Origin of bipedalism in dinosaurs: Overlooking Carrier’s Constraint

Persons and Currie 2017 debunk on old theory
on bipedalism in dinosaurs and introduce a new one that suffers from taxon exclusion while overlooking a very popular theory from the last thirty years: Carrier’s Constraint (Carrier 1987).

From the abstract:
“Bipedalism is a trait basal to, and widespread among, dinosaurs. It has been previously argued that bipedalism arose in the ancestors of dinosaurs for the function of freeing the forelimbs to serve as predatory weapons.”

I never heard of this reason before. Predatory weapons only happen as a result and much later phylogenetically and only sometimes.

“However, this argument does not explain why bipedalism was retained among numerous herbivorous groups of dinosaurs. We argue that bipedalism arose in the dinosaur line for the purpose of enhanced cursoriality.”

The term ‘enhanced’ is pretty vague. Does it mean ‘better’? But can that be proven? The fastest animals on land now are quadrupedal cheetahs. Bipedal Chlamydosaurus does not have greater speed or endurance. Persons and Currie bring up the “tripping on one’s own forefeet” hypothesis and that, IMHO, has some validity.

“Modern facultatively bipedal lizards offer an analog for the first stages in the evolution of dinosaurian bipedalism. Many extant lizards assume a bipedal stance while attempting to flee predators at maximum speed.”

But quadrupedal lizards are just as fast as bipedal ones. Lizards gain no speed when switching to bipedal locomotion as Persons and Currie also note.

Bipedal lizard video marker

Figure 1. Click to play video. Just how fast can quadrupedal/bipedal lizards run? This video documents 11 meters/second in a Callisaurus at the Bruce Jayne lab.

“Bipedalism, when combined with a caudofemoralis musculature, has cursorial advantages because the caudofemoralis provides a greater source of propulsion to the hindlimbs than is generally available to the forelimbs.”

Yes, at first, especially when the forelimbs are lifted from the ground! Persons and Currie stay clear of the bipedal ability of fenestrasaurs including pterosaurs. There, in taxa like Cosesaurus, the driving force switches to the hips.

“That cursorial advantage explains the relative abundance of cursorial facultative bipeds and obligate bipeds among fossil diapsids and the relative scarcity of either among mammals.”

Actually there is no abundance of bipeds anywhere among diapsids, except in the Fenestrasauria (not related to archosaur-line diapsids) and Archosauria + Poposauria. Persons and Currie also stay clear of the inverted bipeds among mammals, the bats, and they are numerous.

None of the so-called ‘reasons’ why are pertinent
without the random evolution of longer hind limbs than forelimbs and the ability to balance over the hind limbs, whether running or standing still. It also helps to have even a small anterior addition to the ilium, according to Shine and Lambeck 1989. The pubic foot of theropods and the prepubis of pterosaurs also provide femoral muscle anchors.


  1. Persons and Currie do not indicate the node at which bipedalism arose in the last common ancestor of bipedal crocs and dinosaurs: Gracilisuchus and Turfanosuchus at the base of the Poposauria. In the large reptile tree (LRT)  Gracilisuchus (Fig. is the last common ancestor of bipedal crocs, like Scleromochlus, and bipedal pro-dinosaurs, like Lewisuchus.
  2. Persons and Currie subscribe to the outdated hypothesis of “Avemetatarsalia” in which former members, like pterosaurs now nest with lepidosaurs and Lagerpeton now nests with chanaresuchids.
  3. Persons and Currie also avoid the likely bipeds, Arizonasaurus and Postosuchus.
  4. Persons and Currie discuss the the likely biped, Eudibamus, but incorrectly ascribe it to the bolosaurs.
  5. Persons and Currie overlooked Carrier’s Constraint, which holds that,“air-breathing vertebrates which have two lungs and flex their bodies sideways during locomotion find it very difficult to move and breathe at the same time, because the sideways flexing expands one lung and compresses the other, shunting stale air from lung to lung instead of expelling it completely to make room for fresh air.” — but is that the reason to go bipedal? or just the first and biggest advantage narrow-gauge bipedal reptiles enjoy?
Click to enlarge. Squamates, tritosaurs and fenestrasaurs in the phylogenetic lineage preceding the origin of the Pterosauria.

Figure 2. Tritosaurs and fenestrasaurs in the phylogenetic lineage preceding the origin of the Pterosauria.

What fenestrasaurs gain by a bipedal configuration

  1. height dominance over conspecific rivals for mating privileges. This is emphasized in Langobardisaurus with its long neck. This is emphasized by Cosesaurus by flapping and leaping, both working to increase height.
  2. Ability to breathe while running for added endurance
Chlamydosaurus, the Austrlian frill-neck lizard

Fig. 3. Chlamydosaurus, the Austrlian frill-neck lizard with an erect spine and elevated tail. At one time some paleontologists did not believe what you can see here, that this lizard can stand bipedally. Such was their bias.

What the lizard, Chlamydosaurus, gains by bipedal configuration

  1. combined with their frightfully opening frill neck, dominance over rivals and interlopers, which they charge bipedally.
  2. better ability to survey the local area for rivals (principally) and predators while on the ground, — but Chlamydosaurus is primarily (90%) arboreal for the same reason and 90% bipedal while on the ground, not just while running, which some paleontologists are not aware of or did not believe (Hone and Benton 2007, 2009).
Figure 1. The origin of dinosaurs to scale. Gray arrows show the direction of evolution. This image includes Decuriasuchus, Turfanosuchus, Gracilisuchus, Lewisuchus, Pseudhesperosuchus, Herrerasaurus, Tawa and Eoraptor.

Figure 4. The origin of dinosaurs to scale. Gray arrows show the direction of evolution. This image includes Decuriasuchus, Turfanosuchus, Gracilisuchus, Lewisuchus, Pseudhesperosuchus, Herrerasaurus, Tawa and Eoraptor. Note the phylogenetic miniaturization.

What Gracilisuchus gained by a bipedal configuration

  1. Gracilisuchus is not much taller bipedally. Remember, archosaurs had no scales at this point. Feather quills would appear on dino backs. Osteoderms appeared along croc backs to support their longer spinal columns. So, standing erect might have just been sexy at first.
  2. Overcoming Carrier’s Constraint: greater endurance by not having to undulate while breathing and so continue breathing while running.

What do bipedal reptiles have in common?

  1. Other than sauropods and other reptiles that adopt a tripodal pose bipedal reptiles are generally small, having experienced phylogenetic miniaturization.
  2. Other than Tanystropheus, bipeds are terrestrial and/or arboreal
  3. Longer hind limbs than forelimbs
  4. Anterior process of the illiim, no matter how small
  5. Typically stronger or more sacral connections to the ilium
  6. Typically a long neck and short torso (but Longisquama (Fig. 2), as a lemur analog, and lemurs themselves break that rule).
Figure 1. The ancestry of Scleromochlus going back to Lewisuchus, Saltoposuchus, Terrestrisuchus, SMNS 12591 and Gracilisuchus.

Figure 1. The ancestry of Scleromochlus going back to Lewisuchus, Saltoposuchus, Terrestrisuchus, SMNS 12591 and Gracilisuchus.

It’s easy to overlook the most obvious.
I have a feeling that this will not be the first time Persons and Currie are going to be reminded of Carrier 1987.

Carrier DR 1987. The evolution of locomotor stamina in tetrapods: circumventing a mechanical constraint. Paleobiology (13): 326–341.
Clemente CJ, Withers PC, Thompson G, Lloyd D 2008. Why Go Bipedal? Locomotion and Morphology in Australian Agamid Lizards.J. Exp. Bio. 211: 2058-2065
Peters D 2000b. A reexamination of four prolacertiforms with implications for pterosaur phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106: 293–336.
Peters D 2007. The origin and radiation of the Pterosauria. In D. Hone ed. Flugsaurier. The Wellnhofer pterosaur meeting, 2007, Munich, Germany. p. 27.
Persons WS and Currie PJ 2017. The functional origin of dinosaur bipedalism: Cumulative evidence from bipedally inclined reptiles and disinclined mammals. Journal of Theoretical Biology, 2017; 420: 1 DOI: 10.1016/j.jtbi.2017.02.032
Shine R and Lambeck R 1989. Ecology of Frillneck Lizards, Chlamydosaurus kingii (Agamidae), in Tropical Australia. Aust. Wildl. res. Vol. 16: 491-500.
Snyder RC 1954. The anatomy and function of the pelvic girdle and hind limb in lizard locomotion. American Journal of Anatomy 95:1-46


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