• Category Archives Extant Reptiles
  • Articles about extant, or living reptiles

  • 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.


  • 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 563

  • New study suggests that group nesting should be the norm – not the exception – in reptiles.

    A colony of _Eumeces fasciatus_ brood their eggs.
    A colony of Eumeces fasciatus brood their eggs.

    Continuing my trend of “catching up,” an article in the November issue of Natural History magazine, talks about a new study in the Quarterly Review of Biology, that finds group nesting to be very common place among extant reptiles.

    That study would be:

    Doody, J.S., Freedberg, S., Keogh, J.S.? 2009. Communal Egg-Laying in Reptiles and Amphibians: Evolutionary Patterns and Hypotheses. Quart. Rev. Biol. Vol.84(3):229-252.

    In the paper, Doody et al (no laughing) did a massive search through the herpetological literature (both technical journals, and hobbyist magazines) to look at instances of communal egg laying in reptiles and amphibians (herps). I’m not being hyperbolic here either, as the paper states:

    In total, our assembled database was gathered from 290 different sources, including 176 different scientific journals, 72 books or book chapters, 29 unpublished reports, and 13 unpublished theses. We also have included a number of reliable personal communications from herpetologists.

    What the authors found was that group gatherings of herps are vastly more common than previously believed. Group egg laying was found to be present in 345 reptile species. Now you might be thinking 345 really isn’t all that much for a group composed of some 8700 species.

    Well then aren’t you a Debbie Downer?

    _Ophisaurus attentuatus_ brooding her eggs.
    Ophisaurus attentuatus brooding her eggs.

    Seriously though, the authors address this by mentioning:

    Although the difficulty in locating nests hampers our ability to determine the actual frequency of communal egg-laying among species, we can better estimate this proportion by dividing the number of known communally egg-laying species by the total number of species, excluding those for which eggs have not been found. We conducted such a calculation for the three families of Australian lizards known to include multiple communally egg-laying species—Gekkonidae, Pygopodidae, and Scincidae—as gleaned from the Encyclopedia of Australian Reptiles database (Greer 2004). Proportions of these lizard families known to lay communally were 4–9%, but, when we exclude species for which nests are not known, these values rise dramatically to 73–100%

    The biggest take home message to get from Doody et al’s review, is just how much we don’t know about extant reptiles.

    …the present review highlights our inadequate knowledge of the nests and/or eggs of reptiles. For instance, the eggs or nests are known in only 7% of Australian lizards of the three families that commonly lay communally (N = 411 oviparous spp.) (Greer 2004).The extent of this knowledge for Australian lizards is probably similar to that for reptile eggs on other continents, particularly South America, Africa, and Asia, where the reproductive habits of reptiles are poorly known. This is in stark contrast to other vertebrates such as birds, for which complete field guides to the eggs and nests are available for several continents

    Indeed, just by doing the brief research run needed to compile this blog post, it was apparent that communalism is much more common in reptiles than anyone ever thought. However, because so many of these reports are either anecdotal, or buried in obscure journals, it is easy to miss all the many cases where it is known.

    This discovery lead the authors to the inevitable follow up question of: “why?” What benefit do mothers gain by nesting communally?

    Numerous hypotheses for why animals nest communally, have been proposed.

    • Saturated habitat (only so many suitable nest sites)
    • Sexual selection (choice of males that live in a particular area)
    • Artifact of grouping for other reasons
    • Attack abatement (easier to hide a bunch of eggs in one site, than in multiple sites. Less chance that your eggs will be the ones that are eaten).
    • Maternal Benefits (save time and energy finding a suitable nest site by “freeloading”)
    • Reproductive success (if the nest site worked once before…)
    • Egg insulation

    The authors reviewed all of these possible reasons for communal egg laying in herps. In the end, they found evidence for both the maternal benefits hypothesis, and the reproductive success hypothesis, though they felt a mixed model better explained things.

    Python brooding her eggs.
    Python brooding her eggs.

    Sadly, though the authors mentioned how a lack of information on the natural history of most reptiles is largely responsible for this sudden revelation about their nesting behavior, they nevertheless make repeated mentions of how “social interactions are generally less complex in reptiles and amphibians than in other tetrapods” or how herp sociality forms “relatively simple systems“.

    The reality is that the old view of simplistic “loner” reptiles that only come together to mate, is not accurate. This is especially true for parental care in reptiles.

    The popular view (among the public, and the scientific community) is that reptiles are? “lay’em and leave’em” types when it comes to reproduction. Despite all the herpetological knowledge to the contrary that has been acquired in the past 50 years, it is still popular to spout the party line about reptiles being “uncaring parents.”

    Zoologist Louis Somma took issue with this view of reptilian (in particular, chelonian and lepidosaurian) parenting. He conducted a literature search to see how often mentions of parental care in reptiles are recorded. In the end he wound up finding 1400 references to parental care in reptiles (Somma 2003)!

    Somma’s survey covered various aspects of parental care. He found reported evidence of nest building and / or guarding in tortoises like Manouria emys (McKeown 1999), Gopherus agassizii (Barrett & Humphreys 1986) and 4 other species of chelonian.

    Turning to lepidosaurs, Somma found parental behaviour to be present in 133 species of lizards and 102 species of snakes. Even a species of tuatara (Sphenodon punctatus) is known to guard its nests (Refsnider et al. 2009). Though these numbers appear small compared to the total amount of species that have been described; much like the Doody et al. paper, this is just based off of species whose nesting behaviours we do know. That these taxa all span a wide phylogenetic range, suggests that parental care is more commonplace than initially thought.

    Nest guarding is usually a maternal trait, but some squamates exhibit nest guarding behaviour in both parents, such as some cobra and crotaline snakes (Manthey and Grossman 1997) , as well as tokay geckos (Zaworski 1987).

    Not only active guarding of the nest, but actual brooding of the eggs is also commonly reported in squamates such as various python species (Harlow & Grigg 1984, Lourdais et al. 2007), and skinks (Hasegawa 1985, Somma & Fawcett 1989). Some species are even known to groom their newly hatched young (Somma 1987).

    More interesting still are various reports and observations of parental feeding in some reptile species, such as the skink Eumeces obsoletus (Evans 1959), and the cordylid lizard Cordylus cataphractus (Branch 1998). Not to mention recent evidence of parental feeding in captive crocodylians.

    This leads me to the only reptile group where parental care is well publicized: that of the 23 extant crocodylian species. I could, at this point, list references for parental care in crocodylians. However because this behaviour is so well documented for this group, it would seem unnecessary. It is? better to shed light on the many (MANY) examples of parental care in other reptile species. I also didn’t include related examples like placental evolution in the skink genus Mabuya, or instances of egg binding in captive reptile mothers; due to a lack of appropriate substrate to lay their eggs.

    In the end, the paper by Doody et al. adds to a growing body of evidence which suggests that the “lay’em and leave’em” reptile species of the world, are the exceptions? and not the rule.

    ~ Jura

    Next time: Biomechanics of running suggest “warm-blooded” dinosaurs. Or: why the aerobic capacity model needs to die already.


    Barrett, S.L. & Humphrey, J.A. 1986. Agonistic Interactions Between Gopherus agassizii (Testudinidae)
    and Heloderma suspectum (Helodermatidae). Southwestern Naturalist, 31: 261-263.
    Branch, B.. 1998. Field Guide to Snakes and Other Reptiles of Southern Africa. Third revised edition. Sanibel Island: Ralph Curtis Books Publishing.
    Doody, J.S., Freedberg, S., Keogh, J.S.? 2009. Communal Egg-Laying in Reptiles and Amphibians: Evolutionary Patterns and Hypotheses. Quart. Rev. Biol. Vol.84(3):229-252.
    Evans, L.T. 1959. A Motion Picture Study of Maternal Behavior of the Lizard, Eumeces obsoletus Baird and Girard. Copeia, 1959: 103-110.
    Harlow, P and Grigg, G. 1984. Shivering Thermogenesis in a Brooding Python, Python spilotes spilotes. Copeia. Vol.4:959?965.
    Hasegawa, M. 1985. Effect of Brooding on Egg Mortality in the Lizard Eumeces okadae on Miyake-jima, Izu Islands, Japan. Copeia, 1985: 497-500.
    Lourdais, O., Hoffman, T.C.M., DeNardo, D.F. 2007. Maternal Brooding in the Children’s Python (Antaresia childreni) Promotes Egg Water Balance. J. Comp. Physiol. B. Vol.177:560-577.
    Manthey, U. and W. Grossman. 1997. Amphibein & Reptilien S?dostasiens. Natur und Tier Verlag, M?nster.
    Mckeown, S. 1999. Nest Mounding and Egg Guarding of the Asian Forest Tortoise (Manouria emys). Reptiles, 7(9): 70-83.
    Refsnider, J.M., Keall, S.N., Daugherty, C.H., & Nelson, N.J. 2009. Does nest-guarding in Female Tuatara (Sphenodon punctatus) Reduce Nest Destruction by Conspecific Females? Journal of Herpetology. vol.43(2):294-299.
    Somma, L.A. 1987. Maternal Care of Neonates in the Prairie Skink, Eumeces septentrionalis. Great Basin Naturalist, 47: 536-537.
    Somma, L.A. & Fawcett, J.D. 1989. Brooding Behaviour of the Prairie Skink, Eumeces septentrionalis, and its Relationship to the Hydric Environment of the Nest. Zoological Journal of the Linnean Society. Vol.95: 245-256.
    Somma, L. 2003. parental Behavior in Lepidosaurian and Testudinian Reptiles: A Literature Survey. Krieger Publishing Company. 174pgs. ISBN: 157524201X
    Zaworksi, J.P. 1987. Egg Guarding Behavior by Male Gekko gecko. Bulletin of the Chicago Herpetological Society, 22: 193.
    <object width=”425″ height=”344″><param name=”movie” value=”http://www.youtube.com/v/JiZAtn8fmtI&hl=en_US&fs=1&”></param><param name=”allowFullScreen” value=”true”></param><param name=”allowscriptaccess” value=”always”></param><embed src=”http://www.youtube.com/v/JiZAtn8fmtI&hl=en_US&fs=1&” type=”application/x-shockwave-flash” allowscriptaccess=”always” allowfullscreen=”true” width=”425″ height=”344″></embed></object>

  • Bone histology says ectothermic goats, but does it?

    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.

    Myotragus balearicus reconstruction.
    Myotragus balearicus reconstruction.

    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.

    For instance:

    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

  • Land lubbing crocs get their day in the sun. Also, there’s a varanid special on NOVA.

    Dr. Paul Sereno stands with others at a meeting for the American Association for the Advancement of Science in Chicago. Note the wheelbarrow like retroarticular processes on the "boar croc."
    Dr. Paul Sereno stands with others at a meeting for the American Association for the Advancement of Science in Chicago. Note the wheelbarrow like retroarticular processes on the "boar croc."

    After spending? a few years collecting and looking at the weirdness that is Gondwanan crocodyliformes, Dr. Paul Sereno has finally started to unveil stuff. With the help of National Geographic comes When Crocs Ate Dinosaurs. It appears to be a special that focuses on the remarkable – and often underrated – diversity seen within this group of animals. The highlight of the program (at least in my opinion) is the focus on all the very un-crocodile like crocodyliformes.

    The National Geographic website has a special section that shows off the various, apparently unnamed, taxa. For now, there are just placeholder names that will likely hurt the eyes and ears of anyone who had to deal with the aftermath of The Land Before Time.

    The artwork is by artist Todd Marshall. I’ve always enjoyed his portrayals of prehistoric reptiles (he tends to get almost too fanciful with dewlaps and spikes though). Sadly the accompanying animations do not do Marshall’s incredible artwork justice.? It will be interesting to see how it all gets integrated into the television show.

    Also airing tonight is a special on NOVA entitled: Lizard Kings. It features the work of Dr. Eric Pianka; a well known and respected lizard ecologist who has focused on monitors for much of his career.? The special looks to be very interesting. Especially given that it appears to have taken years for the film crew to get the footage they needed. As you read this the special has already aired. However, PBS does make their shows avaialable to watch online for free, on their website. The show should also be viewable on Hulu by tomorrow.

    A perentie monitor (_Varanus giganteus_) poses for the camera.
    A perentie monitor (_Varanus giganteus_) poses for the camera.

    I realize that both of these options are only available in the states. To date there seems to be no international options. At best there are some workarounds.

    Still, for those that can get them, both shows should prove to be entertaining.


  • New paper dispells Komodo myth. Also Megalania may have been the world’s largest venomous animal.

    Megalania chasing down Genyornis newtoni. Illustration by Peter Trusler for Wildlife of Gondwana
    Megalania chasing down Genyornis newtoni. Illustration by Peter Trusler for Wildlife of Gondwana

    Just announced today in the journal: PNAS, is a new comprehensive study on Komodo dragon feeding ecology. The comprehensive nature of the paper is the result of the contributions from around 28 individuals from all over Australia, as well as the Netherlands, and Switzerland.

    The paper in question is:

    Fry, B., Wroe, S., Teeuwissed, W., van Osch, M.J.P., Moreno, K., Ingle, J., McHenry, C., Ferrara, T., Clausen, P., Scheib, H., Winter, K.L., Greisman, L., Roelants, K., van der Weerd, L., Clemente, C.J., Giannakis, E., Hodgson, W.C., Luz, S., Martelli, P., Krishnasamy, K., Kochva, E., Kwok, H.F., Scanlon, D., Karas, J., Citron, D.M., Goldstein, E.J.C., Mcnaughtan, J.E., and Norman, J. 2009. A Central role for Venom in Predation by Varanus komodoensis (Komodo Dragon) and the Extinct Giant Varanus (Megalania) prisca. PNAS Early Release. doi:10.1073/pnas.0810883106

    *catches breath*

    The paper is only six pages long, which downplays just how much work must have gone into this project. The authors used Finite Element Analysis, MRIs, and traditional biochemical and dissectional techniques to look deep into the venom apparatus of the living Komodo dragon (V. komodensis).

    For those who may have missed it on the first go around, it has recently been discovered that venom is more widespread among squamates than previously thought (Fry et al 2005). The authors of that paper (a few of whom are on this paper) found the presence of specific glands at the base of the mandible in numerous lizard species. These glands were found to release salivary proteins that were, in fact, venom.

    It was a “primitive” venom for the most part, with little denaturing, or tissue destroying properties, but enough that it seemed to warrant the construction of a new clade of squamates named: Toxicofera (Fry et al 2005, Vidal & Hedges 2009). Though the discovery of incipient venom production in many squamates, was an intriguing surprise, the resultant cladogram has proven problematic, and controversial. The authors found iguanians (iguanas, chameleons, most pet lizards) to be deeply nested within scleroglossa (skinks, snakes, varanids); a view that flies in the face of every morphological study ever done on this group (e.g. Romer 1956, Pianka and Vitt 2003). In order for Toxicofera’s current associations to be valid, iguanians would have to have re-evolved both their temporal bars, as well as a fleshy tongue. While possible (few things in evolution are impossible), it is extremely unlikely; kind of like expecting snakes to re-evolve limbs.

    Despite this contentious relationship, the discovery of venom glands in animals like monitor lizards, was a surprise. This new study by Fry et al is the first to really look at the venom secreting abilities of this gland, and what it means to Komodo dragon ecology.

    Photo by: Jeff Werner [Fauna vol.1 number 3 Mar 98]
    Photo by: Jeff Werner Fauna vol. 1 (3). March 1998.

    It turns out that the mandibular venom gland in V.komodoensishas six different compartments that open between the teeth of the lower jaw. Unlike venomous snakes and helodermatid lizards, the venom does not travel through any grooves in the teeth. Rather, it appears to pool at their base; bathing the teeth of the lower jaw prior to biting a prey animal. It’s a crude method of venom delivery, but one that might explain why Komodo dragons have such thick gums (which the teeth erupt through during a bite).

    According to the authors, the mandibular venom gland of a 1.6m (5.25ft) Komodo dragon has enough fluid to produce 150mg of venom; 30mg of which would be available for delivery. That’s a fair amount of venom, but how does that translate to toxicity?

    Though the delivery method is crude, the venom is fairly potent. According to the authors it only takes 0.1mg/kg of venom in the blood stream to cause pronounced hypotension, and only 0.4mg/kg to cause hypotensive collapse (fainting).

    To put this into perspective, I weigh approximately 76kg (168lbs). It would take approximately 7.6mg of Komodo dragon venom to make me light headed, and 30mg to knock my arse out.

    Hmm, maybe I should reconsider that Komodo island trip?

    Fry et al go on to discuss how V.komodoensis goes about using this venom delivery system during predation. It was at this point that I became a bit hesitant.

    Komodo dragon feeding ecology has been the subject of much misconception. Much like dinosaurs, earlier work on these beasts was more accurate than the work that soon followed. When Komodo dragons were first discovered, they were thought to be scary top predators of their respective habitat. This was quickly downgraded to obligate scavenger; possibly due to the animal’s willingness to eat prekilled meat, but more likely from general incredulity that a large reptile can actively hunt mammals (see table 10-2 of Auffenberg 1981 for examples). It really wasn’t until Dr. Walter Auffenberg spent some 13 months in the wild with Komodo dragons, that this myth was officially dispelled, and some 20 years after for it to become common knowledge. However, once it was discovered that animals lucky enough to escape from an initial V.komodoensis attack were found to die hours/days later, the view of Komodo dragons as “bite and release” predators was born (e.g. Bakker 1986).

    Auffenberg’s work did show that there is something septic about the bite of oras. This was originally attributed to bacterial flora living in the fairly dirty mouths of these predators. Indeed one study (Gillespie et al 2002) found 54 potentially pathogenic bacteria living in the mouths of oras!

    However, and this is the part that always seems to get glossed over: there has never been a reported case of a komodo monitor using this “bite and release” killing strategy. Despite spending over a year living with these animals, Auffenberg never once found an animal bitten, released and then later tracked down after it died. Komodo dragon attacks were quite the opposite in fact. Small, to relatively large prey (goats, boar) were often killed on the spot using violent side to side shaking to snap the neck, while large prey like water buffalo were hamstringed (Achilles tendon severed), followed by abdominal evisceration of the now paralyzed (and often still alive) animal.

    Despite the gruesome detail in which Auffenberg described ora attacks, as well as the sheer lack of evidence for a viper style feeding strategy; one can still read about how Komodo dragons “avoid confrontation with their prey” by allegedly employing this method of killing (for instance).

    So one can forgive my trepidation over what was to be written about next in the Fry et al paper.

    The authors do discuss the alleged “bite and release” hunting style posited for V.komodoensis, but are quick to point out (as I just did) that there has never been a documented case of this hunting strategy being used on dragon prey.Dr. Fry went went one step further in an interview for Science News:

    What’s more, rare sightings of the lizards hunting didn’t fit with this method. Victims typically died quickly and quietly after going into shock, the authors say. “No one’s actually seen a Komodo dragon track a prey for three days until it dies of septicemia,” Fry says. “It’s an absolute fairy tale.”

    This was very comforting to see. One can only hope that the other news outlets don’t miss this point when doing their write ups (Edit: so much for hope).

    Fry et al then went on to dispel the myth that the mouth of dragons contain toxic microflora. Though there have been studies that have shown the presence of potentially pathogenic bacteria in wild oras, none of these studies found a consistent microflora between individuals. In fact, the authors point out that some of the bacteria found in Komodo dragon mouths, were the same bacteria found in the guts of most lizards.

    That venom must be playing an important role in predation was determined by looking at the evolution of venom in squamates. The authors point out that:

    We have shown that in the species that have developed secondary forms of prey capture (e.g., constricting) or have
    switched to feeding on eggs, the reptile venom system undergoes rapid degeneration characterized by significant atrophying of the
    glands, reduction in fang length, and accumulated deleterious mutations in the genes encoding for the venom proteins (9, 26,
    27). This is a consequence of selection pressure against the bioenergetic cost of protein production (28). The robust glands
    and high venom yield in V. komodoensis thus argue for continued active use of the venom system in V. komodoensis.

    So, while the venom of Komodo dragons is not the primary means by which dragons dispatch their prey, it still must play a pretty important role in prey acquisition. Since envenomated prey tend to become docile and quiet (Auffenberg, 1981, and this paper), it may just play a role in initiating shock, and reducing retaliatory actions by prey. It may also serve as a good “failsafe” in the event of a missed kill. Bitten prey that are “lucky” enough to escape an initial attack, tend to find themselves easily preyed upon shortly thereafter. This is similar to hunting tactics seen in Canadian lynx (the only mammalian carnivores known to have a septic bite) when hunting caribou (Auffenberg 1981).

    Komodo dragon FE skull made by the Computation Biomechanics Research Group. UNSW, Sydney Australia.
    Komodo dragon FE skull made by the Computation Biomechanics Research Group. UNSW, Sydney Australia.

    Using Finite Element Analysis, the authors compared the bite and skull strength of V.komodoensis with that of a similar sized saltwater crocodile (Crocodylus porosus). The results they obtained agreed with previous FE work on Komodo dragons (Moreno et al 2008), which found the bite of oras to be remarkably weak on its own, thus requiring the aid of the postcranial musculature in delivering much of the force. Ora skull strength is at its greatest during bite and pull behaviour. This data agrees well with field observations showing oras biting and pulling back on their prey. Coupled with their recurved and serrated teeth, this results in the creation of large, gaping wounds, which would aid in venom delivery as the ora’s venom would be spread throughout; quickly entering the bloodstream and speeding up shock.

    Finally the authors extrapolated their work to the monstrous lacertilian behemoth Varanus (Megalania) prisca. Using the extant phylogenetic bracketing method (Witmer 1995, 1998), they were able to determine the likelihood of venom being present in Megalania. If true, this would make Megalania the largest venomous carnivore to have ever lived.

    I’m not sure I buy this part. As Fry et al mentioned in the paper, the venom apparatus tends to degrade quickly when not used. Megalania was a big animal (over 2,000 kg according to the authors, though Molnar 2004 places it as just under 2,000kg for the largest individuals). Any hole that V(M)prisca would create when attacking its prey, would have been devastating enough without the need for anticoagulating venom.


    Like the other members of this unique varanid lizard clade, the jawbones of V. prisca are also relatively gracile compared with the robust skull and the proportionally larger teeth similarly serrated (Fig. 3).

    I’d be careful about this assumption, as there is only one fairly complete maxilla (upper jaw bone), and portions of the dentary (tooth bearing lower jaw bone), known for Megalania. This makes comparison with extant monitors, rather hard to do. What little skull bones do exist, show that the skull of Megalania was stronger (or at least, less flexible) than that of other monitor lizards (Molnar, 2004).

    As it stands right now, there are frustratingly too few fossils of Megalania (especially the skull) to accurately say one way, or the other in regards to venom delivery.

    Of course that doesn’t make it any less interesting to speculate about. 🙂



    Auffenberg, Walter, 1981, The Behavioral Ecology of the Komodo Monitor, Florida University press, pgs: 406.

    Bakker, R. 1986. The Dinosaur Heresies. William Morrow. New York. ISBN: 0821756087, 978-0821756089 pgs: 481.

    Fry, B.G., Vidal, N., Norman, J.A., Vonk, F.J., Scheib, H., Ryan Ramjan, S.F., Kuruppu, S., Fung, K., Hedges, S.B., Richardson, M.K., Hodgson, W.C., Ignjatovic, V., Summerhayes, R., Kochva, E. 2005. Early Evolution of the Venom System in Lizards and Snakes. Nature. Vol.439:584-588.

    Gillespie, D., Fredekin, T., Montgomery, J.M. 2002. “Microbial Biology and Immunology” in: Komodo Dragons: Biology and Conservation. James Murphy, Claudio Ciofi, Colomba de La Panouse and Trooper Walsh (eds). pgs: 118-126. ISBN: 1588340732/978-1588340733

    Molnar, R.E. 2004. Dragons in the Dust: The Paleobiology of the Giant Monitor Lizard Megalania. Indiana University Press. 210pgs. ISBN: 0253343747/978-0253343741

    Moreno, K., Wroe, S., Clausen, P., McHenry, C., D’Amore, D.C., Rayfield, E.J., Cunningham, E. 2008. Cranial Performance in the Komodo Dragon (Varanus komodoensis) as Revealed by High-Resolution 3-D Finite Element Analysis. J.Anat. Vol.212:736-746.

    Pianka, E.R., and Vitt, L.J. 2003. Lizards Windows to the Evolution of Diversity. U.Cal.Press. 333pgs. ISBN: 0520234014/9780520234017

    Romer, A.S. 1956. The Osteology of the Reptiles. Krieger Publishing 800pgs. ISBN: 089464985X/978-0894649851

    Vidal, N. and Hedges, S.B. 2009 The Molecular Evolutionary Tree of Lizards, Snakes, and Amphisbaenians. Biologies. Vol.332(2-3):129-139.

    Witmer, L. 1995. “The Extant Phylogenetic Bracket and the Importance of Reconstructing Soft Tissues in Fossils” in: Functional Morphology in Vertebrate Paleontology. Jeff Thomason (ed). Cambridge Univ. Press. Cambridge, UK. pgs 19-33. ISBN 0521629217/978-0521629218

    Witmer, L. M. 1998. Application of the Extant Phylogenetic Bracket (EPB) Approach to the Problem of Anatomical Novelty in the Fossil Record. J.Vert.Paleo. Vol.18(3:Suppl.): 87A.

  • 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.



    Bakker, R. 1986. The Dinosaur Heresies. William Morrow. New York. ISBN: 0821756087, 978-0821756089 pps: 209-212.Buckley, G.A., Brochus, C.A., Krause, D.W., Pol.D. 2000. A Pug-Nosed Crocodyliform from the late Cretaceous of Madagascar. Nature. vol.405:941-944.

    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.

    Gilmore, C.W. 1905. The Mounted Skeleton of Triceratops porosus.? Proceedings United States National Museum. Vol.29:433-435.

    Lull, R.S. 1933. A Revision of the Ceratopsia, or Horned Dinosaurs. Memoirs of the Peabody Museum of Natural History. Vol.3:1-175.

    Naish, D. 2001. Fossils Explained 34: Crocodilians. Geology Today. Vol.17(2):71-77.

    Nobre, P.N. and Carvalho, I.S. 2006. Adamantinasuchus navae: A New Gondwanan Crocodylomorpha (Mesoeucrocodylia) from the Late cretaceous of Brazil. Gondwana Research. Vol.10:370-378.

    Paul, G.S., and Christiansen, P. 2000. Forelimb Posture in Neoceratopsian Dinosaurs: Implications for Gait and Locomotion. Paleobiology, 26(3):450-465.

  • T-U-R-T-L-E Power! Part 1: Turtles are Weird.

    As the meme goes: I like turtles!

    They are such a unique group of animals, that one can’t help but be drawn to them. Yet despite their uniqueness, turtles tend to get thrown into the wastebin of “living fossils”. It’s not uncommon to hear documentaries, or books refer to turtles as having been static since their first appearance 200+ million years ago. It’s unfortunate because statements like these tend to downplay just how weird and wonderful turtles really are.

    So why are turtles so weird? Well, as one might expect, it’s all about the shell. The turtle shell is an iconic image. Everyone knows what a turtle basically looks like. Even strange turtles like the mata mata (Chelus fimbriatus) are still recognizable as turtles. Contrary to popular belief, turtles can neither come out of their shells, nor does the shell act as their home. One cannot pull a turtle from its shell. The shell is the result of a phenomenal transformation of the backbone, ribcage, sternum, clavicles and gastralia.

    Turtle shells are different from the armoured “shells” seen on dinosaurs like the ankylosaurs. It is also fundamentally different from the armour seen on armadillos, crocodylians and every other vertebrate out there. In all these other animals, the armour is composed of bony plates that are formed from bone which is made intramembranously in the dermal portions of the body. Turtles are the only animals we know of that develop their armour by using this dermal bone in conjunction with endochondral bones (i.e.. the vertebrae and rib cage).

    A turtle “coming out of its shell.” Image from the Encarta website.

    It is at this point that turtles go from simply being unique, to just being weird. In order for the shell to protect the exposed limbs and head, the shell had to engulf the limb girdles. The rib cage had to actually grow over the pectoral and pelvic girdles. Think about that for a minute. Take a look in the mirror sometime and see how your arms are placed. Our arms, and the arms of every other tetrapod alive today, are set outside the rib cage. In fact, we actually can (and do) rest our arms along the outside of our ribs. Turtles can’t do that. Having one’s ribs on the outside can really hamper the ability to move the arms. The arms can extend, but they cannot bend without banging into the ribs. In order to fix this, turtles had to reverse the way their arms bend. Turtle arms bend towards one another, rather than away as they do in all other tetrapods. Imagine if your arms bent like your legs do, and you get the idea. Protection of the head required another unique innovation. Namely, turtles had to become double jointed. Turtle neck articulation follows a standard “ball and socket” arrangement that is widespread among various extant reptiles. However, within each species there is between one and two vertebrae that feature a “ball” on both sides (Romer, 1956). This biconcavity creates a hinge joint that can bend a full 90°. It is this special joint, more than anything else, that allows turtles to contort their necks in such a manner. For pleurodires, as the name implies, this articulation allows the neck to be tucked to the side of the body under a lip of the carapace. For cryptodires, these double joints allow the head and neck to literally go inside the body cavity; something no other tetrapod can do, and something that is decidedly weird. 🙂 Another issue with having a shell composed of fused ribs and vertebrae, is that flexibility is reduced to zero. This has a huge effect on speed. Turtles cannot extend their stride by bending their spine; a behaviour that all other tetrapods are capable of . The only way to increase stride length is to increase the lengths of the limbs. This puts an immediate limit on turtle speed. While longer limbs could be evolved, they would not be able to fit inside the shell. The only way for a turtle to go faster is to speed up the stride frequency. Turtles were thus forced to give up on ever being speedy. Though there are some chelonian members (e.g. my Terrapene ornata luteola) which put that statement to the test.

    Yet another weird characteristic of turtles is how they have circumvented the issue of breathing while encased in armour.

    Normally, in tetrapods, breathing is achieved through the bellow like pumping of the lungs. This is accomplished by muscles connected to the ribs. These muscles expand the ribcage, allowing air to enter. As turtles no longer had the joints that allow the ribs to move, they lost the muscles that moved them. This creates a problem unique to turtles. How does one get air both in, and out of the body cavity. This is a problem that seems to have been solved multiple times in turtle evolution. Tortoises can “rock” their pectoral girdles back and forth in order to pump the lungs. Many semi-aquatic turtles can use the buoyancy of water to push air out of their lungs, while others can use the weight of their viscera to pull down on the lungs and allow air in. Many, though, have evolved sheets of muscle connected to the lungs, which will either expand, or contract the lungs and allow for respiration. Some, such as box turtles (Terrapene) require a sheets of muscle that will both expand and contract the lungs. In these animals, both inhalation and exhalation, are an energetic process. The upshot to this, is that by having independent muscles for respiration, box turtles are able to breathe even when fully sealed inside their shells (Landberg et al, 2003).

    One strange aspect of chelonians that is rarely brought up, is how incredibly diversified they are. If turtles had died out at the end of the Mesozoic, and all we had to go on were fossils, I doubt we would ever have realized just how “flexible” the turtle bodyplan actually is.

    Despite being encased in a shell both above and below, turtles are capable of chasing down prey (e.g. Trionyx and Apalone). Some are adept excavators; making extensive burrows that can run as long as 9 meters (30ft) and be 3.6m (12ft) deep (Gopherus agassizii). Still others like pancake tortoises (Malacochersis tornieri) are proficient rock climbers. Probably most surprising are musk turtles (Sternotherus). These normally waterbound turtles are quite adept tree climbers. Sternotherus minor has been observed scaling cypress trees up to 2 meters (Orenstein, 2001). Both of these species have relatively small plastrons which give them added flexibility. Still, even stiffened tanks like Leopard tortoises (Geochelone pardalis ) have been observed scaling fences that were blocking their way. The animals would climb up one side and then just topple over the other (Orenstein, 2001).

    Some, such as the big-headed turtle (Platysternon megacephalum) have evolved huge heads with strong jaws for crushing shellfish. Others are efficient filter feeders (Podocnemis unifilis); sieving the water of small food particles.

    Many freshwater turtle species have re-evolved ?gills.? These are areas of thin, permeable skin usually around their cloaca. This allows these species to take in oxygen through the water.

    Lastly, turtles don’t grow old (Congdon, 1992). Unlike most other animals, turtles show little to no signs of age related deterioration. 74 year old three toed box turtles (Terrapene carolina triunguis) were found to be just as reproductively active as turtles some 40 years younger than them. (Miller, 2001).

    So chelonians are weird, but how did they come to be this way? For that, you’ll have to stay tuned.


    Extra geek points to folks who got the reference to the Partners in Kryme song from the first TMNT movie. Id est: the original turtle rap. None of that Vanilla Ice crap.


    Congdon, J. 1992. Senescence in Turtles: Evidence from Three Decades of Study on the E. S. George Reserve. Senescence in Organisms in Natural Populations. American Association of Gerontologists. Washington, D.C.

    Landberg, T., Mailhot, J.D., Brainerd, E.L. 2003. Lung Ventilation During Treadmill Locomotion in a Terrestrial Turtle, Terrapene carolina. J.Exp.Biol. Vol. 206: 3391-3404.

    Miller, J.K. 2001. Escaping Senescence: Demographic Data from the Three-Toed Box Turtle (Terrapene carolina triunguis).

    Orenstein, R. 2001. Turtles, Tortoises and Terrapins: Survivors in Armor. Firefly Books. 304 pps. ISBN 1-55209-605-X

    Romer, A.S. 1956. Osteology of the Reptiles. U.Chicago Press. ISBN: 0-89464-985-x 772pps

  • The return of fruit eating crocodylians.

    Back in 2002, a short paper came out that commented on the observation that captive caimans would eat fruit left in their cage. When I initially read the paper, I found it interesting. In the end, though, I assumed this to just be a fairly anomalous incident.

    Now Darren Naish of Tet Zoo has followed up on this story with further evidence of frugivory in crocodylians.

    As one can see, this observation has been filmed at least once.

    So does this mean that crocodylians are not as completely carnivorous as once thought? It’s hard to say. All observations made so far have been from alligatorids (alligators and caimans). This might be an apomorphic trait to this group. Only more observations will say for sure.

    Another option that Darren pointed out, is that this was a learned trait of these captive animals. In each case, observed animals were found to be sharing their enclosures with herbivorous animals (usually tortoises). This type of operant learning is rather rare, and would be amazing if found to be true.

    However, as evidenced by the comments of St. Augustine Alligator Farm park director, John Brueggen, fruit eating has been observed in wild animals too; so this is not simply a case of bored captives.

    Whatever the case, these observations do illustrate just how adaptaptable crocodylians are as a group.


  • Leopard takes out crocodile

    The Telegraph had a report today on a unique event in Africa. Hal Brindley, an American Wildlife photographer, happened to be in the right place in the right time for this to happen. The shots come from Kruger national park.

    Photo taken by Hal Brindley and destributed by Solo Syndication

    Photo taken by Hal Brindley and destributed by Solo Syndication

    The photos show a leopard (Panther pardus) taking out a Nile crocodile (Crocodylus niloticus). The croc is smaller than the leopard, but the results are no less shocking.

    Leopards, and most big cats in general, don’t go after crocodiles as a normal routine. Not only do the felines run the risk of becoming the hunted, but even if they do win (like this leopard did), there is really not a lot of meat to go around. Much of it is locked away by the bony osteoderms.

    The only time I have seen a leopard take out a crocodile, was in a case where both animals were slowly starving and dehydrating. The crocodile died first, and the leopard took advantage of the free meal.

    As one can see in the photo, this leopard wasn’t exactly starving, so it is hard to imagine desperation being the cause of this attack. The article also mentioned that the leopard attacked the croc on its own turf. Both situations are very unique, and make the fact that the leopard won, all the more amazing.

    There was no run time given for the battle, so I’m not sure how long it lasted. Judging from some of the shots though, it must have been intense.

    Nature is just full of surprises.