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  • Mechanics of bipedalism suggest dinosaurs had to be warm-blooded. Or: Why the aerobic capacity model needs to be retired.

    The old "cold blooded or warm blooded" argument once again rears its ugly head.

    [Editor’s note: A response from the authors can be found here. It answers many of the questions I had about the paper, though I feel the biggest question remains open for debate. I appreciate the authors taking their time to answer my questions, and PLoS ONE for allowing this type of open communication.]

    This post has taken an inordinate amount of time to write up. Mostly because it required finding enough free time to sit down and just type it out.  So I apologize ahead of time for bringing up what is obviously old news, but I felt this paper was an important one to talk about, as it relied on a old, erroneous, but very pervasive, popular and rarely questioned hypothesis for how automatic endothermy (mammal and bird-style “warm-bloodedness”) evolved.

    Back in November, a paper was published in the online journal: PLoS ONE. That paper was:

    Pontzer, H., Allen, V. & Hutchinson, J.R. 2009. Biomechanics of Running Indicates Endothermy in Bipedal Dinosaurs. PLoS ONE.Vol 4(11): e7783.

    Using muscle force data for the hindlimbs of theropods, and applying it to a model based on Pontzer (2005, 2007), the authors were able to ascertain the approximate aerobic requirements needed for large bipedal theropods to move around. Their conclusion was that all but the smallest taxa had to have been automatic endotherms (i.e. warm-blooded).

    Time to stop the ride and take a closer look at what is going on here.

    In 2004, John Hutchinson – of the Royal Veterinary College, London UK – performed a mathematical study of bipedal running in extant taxa. He used inverse dynamics methods to estimate the amount of muscle that would be required for an animal to run bipedally. He then tested his models on extant animals (Basiliscus, Iguana, Alligator, Homo, Macropus, Eudromia, Gallus, Dromaius, Meleagris, and Struthio). The predictive capacity of his model proved to be remarkably substantial and stable (Hutchinson 2004a).  A follow up paper in the same issue (Hutchinson 2004b) used this model to predict bipedal running ability in extinct taxa (Compsognathus, Coelophysis, Velociraptor, Dilophosaurus, Allosaurus, Tyrannosaurus and Dinornis).  Results from this study echoed previous studies on the running ability of Tyrannosaurus rex (Hutchinson & Garcia 2002), as well as provided data on the speed and agility of other theropod taxa.

    The difference between effective limb length and total limb length in the leg of Tyrannosaurus rex

    Meanwhile in 2005, Herman Pontzer – of Washington University in St. Louis, Missouri – did a series of experiments to determine what was ultimately responsible for the cost of transport in animals. To put it another way: Pontzer was searching for the most expensive thing animals have to pay for in order to move around. One might intuitively assume that mass is the ultimate cost of transport. The bigger one gets, the more energy it requires to move a given unit of mass, a certain distance. However experiments on animals found the opposite to be the case. It actually turns out that being bigger makes one “cheaper” to move.  So then what is going on here?

    Pontzer tested a variety of options for what could be happening; from extra mass, to longer strides. In the end Pontzer found that the effective limb length of animals, was ultimately the limiting factor in their locomotion. Effective limb length differs from the entirety of the limb. Humans are unique in that our graviportal stance has us using almost our entire hindlimbs. Most animals, however, use a more crouched posture that shrinks the overall excursion distance of the hindlimb (or the forelimb). By taking this into account Pontzer was able to find the one trait that seemed to track the best with cost of transport in animals over a wide taxonomic range (essentially: arthropods – birds).

    This latest study combines these two technique in order to ascertain the minimum (or approx minimum) oxygen requirements bipedal dinosaurs would need in order to walk, or run.

    As with the previous papers, the biomechanical modeling and mathematics are elegant and robust. However, this paper is not without its flaws. For instance in the paper the authors mention:

    We focused on bipedal species, because issues of weight distribution between fore and hindlimbs make biomechanical analysis of extinct quadrupeds more difficult and speculative.

    Yet this did not stop the authors from applying their work on bipeds, to predicting the maximum oxygen consumption of quadrupedal iguanas and alligators. No justification is ever really given for why the authors chose to do this. Making things even more confusing, just a few sentences later, it is mentioned (ref #s removed to avoid confusion):

    Additionally, predicting total muscle volumes solely from hindlimb data for the extant quadrupeds simply assumes that the fore and hindlimbs are acting with similar mechanical advantage, activating similar volumes of muscle to produce one Newton of GRF. This assumption is supported by force-plate studies in other quadrupeds (dogs and quadrupedal chimpanzees)

    The force plate work cited is for quadrupedal mammals. However, mammals are not reptiles. As Nicholas Hotton III once mentioned (1994), what works for mammals, does not necessarily work for reptiles. This is especially so for locomotion.

    In many reptiles (including the taxa used in this study) the fore and hindlimbs are subequal in length; with the hindlimbs being noticeably longer and larger. Most of the propulsive power in these reptiles comes from the hindlimbs (which have the advantage of having a large tail with which to lay their powerful leg retractor on). The result is that – unlike mammals – many reptiles are “rear wheel drive.”

    The last problem is by far the largest, and ultimately proves fatal to the overall conclusions of the paper. The authors operated under the assumptions of the aerobic capacity model for the evolution of automatic endothermy.

    It is here that we come to the crux of the problem, and the main subject of this post.

    Continue reading  Post ID 516


  • Alligators can shift their lungs and lizard ecology determines movement.

    There were two new papers released today in the Journal of Experimental Biology.

    The first one is the biggest, as it received a news story.


    Uriona, T.J., and Farmer, C.G. 2008. Recruitment of the diaphragmaticus, ischiopubis and other respiratory muscles to control pitch and roll in the American alligator (Alligator mississippiensis). J. Exp. Biol. Vol. 211: 1141-1147 doi: 10.1242/jeb.015339

    Abstract

    We used electromyography on juvenile American alligators to test the hypothesis that the following muscles, which are known to play a role in respiration, are recruited for aquatic locomotion: M. diaphragmaticus, M. ischiopubis, M. rectus abdominis, M. intercostalis internus, and the M. transversus abdominis. We found no activity with locomotion in the transversus. The diaphragmaticus, ischiopubis, rectus abdominis and internal intercostals were active when the animals executed a head-down dive from a horizontal posture. Weights attached to the base of the tail resulted in greater electrical activity of diaphragmaticus, ischiopubis and rectus muscles than when weights were attached to the head, supporting a role of this musculature in locomotion. The diaphragmaticus and rectus abdominis were active unilaterally with rolling maneuvers. Although the function of these muscles in locomotion has previously been unrecognized, these data raise the possibility that the locomotor function arose when Crocodylomorpha assumed a semi-aquatic existence and that the musculoskeletal complex was secondarily recruited to supplement ventilation.

    Scientists at the University of Utah have discovered the unique internal subtleties that allow crocodylians to sink, rise, pitch and roll; all without disturbing the water (much). It turns out that the main muscles used for breathing, are also used to actually shift the lungs within the body!

    That’s just crazy awesome. Uriona & Farmer’s work raises the question of how prevalent this ability is in other semi-aquatic animals (e.g. seals, terrapins, manatees). By shifting the lungs further back in the body, crocodylians are able to change their local density. This allows the front, or back of the animal to rise and sink separately from the rest of the body. So too does moving the lungs from side to side allow for rolling in the water. All of this can occur without the need to move any external body parts. This means no extra turbulence gets created in the water, thus allowing crocodylians to better sneak up on their fishy, or fleshy prey.




    Baby crocodiles exhibiting their unique pulmonary powers.

    If anything, it sure speaks to why crocodyliformes have held dominion over the semi-aquatic niche for over 200 million years. Uriona and Farmer do suggest that the ability of these respiratory muscles to do this might not be an exaptation. Rather, this might have been the initial impetus behind the evolution of these muscles. Only later would they have been exapted to help with breathing on land. Though the authours provide some good parsimonious reasons for why this may be (basically it would take less evolutionary steps to accomplish than the other way around), it doesn’t really jive with the fossil evidence. Part of the reason why the crocodylian diaphragm works, is because the pubis (the forepart of the hip bone in most animals, and the part that juts out so prominently in theropod dinosaurs), is mobile. This mobility occurred early on in crocodyliforme evolution, with the crocodylomorph Protosuchus having a pubis that was almost mobile. The problem arises when one looks at this early crocodylomorph. Protosuchus was obviously terrestrial. If Protosuchus was evolving a mobile pubis already, then it was doubtful that it was being used to allow lung shifting in the body (an ability that is helpful when underwater, but pretty pointless on land). Furthermore, Crocodylia proper is the umpteenth time that crocodyliformes have returned to a semi-aquatic existence. It is doubtful that all the numerous land outings that occurred during crocodyliforme evolution, would have retained the ability to move the lungs to and fro. It seems far more likely that this was an ability that evolved in Crocodylia, or somewhere close by on the evolutionary tree, in some taxa that was still semi-aquatic.


    Protosuchus

    Protosuchus richardsoni. An example of an early crocodylomorph.

    Of course it is also possible that crocodyliforme phylogeny is just all f-ed up. With the amount of convergence rampant in that lot, this remains a distinct possibility.

    Either way this is a cool discovery, and one worthy of adding to the crocodylian pages.

    The second paper also comes from the Journal of Experimental Biology. This one involves lizards.


    McElroy, E.J., Hickey, K.L., Reilly, S.M. 2008. The correlated evolution of biomechanics, gait and foraging mode in lizards. J. Exp. Biol. Vol. 211: 1029-1040. doi: 10.1242/jeb.015503

    Abstract

    Foraging mode has molded the evolution of many aspects of lizard biology. From a basic sit-and-wait sprinting feeding strategy, several lizard groups have evolved a wide foraging strategy, slowly moving through the environment using their highly developed chemosensory systems to locate prey. We studied locomotor performance, whole-body mechanics and gaits in a phylogenetic array of lizards that use sit-and-wait and wide-foraging strategies to contrast the functional differences associated with the need for speed vs slow continuous movement during foraging. Using multivariate and phylogenetic comparative analyses we tested for patterns of covariation in gaits and locomotor mechanics in relation to foraging mode. Sit-and-wait species used only fast speeds and trotting gaits coupled with running (bouncing) mechanics. Different wide-foraging species independently evolved slower locomotion with walking (vaulting) mechanics coupled with several different walking gaits, some of which have evolved several times. Most wide foragers retain the running mechanics with trotting gaits observed in sit-and-wait lizards, but some wide foragers have evolved very slow (high duty factor) running mechanics. In addition, three evolutionary reversals back to sit-and-wait foraging are coupled with the loss of walking mechanics. These findings provide strong evidence that foraging mode drives the evolution of biomechanics and gaits in lizards and that there are several ways to evolve slower locomotion. In addition, the different gaits used to walk slowly appear to match the ecological and behavioral challenges of the species that use them. Trotting appears to be a functionally stable strategy in lizards not necessarily related to whole-body mechanics or speed.

    I haven’t had a chance to read much more than what was written already. I do take a bit of offense to the authours referring to scleroglossan foraging technique as “slow,” but what are you going to do?

    I do find it interesting that lizards seem to have lost the ability to “walk” numerous times. That almost seems bizarre. The study points out that ecology produces heavy pressures on lizards in terms of their locomotion type. This is extremely pertinent given how often one hears the old (and wrong!) adage about “reptiles” being incapable of intense aerobic activity.

    According to the above study (among others), it all depends on the animals being tested.

    There we go. Two really cool papers on reptiles, being released in one day.

    ~Jura

    Yes, I know. I used jive. I’m sorry.