00a03). The <b>lepidosaurs<\/b> include modern lizards, snakes, and tuataras. The <b>archosaurs<\/b> include modern crocodiles and alligators, and the extinct pterosaurs (\u201cwinged lizard\u201d) and dinosaurs (\u201cterrible lizard\u201d). Clade Dinosauria includes birds, which evolved from a branch of dinosaurs.<\/p>\n<div id=\"attachment_2550\" style=\"width: 1034px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" aria-describedby=\"caption-attachment-2550\" class=\"size-large wp-image-2550\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/1223\/2017\/02\/06212953\/Figure_29_04_03-1024x636.png\" alt=\"The trunk of the amniote phylogenetic tree is the ancestral amniote. Initially, the tree branches into diapsids, anapsids, and synapsids. Synapsids give rise to mammals, which are therapsids. Anapsids are all extinct. Diapsids are subdivided into two groups, lepidosaurs and archosaurs. Lepidosauria includes plesiosaurs, ichthyosaurs, Sphenodontia and Squamata, which includes lizards and snakes. Archosauria branches into Crocodilia, pterosaurs, dinosaurs, and birds.\" width=\"1024\" height=\"636\" \/><\/p>\n<p id=\"caption-attachment-2550\" class=\"wp-caption-text\">Figure\u00a03.\u00a0This chart shows the evolution of amniotes. The placement of Testudines (turtles) is currently still debated.<\/p>\n<\/div>\n<div class=\"textbox exercises\">\n<h3>Practice Question<\/h3>\n<p>Members of the order Testudines have an anapsid-like skull with one opening. However, molecular studies indicate that turtles descended from a diapsid ancestor. Why might this be the case?<\/p>\n<p><textarea aria-label=\"Your Answer\" rows=\"2\"><\/textarea><\/p>\n<div class=\"qa-wrapper\" style=\"display: block\"><span class=\"show-answer collapsed\" style=\"cursor: pointer\" data-target=\"q540837\">Show Answer<\/span><\/p>\n<div id=\"q540837\" class=\"hidden-answer\" style=\"display: none\">The ancestor of modern Testudines may at one time have had a second opening in the skull, but over time this might have been lost.<\/div>\n<\/div>\n<\/div>\n<h2><strong>Check Your Understanding<\/strong><\/h2>\n<p>Answer the question(s) below to see how well you understand the topics covered in the previous section. This short quiz does\u00a0<strong>not<\/strong>\u00a0count toward your grade in the class, and you can retake it an unlimited number of times.<\/p>\n<p>Use this quiz to check your understanding and decide whether to (1) study the previous section further or (2) move on to the next section.<\/p>\n<p>\t<iframe id=\"lumen_assessment_4986\" class=\"resizable\" src=\"https:\/\/assessments.lumenlearning.com\/assessments\/load?assessment_id=4986&#38;embed=1&#38;external_user_id=&#38;external_context_id=&#38;iframe_resize_id=lumen_assessment_4986\" frameborder=\"0\" style=\"border:none;width:100%;height:100%;min-height:400px;\"><br \/>\n\t<\/iframe><\/p>\n\n\t\t\t <section class=\"citations-section\" role=\"contentinfo\">\n\t\t\t <h3>Candela Citations<\/h3>\n\t\t\t\t\t <div>\n\t\t\t\t\t\t <div id=\"citation-list-2541\">\n\t\t\t\t\t\t\t <div class=\"licensing\"><div class=\"license-attribution-dropdown-subheading\">CC licensed content, Original<\/div><ul class=\"citation-list\"><li>Introduction to Amniotes. <strong>Authored by<\/strong>: Shelli Carter and Lumen Learning. <strong>Provided by<\/strong>: Lumen Learning. <strong>License<\/strong>: <em><a target=\"_blank\" rel=\"license\" href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/\">CC BY: Attribution<\/a><\/em><\/li><\/ul><div class=\"license-attribution-dropdown-subheading\">CC licensed content, Shared previously<\/div><ul class=\"citation-list\"><li>Modification of Amniote. <strong>Provided by<\/strong>: Wikipedia. <strong>Located at<\/strong>: <a target=\"_blank\" href=\"https:\/\/en.wikipedia.org\/wiki\/Amniote\">https:\/\/en.wikipedia.org\/wiki\/Amniote<\/a>. <strong>License<\/strong>: <em><a target=\"_blank\" rel=\"license\" href=\"https:\/\/creativecommons.org\/licenses\/by-sa\/4.0\/\">CC BY-SA: Attribution-ShareAlike<\/a><\/em><\/li><li>Biology. <strong>Provided by<\/strong>: OpenStax. <strong>Located at<\/strong>: <a target=\"_blank\" href=\"http:\/\/cnx.org\/contents\/185cbf87-c72e-48f5-b51e-f14f21b5eabd@10.8\">http:\/\/cnx.org\/contents\/185cbf87-c72e-48f5-b51e-f14f21b5eabd@10.8<\/a>. <strong>License<\/strong>: <em><a target=\"_blank\" rel=\"license\" href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/\">CC BY: Attribution<\/a><\/em>. <strong>License Terms<\/strong>: Download for free at http:\/\/cnx.org\/contents\/185cbf87-c72e-48f5-b51e-f14f21b5eabd@10.8<\/li><\/ul><\/div>\n\t\t\t\t\t\t <\/div>\n\t\t\t\t\t <\/div>\n\t\t\t <\/section>","protected":false},"author":17,"menu_order":5,"template":"","meta":{"_candela_citation":"[{\"type\":\"original\",\"description\":\"Introduction to Amniotes\",\"author\":\"Shelli Carter and Lumen Learning\",\"organization\":\"Lumen Learning\",\"url\":\"\",\"project\":\"\",\"license\":\"cc-by\",\"license_terms\":\"\"},{\"type\":\"cc\",\"description\":\"Modification of 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