8/19/2013. Topic 4: The Origin of Tetrapods. Topic 4: The Origin of Tetrapods. The geological time scale. The geological time scale.

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1 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. Topic 4: The Origin of Tetrapods 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? The geological time scale 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 Eon Phanerozoic Precambrian Era Cenozoic Mesozoic Paleozoic Proterozoic Archaen Hadean MYA Precambrian accounts for ~88% of Earth history We will concentrate on Phanerozoic 1 st multicellular organisms 1 st unicellular organisms Age of oldest rocks The Paleozoic Era The Mesozoic Era Period Permian Carboniferous Devonian Silurian Ordovician Cambrian MYA Events 1 st 1 st reptiles, amphibians specialize 1 st amphibians 1 st jawed fishes 1 st jawless fishes & land plants 1 st vertebrates Period Cretaceous Jurassic Triassic MYA Events - Mass extinction at end of Cretaceous - 1 st modern Squamata - 1 st Urodela, Anura - 1 st Rynchocephalia - High reptile diversity - 1 st birds -1 st Angiosperms, dinosaurs, mammals - 1 st Testudines, 1

2 8/19/2013 The Cenozoic Era Period What are stem and crown groups? Epoch MYA Holocene Recent Quanternary Pleistocene Tertiary Pliocene Miocene 5-24 Oligocene Eocene Paleocene 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 (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), (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

3 Sarcopterygian lineages Where do herps fit into the vertebrate phylogeny? 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 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 Early tetrapod relationships are contentious How do these two hypotheses differ? How are they the same? Why might we prefer one over the other? Traditional hypothesis Inclusion of only some taxa, assumption of Temnospondyl-Lissamphibian association Laurin & Reisz 1997 No a priori assumptions, many more taxa included Pough et al Fig 2-8 1

5 Laurin & Reisz 1997 are: Sister to Lissamphibia Amniotes (including reptiles) are: Sister group to Lissamphibia+ Elpistostegidae (Panderichthys) Outgroup to a but closer than Dipnoi Dorsoventrally, no dorsal fin, dorsal eyes water Tetrapod-like body but with fins Developed frontal bones a synapomorphy with stegocephalia Pough et al Fig 2-8 Pough et al Fig 2-3 Acanthostega & Ichthyostega from Devonian (360mya) of Greenland and Russia Four limbs with digits, but also a branchial apparatus with gills Acanthostega & Ichthyostega 6-8 digits per limb What does this mean for ancestral number of digits? Pough et al Fig 2-2 Vertebrate Life Fig 10-3 Carboniferous Few centimeters to over 1m Limbed, posture more upright than Acanthostega/Ichthyostega Some aquatic, others (below) terrestrial Typically Amniota We will come back to these Mammals, birds, snakes & lizards, turtles, crocidilians Next slide Pough et al Fig 2-7 1

6 Carboniferous & Permian Diverse Microsauria & small Aisopoda Nectridea & newt-like Lissamphibia Urodela, Anura, Gymnophiona Extant (finally) Pough et al Fig 2-4b,c; 4-14, 4-15 Photos: KP Bergmann, Pough et al. Fig 3-8 transition from water to land? Obligate Aquatic Terrestrial What is involved in this major evolutionary transition? This transition involved aspects of structure and function associated with: 2

7 Body support Moving onto land means coping with In the water, the body is supported by the water, on land by limbs Coping with Small in water Large on land Actinopterygii Ground reaction forces Pectoral girdle became from the head Minimizes forces exerted on skull and brain, increases head Panderichthys Ichthyostega Benton 1997 pg 183; Benton Fig 4-2a; Pough et al 2004 Fig 2-2 Ground reaction forces Limb girdles became more to the vertebral column Allows limbs to become support structures and transfer forces to the axis Protopterus (African lungfish) Locomotion 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 Locomotion Limbs change dramatically: More More Switch from extrinsic to intrinsic limb musculature Locomotion Vertebral morphology becomes more Zygapophyses limit excessive intervertebral forces, especially Panderichthys & Acanthostega Acanthostega Vertebrate Life Fig 10-3 Kardong & Zalisko 2002: Fig 5.8c, 5.11b 1

8 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 Respiration Gills only to gills & lungs to lungs only Both gills and lungs are ancestral because lungs arose from Acanthostega still had gills Ventilation increasingly powered by of the Excretion Trend from excreting ammonia to to Ammonia is most toxic but also most water soluble Ammonia is voided quickly through Urea and Uric Acid are concentrated and excreted by Water balance Fishes live in excess water Lissamphibia live in moist environments, but more water- Mucous glands limit evaporation from an amphibian s body Lissamphibia have various behaviors and postures that limit water loss Hearing 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 1 st 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 Feeding is prevalent in fishes, impossible on land On land, jaws and the tongue play a more important role Kardong & Zalisko Fig 5.29d Pough et al 2004, Fig

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