Origin of Tetrapods - indiana.edu

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Department of Geological Sciences | Indiana University (c) 2011, P. David Polly G404 Geobiology Eurypos , early Permian temnospondyl (painting by Douglas Henderson, 1990) Reading: Benton Chapters 4 and 5 and life in the Carboniferous and Permian Origin of Tetrapods

Transcript of Origin of Tetrapods - indiana.edu

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

G404 Geobiology

Eurypos , early Permian temnospondyl (painting by Douglas Henderson, 1990)

Reading: Benton Chapters 4 and 5

and life in the Carboniferous and Permian

Origin of Tetrapods

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

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Osteichthyes

Tetrapoda

Sarcopterygia

Acti

no

pte

rygia

Co

elo

ca

nth

s

Dip

no

an

s (

lun

gfi

sh

)

Osteolepis

Eusthenopteron

Pandericthyes

Icthyostega

Acanthostega

De

rive

d t

etr

ap

od

s

After Coates and Ruta, 2007. Fins into Limbs.

Phylogeny of Bony FishCrown coelocanths

and dipnoansStem

TetrapodaCrown

Tetrapoda

Tetrapoda: vertebrates more closely related to living amphibians and amniotes than to their nearest living relatives

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

G404 Geobiology

Osteichthyes

Tetrapoda

Sarcopterygia

Acti

no

pte

rygia

Co

elo

ca

nth

s

Dip

no

an

s (

lun

gfi

sh

)

Osteolepis

Eusthenopteron

Pandericthyes

Icthyostega

Acanthostega

De

rive

d t

etr

ap

od

s

Synapomorphies

• muscular pectoral and pelvic limbs with substantial limb bones• one or more squamosal bones• one or more splenial bones

• flattened head• enlarged ribs• humerus with anterior keel for muscle attachment

• carpus and tarsus• up to eight digits• broad contact between jugal and quadratojugal

• reduction to five digits

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Coates, M. I., M. Ruta, and M. Friedman. 2008. Ever since Owen: changing perspectives on the early evolutio of tetrapods. Annual Review of Ecology, Evolution, and Systematics, 39: 571-592.

Transformation of thelimb during origin of TetrapodaTransformation of early tetrapod limb involves modification of bones, loss of some bones, addition of other bones

Formation and growth of these bones is regulated by gene expression during development. Changes in the regulation result in evolutionary changes in the adult.

In earliest development, the precursors of the bones are similar in tetrapods and their closest relatives.

Developmental biology (ontogeny) is an important aid to paleontologists for identifying or confirming homologies in radically transformed groups

Sauripterusfinned tetrapod

Baramedafinned tetrapod

Tiktaalikfinned tetrapod

Eusthenopteronfinned tetrapod

Gogonasusfinned tetrapod

Sterropterygionfinned tetrapod

Rhizodopsisfinned tetrapod

Acanthostegalimbed tetrapod

Tulerpetonlimbed tetrapod

Greererpetonlimbed tetrapod

Westlothianalimbed tetrapod

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

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Coates, M. I., M. Ruta, and M. Friedman. 2008. Ever since Owen: changing perspectives on the early evolutio of tetrapods. Annual Review of Ecology, Evolution, and Systematics, 39: 571-592.

Skeletal transformations in early tetrapodsLobe-fin bones include humerus, radius and ulna. These are transformed into weight bearing limbs.

Hyomandibula attaches hyoid arch (the support skeleton between jaws and gill arches) to neurocranium. Transformed into massive stapes at otic region of braincase.

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

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Challenges for a fish out of waterRespiration. Fine structure of gills collapse in air, reducing surface area and inhibiting gas exchange. Solution: cutaneous respiration, lung respiration.

Support against gravity. Original vertebrate skeleton not really evolved for supporting the body off the ground. Solution: derived limbs and vertebral column.

Sensory perception. Ever see a fish with ears? Solution: transformation of hyomandibula to stapes, reorganization of skull for lateral sight, improvements to olfactory system.

Reproduction 1. Fish typically spawn. Solution: internal fertilization.

Reproduction 2. Egg desiccation. Solution: amniotic membrane surrounding embryo in egg to prevent desiccation.

Communication. Ever hear a fish scream? Solution: vocalizations in conjunction with hearing, new olfactory chemical signals, new visual signals.

Food. Fish are typically predatory and have prey capture strategiesthat often involve sucking prey into mouth with water. Solution: reorganization of jaws, neck, new dietary types.

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

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Phylogeny of basal tetrapods

Coates, M. I., M. Ruta, and M. Friedman. 2008. Ever since Owen: changing perspectives on the early evolutio of tetrapods. Annual Review of Ecology, Evolution, and Systematics, 39: 571-592.

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

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Laurin, 2010. How Vertebrates Left the Water.

Structure of the tetrapod middle earInner ear housed in prootic and opisthotic bones of braincase

Hyomandibula transformed into stapes, which connects opening in wall of braincase to tympanum in skin

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

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Early development of the centrum in the chick (from Patten, 1958. Foundations of Embryology.)

Early vertebral development

Somites

Vertebrae are modeled around notochord from somite tissue

Each vertebra develops between somites, receiving tissue from one in front and one behind

Directly related to evolutionary transformations in vertebrae of early tetrapods

9 day mouse embryo, before skeletal development begins

Neural tube

Heart

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(tree from Coates, et al. 2008. Annual Review of Ecology, Evolution, and Systematics, 39: 571-592)

Vertebral evolution in early tetrapodsLissamphibians retain intercentrum, amniotes retain pleurocentrum

Neural arch

Intercentrum

Pleurocentrum

Mastodontosaurus

Eryops

Icthyostega

Seymouria

Amniote

Anterior

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

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Stayton, C.R. and M. Ruta. 2006. Palaeontology, 49: 307-337.

TemnospondylsClose relatives to crown-group Amphibians

Long-lived group, Early Carboniferous (ca. 330 mya) to Early Cretaceous (ca. 120 mya)

Origin of crown-group amphibia associated with stereospondyls

Possess stereospondylous form of vertebrae, with the body composed of the intercentrum like in living amphibians

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Amniota and skull fenestrae Names of fenestrae:Temporal fenestraSupratemporal fenestraInfratemporal fenestraAntorbital fenestraMandibular fenestration

Diapsida

Amniota

Archosauria

Dinosauria

Synapsida

Lepidosauria

Saurischian

Aves

Ornithischian

Living Mammals

Living Lizards and Snakes Living Crocodiles

and Alligators

Bird-hipped dinosaurs

Lizard-hipped dinosaurs Living Birds

Reptilia

Crurotarsi

Turtles

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

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from Carroll, 2009. The Rise of Amphibians

Non-amniote tetrapod skull: Proterogyrinus

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(from Romer, 1966, Vertebrate Paleontology)

Amniote Skull: Early Anapsid Reptile

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Protogyrinus

DiadectesCaptorhinus

Youngina

Kuhneosaurus

EdmontosaurusArchaeopteryx

Titanophoneus Dimetrodon

Diapsida

Amniota

Archosauria

Dinosauria

Synapsida

Lepidosauria

Aves

Cladogram of selected amniotesand outgroups

Outgroups to Amniota

1. Supratemporal fenestra2. Infratemporal fenestra

1. Single temporal fenestra not bounded by quadratojugal

Synapomorphies of Diapsida

Node

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Protogyrinus

DiadectesCaptorhinus

Youngina

Kuhneosaurus

EdmontosaurusArchaeopteryx

Titanophoneus Dimetrodon

Amniota

AmniotaReduction or loss of the posterior bones of the cranium associated with origin of the neck: tabular, postparietal, supratemporal, infratemporal.

Outgroup state: large tabular, postparietal, supratemporal, infratemporal

Ingroup state: small or absent tabular, postparietal, supratemporal, infratemporal, often positioned on posterior of cranium

Non-amniotes Amniotes

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

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SynapsidaOpening of the synapsid fenestra between the jugal, squamosal and post-orbital bones, not bounded by quadratojugal. (Note is in a different place than in diapsids).

Surangular, articular, and quadratojugal reduced in size.

Outgroup state: No opening exclusively between squamosal, jugal, and postorbital. Surangular and quadratojugal major parts of skull.

Ingroup state: Opening between postorbital, squamosal and jugal. Quadratojugal and surangular reduced and moved posteriorly.

SynapsidsNon-synapsids

Protogyrinus

DiadectesCaptorhinus

Youngina

Kuhneosaurus

EdmontosaurusArchaeopteryx

Titanophoneus Dimetrodon

Synapsida

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

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(from Romer, 1966, Vertebrate Paleontology)

Synapsid Amniote Single temporal fenestra bounded by postorbital, jugal, and squamosal

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

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(from Romer, 1966, Vertebrate Paleontology)

Synapsid Amniote

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Romer, 1966, Vertebrate Paleontology

Synapsids - dominant land vertebrates of the Permian

Gauthier, Kluge, and Rowe, 1988. Amniote phylogeny.

Edaphosaurus

“Pelycosaurs”

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Romer, 1966, Vertbrate Paleontology.

Derived synapsids

Gauthier, Kluge, and Rowe, 1988. Amniote phylogeny.

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

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Hopson, in Wake et al., Hyman’s Comparative Anatomy.

Transformations of the middle ear in synapsidsQuadrate and articular become smaller as muscular focus shifts anteriorly to dentary

Quadrate and articular join the stapes (formerly hyomandibula) as small bones in the middle ear

Angular bone holds tympanum, becomes tympanic annulus

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DiapsidaOpening of temporal fenestra between jugal, quadratojugal, and squamosal (postorbital sometimes involved).

Opening of second fenestra between parietal, postfrontal, and squamosal (postorbital sometimes involved).

Outgroup state: No fenestrae in diapsid positions.

Ingroup state: two temporal fenestrae, one between jugal, quadratojugal and squamosal, one between parietal, postfrontal and squamosal.

Non-diapsids

Diapsids

Protogyrinus

DiadectesCaptorhinus

Youngina

Kuhneosaurus

EdmontosaurusArchaeopteryx

Titanophoneus Dimetrodon

Diapsida

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

G404 Geobiology

(from Romer, 1966, Vertebrate Paleontology)

Early Diapsid Amniote

Double temporal fenestra: top bounded by parietal, postfrontal, postorbital, and squamosal; bottom bounded by postorbital, jugal, quadratojugal, and squamosal (postorbital sometimes inserts between, as in this skull)

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(from Romer, 1966, Vertebrate Paleontology)

Lepidosaur Diapsid

Loss of quadratojugal opens lower temporal fenestra on inferior side (allows for more mobility in lower jaw)

Department of Geological Sciences | Indiana University (c) 2011, P. David Polly

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(from Romer, 1966, Vertebrate Paleontology)

Ornithischian Dinosaurs

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(from Romer, 1966, Vertebrate Paleontology)

Avian Theropod Dinosaur

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Pangeathe Late Permian

(260 mya) Single continentMassive global extinction

Reconstruction by Ron Blakeyhttp://jan.ucc.nau.edu/~rcb7/index.html

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60% 57% 82% 53%

47%

Permo-Triassic extinctionca. 251 million years ago

Marine diversity

Tetrapod diversity

83%

Bernard et al, 2010. Acta Palaeontologica Polonica, 55: 229-239

Nearly 95% of the Earth’s species became extinct.

Eruption of Siberian traps peaked 251 mya, covering at least 1.6 million square km, an area the size of Europe, with 400 to 3000 m of flood basalt, lasting 600,000 years.

Oxygen isotope data suggest rapid global rise in temperature of 6C, which combined with Pangea continent configuration, reduces ocean circulation and dissolved oxygen to create anoxic conditions on the floor.

Carbon isotope excursions indicate that CO2 increased in atmosphere through production by the Siberian Traps, which raised global temperature enough to melt gas hydrate deposits, which further increased atmospheric CO2 and temperature... “runaway greenhouse effect”.

Tetrapods hard hit, with the dicyondont Lystrosaurus being one of the few found in fossil record for millions of years after extinction. Forest communities absent until Middle Triassic.

Siberian traps - basalt formations left by surficial lava flow

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Scientific papers for further readingBenton, M.J. and R.J. Twitchett. 2003. How to kill (almost) all life: the end-Permian extinction event. Trends in Ecology and Evolution, 18: 358-365.

Coates, M. I., M. Ruta, and M. Friedman. 2008. Ever since Owen: changing perspectives on the early evolution of tetrapods. Annual Review of Ecology, Evolution, and Systematics, 39: 571-592.

Smith, R.M.H. and P.D. Ward. 2001. Pattern of vertebrate extinctions across an event bed at the Permian-Triassic boundary in the Karoo Basin of South Africa. Geology, 29: 1147-1150.

Stayton, C.R. and M. Ruta. 2006. Geometric morphometrics of the skull roof of stereospondyls (Amphibia: Temnospondyli). Palaeontology, 49: 307-337.