• Tag Archives dinosaurs
  • The return of the scaly T. rex to modern paleo-art

    Tyrannosaurus rex walking towards camera. Art by John Sibbick.

    [NOTE: Post has been updated to include a section on scale size]

    This has certainly been an interesting year. Two papers dropped in the past three months that have put the brakes on a recent trend in paleo-art. That trend? Why the feather-coated T. rex of course.

    First, in March, we saw the release of a paper detailing a new species of Daspletosaurus and its relationship to D. torosus.

    Carr, T.D., Varricchio, D.J., Sedlmayr, J.C., Roberts, E.M., Moore, J.R. 2017. New Tyrannosaur with Evidence for Anagenesis and Crocodile-Like Facial Sensory System. Scientific Reports. 7(44942):1–11.

    In this paper, Carr et al. argue for the designation of a new Daspletosaurus species, D. horneri. The authors argue, based on skull shape and chronostratigraphic position, that D. horneri was the direct ancestor to D. torosus. I thought that the authors put forth a compelling argument for this anagenic event and backed up their position well. Interestingly, this part of the paper should have been the most controversial. As anyone who has read anything from Horner and Scanella over the past eight years can attest, arguing for a direct ancestor-descendant relationship for dinosaurs is difficult to do and even harder to win over others in the field. So it is somewhat surprising to see a case for anagenesis in Daspletosaurus taken so well by the palontological community. All the more so given that it involves a tyrannosaur, the poster children for “cool guy” dinosaurs.

    Instead, the most controversial part of the paper wound up being their soft-tissue reconstruction of the face for D. horneri. The author responsible for the soft-tissue reconstruction was Jayc Sedlmayr of Louisiana State University. Sedlmayr did his doctorate on osteological correlates for vasculature in extant archosaurs (birds & crocs). He is the seminal alumnus of the WitmerLab and thus is well within his wheelhouse for this type of soft-tissue reconstruction. Sedlmayr borrowed heavily from the work of another WitmerLab alumnus, Tobin Hieronymus, whose PhD work involved osteological correlates for integument on the skulls of animals. Although the skin is often well away from the underlying bones on most of the body, there are exceptions when it comes to the skull. There, areas that are not heavily muscled, tend to show intimate connections between the skin and the underlying bone. Hieronymus used these connections to determine how different integumentary appendages (scales, hair, feathers) affect the underlying bone (Hieronymus & Witmer 2007; Hieronymus et al. 2009). The authors found that the surface texture along the skull of D. horneri was “hummocky”. That is, it was covered in lots of closely packed ridges. According to Hieronymus & Witmer (2007), this texture correlates to scales as the overlying integumentary appendage. Thus, according to the authors, D. horneri had a scaly face (this is grossly oversimplified as the authors were able to piece together a variety of different integument variants along the skull, but you get the idea).

    Scaly tyrannosaur cannonball one had been shot.

    Then two weeks ago, we saw the release of another paper on tyrannosaur integument. However, unlike the previous paper, this one was specifically dedicated to integumentary types in tyrannosaurids.

    Bell, P.R., Campione, N.E., Persons, W.S., Currie, P.J., Larson, P.L., Tanke, D.H., Bakker, R.T. 2017. Tyrannosauroid Integument Reveals Conflicting Patterns of Gigantism and Feather Evolution. Biology Letters. 13:20170092

    In this paper, the authors set out to survey all known instances of “skin” impressions for tyrannosaurids. Their list of taxa included Albertosaurus, Tarbosaurus, Daspletosaurus, and Gorgosaurus. Their results pretty definitively indicated that scales were the predominant integumentary appendage on tyrannosaurids. The authors then went on to speculate why that would be if earlier tyrannosauroids had filamentous integument. They performed an ancestral character state reconstruction based on Parsimony and Bayesian-based trees from Brussatte and Carr 2016. Their results found that filaments came out strongly as the ancestral character for tyrannosauroids, but by no later than Tyrannosauridae proper, a reversion to scales had taken effect. The authors attributed this to body size evolution. Namely, larger tyrannosauroids reverted to scales over protofeathers.

    Cannonball number 2 had just been shot.

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  • New study shreds the dinosaur family tree (and exposes double-standards in Phylogenetic Nomenclature)

    Figurative illustration of the new phylogeny by Baron et al. 2017

    Most folks who visit my site by now have seen the big dinosaur news that has hit the interwebs. A new study from Matthew Baron, David Norman and Paul Barrett from University of Cambridge and the Natural History Museum of London, has seriously challenged the classic interpretation of dinosaur phylogeny.

    Baron, M.G., Norman, D.B., Barrett, P.M. 2017. A New Hypothesis of Dinosaur Relationships and Early Dinosaur Evolution. Nature. 543:501–512.

    Classical dinosaur phylogenetics

    Although originally thought of as two unrelated branches of Reptilia that grew to immense size during the Mesozoic (e.g., Charig et al. 1965), for the last 43 years the group, Dinosauria, has been considered monophyletic (i.e., sharing a single origin) with the subgroups, Saurischia & Ornithischia, forming the first major branches within the group (Bakker et al. 1974). Saurischians, or “reptile hips” were aligned together by their similar hip shapes, skull characters (e.g., open antorbital fenestrae), and inferred soft tissues (e.g., air sacs). Ornithischians, or “bird hips” shared a hip structure that was superficially similar to that of birds, with a pubis that pointed caudally rather than rostrally, along with a variety of unique skull characters such as a neomorphic bone known as the predentary.

    Study after study showed that this relationship was sound, and so it stayed that way. The problem with getting the same answer over and over again is that one eventually stops questioning it. Consistent results become  common knowledge, and may even graduate to dogma. That’s not so bad if that common knowledge is true, but all too often many of these “obvious” cases wind up being just so stories upon closer inspection.

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  • Modern-day paleo myths: Dinosaurs as lizards


    In this day and age there are no shortage of books, websites, and videos dedicated to debunking classic paleo myths. The majority of this mythbusting focuses on myths about dinosaurs. As the poster children for paleontology, this isn’t that surprising. With so many takes on this subject it comes as no surprise that all of the classic dinosaur myths have long since been debunked, such as dinosaurs as low-energy tail draggers, walking around like Godzilla, being evolutionary failures, inferiority to mammals, being pee brained monsters, etc.

    However, as quickly as these classic dinosaur myths have been eradicated, new ones have come and taken their place. These myths/misconceptions are routinely cited today without any question despite being just as erroneous as the myths that preceded them.

    This is the start of a new series I want to cover on the site: dispelling modern myths in vertebrate paleontology. Given the bent of my website, these myths/misconceptions will largely stay focused on reptile-related animals, though I am open to taking the occasional foray into other animal groups if the myths are egregious enough (which is to say that suggestions are welcomed).

    The seminal installment for this series is one that I see mentioned time and again:

    “Dinosaurs were once thought of as big lizards.”

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  • Jurassic World and the case of the droopy-tailed Stegosaurus

    As I write this the US premiere of Jurassic World is just around the corner. I had gone back and forth regarding this post given that we currently know very little about the film and as such the interpretations written about here and elsewhere may well be pointless by the time the film premieres.

    Ultimately I decided to post this anyway since the overall thrust of the article should remain true regardless of how the film pans out.

    Now there has been a lot of buzz around Jurassic World since it was first announced last year. The buzz has been mixed, but fairly positive. I suspect this was, in part, because everyone was happy to hear that the godawful military dinosaur idea was shelved in favour of a more “traditional” JP franchise storyline. Nonetheless the movie has still drawn its fair share of detractors, including myself. Most of the people who are unhappy with the film are either paleontologists, or hardcore dinophiles. Many of the problems leveled at the film have to do with the portrayal of the extinct animals. The problems are actually myriad ranging from pterosaurs capable of picking up humans using grasping feet, mosasaurs that are twice the size of blue whales, sauropods covered in elephant skin rather than scales (a problem not unique to Jurassic World), everything about Velociraptor, and of course Indominus rex.  My biggest beef with the film is that the dinosaurs are not being shown as dinosaurs so much as monsters. However, after The Lost World: Jurassic Park came out it became pretty evident that Spielberg’s original vision of portraying dinosaurs as animals had been shelved in favour of the more entertainment-friendly movie monster approach. However, for what seems like a majority of the detractors, the biggest gripe with the film has to do with a lack of  feathers on pretty much all the dinosaurs. This seems to be a common theme these days with a particularly vocal group of dinophiles and paleontologists strongly pushing for the feathering of every dinosaur in sight and insisting that all media that portrays scaly (erroneously called: “naked”) dinosaurs is inaccurate. Never mind the fact that a feathered, pack-hunting, 2 meter tall Velociraptor mongoliensis is still every bit as inaccurate as a scaly one.

    Anyway, I digress. Dealing with the overwhelming amount of internet drama surrounding Jurassic World (and the media depiction of dinosaurs in general) is a topic for another day. My reason for writing this post is centered around one particular criticism that popped up a few weeks ago.

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  • “Feathers” on the big, “feathers” on the small, but “feathers” for dinosaurs one and all?


    Yutyrannus artwork by Brian Choo. Sciurumimus artwork by Arkady Rose

    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 theropod 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 guy—dubbed: Sciurumimus albersdoerferi (Albersdörfer’s squirrel mimic)—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, Sciurumimus albersdoerferi was 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.

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  • Get in on the deal: Indiana University Press one day sale.

    I apologize ahead of time for what will likely sound like spam, but:

    Just a quick post to remind folks that today, and only today, Indiana University Press is offering a 60% off sale on all their books. That includes their famed Life of the Past series.

    So if you have yet to get your copy of The Complete Dinosaur, or have been itching to snag the most comprehensive book ever written on Deinosuchus, ankylosaurs, or mosasaurs, but didn’t have the necessary funds; now is your chance to get them for cheap.

    Just remember, the sale ends today.


  • New study shows that gators are one-way breathers too.

    I would be remiss not to talk about this amazing discovery published last week in Science.

    Farmer,C.G. & Sanders,K. 2010. Unidirectional Airflow in the Lungs of Alligators. Science. vol.327:338-340

    The anatomical similarities of alligators and birds has been known for quite some time (at least 100 years), and this anatomical similarity extends down into the lungs. Though alligators lack the pneumatic carvings of the post-cranial skeleton (air sacs) that are seen in birds, saurischian dinosaurs and pterosaurs; their lungs and bronchi do share the same structural features.

    Birds have a unique lung design that allows air to pass through it in a single direction. Unlike mammals, there is no “dead end” to the avian lung. This provides the benefit of a constant supply of highly oxygenated air to the lung tissue; which allows for more efficient gas exchange. Up until last week, this lung design was thought to be a hallmark of birds, and possibly saurischian dinosaurs, and pterosaurs.

    Well it turns out that this unique avian synapomorphy is a heck of a lot older than we thought.

    Dr. Colleen Farmer, and Kent Sanders M.D. of the University of Utah, considered the uncanny anatomical similarities of the avian and crocodylian lung, and wondered if these similarities extended to the physiology too. In other words: If it looks like a unidirectional lung, does it also function like one?

    Farmer & Sanders set to work by removing the lungs of four dead alligators donated to her lab. They pumped air through them, and monitoring the direction in which it traveled (using flowmeters). They then surgically inserted flowmeters into anesthetized alligators, and measured the airflow direction in living animals. Lastly, to drive the point across completely, they filled up an excised lung with fluid that contained fluorescent beads, and proceeded to pump the water in and out. This last test was recorded, and three movies of it, were made available to the public. They can be viewed here. Three was probably overkill though, as once you’ve seen fluorescent beads move one way in a gator lung, you’ve seen them all. : )

    The results showed conclusively that alligator lungs pump air through them in one direction only. The repercussions of this find are actually pretty enormous. For starters, the similarity in anatomy and physiology of avian and crocodylian lungs, suggests that they are homologous. This would mean that both groups inherited these lungs from a common ancestor. This means that it was highly likely that all dinosaurs, pterosaurs, rauisuchians, aetosaurs, phytosaurs and the myriad of other archosaurs that graced this planet some 200 million years ago, housed this particular flow-through style lung.

    It also helps put to rest arguments about air sac functions. It has long been argued that the presence of a unidirectional lung, necessitates the presence of air sacs to “pump the air in.” (air sacs offer zero, or next to zero gas exchange potential, so there is no actual breathing going on in them). A lack of air sacs in ornithischian dinosaurs, has been used to suggest that their pulmonary physiology was more like mammals and lizards, than it was like birds (Ruben et al 1999). Data from previous research (O’Connor & Claessens 2005) has cautioned that the presence of air sacs does not guarantee the existence of a flow through system. These latest data now show us that a flow-through system can, and likely did, evolve without the “need” for an air sac pump.

    CT scan of alligator, with 3D reconstruction of lungs. For more details on what the colours mean, click the picture.

    Exactly how all of this works, is still not understood. The “hepatic piston” diaphragmatic pump of crocodylians is well known, and is likely the ultimate driver of respiration in these animals, but the nuts & bolts of how all this unidirectional flow takes place (the fluid dynamics of the lung) remains a mystery. One question that would be worthy of a follow up study (which the author’s have hinted at doing) is whether, or not a cross-current, or counter-current system (where deoxygenated blood flows perpendicular, or opposite the direction of highly oxygenated air) is present in crocodylians too. A cross-current system is found in birds. Is that unique to them, or was this also a phylogenetic “hand-me-down?” Hopefully now, with this new discovery, future research will be done on the crocodylian lung, to further understand how it actually works.

    Ultimately that is the biggest piece of news to come out of this paper. For well over 100 years, the crocodylian lung was just assumed to be a “dead-end” space that worked in a manner similar to that of mammals. It wasn’t until someone actually thought “what do we really know about this structure” did we find something quite the opposite taking place. This is hardly the first time that this has happened either (for instance). As I have mentioned (ranted/harped on) before, reptiles tend to get the short end of the stick when it comes to a lot of biological and paleontological studies (especially if they involve comparison between broad animal groups [classes]). I’m always amazed (though rarely surprised) when a study that actually looks into commonly held assumptions about these critters, finds said assumptions to be quite off the mark. Here’s hoping that we continue to see future studies like this, go on.

    In the end, all of this brings us closer to the truth about how life really works; which is why we do all of this stuff in the first place.



    Farmer,C.G. & Sanders,K. 2010. Unidirectional Airflow in the Lungs of Alligators. Science. vol.327:338-340

    O’Connor, P.M.& Claessens, A.M. 2005. Basic Avian Pulmonary Design and flow-Through Ventilation in Non-Avian Theropod Dinosaurs. Nature. Vol. 436:253-256.

    Ruben, J.A., Dal Sasso, C., Geist, N.R., Hillenius, W.J., Jones, T.D. 1999. Pulmonary Function and Metabolic Physiology of Theropod Dinosaurs. Science. Vol.283(5401):514-516.

  • 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.

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  • New paper says dinosaurs were endomorphs.

    From left to right: Endomorphic Jay Cutler, Mesomorphic Arnold Schwarzenegger, and Ectomorphic poster-child Frank Zane
    From left to right: Endomorphic Jay Cutler, Mesomorphic Arnold Schwarzenegger and Ectomorphic poster-child Frank Zane

    Endo-what now? Allow me to explain.

    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.

    So what am I rambling on about?

    Grab a calculator and come along for the ride.

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  • Sprawling crocodylians walk straight even if there isn’t much O2 to go around.

    Photo of estuarine crocodile by: D. Parer and E. Parer-Cook
    Photo of estuarine crocodile by: D. Parer and E. Parer-Cook

    Two new papers have recently hit the journal circuit. Both of them involve using living crocodylians to gain a better understanding of paleo-life.

    The first one comes from Denver Museum of Natural History paleontologist, Dr. Kenneth Carpenter:

    Carpenter, K. 2009. Role of Lateral Body Bending in Crocodylian Track Making. Ichnos. Vol.16:202-207. doi:10.1080/10420940802686137.

    The study used an adult Caiman sclerops (first use of a large adult reptile for a locomotion study; at least as far as I know) placed in a small room with two 30cm walls placed on either side of it. This restricted any lateral movement, and “funneled” the animal out the singular opening. At this opening, a camera was placed. It would photograph the animal as it left the room. The room itself, had a smoothed mud covering. This muddy floor would record the tracks of the C.sclerops as it walked by.? Several runs were done, and photographs were taken for each run.

    This is the first study I have seen that gave a front view shot of an adult crocodylian as it walked along. As Carpenter mentioned in the paper:

    This front view is in contrast to most photographic studies which only capture pro?le and top views….

    Carpenter also mentioned the potential of there being an ontogenetic change in limb stance as animals move from hatchling to adult. This is something that I have hinted at previously Hatchling crocodylians seem to have weaker femoral adductors than adults. This is understandable given the greater weight that adult femora need to bear. This can result in a skewed view of crocodylian erect stance; with most authors tending to underestimate the degree of “parasagittality.”

    That said, I was surprised to read that Carpenter had found the adult Caiman sclerops to have a hip adduction angle of approximately 65? from the horizontal. Judging from figure1B, the hindlimb appears to be much closer to the midline than the forelimb. Fig1D seems even closer to, if not 90?. It is important to point out that much of the hindlimb is blocked by the body in this shot, as the animal is fully laterally extended. A concurrent shot from behind would have been very useful here; as would an x-ray series of shots throughout the walk phase (for instance: see this long video of a Crocodylus acutus walk cycle. Pay special attention to the position of the femur).

    Alas, that is not what the paper is about.

    The paper is about how lateral movements during locomotion, have substantial effect on trackways. Dr. Carpenter points out how, despite the semi-erect stance of the forelimbs, the track evidence would suggest an animal with a much narrower (parasagittal?) stance. This has bearing on how prehistoric reptiles, in particular: quadrupedal dinosaurs, may have stood.

    One might rightfully ask if we should expect dinosaurs to have had any lateral movement to their walking cycle at all. Carpenter points out that lateral body bending, though not quite as exaggerated as that of crocs, is present in most tetrapods. Birds seem to be the sole exception, with their extremely stiff thorax. However birds are also obligate bipeds, and the avian thorax is much shorter and stiffer than that of dinosaurs.

    So it would seem to be a likely bet that quadrupedal dinosaurs likely exhibited some degree of lateral body bending.

    Triceratops pic from britannica.com, but originally from: Mounted Skeleton of Triceratops elatus? by Henry Fairfield Osborn, American Museum Novitiates, Sept. 6, 1933
    Triceratops pic from britannica.com, but originally from: Mounted Skeleton of Triceratops elatus? by Henry Fairfield Osborn, American Museum Novitiates, Sept. 6, 1933

    Carpenter’s work rightfully asks us to caution reconstructions of stance based largely off of trackway evidence. A fine case study that the paper brings up, is ceratopians. This group, more than any other, has received considerable attention for how the forelimbs were oriented. Early work on ceratopians, favoured a hefty sprawl to the forelimbs (e.g.? Gilmore 1905, or Lull 1933). This was critically evaluated during the heyday of the dinosaur renaissance. Authors such as Bakker (1986), Paul and Christiansen (2000), instead favoured a fully erect stance. A large portion of the data supporting this assertion, was trackway based. The results of this study call into question that view. However this was not the first paper to have done so. Thompson and Holmes (2007) also questioned the “erect ceratopid” view, using a half scale model of a Chasmosaurus irvinensis forelimb. Their results come closer to the results from this paper. Though Thompson and Holmes felt that there was no real modern analogue to ceratopian forelimb mechanics.

    In the end, Dr. Carpenter reminds future researchers of the importance in incorporating the entire animal when analyzing trackways.

    The second paper comes from the Journal of Experimental Biology.

    Owerkowicz, T., elsey, R.M. and Hicks, J.W. 2009. Atmopsheric Oxygen Level Affects Growth Trajectory, Cardiopulmonary Allometery and Metabolic Rate in the American Alligator (Alligator mississippiensis). J.Exp.Biol. Vol.212:1237-1247. doi:10.1242jeb.023945.

    The authors embarked on a study of how previous paleo-atmospheric oxygen levels might have affected the lives of animals that would have been alive through these times. According to Owerkowicz et al, crocodylians were chosen because:

    Given their phylogenetic position and highly conserved morphology throughout their evolutionary history, crocodilians are often thought to retain many characteristics of basal archosaurs.

    I do take some issue with this, as prior reviews on crocodylomorph diversity (Naish 2001) coupled with many new discoveries ( Buckley et al 2000,? Clark et al 2004, Nobre & Carvalho 2006)? continually cast doubt on the old view that crocodylians have survived “unchanged” for some 200 million years. Nevertheless, the results of the study are both interesting, and relevant to reconstructions of how paleo-life would have adapted to these wildly different paleo-atmospheres.

    Owerkowicz et al raised groups of hatchling American alligators (Alligator mississippiensis) under three different atmospheric conditions. A hypoxic (12% O2) condition reminiscent of paleo-atmospheric models for the late Triassic/Early Jurassic periods. Current atmospheric conditions (21% O2), and a hyperoxic (30% O2) condition reminiscent of paleo-atmospheric models for the Carboniferous and Permian periods.

    The results were interesting, though not too surprising. As expected, hypoxic alligator hatchlings were smaller than their normal and hyperoxic counterparts. However, the degree of growth stunting is pretty surprising. Hypoxic hatchlings were about 12% shorter and 17% smaller than normal hatchlings.

    Baby alligators pic from REPTILES mag. December 94. Author unknown.
    Baby alligators pic from REPTILES mag. December 94. Author unknown.

    Surprisingly, hatching time did not change under any conditions. This suggests a degree of “hard wired” embryological development inside the egg. In the case of the hypoxic hatchlings, they came out “almost done.” While all three groups had remnants of a yolk sac upon hatching, the hypoxic hatchlings actually had the yolk sac still protruding (normal and hyperoxic hatchlings just showed distended bellies). In some cases, the yolk sac was larger around than the hind legs, thus making movement clumsy and cumbersome.

    Other interesting results from this study, included notable changes to the cardiopulmonary system. Hypoxic hatchling lungs were actually smaller than the lungs of normal hatchlings; which appears counterintuitive. The heart, meanwhile, showed distinct hypertrophy in hypoxic animals. The authors believe that lack of lung growth in hatchlings may have been due to the fact that lung function does not start until after hatchlings have hatched.? The heart, on the other hand, is hard at work circulating blood just as soon as it is formed; so it would have experienced the challenges of hypoxia at a very early stage.? Bolstering this hypothesis from the authors was the fact that three months after hatching, hypoxic alligators showed a distinct increase in lung growth rate (the lungs appeared to be “catching up” to the heart).? Hypoxic alligators showed shrunk livers as well. No real explanation for this was given, but it was mentioned that reduced liver mass seems to be a common trait in animals raised in hypoxic conditions. It appears to have some bearing on overall metabolic rate.

    Hyperoxic hatchlings exhibited “typical” organ growth rates.? Where hyperoxic animals excelled was in breathing and metabolic rate.

    Breathing rates were smaller in this group, while metabolism and growth rate were all larger. The explanation by the authors was that these hyperoxic animals were receiving such high amounts of oxygen in each breath, that they were actually hitting saturation at much shallower breaths; hence the shallow breathing. The higher metabolic rate is believed? due to a lack of right-left shunting in the crocodylian heart. This shunting is usually caused by low oxygen levels (like that experienced in diving), and tends to result in metabolic depression to conserve available oxygen stores.? Since these alligators lungs were constantly saturated with oxygen, right-left shunting never occurred, resulting in an elevated metabolism.

    Incidentally, Owerkowicz et el give mention of a cardiac shunt known in embryological birds (via the ductus arteriosis). Though only analogous, one can’t help but wonder what this might have meant for all those dinosaurs that lie between these two groups.

    Interestingly, hypoxic alligator hatchlings also showed a higher standard metabolic rate. Though these animals would voluntarily eat less than their normal and hyperoxic counterparts, their metabolism was more like hyperoxic hatchlings than they were normal hatchlings.? Owerkowicz et al believe the reason for the increased metabolism was due to the higher cost of breathing in these animals. Despite taking “normal” breaths, hypoxic hatchlings were taking in a larger tidal volume than their normal and hyperoxic siblings. The heart was also working harder to deliver enough oxygen to tissues.

    Finally the authors give mention of growth rates in hyperoxic animals. Basically, it is faster. The authors mention that this might be caused by the persistently elevated metabolic rate, or perhaps from channeling saved energy from breathing (which is one of the main energetic costs in reptiles) into biomass.? It could be a mix of both, but I’m more inclined to think that it comes more from channeling energy reserves into other parts of the body. A high metabolism means nothing, if there is not enough free energy to go around. Just look at the hypoxic gators from this study. Despite their high metabolism, they grew slower than their peers.

    The results of this study showed how modern animals can acclimate to different atmospheric conditions. They don’t show how animals would adapt and evolve in these conditions, but they do hint at the general directions, and help give us a clearer picture of what life was like millions of years ago.



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    Clark.J.M., Xu, X., Forster, C.A., Wang, Y. 2004. A Middle Jurassic ‘Sphenosuchian’ from china and the Origin fo the Crocodylian Skull. Nature. Vol.430:1021-1024.

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