Reptilian Rants

30 June, 2008

The old grey sauropod just ain’t what she used to be.

Filed under: Uncategorized — Jura @ 18:31

Actually, I’ve never thought that sauropods were grey. Mammals in general tend to be rather bland in their colour schemes. Reptiles don’t have that problem. With xanthaphores (yellow pigmented cells), erythrophores (red pigmented cells) iridophores (iridescent cells) and melanophores (dark pigmented cells), the range of colour available to reptiles, and by extension - dinosaurs, is quite vast.

That said, I always pictured sauropods as either a brownish green colour, or maybe a very pale blue (blue is generally rare in tetrapods, hence the thought of it being a weak blue).

But I digress.

I grew up during an interesting time for dinosaur research. Unlike the majority of paleontologists working right now, I didn’t grow up learning about dinosaurs being slow, and sluggish mistakes of nature. I also didn’t grow up with the “hummingbirds on crack” version of dinosaurs that is currently pervading popular culture. Rather, I grew up during that strange transitory phase of the Dinosaur Renaissance where dinosaurs were sometimes viewed as sluggish beasts, and other times as racecars of the Mesozoic.

The result, I think, has been a slightly detached and objective look at how perceptions of dinosaurs have changed over time.

Image borrowed from the Old Dinosaur Books site

A “Brontosaurus” getting attacked by Allosaurus during a sojourn on land to lay her eggs. Ah, the classics.

One book I remember fondly was the Golden Book of Dinosaurs (shown above). It featured these beautiful drawings of dinosaurs living life as best we thought at the time. One picture that really stuck in my head, was a shot of two Brachiosaurus; one on land and the other so deep in a lake that one could only make out the crest on the head. I found that page to be so immersive and atmospheric. My knowledge of physics was not so good at the time, so it never dawned on me that this poor sauropod was basically breathing through a straw, with its lungs separated by at least 2 atmospheres from the air entering (as best it could) the nostrils.

Then, around the early nineties when Jurassic Park the book came out, I started to note a distinct change in how dinosaurs were being portrayed. No longer were sauropods swamp bound behemoths. Now they were fully terrestrial titans that could not only support their weight on all four legs, but could even do so on 2 (well 3, if one counts the tail). It was around this time that Robert Bakker’s infamous “Dinosaur Heresies” started making the rounds.

Predatory Dinosaurs of the World. Available on Amazon

Now, admittedly, Heresies came out around 1986, and the changing view of dinosaurs actually started in the seventies. However, it wasn’t until the early nineties that the full effects of Bakker’s work could truly be appreciated. If anything this gives one an idea of the kind of inertia one must deal when it comes to getting scientific ideas out into the public.

Again I digress.

It was around the early nineties when I first read The Dinosaur Heresies. The first few chapters were amazing. I had never seen dinosaurs portrayed this way. They walked better and were more active. In many ways they better fit the concept I had in my head all along.

Then I came up to the end of chapter 3. The thesis of this chapter was to explain why reptiles should not be viewed as inferior to mammals. In order to do so Bakker explained all the various ways in which extant reptiles outshine extant mammals. The end of the chapter features a beautifully drawn shot of the “panzer croc” Pristichampsus snatching a Hyracotherium (formerly Eohippus). The caption read:

Pristichampsus hunted during the Eocene Epoch, about 49 million years ago, but it was very rare, much rarer than big mammalian predators, proof that cold-bloodedness was a great disadvantage.

That’s when the real thesis of the book hit me. The argument wasn’t: “Dinosaurs weren’t slow and stupid, because of the following.”

Rather the argument was: “Dinosaurs weren’t cold-blooded”, because the facts show the following.”

In order to pull dinosaurs out of the mire, Bakker had to change their fundamental thermophysiology. The general concept that cold-bloodedness is inferior to warm-bloodedness, remained the same. This despite Bakker’s initial attempt to explain how “cold-blooded” reptiles outshine “warm-blooded” mammals.

Bakker’s book was just the start. From there, we had Adrian Desmond’s “The Hot Blooded Dinosaurs” and Gregory S. Paul’s infamous: “Predatory Dinosaurs of the World.” Each new book taking the “dinosaurs can’t be cold-blooded” argument a little further. By the time we hit Predatory Dinosaurs of the World, Tyrannosaurus rex was running along at 40mph, dromaeosaurs were practically flapping around, and every species of dinosaur was reaching adult size by between 4-10 years of age.

Sadly it was at this point that Jurassic Park was written. As hardcore fans know, it was Greg Paul’s erroneous sinking of Deinonychus antirrhopus into Velociraptor that gave us the JP “raptors.” It was also at this point that the pendulum of dinosaur physiology officially swung the other way.

The thing that had always bugged me about this view of dinosaurs was the sheer lack of supporting data for it. The assumption was always that dinosaurs were so vastly different from “typical reptiles” that they had to have been doing something different. Yet when one looked at the actual data, dinosaurs came out looking slightly odd at best. For the most part dinosaurs fit the reptile mold quite well. It was these elusive “classic reptiles” that didn’t appear to exist.

Most reptiles don’t fit the “typical reptile” mold at all. Yet despite numerous papers over the past 30 years depicting reptiles doing things normally thought un-reptile like (e.g. caring for their young, competing with large mammals, etc), most of this was dutifully ignored in favour of an older, more outdated view.

It was a problem that Nicholas Hotton III (1980) aptly called: The “endothermocentric fallacy.” Basically, the assumption that being an endotherm is inherently superior to being an ectotherm. Part of that superiority included the ability of endotherms to do everything faster and “better” than similar sized ecotherms. This problems with this way of thinking warrants an entire blog post to itself.  So rather than get bogged down with this particular, I’ll touch more on the endothermocentric fallacy at a later date. For now all that one needs to keep in mind is that the thinking of the time was that if dinosaurs were going to be active at all, then they had to be endotherms.

By the late nineties we had the first evidence of feathers in a small branch of the theropods (Maniraptora). Birds were officially adopted into the dinosaur family tree and the fully endothermic concept of Dinosauria was completely entrenched.

The funny thing, of course, is that this dogmatic view of dinosaur metabolism was just as bad as the early 20th century’s “cold-blooded” swamp bound view. Sure dinosaurs were more active now, but the data supporting it was just as nebulous as the stuff that was used to keep dinos in the swamp.

Enter the 21st century, and the late…um, 0’s (does anyone have a name for this decade yet?). Biomechanic work on dinosaurs has started to reveal amazing insights into the physical limits of what dinosaurs could do, and the results have started to pull the pendulum back again.

Work by John Hutchinson and Mariano Garcia (2002) on T.rex showed that not only could T.rex not hit 40mph, but it technically couldn’t run either. A biomechanic assessment of theropod forelimbs by Ken Carpenter (2002) has shown that the “bird-like” dromaeosaurs could not fold their arms up like birds after all.

Work by Rothschild and Molnar (2005) on sauropod stress fractures shows no signs of rearing activity in sauropods, while work by Kent Stevens and J. Michael Parrish (2005) has pulled the swan-like curve out of sauropod necks; placing things far more horizontally.

Work by Gregory Erickson and others (2001) on micro-slices of dinosaur bone, has indicated that very few dinosaurs hit adult size in less than 15 years.

Now we have a new study by Lehman and Woodward (2008) which follows up on Erickson et al’s work, and actually shows that even this toned down version of dinosaur growth, is probably too fast as well. Lehman and Woodward focused on sauropods and studies on their bone microstructure. What they did was compare bone growth data to a well used equation for growth in animals.

Bertalanffy growth equation

Deemed the Bertalanffy equation; it states that the mass at any given age is an exponential function limited by the asymptote of adult body mass. This equation has been used extensively in studies on bird and elephant growth among others. An example of the equation is given to the right for fish.

When the authours did this, they discovered something quite interesting. Instead of taking 15 years to reach adult mass, sauropods like Apatosaurus excelsus took closer to 70 years!!.

Other sauropods measured took between 40 and 80 years! This is a substantial decrease in growth rate estimated before. Mind you this is data taken, in some cases, from the same piece of bone that Erickson et al had used. So one can’t suggest anomalous bones being used as the reason behind the surprising results. The authours also went to great lengths to take into account differences in mass estimations as well as allometric growth of body parts. In each case, the changes had little affect on the overall outcome (in many cases, it made growth go even slower).

Now keep in mind, we are talking about the time it took sauropods to reach full adult size. This is not the time taken to reach sexual maturity. Earlier studies by Erickson et al (2007) had already discovered that dinosaurs didn’t wait to grow up before engaging in sex, so there is no issue here of 80 year old sauropods finally “doing the nasty.”

What this does show is that growth in dinosaurs might not be as determinate as initially thought. An 80 year old sauropod might just have been close to the edge of its lifespan at this point (though the possibility of bicentennial sauropods does still exist). It also shows that dinosaurs had growth rates far closer to the realm of reality (before it was hard to imagine how an Apatosaurus excelsus was able to pound down enough food daily to add 13.6 kg of new mass a day. Especially given their small mouths).

Thermophysiologically what does this all mean? Were dinosaurs “cold-blooded” after all?

That’s one of those questions that will never be fully answered (short of a time machine). What this does do, is pull dinosaurs ever further away from the “definitely warm-blooded” category, and push them right back into the middle again. When/if the dust settles on this metabolism debate, I suspect that dinosaurs will probably remain in the middle somewhere.

Of course while all of this is going on with dinosaurs, we have other studies like those of Tumarkin-Deratzian (2007) showing the existence of fibrolamellar bone growth in wild alligators, that are finally moving the rusty pendulum of reptile metabolism out of the “classic reptile” category, and much closer to the middle.

So in the end dinosaurs will still probably wind up being “good reptiles.” Thankfully the exact definition of what that entails will have probably changed by then.

~ Jura


References

Carpenter, K. 2002. Forelimb Biomechanics of Nonavian Theropod Dinosaurs in Predation. Concepts of Functional Engineering and Constructional Morphology. Vol. 82(1): 59-76.
Erickson, G.M., Curry Rogers, K., Varricchio, D.J., Norell, M.A., Xu, X. 2007. Growth Patterns in Brooding Dinosaurs Reveals the Timing of Sexual Maturity in Non-Avian Dinosaurs and Genesis of the Avian Condition. Biology Letters Published Online. doi: 10.1098/rsbl.2007.0254
Erickson, G.M., K. C. Rogers, and S.A. Yerby. 2001. Dinosaurian Growth Patterns and Rapid Avian Growth Rates. Nature 412: 429–433.
Hotton, N., III. 1980. An Alternative to Dinosaur Endothermy: The Happy Wanderers. In A Cold Look at the Warm-Blooded Dinosaurs (R.D.K. Thomas and E.C. Olson Eds.), pp. 311-350, AAAS, Washington, DC
Hutchinson, J.R., Garcia, M. 2002. Tyrannosaurus was not a fast runner. Nature 415: 1018-1021.
Lehman, T.M., and Woodward, H.N. 2008. Modeling Growth Rates for Sauropod Dinosaurs. Paleobiology. Vol. 34(2): 264-281.
Rothschild, B.M., and Molnar, R.E. 2005. Sauropod Stress Fractures as Clues to Activity. In Thunder Lizards: The Sauropodomorph Dinosaurs. (Virginia Tidwell and Kenneth Carpenter eds). Indiana University Press. pp 381-394.
Stevens, K.A., and Parrish, J.M. 2005. neck Posture, Dentition, and Feeding Strategies in Jurassic Sauropod Dinosaurs. In In Thunder Lizards: The Sauropodomorph Dinosaurs. (Virginia Tidwell and Kenneth Carpenter eds). Indiana University Press. pp 212-232.
Tumarkin-Deratzian, A.R. 2007. Fibrolamellar bone in adult Alligator mississippiensis. Journal of Herpetology. Vol. 41. No.2:341-345.

29 June, 2008

So, why go bipedal?

Filed under: Uncategorized — Jura @ 1:24

As human beings this might come off as somewhat of a “duh” question.

“To free our hands up, of course.”

Ah, but like most things in life, the common sense answer is not the right one.

Consider all the bipedal animals alive today. We have humans, obviously; kangaroos, birds, a few lemurs, and a whole swath of lizards. Of these, the “free up the forelimbs” argument really only holds true for humans, lemurs and birds. What of kangaroos and all those lizards? Exactly what does one benefit from when going bipedal?

Do you go faster, run longer, or gain a better vantage point?




C.kingii during a typical foraging run.

All these questions were asked, and somewhat answered in a recent paper by Clemente et al.

Clemente, C.J., Withers, P.C., Thompson, G., Lloyd, D. 2008. Why Go Bipedal? Locomotion and Morphology in Australian Agamid Lizards.J. Exp. Bio. 211: 2058-2065

Abstract:

Bipedal locomotion by lizards has previously been considered to provide a locomotory advantage. We examined this premise for a group of quadrupedal Australian agamid lizards, which vary in the extent to which they will become bipedal. The percentage of strides that each species ran bipedally, recorded using high speed video cameras, was positively related to body size and the proximity of the body centre of mass to the hip, and negatively related to running endurance. Speed was not higher for bipedal strides, compared with quadrupedal strides, in any of the four species, but acceleration during bipedal strides was significantly higher in three of four species. Furthermore, a distinct threshold between quadrupedal and bipedal strides, was more evident for acceleration than speed, with a threshold in acceleration above which strides became bipedal. We calculated these thresholds using probit analysis, and compared these to the predicted threshold based on the model of Aerts et al. Although there was a general agreement in order, the acceleration thresholds for lizards were often lower than that predicted by the model. We suggest that bipedalism, in Australian agamid lizards, may have evolved as a simple consequence of acceleration, and does not confer any locomotory advantage for increasing speed or endurance. However, both behavioural and threshold data suggest that some lizards actively attempt to run bipedally, implying some unknown advantage to bipedal locomotion.

The conclusions reached, were interesting and quite unexpected.

I’d say the most surprising part would have to be the discovery that both endurance and speed were found to be inconsequential. Those were the two forces that I figured would have driven the push towards bipedalism. Apparently this is not the case.

In fact, the only correlate that the authours found was that a switch to bipedalism resulted in an increase in acceleration. Short of that, the authours viewed bipedalism as more of a side effect of speedy locomotion, rather than anything else.

As one authour put it: “The lizards were pulling a wheelie.”

There are some gripes (niggles if you will) with the paper. For one, the authours assert that a switch to bipedalism allowed birds to incorporate their forelimbs into wing design. While being bipedal certainly allowed for this, it could not have been the cause. Birds descended from dinosaurs, and very, very very few dinosaurs had wings. Theropods were sporting freed forelimbs for some 80 million years, or so (probably longer given the proposed ancestors of dinosaurs). Wings were not the cause, simply a benefit. Something else had to have spurred the evolution of bipedality.

Side note: What the heck were theropods doing with those forelimbs anyway? Most paleo artists tend to draw theropods with their arms tucked to the side, yet work by Carpenter (2002) has shown that there was some considerable range of motion in theropod forelimbs. They weren’t brachiating from tree to tree, or anything, but they could certainly do a heckuvalot more than just tuck their arms to the side. Even T.rex with its embarrassingly short forearms, had a surprisingly large range of motion to them. So what’s up paleo art guys? Let’s see some theropods putting their arms to use.

The largest SNAFU in the paper comes from the cladogram that the authours chose. They chose to go with the broken molecular tree used by Townsend et al, which asserts that Iguanians are actually Scleroglossan lizards. This might sit all fine and well when looking at molecules, but it utterly falls apart upon a morphological assessment. In order for Iguanians to fit in the Scleroglossan family tree, they had to have undergone a tonne of morphological reversals, including the re-softening of the tongue and the re-evolution of both postorbital bars (surprisingly, the latter is actually not out of the realm of impossibility as tuataras have apparently done just that).

Due to this tree choice, the authours erroneously concluded that bipedalism evolved only once in the lacertilian tree and was lost a multitude of times, with a putative re-acquirement in varanids.

Another minor complaint comes from the very slight use of Chlamydosaurus kingii; the only lizard known to be a “true” biped (see: Shine & Lambeck 1989). Given that the authours were trying to spot differences between bipeds and quadrupeds, I can understand the use of lizards like Ctenophorus, with their greater spectrum of gaits. However, in doing so they should have qualified their conclusions better in regards to how lizards obtain a bipedal stance. In Chlamydosaurus, bipedal trotting is attained from a standing start. Quadrupedal stance is only seen when stopping to eat. Furthermore, foraging runs and escape runs use two different gaits, with the latter gait more akin to that of other facultatively bipedal lizards. Judging from the stats given in the paper, it seems apparent that the C.kingii used in this study were mostly running away.

The final complaint comes from the supplementary movies given.

Frankly the movies are just too short. I have to go super slow-mo just to see anything. So that’s a bummer.

Overall the paper is rather good. The authours discovered that Lophognathus gilberti runs bipedally 95% of the time. This suggests that C.kingii is not the only truly bipedal lizard out there.

The authours also observed that, despite any advantage in speed, or endurance, some lizards intentionally push their center of mass towards their hips early on in the running phase in order to more quickly obtain a bipedal gait. The reasons behind this are unclear, but do suggest that bipedalism confers some advantage not discovered during this experiment.

One advantage alluded to, but never really elaborated on, was the faster acceleration noted in bipeds. Though maximum speed was no different than in a quadruped, this speed was obtained faster. Ecologically I could see this being very advantageous. When one is trying to avoid a predator, maximum top speed is probably less important than reaching that top speed as fast as possible.

If one lizard has a top speed of 12km/hr and another has a top speed of 8km/hr, and the goal (burrow) is only 50 meters away, then that extra speed isn’t going to mean much. Especially if the slower lizard is able to hit its top speed faster.

Apparently lizards “pull a wheelie” because in their ecosystem it pays to be a drag racer, rather than a Daytona 500 car.

~Jura


References

Carpenter, K. 2002. Forelimb Biomechanics of Nonavian Theropod Dinosaurs in Predation. Concepts of Functional Engineering and Constructional Morphology. Vol. 82(1): 59-76.

Shine, R., & Lambeck, R. 1989. Ecology of Frillneck Lizards, Chlamydosaurus kingii (Agamidae), in Tropical Australia. Aust. Wildl. res. Vol. 16: 491-500.

Townsend T, Larson A, Louis E, Macey JR. 2004. Molecular Phylogenetics of Squamata: The Position of Snakes, Amphisbaenians, and Dibamids, and the Root of the Squamate Tree. Syst Biol. Vol. 53(5):735-57.

23 June, 2008

Stuff is coming

Filed under: Uncategorized — Jura @ 0:19

I know what your thinking, and I have no idea what happened to May either.  Things have been busy for me IRL, but I didn’t want to leave the site hanging.

Anyway, I have some time off coming,  so I’ll be posting something new soon.

Till then: R.I.P. George Carlin. You will be missed, but never replaced.

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