were the necks of apatosaurus and brontosaurus adapted for combat? michael p. taylor 1*, mathew j....

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Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1* , Mathew J. Wedel 2 , Darren Naish and 3 Brian Engh 4 . 1 Department of Earth Sciences, University of Bristol, UK. 2 College of Osteopathic Medicine of the Pacific and College of Podiatric Medicine, Western University of Health Sciences, Pomona, California, USA. 3 National Oceanography Centre, Southampton, University of Southampton, UK. 4 1522 Charles Avenue, Arcata, California 95521, USA. [email protected] Artwork: Bob Bakker

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Page 1: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Were the necks ofApatosaurusand Brontosaurusadapted for combat?

Michael P. Taylor1*,Mathew J. Wedel2,Darren Naish and3

Brian Engh4.

1Department of Earth Sciences, University of Bristol, UK.

2College of Osteopathic Medicine of the Pacific and College of Podiatric Medicine, Western University of Health Sciences, Pomona, California, USA.

3National Oceanography Centre, Southampton, University of Southampton, UK.

41522 Charles Avenue, Arcata, California 95521, USA.

[email protected]

Artwork: Bob Bakker

Page 2: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Apatosaurus louisae (Carnegie Museum)

Page 3: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Apatosaurus louisae (Carnegie Museum)

Page 4: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Brontosaurus excelsus in Zallinger's mural

Page 5: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Brontosaurus excelsus in Zallinger's mural

Page 6: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Brontosaurus excelsus in Zallinger's mural

Page 7: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Zallinger's Brontosaurus (1947)

Page 8: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Knight's Brontosaurus (1897)

Page 9: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

What exactly isan apatosaurine?

Tschopp et al. (2015), Figure 115.Reduced consensus tree obtainedby equal weighting. Fifteen OTUswere deleted a posteriori. Numbersat the nodes indicate the number ofchanges between the two branchesdeparting from the node (for theapomorphy count).

Page 10: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

What exactly isan apatosaurine?

Tschopp et al. (2015), Figure 115.Reduced consensus, apatosaurine section.

Page 11: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

What exactly isan apatosaurine?

Tschopp et al. (2015), Figure 115.Reduced consensus, apatosaurine section.

Page 12: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

What exactly isan apatosaurine?

Tschopp et al. (2015), Figure 115.Reduced consensus, apatosaurine section.

Page 13: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

What exactly isan apatosaurine?

Tschopp et al. (2015), Figure 115.Reduced consensus, apatosaurine section.

Page 14: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

AMNH 460 … whatever it is

Page 15: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

What exactly isan apatosaurine?

Tschopp et al. (2015), Figure 115.Reduced consensus, apatosaurine section.

Page 16: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

NSMT-PV 20375 … whatever is is

Page 17: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Diplodocus and Apatosaurus (Carnegie Museum)

Page 18: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Diplodocus and Apatosaurus (Carnegie Museum)

Page 19: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Tokyo apatosaur

Apatosaurus ajax

Brontosaurus excelsus

Kaatedocus

Diplodocus

Haplocanthosaurus

Camarasaurus

Page 20: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Tokyo apatosaur

Apatosaurus ajax

Brontosaurus excelsus

Kaatedocus

Diplodocus

Haplocanthosaurus

Camarasaurus

Page 21: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Tokyo apatosaur

Apatosaurus ajax

Brontosaurus excelsus

Kaatedocus

Diplodocus

Haplocanthosaurus

Camarasaurus

Page 22: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Tokyo apatosaur

Apatosaurus ajax

Brontosaurus excelsus

Kaatedocus

Diplodocus

Haplocanthosaurus

Camarasaurus

Page 23: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

BrontosaurusexcelsusholotypeYPM 1980

Cervical ?8

Page 24: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Umbaran starfighter

Page 25: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

BrontosaurusexcelsusholotypeYPM 1980

Cervical ?8

Cervical ribsrobust

Page 26: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

BrontosaurusexcelsusholotypeYPM 1980

Cervical ?8

Cervical ribsrobust

Diapophyses andparapophysesrobust

Page 27: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

BrontosaurusexcelsusholotypeYPM 1980

Cervical ?8

Cervical ribsrobust

Diapophyses andparapophysesrobust

Cervical ribsdisplacedventrally

Page 28: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

BrontosaurusexcelsusholotypeYPM 1980

Cervical ?8

Cervical ribsrobust

Diapophyses andparapophysesrobust

Cervical ribsdisplacedventrally

Diapophyses andparapophysesprojectventrolaterally

Page 29: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

BrontosaurusexcelsusholotypeYPM 1980

Cervical ?8

Cervical ribsrobust

Diapophyses andparapophysesrobust

Cervical ribsdisplacedventrally

Diapophyses andparapophysesprojectventrolaterally

Cervical ribs haveanteroventral knobs

Page 30: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Knight's Brontosaurus (1897)

Page 31: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department
Page 32: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department
Page 33: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department
Page 34: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department
Page 35: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Why this big, heavy, weird neck?expensive to build, maintain, and operate.

Page 36: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department
Page 37: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department
Page 38: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Taylor and Wedel (2013a:26) just gave up

Page 39: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Taylor and Wedel (2013a:26) just gave up

Page 40: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

No sex, please, we're sauropods

Page 41: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

But apatosaurs are different

Emiliano Troco

Page 42: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

But apatosaurs are different

John Conway

Page 43: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Sexual combat like necking giraffes?

Page 44: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Giraffes have crass, uncharismatic,Highly fused mammalian skulls ...

Page 45: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

… Whereas apatosaurs have elegant, fragile skulls

Page 46: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Sexual combat like necking giraffes?

Page 47: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

So what were they doing?

?

Page 48: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

1. Ventral displacement of cervical ribs

Improved mechanical advantage of hypaxial muscles.

Page 49: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Stronger ventral movements of the neck

Page 50: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

2. Ventrolaterally directed parapophyseal rami

Oriented to resist ventral impacts

Page 51: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Provided soft-tissueprotection forthe trachea,oesophagus,and majorblood vessels.

3. Ventral troughbetween thecervical ribs

Page 52: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

4. Ventrolateral processes on the cervical ribs

Apatosaurus louisae holotype CM 3018 cervicals 10–12.(Gilmore 1936:plate XXIV)

Page 53: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Ventral midline callositieson ostrich torsos

Slack (2002: figure 4)

(Note the hilarious three-toed ostrich feet)

Page 54: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

4. Ventrolateral processes on the cervical ribs

Maybe instead of this ...

Page 55: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

4. Ventrolateral processes on the cervical ribs

… this? … this?

Page 56: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Keratinous horn in rhinoceroses

Page 57: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

4. Ventrolateral processes on the cervical ribs

Maybe instead of this ...

Page 58: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

4. Ventrolateral processes on the cervical ribs

… this?

Page 59: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Summary

1. Ventral displacement of cervical ribs Improved mechanical advantage of hypaxial muscles. Stronger ventral movements of the neck

2. Ventrolaterally directed parapophyseal rami Oriented to resist ventral impacts

3. Ventral trough between the cervical ribs Provided soft-tissue protection for the trachea, blood vessels.

4. Ventrolateral processes on the cervical ribs Calloused lumps or spikes on ventral surface

⇒ combat by crashing necks ventrally.

Page 60: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department
Page 61: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department
Page 62: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department
Page 63: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department
Page 64: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Male southern elephant seals (Mirounga leonina)

Page 65: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Male northern elephant seals (Mirounga angustirostris)

Page 66: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Male northern elephant seals (Mirounga angustirostris)

Page 67: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Male southern elephant seal (Mirounga leonina)

Page 68: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Northern elephant seal (Mirounga angustirostris)

University Museum of Zoology, Cambridge

Page 69: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Northern elephant seal (Mirounga angustirostris)

University Museum of Zoology, Cambridge

Page 70: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Sea lion (Zalophus californianus)

Page 71: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Sea lion (Zalophus californianus)

Page 72: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Huge soft-tissue envelope

Page 73: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Extremely inexact analogue

Page 74: Were the necks of Apatosaurus and Brontosaurus adapted for combat? Michael P. Taylor 1*, Mathew J. Wedel 2, Darren Naish and 3 Brian Engh 4. 1 Department

Brian Engh