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A new approach to determining the habitual neck posture of sauropods ased on the behaviour of extant animal Michael P. Taylor 1 Mathew J. Wedel 2 Darren Naish 1 aeobiology Research Group, School of Earth and Environmental Sciences, iversity of Portsmouth, Burnaby Building, Burnaby Road, Portsmouth PO1 3QL, artment of Anatomy and College of Podiatric Medicine, Western University o alth Sciences, 309 E. Second Street, Pomona, California 91766-1854, USA.

A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

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Page 1: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

A new approach to determiningthe habitual neck posture of sauropods

based on the behaviour of extant animals

Michael P. Taylor1

Mathew J. Wedel2

Darren Naish1

1Palaeobiology Research Group, School of Earth and Environmental Sciences, University of Portsmouth, Burnaby Building, Burnaby Road, Portsmouth PO1 3QL, UK

2Department of Anatomy and College of Podiatric Medicine, Western University of Health Sciences, 309 E. Second Street, Pomona, California 91766-1854, USA.

Page 2: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Historical background

The earliest sauropod reconstructions had horizontal necks

Camarasaurus (Ryder 1877)Reprinted by Osborn and Mook (1921: plate LXXXII)

Page 3: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Historical background

The earliest sauropod reconstructions had horizontal necks

Brontosaurus (Marsh 1883: plate I)

Page 4: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Historical background

The earliest sauropod reconstructions had horizontal necks

Diplodocus carnegii (Hatcher 1901: plate VIII)

Page 5: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Historical background

Later sauropod reconstructions had raised necks

Camarasaurus (Osborn and Mook 1921: plate LXXXIV)

Page 6: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Historical background

Later sauropod reconstructions had raised necks

Brachiosaurus brancai (Janensch 1950b: plate VIII)

Page 7: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Historical background

Later sauropod reconstructions had raised necks

Brachiosaurus brancai(Paul 1988:fig. 1)

Page 8: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Historical background

Later sauropod reconstructions had raised necks

?!

Brachiosaurus brancai(Paul 1988:fig. 1)

Page 9: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Historical background

Later sauropod reconstructions had raised necksLater sauropod reconstructions had raised necks

Brontosaurus (Zallinger's 1947 mural at YPM)

Page 10: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work has reverted to horizontal

Necks considered low and inflexibleNecks considered low and inflexible

Leicester City Museum Leicester City Museum Cetiosaurus oxoniensisCetiosaurus oxoniensis

Page 11: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Necks considered low and inflexible

Martin (1987: fig. 2)

Page 12: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Osteological neutral pose (ONP)

Stevens and Parrish (2005b: fig. 10.3A) – same specimen

Page 13: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Osteological neutral pose (ONP)

Stevens and Parrish (2005a: fig. 6.5B) – Diplodocus carnegii

Page 14: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

(Giraffes are different: ONP gives rise to an erect neck)

Stevens and Parrish (2005a: fig. 6.2) – Giraffe

Page 15: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Sauropod necks considered low and inflexible

Stevens and Parrish (2005a: fig. 6.14) – Diplodocus carnegii

Page 16: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Necks considered low and inflexible

Stevens and Parrish (1999: fig. 2B) – Diplodocus carnegiiDinoMorph

Page 17: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Necks considered low and inflexible

Stevens and Parrish (1999: 798, 800)

Page 18: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Necks considered low and inflexible

Stevens and Parrish (1999: 798, 800)

Page 19: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Necks considered low and inflexible

Stevens and Parrish (1999: 798, 800)

Page 20: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Necks considered low and inflexible

Claim: ONP is sub-horizontal in all sauropods

Page 21: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Necks considered low and inflexible

Claim: ONP is sub-horizontal in all sauropodsClaim: ONP is the habitual posture

Page 22: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Neck held high

Christian and Dzemski (2007) – stress patterns

Page 23: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Neck held high

Christian and Dzemski (2007) – stress patterns

Similar findings by ...

Page 24: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Neck held high

Christian and Dzemski (2007) – stress patterns

Similar findings by ...Christian and Heinrich (1998)

Page 25: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Neck held high

Christian and Dzemski (2007) – stress patterns

Similar findings by ...Christian and Heinrich (1998)

Christian (2002a)

Page 26: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Neck held high

Christian and Dzemski (2007) – stress patterns

Similar findings by ...Christian and Heinrich (1998)

Christian (2002a)Christian (2002b)

Page 27: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Neck held high

Christian and Dzemski (2007) – stress patterns

Similar findings by ...Christian and Heinrich (1998)

Christian (2002a)Christian (2002b)

Christian (2004)

Page 28: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Recent work

Neck held high

Christian and Dzemski (2007) – stress patterns

Similar findings by ...Christian and Heinrich (1998)

Christian (2002a)Christian (2002b)

Christian (2004)

One-man crusade?

One-man crusade?

Page 29: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

The state of the art

Low necks popularised by the BBC (and AMNH)

Page 30: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

The state of the art

Low necks popularised by the BBC (and AMNH)

All based on ONP

All based on ONP

Page 31: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

But

Page 32: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

ButWhat do extant animals do?

Page 33: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Habitual posture of extant animals

Vidal et al (1986) on “unrestrained awake animals”

Cat, sitting and standing (Vidal et al. 1986: fig. 3)

Page 34: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Habitual posture of extant animals

Vidal et al (1986) on “unrestrained awake animals”

But what about nice, cuddly bunny-rabbits?

Page 35: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Habitual posture of extant animals

Nice, cuddly bunny-rabbits have vertical necks, too.

Rabbit in semi-alert posture (Vidal et al. 1986: fig. 4)

Page 36: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Habitual posture of extant animals

And so do rats and guinea pigs

Rat and guinea pig (Vidal et al. 1986: fig. 5B-C)

Page 37: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Habitual posture of extant animals

And so do rats and guinea pigs

Rat and guinea pig (Vidal et al. 1986: fig. 5B-C)

Is it just mammals?

Is it just mammals?

Page 38: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Habitual posture of extant animals

What about the humble chicken?

Page 39: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Habitual posture of extant animals

Chicken necks habitually bend backwards!

(Vidal et al. 1986: fig. 7)

Page 40: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Habitual posture is not ONP

Vidal et al.'s (1986) key findings

Page 41: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Habitual posture is not ONP

Cervicodorsal joint is maximally extended

Vidal et al.'s (1986) key findings

Page 42: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Habitual posture is not ONP

Cervicodorsal joint is maximally extended

Craniocervical joint is maximally flexed

Vidal et al.'s (1986) key findings

Page 43: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Habitual posture is not ONP

Cervicodorsal joint is maximally extended

Craniocervical joint is maximally flexed

Median part of neck is mostly held rigid

Vidal et al.'s (1986) key findings

Page 44: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Habitual posture is not ONP

Cervicodorsal joint is maximally extended

Craniocervical joint is maximally flexed

Median part of neck is mostly held rigid

The neck is raised and lowered at end joints

Vidal et al.'s (1986) key findings

Page 45: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Cold-blooded/sprawling amniotes

Alligator (Claessens, unpublished photo)

Page 46: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Cold-blooded/sprawling amniotes

Unspecified lizard (Vidal et al. 1986: fig. 8A)Varanid lizard (Brainerd and Owekowicz 2006: fig. 4)

Page 47: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Cold-blooded/sprawling amniotes

Turtle (Landberg et al. 2003:fig. 8)

Page 48: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Non-amniote tetrapods

Salamander (Simons et al. 2000:fig. 4)

Page 49: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Neck posturemapped ontocladogram

Page 50: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Neck posturemapped ontocladogram

Page 51: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Neck posturemapped ontocladogram

Upright posture

Page 52: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Neck posturemapped ontocladogram

Upright posture

Homeothermy

Page 53: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Diplodocus carnegii(based on Hatcher 1901)

(Skull orientation basedon occipital condyle)

Page 54: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Dicraeosaurus deathmatch!

Janensch (1936: plate 16) vs. Wilson (2002: fig. 1)

Page 55: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Dicraeosaurus deathmatch!

Janensch (1936: plate 16) vs. Wilson (2002: fig. 1)

Page 56: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Also ... what does “neutral” actually mean?

Modified from Stevens and Parrish (2005b: fig. 6.8)Neutral pose

Page 57: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Also ... what does “neutral” actually mean?

Modified from Stevens and Parrish (2005b: fig. 6.8)Ventrally deflected by 8°

Page 58: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

Modified from Stevens and Parrish (2005b: fig. 6.8)Dorsally deflected by 8°

Also ... what does “neutral” actually mean?

Page 59: A new approach to determining the habitual neck posture of sauropods based on the behaviour of extant animals Michael P. Taylor 1 Mathew J. Wedel 2 Darren

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