In another example of slow-cooked science, this paper was the culmination of over three years worth of work collecting data on tegus. For the study, the authors looked at adult black and white tegus (Salvatore merianae). Tegus are an interesting group of lizards. They are the largest members of the family Teiidae and are often referred to as the monitor lizards of the new world, due to their convergent lifestyles (highly predaceous, active foragers). Besides their varanid-like demeanor, tegus are also known for their enormous jowls, especially in the males. The jowls hold the pterygoideus muscles, the big jaw snappers, which have been shown to increase in size for males during the breeding season (Naretto et al. 2014). As reptiles, tegus have been assumed to follow the standard ectothermic lifestyle of requiring external sources of heat to warm their bodies and maintain stable body temperatures. Looking at the natural history of the animals, tegus appear to fit the mold pretty well. They have distinctive winter and summer activity levels. In the summer, the animals regularly maintained body temperatures of 32â€“35Â°C, and in the winter they let their body temperatures drop to the temperature of their burrows (15â€“20Â°C). This is all fine and good for a bradymetabolic, ectothermic lizard, but when the researchers tracked body temperatures over time they discovered something completely unexpected.
This year has seen the discovery of two big deal dinosaur specimens. At least they are a big deal in regards to dinosaur integument and, possibly, metabolism.
First off from a few months ago we had the announcement the theropodÂ Yutyrannus hauli, the “beautiful feathered tyrant.”
Xu, X., Kebai, W., Ke, Z., Qingyu, M., Lida, X., Sullivan, C., Dongyu, H., Shuqing, C., Shuo, W. 2012. A Gigantic Feathered Dinosaur from the Lower Cretaceous of China. Nature. Vol.484:92-95
This was not just a single fossil, but a collection of three fossils (one might be tempted to call it a family group, but that would only be speculation). As with all other dinosaur fossils that have been found to have filamentous integument, these guys come from Liaoning, China. They are suspected to have come from the Jehol Group in the Yixian formation. I say suspected because the complete three specimen set was a purchase from a fossil dealer, an all too common occurrence for Chinese fossils. As such the provenance information is unknown. A lot of Chinese fossil dealers don’t like to give away the location of their find due to the potential loss of other profitable specimens. This current trend in China is a good example of what happens when capitalism comes into play with fossil collecting (something that the U.S. has been mostly, but not entirely, able to avoid). So it is currently uncertain whether these fossils are from the Yixian. However given that all the others guys are too it is probably a good bet. Given the sketchy nature in which many Yixian fossils are collected, coupled with the possibly large consequences of the find, one should naturally be skeptical of the fossil. Had it been one individual on multiple slabs I would question its validity as a real thing. However since Y.huali is known from three individuals, and the filaments seem to follow a consistent pattern around the body (compare that to the helter-skelter nature of Tianyulong‘s preservation), forgery seems unlikely. These guys are probably the real deal. This has some potentially far reaching consequences to interpretations of Late Cretaceous coelurosaurs and the Jehol Biota itself (more on this in a bit).
The second announcement came just a few weeks ago. This was the discovery of a potentially new, miniscule theropod from Bavaria Germany.
Rauhut, O.W.M., Foth, C., Tischlinger, H., Norell, M.A. 2012. Exceptionally Preserved Juvenile Megalosauroid Theropod Dinosaur with Filamentous Integument from the Late Jurassic of Germany. PNAS Early Edition:1203238109v1-201203238.
The specimen is exceptionally well preserved. So well preserved in fact that it actually looks like a plastic toy. While this degree of preservation warrants importance all its own, the main interest behind this new guyâ€”dubbed: Sciurumimusalbersdoerferi (AlbersdÃ¶rfer’s squirrel mimic)â€”is the apparent presence of filamentous integument on the body coupled with its apparent placement among much more basal theropods. This discovery has far reaching consequences for theropod integument interpretations. Note: As with Y.hauli, Sciurumimusalbersdoerferiwas also purchased from a private collector. I don’t suspect forgery here either as this was in Germany, where fossil dealing is neither a big problem nor a lucrative business. The exceptional detail on the specimen would also require a substantial amount of theropod knowledge to pull off. Anyone having that amount of knowledge is more likely to be a real paleontologist than a get rich quick forger.
[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:
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.
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.
Once again the blog has taken a backseat to my real life work. It’s unfortunate too as there have been at least three really interesting news stories / technical papers that I feel the need to tackle. The first story I want to talk about is the news of the ancient Mediterranean goat: Myotragus balearicus, and its alleged “reptilian” physiology.
On the outset M.balearicus appears like your standard goat; complete with horns, hooves and (likely) a penchant for eating practically anything. The part that makes M.balearicus stick out the most? is that it was a native inhabitant of small islands in the Mediterranean.? Modern goats reach islands through human intervention. There, they become invasive elements that often damage the native flora and fauna. Without human intervention, it is hard for goats – and indeed? most mammals – to become established on islands. Both getting to the islands, and surviving on them tend to require animals that are more metabolically adaptable. Despite their catholic diets, goats are still limited by the “always on” nature of mammalian metabolism.
At least, that’s what we thought.
Researchers at the Institute of Paleontology at the Autonomous University of Barcelona, looked at microslices of the bones in this goat. What they found was a pattern of bone deposition that is unusual for ungulates. Rather than have layers of bone strewn about in an interwoven pattern, the bone of M.balearicus was laid down evenly in concentric layers. The latter formation is often assumed to be a hallmark of reptiles and other “slow growing” animals. With this in mind, the authors suggest that M.balearicus had evolved a more plastic metabolism.
These findings lend support to the model that posits a shift in life history strategies to a lower end of the growth rate spectrum, in areas where mortality remains low.
The results, while interesting, bother me a bit, as they rely on certain views on reptile growth strategies that are known to be false.
Ectotherm vertebrates have slow and flexible growth rates…
Ectotherms are characterized by lamellar-zonal bone throughout the cortex.
True zonal bone with growth marks deposited seasonally throughout ontogeny is a general ectotherm characteristic. In ectotherms, the bone matrix consists of slow growing lamellar bone.
While it is true that there are ectotherms that grow in a cyclical manner like this (especially animals from temperate regions), this is not a given for all ectotherms. In fact, it has since been well documented that fibrolamellar bone deposition occurs normally in crocodylians, as well as turtles (Reid, 1997).
It is a tad strange, as the authors do cite the Turmarkin-Deratzian gator paper, but they erroneously use it as an example of slow growth and contrast it with the fast fibrolamellar growth seen in most ungulates.? There is even a figure in the paper that shows, and even labels fibrolamellar growth in a crocodile, yet appears to get completely glossed over when it comes time to talk physiology.
Which brings me to my next point. The authors argue that the presence of lamellar zone bone in M.balearicus is suggestive of an ectotherm-like growth strategy. But does lamellar zone bone really indicate slow growth?
Work by Tomasz Owerkowicz on varanids (Owerkowicz 1997),? found that even the sedentary animals in his control group, could lay down bone at the same rate as his sedentary mammals (Morell 1996). Presumably this bone was lamellar zonal, though without the figures on hand, I can’t say for sure.
A more prominent example comes from Lieberman and Crompton (1998), who did a stress study on goats and opossums. The authors were looking at the remodeling response of bone to stress, and accidentally came across an interesting growth difference between these two taxa. They found that their opossums grew at a significantly faster rate than their goats, despite both taxa being of a developmentally equivalent stage. The interesting part is that the goat’s were depositing fibrolamellar bone, while the opossums were producing lamellar bone.
So no, lamellar bone need not be a hallmark of slow growth. Rather, it might be a response of the bones to specific stresses. Lamellar zonal bone is structurally stronger than fibrolamellar bone, so there might have been a more functional need for this type of bone.
Lastly, I have some issues with the final conclusions asserted by the authors in their closing comments:
The reptile-like physiological and life history traits found in Myotragus were certainly crucial to their survival on a small island for the amazing period of 5.2 million years, more than twice the average persistence of continental species. Therefore, we expect similar physiological and life history traits to be present in other large insular mammals such as dwarf elephants, hippos, and deer. However, precisely because of these traits (very tiny and immature neonates,low growth rate, decreased aerobic capacities, and reduced behavioral traits), Myotragus did not survive the arrival of a major predator, Homo sapiens, some 3,000 years ago.
Now I’m sure that there was a need to inject some melodrama at the end (as is typical for many papers), but the assertion that a “reptile-like physiological life history” must also incorporate a small aerobic scope, small neonates and reduced behavioural repertoire, is just uncalled for. All of these are frustratingly common misconceptions about reptiles, and bradymetabolic animals in general. Further, none of these assertions are based on any facts for M.balearicus. The only assertion that could really be tested is the small neonate one, and that appears to be falsified, as data on newborn M.balearicus show that newborns were large and precocial animals; pretty standard fare for an ungulate.
Overall the results of this study are interesting, and I look forward to seeing if pygmy elephants and hippos also display this apparent “slow growing” bone type. Comparing M.balearicus to reptiles based off this one similarity appears unjustified, and only goes to further perpetuate some common reptile misconceptions.
Needless to say, Myotragus balearicus was probably not “cold-blooded,” despite what the news headlines would have one believe.
Next up: Destroying the “uncaring parent” myth.
Lieberman, D.E., & Crompton, A.W. 1998. “Responses of Bone to Stress: Constraints on Symmorphosis.” Principles of Animals design: The Optimization and Symmorphosis Debate. Weibel, E.R., Taylor, R.C., and Bolis, L. (eds). Cambridge U. Press. Pgs: 78-86. ISBN: 0521586674
Morell, V. 1996. A Cold, Hard Look at Dinosaurs. Discover. December. Available online.
Owerkowicz, T. 1997. Effects of Exercise and Diet on Bone-Building: A Monitor Case. Journal of Morphology. V. 232(3): 306
Reid, R.? 1997. ?Dinosaurian Physiology: The Case for ?Intermediate? Dinosaurs.?? The Complete
Dinosaur. Farlow, J.O. and Brett-Surman, M.K. (eds.)? Indiana U. Press. Pgs: 449 – 473. ISBN: 0253333490
If one studies physical fitness (academically, or practically), then one is bound to come across the three main human body types. The endomorph, mesomorph and ectomorph.
Endomorphs are characterized by their ability to easily gain weight (be it fat, or muscle).
Ectomorphs are characterized by their ability to easily lose weight (fat, or muscle)
Mesomorphs are the middle ground group that appear to have the most malleable bodies.
In general, endomorphs have lower metabolisms than the other two, while ectomorphs tend to “run hot” all the time. Few people are all one way, or the other, but a notable dominance of one type, or another is usually prevalent.
The endo/ecto part can get confusing; especially if one is used to these prefixes in the context of endotherm/ectotherm. The names seem to be reversed from what one might normally hear (ectomorphs being more “warm-blooded” than endomorphs etc). The names have nothing to do with thermophysiology. They were coined after the germinative layers of the body during embryonic development. Endoderm forms the digestive tract, and endomorphs are usually stereotyped as fat. Ectotoderm forms the skin, and ectomorphs are usually stereotyped as being “all skin and bones.”
The reason I went with these specific bodybuilders (Jay Cutler, Arnold Schwarzenegger and Frank Zane) was partly to buck these stereotypes, but also to point out something that the news outlets are missing. Namely that having a lower metabolic state, does not mean one is a “couch potato” or has “forgone exercise.” Bigger, means more massive. That may mean fat, but as one can see above, it also can mean muscle and bone. Dinosaurs were not fatter than mammals. They were bigger.