8/19/2013 Topic 5: The Origin of Amniotes Where do amniotes fall out on the vertebrate phylogeny? What are some stem Amniotes? What is an Amniote? What changes were involved with the transition to dry habitats? What are the three main groups of Amniotes? How are Amniote taxa related? Where are Amniotes on the vertebrate phylogeny? (3) – _________ (4) – _________ What is the sister group of Amniota? What taxa comprise the Amniota? Pough et al 2004, Fig 2-1 What are some stem Amniotes? Palaeontological tree: Anthracinosauria, Seymouriamorpha, and Diadectomorpha as extinct outgroups to Amniotes Split between Amniota and Amphibia ~360 Mya What are some stem Amniotes? “_________________” Anthracinosauria Seymouriamorpha Diadectomorpha Diverse and terrestrial Mainly predators Benton 1997 Fig 4.18 Benton 1997 Fig 4.21 The Amniotic Egg What is an Amniote? Synapomorphies ______________ & associated membranes __________ fertilization ____________ maxillary teeth Various other skull and limb characters Contains extraembryonic membranes (4): Amnion Chorion Allantois Yolk sac Pough et al 2004, Fig 2-1 1 8/19/2013 The Amniotic Egg __________ Surrounds embryo Forms space filled with amniotic fluid ______ exchange _________ embryo The Amniotic Egg Amnion ___________ Dessication Concussion Chorion Allantois Yolk sac Allantois Yolk sac The Amniotic Egg The Amniotic Egg Amnion Chorion ____________ Sac formed from posterior part of GI tract Collects ___________ during development, growing as it fills Involved in formation of _______________ in placental mammals Amnion Chorion Allantois _____________ Filled with yolk, which is the nutrient source for the embryo Shrinks as yolk is used up Yolk sac What changes were involved with the transition to dry habitats? The Amniotic Egg Outer-most membrane Forms placenta in placental mammals What about the shell? The other membranes The shell is deposited by the _____________ Can be calcified and hard or leathery and soft Involved in: Form outside embryo Connected to embryo Amphibians began moving away from the water Many are terrestrial However: _____________ _____________ of the embryo Photo © KP Bergmann Skin is still highly _______________ to gases and water Reproduction is mainly limited to __________ Photos © KP Bergmann, PJB 2 8/19/2013 What changes were involved with the transition to dry habitats? Amniote lifecycle ______________ of water Many live in very dry habitats Facilitated mainly by adaptation of the: What changes were involved with the transition to dry habitats? Adaptations of the Egg Amnion, Chorion, Shell Protect embryo from _____________ Forces are more jarring on land Protect embryo from _____________ Less permeable to water ________ (amniotic) ________ Photo © KP Bergmann Photos © PJB What changes were involved with the transition to dry habitats? Adaptations of the skin: ________________ Three different amniote conditions Deposition of __________ in epidermis What are the three main groups of Amniotes? Evolved independently in several groups ________, feathers, hair All act to limit water loss Different patterns of temporal fenestration Temporal region of the skull is posterior to the orbit A fenestra is an opening without a structure running through it (L. – “window”) An~ none Syn~ one Di~ two Photo © PJB What are the three main groups of Amniotes? _____________ _____________ No temporal fenestrae What are the three main groups of Amniotes? Synapsida – _____________ and stem mammals Anapsida – ______________ and stem turtles Diapsida – Archosauria (birds & crocodilians), Lepidosauria (squamates and rhynchocephalians) and various stem groups One temporal fenestra Surrounded by postorbital, jugal and squamosal bones _____________ Two temporal fenestrae Lower one Upper one Homologous to synapsid Synapsida Anapsida Diapsida Surrounded by postorbital, squamosal and parietal bones Benton 1997 Fig 5.18 3 8/19/2013 What are the three main groups of Amniotes? Biodidac How are Amniote taxa related? Which condition is ancestral? Derived? What implications do the alternative placements of Testudines (A or B) have on the evolution of the anapsid condition? What kinds of data might be brought to bear on this problem? Synapsida Anapsida Diapsida Pough et al 2004, Fig 2-1 1 8/19/2013 How are Amniote taxa related? Which condition is ancestral? ___________ ________________ ________________ Derived? A: ______________ B: ______________ Synapsida Anapsida Diapsida Pough et al 2004, Fig 2-1 How are Amniote taxa related? What implications do the placements of Testudines (A or B) have on the evolution of the anapsid condition? A: B: Synapsida Anapsida Diapsida Pough et al 2004, Fig 2-1 2 8/19/2013 How are Amniote taxa related? What kinds of data might be brought to bear on this problem? _________________ _________________ _________________ Each has been used and says something slightly different… Synapsida Anapsida Diapsida Pough et al 2004, Fig 2-1 How are Amniote taxa related? Traditional hypothesis: A Mainly based on extant taxa and ____________ data Includes only some fossil taxa Assumes that all ____________ are related Synapsida Anapsida Diapsida Pough et al 2004, Fig 2-1 3 8/19/2013 How are Amniote taxa related? Traditional hypothesis: A How are Amniote taxa related? Hypothesis B: Based on extensive fossil sampling and no a priori assumptions of relationship Testudines have a ___________________ anapsid condition ______________ with “Parareptiles” Sister to Sauropterygia Basal anapsids “Parareptiles” Pareisaurs Procolophids Pough et al. Fig 2-9 Benton Fig 5.15 How are Amniote taxa related? Hypothesis B: How are Amniote taxa related? What do __________ data have to say? Limitation: no ______ _____ can be included Testudines are placed within ____________ (support for B) BUT: Testudines cluster with the ___________, not the Lepidosauria Who are the sauropterygians? Plesiosaurs Ichthyosaurs Aquatic Only ___________ temporal fenestrae B Hypothesis C? C Testudines A Testudines Modified from Pough et al. Fig 2-9 Benton Fig. 6.7 How are Amniote taxa related? How are Amniote taxa related? B So, where do Testunides fit in? ________________ Within the _______ Molecular and morphological data suggest convergence with “Parareptiles” C Testudines Who are the Amniota? _____________ _____________ _____________ Crocodilians Birds _____________ Rhynchocephalia Squamata A Snakes Lizards Amphisbaenia Testudines Modified from Pough et al. Fig 2-9 Pough et al 2004, Fig 2-1 1 8/19/2013 How are Amniote taxa related? Mammalia synapomorphies: Synapsid Atlas vertebra 3+ sacral vertebrae ___ cervical vertebrae _________ Mammary Glands Etc. How are Amniote taxa related? Mammalia Not dealt with further in this course Photos PJ Bergmann Pough et al 2004, Fig 2-1 How are Amniote taxa related? Reptilia (5) Includes the remaining amniotes to be monophyletic Without birds is ______________ Many obscure synapomorphies How are Amniote taxa related? Archosauria synapomorphies: They have _________ (even birds – on feet) Muscular Diaphragm 2 ventricles in heart ______________ No urinary bladder No vomeronasal organ Pough et al 2004, Fig 2-1 Pough et al 2004, Fig 2-1 How are Amniote taxa related? Aves (Birds) Birds are dinosaurs, & dinosaurs are Archosaurs How are Amniote taxa related? Reptilia • Aves have many synapomorphies: • No teeth • Wings • Feathers (shared with some dinos) Lots of diversity ~8000 spp. without birds Online Reptile database: http://www.reptiledatabase.org/ Testudines 300 Rhynchocephalia 2 Squamata 7,750 Crocodylia 23 Aves 8,700 TOTAL ~ 17,000 • Not covered further in this course Photos © PJB & KP Bergmann Nature 421, 335 - 340 (2003) 2