8/19/2013 Topic 4: The Origin of Tetrapods Next two lectures will deal with: Origin of Tetrapods, transition from water to land. Origin of Amniotes, transition to dry habitats. The geological time scale Organizes the history of the earth Based on geological and biological/fossil criteria Allows us to consider “________________” Time scale over which geological and ___________________ phenomena occur Organized into hierarchical ________, ________, ______________, and ______________ The Paleozoic Era Period Permian 245-286 286-320 Devonian 360-408 Silurian 408-438 Ordovician Cambrian 438-505 505-570 What is the geological time scale and why is it important? Where do herps fit in the vertebrate phylogeny? What are the evolutionary origins of tetrapods? What changes were involved in the transition from water to land? The geological time scale Eon Era MYA Cenozoic 0-65 Mesozoic 65-245 Paleozoic 245-570 Proterozoic 570-2500 Phanerozoic Precambrian Precambrian accounts for ~88% of Earth history We will concentrate on Phanerozoic 1st multicellular organisms Archaen 2500-3800 1st unicellular organisms Hadean 3800-4600 Age of oldest rocks The Mesozoic Era MYA Carboniferous Topic 4: The Origin of Tetrapods Events Period MYA Events 65-144 - Mass extinction at end of Cretaceous - 1st modern Squamata 1st _________________ Cretaceous 1st reptiles, amphibians specialize - 1st Urodela, Anura - 1st Rynchocephalia - High reptile diversity - 1st birds 1st amphibians 1st jawed fishes 1st Jurassic 144-208 Triassic 208-245 jawless fishes & land plants 1st vertebrates -1st - 1st Angiosperms, dinosaurs, mammals Testudines, ___________ 1 8/19/2013 The Cenozoic Era Period What are stem and crown groups? Epoch MYA Holocene Recent Quanternary Pleistocene 0.01-1.5 Tertiary Pliocene 1.5-5 Miocene 5-24 Oligocene 24-37 Eocene 37-58 Paleocene 58-65 Comments Crown groups: Are monophyletic Contain all extant taxa Can contain extinct taxa _________________ – contains crown group plus closely related extinct taxa. Last common ancestor is older than that for its extant members 1st hominines -1st Gymnophiona - 1st pacental mammals Stem groups: _________________ – smallest monophyletic group to contain the last common ancestor of all extant members of a taxon Evolution of humans What are stem and crown groups? Modern humans Where do herps fit into the vertebrate phylogeny? Are also monophyletic Contain all extant & extinct taxa Phylogeny of jawed vertebrates What are outgroups of Tetrapoda (3)? Actinopterygii (6) Actinistia (7) Dipnoi (8) ____________________ (6) are ray-finned fishes Extremely diverse (~25,000 spp.) Highly derived Fins have extrinsic muscle Pough et al 2004, Fig 2-1 Benton Fig 9.11 Where do herps fit into the vertebrate phylogeny? Sarcopterygian lineages ________________ (2) Lobe-finned fishes Contain Actinistia (7), Dipnoi (8), Tetrapoda (3) _________________ Fins/limbs are supported by bone and contain ___________ muscle Teeth have enamel Pough et al 2004, Fig 2-1 Benton Fig 9.11 _________________ What kind of characters are these? Coelacanth, Latimeria 2 extant species Symmetrical, 3-lobed ________ “living fossil” Lung fishes 3 extant genera Gondwanan distribution 1 in S. America, 1 in Africa, 1 in Australia Vertebrate Life Fig 8-35 2 8/19/2013 Sarcopterygian lineages Tetrapoda Includes all modern amphibians and amniotes, and their last common ancestor. _________________ _________________ Fully developed _____________ girdle Pectoral girdle free from the skull (not the case in non-tetrapods) Discrete shaft of ______________ Where do herps fit into the vertebrate phylogeny? This phylogeny includes only extant taxa Many species of early tetrapods and stegocephalians are extinct Therefore, must look at both stem and crown Amphibia Pough et al 2004, Fig 2-1 Stem and Crown Amphibia Most groups of Amphibians and all early tetrapods are ____________ Benton 1997, Fig 4-21 3 8/19/2013 Early tetrapod relationships are contentious How do these two hypotheses differ? How are they the same? Why might we prefer one over the other? Stegocephali Tetrapoda Lepospondyls Temnospondyls Laurin & Reisz 1997 Traditional hypothesis Inclusion of only some taxa, assumption of Temnospondyl-Lissamphibian association No a priori assumptions, many more taxa included Pough et al. 2004 Fig 2-8 1 8/19/2013 What are the evolutionary origins of tetrapods? What are the evolutionary origins of tetrapods? Elpistostegidae (Panderichthys) Laurin & Reisz 1997 Lepospondyls are: __________________ Sister to Lissamphibia Amniotes (including reptiles) are: Sister group to Lissamphibia+Lepospondyls Outgroup to Stegocephalia but closer than Dipnoi Dorsoventrally _____________, no dorsal fin, dorsal eyes _________________ water Tetrapod-like body but with fins Developed frontal bones a synapomorphy with stegocephalia Stegocephali Tetrapoda Lepospondyls Pough et al. 2004 Fig 2-8 Temnospondyls What are the evolutionary origins of tetrapods? Stegocephali Lepospondyls Tetrapoda Temnospondyls ___________ from Devonian (360mya) of Greenland and Russia Four limbs with digits, but also a branchial apparatus with gills ________________ Pough et al. 2004 Fig 2-2 What are the evolutionary origins of tetrapods? Acanthostega & Ichthyostega Stegocephali Lepospondyls Tetrapoda Temnospondyls Vertebrate Life Fig 10-3 What are the evolutionary origins of tetrapods? Carboniferous Few centimeters to over 1m Limbed, posture more upright than Acanthostega/Ichthyostega Some aquatic, others (below) terrestrial Typically ________________ Pough et al. 2004 Fig 2-7 6-8 digits per limb What does this mean for ancestral number of digits? Amniota Stegocephali Lepospondyls Tetrapoda Temnospondyls _______________ Temnospondyls Pough et al. 2004 Fig 2-3 What are the evolutionary origins of tetrapods? Acanthostega & Ichthyostega Stegocephali Lepospondyls Tetrapoda Temnospondyls We will come back to these Mammals, birds, snakes & lizards, turtles, crocidilians Next slide Stegocephali Lepospondyls Tetrapoda Temnospondyls 1 8/19/2013 What are the evolutionary origins of tetrapods? What are the evolutionary origins of tetrapods? Lepospondyls Carboniferous & Permian Diverse Microsauria – _________ & small Aisopoda – ___________ Nectridea – _________ & newt-like Pough et al. 2004 Fig 2-4b,c; 4-14, 4-15 What changes were involved in the transition from water to land? Obligate Aquatic What is involved in this major evolutionary transition? Stegocephali Lepospondyls Tetrapoda Temnospondyls Urodela, Anura, Gymnophiona Extant (finally) Photos: KP Bergmann, Pough et al. Fig 3-8 What changes were involved in the transition from water to land? This transition involved aspects of structure and function associated with: Terrestrial Lissamphibia ____________________ ____________________ ____________________ ____________________ ____________________ ____________________ ____________________ ____________________ 2 8/19/2013 What changes were involved in the transition from water to land? What changes were involved in the transition from water to land? Body support Moving onto land means coping with _____________ In the water, the body is supported by the water, on land by limbs Coping with ______________ __________________ Ground reaction forces Actinopterygii Pectoral girdle became ____________ from the head Minimizes forces exerted on skull and brain, increases head ____________ Small in water Large on land Ichthyostega Panderichthys Benton 1997 pg 183; Benton Fig 4-2a; Pough et al 2004 Fig 2-2 What changes were involved in the transition from water to land? What changes were involved in the transition from water to land? Ground reaction forces Limb girdles became more ______________ to the vertebral column Allows limbs to become support structures and transfer forces to the axis Locomotion Protopterus (African lungfish) A switch from ____________ to walking and running Terrestrial locomotion is much harder on the ___________ Fin-based swimming is replaced by pushing off a substrate with the limbs Lateral undulation plays a role in both ________________ less of an issue on land Acanthostega Benton 1997 Fig 3.22; Pough et al 2004 Fig 2-2a What changes were involved in the transition from water to land? Locomotion Locomotion Limbs change dramatically: More ____________ More ________________ Switch from extrinsic to intrinsic limb musculature Panderichthys & Acanthostega What changes were involved in the transition from water to land? Vertebral morphology becomes more ______________ Zygapophyses limit excessive intervertebral forces, especially _____________ Acanthostega Vertebrate Life Fig 10-3 Kardong & Zalisko 2002: Fig 5.8c, 5.11b 1 8/19/2013 What changes were involved in the transition from water to land? What changes were involved in the transition from water to land? Body shape Very _________________ in fishes Streamline but dorsoventrally flattened in Panderichthys Allows invasion of shallow water Also dorsal eyes, no dorsal fin Decrease in hydrodynamics in terrestrial forms What changes were involved in the transition from water to land? Trend from excreting ammonia to __________ to ___________ Ammonia is most toxic but also most water soluble Both gills and lungs are ancestral because lungs arose from ____________ Acanthostega still had gills Ventilation increasingly powered by __________ of the ______________ Water balance Lissamphibia live in moist environments, but more water_________________ Ammonia is voided quickly through ___________ Mucous glands limit evaporation from an amphibian’s body Urea and Uric Acid are concentrated and excreted by _______________ Lissamphibia have various behaviors and postures that limit water loss Sound travels differently in air than in water Other organisms are similar density to water In air there is a differential between densities Hyomandibula bone of the 1st gill arch gives rise to the _______________, an inner ear bone that connects to the ____________________ This allows for better conduction of vibrations to the inner ear Kardong & Zalisko Fig 5.29d Fishes live in excess water Hearing Gills only to gills & lungs to lungs only What changes were involved in the transition from water to land? What changes were involved in the transition from water to land? Excretion Respiration What changes were involved in the transition from water to land? Feeding ______________ is prevalent in fishes, impossible on land On land, jaws and the tongue play a more important role Pough et al 2004, Fig 11-19 2
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