Shark week 4: megalodon tooth colonized by boring clams
November 9, 2022
Alert readers probably noticed that I titled the first post in this series “Matt’s first megalodon tooth“, implying that there would be other megalodon teeth to follow. Here’s my second one.
At first glance, this is a pretty jacked-up megalodon tooth. It is pocked with circular and ovoid craters, and has a big fat hole drilled right through it. Hardly collector grade! And in fact that’s what first caught my attention about this tooth — it’s a 6-incher that was being offered for an enticingly low price. But I got even more excited when I clicked past the thumbnail image on the sale site and saw precisely how this tooth was damaged. This is not random, senseless taphonomic battery (ahem); this tooth was colonized by a bunch of boring clams.
Like Adam Savage — and, I suspect, most collectors-of-things — I am fascinated by objects and the stories that they tell. And this tooth tells several stories. First, it’s a huge tooth from a huge shark, a truly vast, multi-ton animal heavier than a T. rex and longer than my house. Second, it’s a fossil that’s millions of years old, evidence of an extinct species from a vanished ecology, one where gigantic sharks and macroraptorial sperm whales hunted small baleen whales, early seals and sea lions, and manatees and sea cows. And third, it’s a relic of another, entirely different ecology, one in which this shed tooth sank to the sea floor and was colonized by a host of smaller organisms, including most obviously hole-boring clams. In effect, this one tooth was a miniature reef, supporting multiple species of invertebrates. The traces left by those invertebrates are themselves ichnofossils, so this tooth is a body fossil with ichnofossils dug out of it. It’s turtles all the way down!
Can we figure out what any of those invertebrates were? Just a few years ago that would have been a challenging task for a non-specialist, but fortunately in 2019 Harry Maisch and colleagues published a really cool paper, “Macroborings in Otodus megalodon and Otodus chubutensis shark teeth from the submerged shelf of Onslow Bay, North Carolina, USA: implications for processes of lag deposit formation”. That paper is very well illustrated, and the figures basically serve as a field guide for anyone who wants to identify similar traces in rocks or teeth of equivalent age. I will take up that sword in a future post.
Incidentally, this is now the biggest tooth in my little collection, just slightly — but noticeably — bigger than my first megalodon tooth: 157mm on the long side, vs 155mm, and 112mm max root width, vs 107mm.
Bonus goofy observation: I strongly suspect that no other megalodon tooth in the world beats this one in simulating a Star Trek phaser.
Reference
Shark week 2: great white teeth redux
November 6, 2022
Cast (white) and fossil (gray) great white shark teeth, lingual (tongue) sides.
Something cool came in the mail today: a fossil tooth of a great white shark, Carcharodon carcharias. The root is a bit eroded, but the enamel-covered crown is in great shape, and it’s almost exactly the same size as my cast tooth from a modern great white.
The labial (outer or lip-facing) sides of the same teeth.
I got this for a couple of reasons. One, I wanted a real great white shark tooth to show people alongside my megalodon tooth (for which see the previous post). Extant great whites are quite rightly protected, and their teeth are outside my price range when they are available at all. Fortunately there are zillions of fossil great white teeth to be had.
Also, the cast great white tooth has been kind of a disappointment. It’s so white that it’s actually a letdown, visually. Tactilely it’s great, with all kinds of subtle features on the crown especially, but those features are almost impossible to see or photograph. In the photo above, you can make out some of the long, smooth wrinkles in the enamel of the cast tooth, but the median ridge, which is dead obvious on the fossil tooth, only shows up under very low-angle lighting on the cast. The fossil tooth is just a more interesting and more informative specimen, material and origin aside. Now that I have it, I might try either staining or painting the cast tooth, to see if I can rehabilitate it as a visual object.
This fossil tooth is also noticeably thicker than the cast tooth. I don’t know if that’s serial, individual, population, or evolutionary variation. In the last post I contrasted the proportional thinness of the cast tooth with the robustness of the megalodon tooth; this fossil tooth might fare a little better if subjected to the same comparison. I should have thought to do that when I was taking these photos.
Speaking of comparisons, here’s megalodon to remind everyone who’s boss. There’s no scale bar here, but the cast great white tooth is 65mm from the tip of the crown to the tip of the longer root, and the meg tooth is 155mm between the same points.
Now I have a gleam in my eye of assembling a couple of sets of fossil teeth: one to illustrate the evolution of the modern great white from its less-serrated ancestors, like this diagram from the great white Wikipedia page, and one to illustrate the evolution of megalodon from its side-cusped ancestors, like this diagram from the megalodon page — presuming that current hypotheses for the two lineages are accurate. If I ever get either set done, I’m sure I’ll yap about it here.
Matt’s first megalodon tooth
November 5, 2022
I got this thing a while back. I’d always wanted one, and it really does spark joy.
First up: what should we call this critter? AFAIK, the species name has never been in doubt, it’s always been [Somegenus] megalodon. That genus has variously been argued to be Carcharodon (same as the extant great white shark, Carcharodon carcharias), Carcharocles, Otodus, Megaselachus, and probably others. From my limited reading, the current consensus seems to be converging on Otodus, for reasons that seem reasonable to me, but I’m hardly an expert on this problem. It’s not that I think it’s unimportant, more that the generic identity of [Somegenus] megalodon has been historically labile, and as a non-expert I hesitate to come down firmly behind any of the hypotheses. If it’s still Otodus megalodon in another decade, I might take a stand. If you want to do a deep dive on this, check out Kent (2018: 80-85). In the meantime, I’m going to refer to it informally as ‘megalodon’, without italics. Although the actual genus name Megalodon was tragically wasted a fossil clam (true story), I’m confident that no-one, scientist or layperson, will misunderstand when I refer to the humongous extinct megatoothed shark as megalodon.
With that out of the way: wow, that’s a big freakin’ tooth! Here it is again with a scale bar.
The serrations on the sides are very cool. The edges are worn a bit in places, and that plus the visible notch on one side of the tooth (upper left in the photo above) suggests that this tooth was used, as opposed to being a replacement tooth that rotted out of the jaw before it ever had a chance to be deployed. Where ‘used’ means ‘used to punch and then tear immense holes in other animals’. Pretty wild to think about ancient whales dying on this very tooth.
I use this thing at outreach events, and I got a cast tooth of a modern great white shark for comparison. Those great white teeth are 10 bucks at Bone Clones, so I got a bunch of them and gave them to nieces and nephews as stocking stuffers.
Here’s a labeled version. From what I’ve been able to determine (i.e., shark people, please correct me if I’m wrong!), most shark teeth ‘lean’ away from the body midline. Upper teeth of megalodon tend to be very wide, with wide, shallow angles at the base, whereas lower teeth are more dagger-shaped and have a more pronounced basal angle. I’m pretty sure this meg tooth is a lower, and we’re looking at the lingual (tongue) side in this photo (more on that in a bit), so the tooth is facing the same way we are. I think that makes it a left lower tooth. The great white tooth is a probably a left upper, although great whites apparently have one tooth position that leans mesially instead of distally, so I could have that one wrong-sided. The ‘bourlette’ is an area of exposed orthodentine between the root and the enamel that covers the tooth crown (Kent 2018: 86). This tooth is not in perfect shape, there’s been some peeling of the enamel just above the bourlette.
I think this photo makes the size-comparison point even more clearly.
Worth noting: if the hypothesis that megalodon belongs in Otodus is correct, the similarities between megalodon and the great white shark are convergent; megalodon teeth are Otodus teeth that lost their side-cusps, and great white teeth are basically wider, serrated mako teeth. That level of convergence shouldn’t be surprising to anyone who has seen a thylacine skull. Still, this photo makes it very obvious why Louis Agassiz assigned megalodon to Carcharodon, the great white shark genus, when he named the species back in 1843: the two look a lot alike. (Also: Agassiz didn’t have all the transitional fossils that we do now.)
Boomerang thought, added in post: at least, megalodon teeth look a lot like the upper teeth of great whites. The lower teeth of great whites are much narrower and more mako-esque.
A couple of features worth noting here. The mesial margin has a little wrinkle, which cannot be damage because the serrations follow the in-folded contour. This seems to be a minor developmental anomaly that is pretty common in megalodon teeth. The distal margin has a distinct notch, also mentioned above, which probably represents feeding damage sustained in life.
Arguably this side-view is even more striking; the megalodon tooth is 2.38 times the length of the great white tooth (155mm vs 65mm on the long side), but more than three times as thick (29mm vs 9mm max thickness), and the blade of the tooth stays proportionally thicker over more of its length. This tooth was built to do some work.
Am I fanboying here? Sure, a little (and not for the first time). Giant extinct monsters are exciting, and I’m happy to celebrate that while also wanting to know more about how they lived.
The thing that surprised me the most while reading up on shark teeth is how they are oriented in the jaws. I’d always assumed that the convex faces (toward the bottom of the above photo) faced outward (labial or lip-facing), and the flat faces (toward the top of the above photo) faced inward (lingual or tongue-facing), but it’s actually opposite. In the photo above, the labial or outward faces are up, and the lingual or inward faces are down. I’m sure this is old hat to shark people, but it hurts my head. Most teeth I know of have their convex faces outward, like human incisors and tyrannosaur premaxillary teeth. Plus, instinctively it seems like predator teeth should curve toward the back of the mouth, but with their flat labial faces and convex lingual faces, most shark teeth seem to curve toward the front (I realize that they may have been placed in the jaws so that they still pointed backwards overall). I was so surprised by this that I did a lot of checking before bringing it up in this post, but it’s clear even in really good photos of live great white sharks with their mouths open. There’s no bigger story here, just me confronting my own misapprehension about animal morphology. Still seems weird.
If you want to know more about how megalodon lived, I’ve included links below to some papers on its size (Shimada 2019, Shimada et al. 2020, Cooper et al. 2020, 2022, Perez et al. 2021), breeding habits and life history (Miller et al. 2018, Shimada et al. 2021, 2022), evolution (Shimada et al. 2016, Kent 2018, Perez et al. 2018), and paleobiology (Maisch et al. 2019, Ballell and Ferron 2021, Miller et al. 2022, Sternes et al. 2022). This is a highly idiosyncratic collection based on like one evening of messing around on Google Scholar. I’m sure I missed tons of important work, so feel free to recommend more refs in the comments.
Oh, like virtually everything else on this site, these photos are freely available under the CC-BY license, so if you want to use them, modify them, etc., go nuts.
References
- Ballell, A. and Ferrón, H.G. 2021. Biomechanical insights into the dentition of megatooth sharks (Lamniformes: Otodontidae). Scientific Reports 11(1): 1-9.
- Cooper, J.A., Pimiento, C., Ferrón, H.G. and Benton, M.J. 2020. Body dimensions of the extinct giant shark Otodus megalodon: a 2D reconstruction. Scientific Reports 10(1): 1-9.
- Cooper, J.A., Hutchinson, J.R., Bernvi, D.C., Cliff, G., Wilson, R.P., Dicken, M.L., Menzel, J., Wroe, S., Pirlo, J. and Pimiento, C. 2022. The extinct shark Otodus megalodon was a transoceanic superpredator: Inferences from 3D modeling. Science Advances 8(33): p.eabm9424.
- Kent, B.W. 2018. The cartilaginous fishes (chimaeras, sharks, and rays) of Calvert Cliffs, Maryland, USA. The Geology and Vertebrate Paleontology of Calvert Cliffs, Maryland, USA. Smithsonian Contributions to Paleobiology 100: 45-157.
- Maisch IV, H.M., Becker, M.A. and Chamberlain Jr, J.A. 2020. Macroborings in Otodus megalodon and Otodus chubutensis shark teeth from the submerged shelf of Onslow Bay, North Carolina, USA: implications for processes of lag deposit formation. Ichnos 27(2): 122-141.
- Miller, A., Gibson, M. and Boessenecker, R. 2021. A megatoothed shark (Carcharocles angustidens) nursery in the Oligocene Charleston Embayment, South Carolina, USA. Palaeontologia Electronica 24(2): 1-19.
- Miller, H.S., Avrahami, H.M. and Zanno, L.E. 2022. Dental pathologies in lamniform and carcharhiniform sharks with comments on the classification and homology of double tooth pathologies in vertebrates. PeerJ 10: e12775.
- Perez, V.J., Godfrey, S.J., Kent, B.W., Weems, R.E. and Nance, J.R. 2018. The transition between Carcharocles chubutensis and Carcharocles megalodon (Otodontidae, Chondrichthyes): lateral cusplet loss through time. Journal of Vertebrate Paleontology 38(6): e1546732.
- Perez, V.J., Leder, R.M. and Badaut, T. 2021. Body length estimation of Neogene macrophagous lamniform sharks (Carcharodon and Otodus) derived from associated fossil dentitions. Palaeontologica Electronica 24(1): a09.
- Shimada, K. 2019. The size of the megatooth shark, Otodus megalodon (Lamniformes: Otodontidae), revisited. Historical Biology 33(7): 904-911.
- Shimada, K., Chandler, R.E., Lam, O.L.T., Tanaka, T. and Ward, D.J. 2016. A new elusive otodontid shark (Lamniformes: Otodontidae) from the lower Miocene, and comments on the taxonomy of otodontid genera, including the ‘megatoothed’clade. Historical Biology 29(5): 704-714.
- Shimada, K., Becker, M.A. and Griffiths, M.L. 2020. Body, jaw, and dentition lengths of macrophagous lamniform sharks, and body size evolution in Lamniformes with special reference to ‘off-the-scale’gigantism of the megatooth shark, Otodus megalodon. Historical Biology 33(11): 2543-2559.
- Shimada, K., Bonnan, M.F., Becker, M.A. and Griffiths, M.L. 2021. Ontogenetic growth pattern of the extinct megatooth shark Otodus megalodon—implications for its reproductive biology, development, and life expectancy. Historical Biology 33(12): 3254-3259.
- Shimada, K., Maisch IV, H.M., Perez, V.J., Becker, M.A. and Griffiths, M.L. 2022. Revisiting body size trends and nursery areas of the Neogene megatooth shark, Otodus megalodon (Lamniformes: Otodontidae), reveals Bergmann’s rule possibly enhanced its gigantism in cooler waters. Historical Biology: 1-10.
- Sternes, P.C., Wood, J.J. and Shimada, K., 2022. Body forms of extant lamniform sharks (Elasmobranchii: Lamniformes), and comments on the morphology of the extinct megatooth shark, Otodus megalodon, and the evolution of lamniform thermophysiology. Historical Biology 1-13.
Giraffe and human metacarpals compared
May 30, 2020
Here’s one of my most prized possessions: a cannon bone from a giraffe. I got it last fall from Necromance, a cool natural history store in LA. Originally they had a matched pair on display in the front window. Jessie Atterholt got one of them last summer, and I got the other a few months later.
The cannon bones of hoofed mammals consist of fused metacarpals (in the forelimbs) or metatarsals (in the hindlimbs). In this case, the giraffe cannon bone in the top photo is the one from the right forelimb, consisting of the fused 3rd and 4th metacarpals, which correspond to the bones in the human hand leading to the middle and ring fingers. Only my third metacarpal is traced in the top photo. For maximum homology goodness I should have traced MC4, too, but I’m lazy.
I didn’t know that this was a right forelimb cannon bone when I got it. In fact, I only figured that out this afternoon, thanks to the figures and text descriptions in Rios et al. (2016), which I got free through Palaeontologia Electronica (you can too). The weirdly large and perfectly circular holes at the ends of my cannon bone were clearly drilled out by somone, I guess maybe for mounting purposes? At first I thought it might have been to help the marrow cook out of the shaft of the bone during simmering and degreasing, but none of the drilled holes intersect the main marrow cavity, they’re just in the sponge of trabecular bone at the ends of the element.
This post is a sequel to one from last year, “Brachiosaurus and human metacarpals compared“, which featured metacarpal 3 from BYU 4744, the partial skeleton of Brachiosaurus from Potter Creek, Colorado. I know what everyone’s thinking: can we make these two high-browsing giants throw hands?
Yes, yes we can. The giraffe cannon bone is 75.5cm long, and the brachiosaur metacarpal is 57cm long, or 75.5% the length of the giraffe element. I scaled the two bones correctly in the above image. My hands aren’t the same size because they’re at different distances from the camera, illustrating the age-old dictum that scale bars are not to be trusted.
The Potter Creek brachiosaur is one of the largest in the world–here’s me with a cast of its humerus–but ‘my’ giraffe is not. World-record giraffes are about 19 feet tall (5.8m), and doing some quick-and-dirty cross-scaling using the skeleton photo above suggests that the metacarpal cannon bone in a world-record giraffe should be pushing 90cm. So the giraffe my cannon bone is from was probably between 15.5 and 16 feet tall (4.7-4.9m), which is still nothing to sniff at.
I don’t know how this bone came to be at Necromance. I assume from an estate sale or something. I only visited for the first time last year, and at that time they had three real bones from giraffes out in the showroom: the two cannon bones and a cervical vertebra. They might have put out more stuff since–it’s been about six months since I’ve been there–but all of the giraffe bones they had at that point have been snapped up by WesternU anatomists. Jessie and I got the cannon bones, and Thierra Nalley got the cervical vertebra, which is fair since she works on the evolution of necks (mostly in primates–see her Google Scholar page here). I don’t know if there are any photos of Thierra’s cervical online, but Jessie did an Instagram post on her cannon bone, which is nearly as long as her whole damn leg.
There will be more anatomy coming along soon, and probably some noodling about sauropods. Stay tuned!
Reference
Nature’s CT machine, part 2: an apatosaurine in the Salt Wash
January 28, 2020
In the last post, we looked at some sauropod vertebrae exposed in cross-section at our field sites in the Salt Wash member of the Morrison Formation. This time, we’re going to do it again! Let’s look at another of my faves from the field, with Thuat Tran’s hand for scale. And, er, a scale bar for scale:
And let’s pull the interesting bits out of the background:
Now, confession time. When I first saw this specimen, I interpeted it as-is, right-side up. The round thing in the middle with the honeycomb of internal spaces is obviously the condyle of a vertebra, and the bits sticking out above and below on the sides frame a cervical rib loop. I figured the rounded bit at the upper right was the ramus of bone heading for the prezyg, curved over as I’ve seen it in some taxa, including the YPM Barosaurus. And the two bits below the centrum would then be the cervical ribs. And with such big cervical rib loops and massive, low-hanging cervical ribs, it had to an apatosaurine, either Apatosaurus or Brontosaurus.
Then I got my own personal Cope-getting-Elasmosaurus-backwards moment, courtesy of my friend and fellow field adventurer, Brian Engh, who proposed this:
Gotta say, this makes a lot more sense. For one, the cervical ribs would be lateral to the prezygs, just as in, oh, pretty much all sauropods. And the oddly flat inward-tilted surfaces on what are now the more dorsal bones makes sense: they’re either prezyg facets, or the flat parts of the rami right behind the prezyg facets. The missing thing on what is now the right even makes sense: it’s the other cervical rib, still buried in a projecting bit of sandstone. That made no sense with the vert the other way ’round, because prezygs always stick out farther in front than do the cervical ribs. And we know that we’re looking at the vert from the front, otherwise the backwards-projecting cervical rib would be sticking through that lump of sandstone, coming out of the plane of the photo toward us.
Here’s what I now think is going on:
I’m still convinced that the bits of bone on what is now the left side of the image are framing a cervical rib loop. And as we discussed in the last post, the only Morrison sauropods with such widely-set cervical ribs are Camarasaurus and the apatosaurines. So what makes this an apatosaurine rather than a camarasaur? I find several persuasive clues:
- If we have this thing the right way up, those prezygs are waaay up above the condyle, at a proportional distance I’ve only seen in diplodocids. See, for example, this famous cervical from CM 3018, the holotype of A. louisae (link).
- The complexity of the pneumatic honeycombing inside the condyle is a much better fit for an apatosaurine than for Camarasaurus–I’ve never seen that level of complexity in a camarasaur vert.
- The bump on what we’re now interpreting as the cervical rib looks suspiciously like one of the ventrolateral processes that Kent Sanders and I identified in apatosaurine cervicals back in our 2002 paper. I’ve never seen them, or seen them reported, in Camarasaurus–and I’ve been looking.
- Crucially, the zygs are not set very far forward of the cervical ribs. By some rare chance, this is pretty darned close to a pure transverse cut, and the prezygs, condyle (at its posterior extent, anyway), and the one visible cervical rib are all in roughly the same plane. In Camarasaurus, the zygs strongly overhang the front end of the centrum in the cervicals (see this and this).
But wait–aren’t the cervical ribs awfully high for this to be an apatosaurine? We-ell, not necessarily. This isn’t a very big vert; max centrum width here is 175mm, only about a third the diameter of a mid-cervical from something like CM 3018. So possibly this is from the front of the neck, around the C3 or C4 position, where the cervical ribs are wide but not yet very deep. You can see something similar in this C2-C5 series on display at BYU:
Or, maybe it’s just one of the weird apatosaurine verts that has cervical rib loops that are wide, but not very deep. Check out this lumpen atrocity at Dinosaur Journey–and more importantly, the apatosaur cervical he’s freaking out over:
UPDATE just a few minutes later: Mike reminded me in the comments about the Tokyo apatosaurine, NSMT-PV 20375, which has wide-but-not-deep cervical ribs. In fact, C7 (the vertebra on the right in this figure) is a pretty good match for the Salt Wash specimen:
NSMT-PV 20375, cervical vertebrae 3, 6 and 7 in anterior and posterior views. Modified from Upchurch et al. (2005: plate 2).
UPDATE the 2nd: After looking at it for a few minutes, I decided that C7 of the Tokyo apatosaurine was such a good match for the Salt Wash specimen that I wanted to know what it would look like if it was similarly sectioned by erosion. In the Salt Wash specimen, the prezygs are sticking out a little farther than the condyle and cervical rib sections. The red line in this figure is my best attempt at mimicking that erosional surface on the Tokyo C7, and the black outlines on the right are my best guess as to what would be exposed by such a cut (or pair of cuts). I’ve never seen NSMT-PV 20375 in person, so this is just an estimate, but I don’t think it can be too inaccurate, and it is a pretty good match for the Salt Wash specimen.
Another way to put it: if this is an apatosaurine, everything fits. Even the wide-but-not-low-hanging cervical ribs are reasonable in light of some other apatosaurines. If we think this is Camarasaurus just because the cervical ribs aren’t low-hanging, then the pneumatic complexity, the height of the prezygs, and the ventrolateral process on the cervical rib are all anomalous. The balance of the evidence says that this is an apatosaurine, either a small, anterior vert from a big one, or possibly something farther back from a small one. And that’s pretty satisfying.
One more thing: can we take a moment to stand in awe of this freaking thumb-sized cobble that presumably got inside the vertebra through one its pneumatic foramina and rattled around until it got up inside the condyle? Where, I’ll note, the internal structure looks pretty intact despite being filled with just, like, gravel. As someone who spends an inordinate amount of time thinking about how pneumatic vertebrae get buried and fossilized, I am blown away by this. Gaze upon its majesty, people!
This is another “Road to Jurassic Reimagined, Part 2″ post. As before, Part 1 is here, Part 2 will be going up here in the near future. As always, stay tuned.
References
- Upchurch, P., Tomida, Y., and Barrett, P.M. 2005. A new specimen of Apatosaurus ajax (Sauropoda: Diplodocidae) from the Morrison Formation (Upper Jurassic) of Wyoming, USA. National Science Museum Monographs No. 26. Tokyo. ISSN 1342-9574.
- Wedel, M.J., and Sanders, R.K. 2002. Osteological correlates of cervical musculature in Aves and Sauropoda (Dinosauria: Saurischia), with comments on the cervical ribs of Apatosaurus. PaleoBios 22(3):1-6.
When I visited Dinosaur National Monument in October with Brian Engh and Yara Haridy, we spent a decent amount of time checking out DNM 28, a skull and associated bits of Camarasaurus. In particular, I got some shots of the axis (the second cervical vertebra behind the head), and it got me thinking about pneumaticity in this unusual element. Why I failed to get a full set of orthogonal shots is quite beyond my capacity, but we can roll with what I have. Before we go on, you might want to revisit Tutorial 36 to brush up on the general parts of the atlas-axis complex.
Here’s the axis in left lateral view (so, anterior to the left).
And a labeled version of the same. A few things to note:
- One oddity of sauropod axes (and of axes of most critters) is that not only are the articular facets of the prezygapophyses not set forward of the neural arch, they’re set backward, well behind the forward point of the arch.
- The dens epistrophei or odontoid process is sticking out immediately below the neural canal. This is the tongue of bone that articulated with the atlas (first cervical vertebra) in life.
- Check out the prominent epipophysis above the postzygapophysis, which anchored the long dorsal neck muscles. (For more on epipophyses, see these posts, and especially this one.)
- The diapophysis and parapophysis articulated with a cervical rib, which is not shown here. In fact, I don’t remember seeing it in the drawer that this vert came from. The atlantal and axial cervical ribs are small, apparently fused late in life if they fused at all, and are easily lost through taphonomic processes.
- At least three pneumatic features are visible in this lateral view: the lateral fossa on the centrum, which is referred to as the “pleurocoel” in a lot of older literature; a ventral fossa that lies between the parapophysis and the midline ventral keel; and a fossa on the neural arch, behind the postzygodiapophyseal lamina. In the nomenclature of Wilson et al. (2011), this is the postzygocentrodiapophyseal fossa.
“Postzygocentrodiapophyseal fossa” is a mouthful, but I think it’s the only way to go. To be unambiguous, anatomical terminology needs to references specific landmarks, and the schemes proposed by Wilson (1999) for vertebral laminae and Wilson et al. (2011) for vertebral fossae are the bee’s knees in my book.
Nomenclatural issues aside, how do we know that these fossae were all pneumatic? Well, they’re invasive, there’s no other soft-tissue system that makes invasive fossae like that in archosaur vertebrae (although crocs sometimes have shallow fossae that are filled with cartilage or fat), and the same fossae sometimes have unambiguous pneumatic foramina in other vertebrae or in other sauropods.
Most of the features labeled above are also visible on the right side of the vertebra, although the ventral fossa is a little less well-defined in this view, and I can’t make out the prezyg facet. Admittedly, some of the uncertainty here is because of my dumb shadow falling across the vertebra. Specimen photography fail!
The paired ventral fossae are more prominent in this ventral view, on either side of the midline ventral keel (anterior is to the top).
And here’s a labeled version of the same ventral view.
Finally, here’s the posterior view. It’s apparent now that the neural spine is a proportionally huge slab of bone, like a broad, tilted shield between the postzygapophyses (which are also quite large for the size of this vertebra). The back side of the neural spine is deeply excavated by a complex fossa with several subfossae (kudos again to Jeff Wilson [1999] for that eminently useful term).
Here’s the same shot with some features of interest labeled. If I’ve read Wilson et al. (2011) correctly, the whole space on the back side of the neural spine and above the postzygs could be considered the spinopostzygapophyseal fossa, but here I’ve left the interspinous ligament scar (ILS) unshaded, on the expectation that the pneumatic diverticula that created that fossa were separated on the midline by the interspinous ligament. I might have drawn the ILS too conservatively, conceivably the whole space between the large deeply-shadowed subfossae was occupied by the interspinous ligament.
I’m particularly interested in those three paired subfossae, which for convenience I’m simply calling A, B, and C. Subfossa A may just be the leftover space between the spinopostzgyapophyseal laminae laterally and the interspinous ligament medially. I think subfossa B is invading the ramus of bone that goes to the epipophysis and postzygapophysis, but I didn’t think to check and see how far it goes (that might require CT anyway).
Subfossa C is the most intriguing. Together, those paired fossae form a couple of shallow pits, just on either side of the midline, and aimed straight forward. They can’t be centropostzygapophyseal fossae, which used to be called peduncular fossae, because they’re not in the peduncles on either side of the neural canal, they’re up above the lamina that connects the two postzygapophyses. Could they be ligament attachments? Maybe, but I’m skeptical for at least four reasons:
- Although interspinous ligament attachments often manifest as pits in the cervical vertebrae of birds, in sauropods they usually form rugosities or even spikes of bone that stick out, not inward. Furthermore, these pits are smooth, not rough like the interspinous ligament scars of birds.
- The interspinous ligament in tetrapods is typically a single, midline structure, and these pits are paired.
- Similar pits in front of the neural spine are present in some sauropod caudals, and they appear to be pneumatic (see Wedel 2009: p. 11 and figure 9).
- Pits at the base of the neural spine seem to be fairly uncommon in sauropod vertebrae. If they were attachment scars from the universally-present interspinous ligaments, we should expect them to be more prominent and more widespread.
But if these paired pits are not ligament scars, what are they? Why are they present, and why are they so distinct? Sometimes (often?) subfossae and accessory laminae look like the outcome of pneumatic diverticulum and bone reacting to each other (I almost wrote ‘playing together’), in what looks like a haphazard process of adaptation to local loading. But the symmetry of these pits argues against them being incidental or random. They don’t seem to be going anywhere, so maaaybe they are the first hoofbeats of the embossed laminae and “unfossae” that we see in the vertebrae of more derived sauropods (for which see this post), but again, their symmetry in size and placement isn’t really consistent with the “developmental program gone wild” appearance of “unfossae”. I really don’t know what to make of them, but if you have ideas, arguments, or observations to bring to bear, the comment field is open.
In summary, sauropod axes are more interesting than I thought, even in a derpasaurus like Cam. I’ll have to pay more attention to them going forward.
References
- Wedel, M.J. 2009. Evidence for bird-like air sacs in saurischian dinosaurs. Journal of Experimental Zoology 311A:611-628.
- Wilson, J. A. 1999. A nomenclature for vertebral laminae in sauropods and other saurischian dinosaurs. Journal of Vertebrate Paleontology 19(4): 639-653. DOI: 10.1080/02724634.1999.10011178
- Wilson, J. A., Michael, D. D., T. Ikejiri, E. M. Moacdieh, and J. A. Whitlock. 2011. A nomenclature for vertebral fossae in sauropods and other saurischian dinosaurs. PLoS One 6(2): e17114. DOI: 10.1371/journal.pone.0017114
Worn and unworn Camarasaurus teeth in the collections at Dinosaur National Monument
November 2, 2019
Unworn:
Worn:
Spent some time last week just admiring these things. They’re pretty cool.
EDIT: in answer to Mike’s question in the first comment below, here’s a photo of some more worn teeth, showing that the level of wear in the one shown above is not unusual. Also, all of these worn teeth still had full roots, with no sign of the root resorption that would have preceded shedding of the tooth, so they were evidently going to be used for a while yet, probably a few months at least — BUT see the very useful comment from Jens Kosch below on the likely rapidity of tooth replacement in Camarasaurus.
Brachiosaurus and human metacarpals compared
October 27, 2019
Nothing too serious here, just a fun shot I got while in the collections at BYU this past week. The Brachiosaurus element is metacarpal 1 (thumb column) from BYU 4744, the Potter Creek material. I highlighted my own metacarpal 3. There is a metacarpal 3 from this specimen, but I didn’t see it on the shelf. According to D’Emic and Carrano (2019), the MC3 is 60cm long, vs 57cm for this MC1. So this photo could have been 3cm more impressive!
Oh, ignore the tag on the left that says “radius”. You could be forgiven for thinking that the bone I have my hand on is a radius, but the radius from this individual is 1.34 meters long, or about two-and-a-third times the length of this metacarpal.
Reference
D’Emic, M.D. and Carrano, M.T., 2019. Redescription of Brachiosaurid Sauropod Dinosaur Material From the Upper Jurassic Morrison Formation, Colorado, USA. The Anatomical Record.
Vertebral orientation: Varanus komodoensis would like a word
September 25, 2018
I am still building up to a big post on vertebral orientation, but in the meantime, check out this caudal vertebra of a Komodo dragon, Varanus komodoensis. This is right lateral view–the vert is strongly procoelous, and the articular ends of the centrum are really tilted relative to the long axis. I find this encouraging, for two reasons. First, it helped me clarify my thinking on how we ought to orient vertebrae, which Mike wrote about here and here. And second, it gives me some hope, because if we can figure out why tilting your articular surfaces makes functional sense in extant critters like monitors, maybe we can apply those lessons to sauropods and other extinct animals.
This is LACM Herpetology specimen 121971. Many thanks again to Neftali Camacho for access and assistance, and to Jessie Atterholt for basically doing all the other jobs while I was faffing about with this Komodo dragon.
Xenoposeidon comes to California
September 5, 2018
No time right now for me to dig into the interesting and important discussion on how we should orient vertebrae (here and here so far) – that will be coming soon. In the meantime, here’s something else.
As printed, in one of WesternU’s 3D printers.
Coming off the tray.
Cleaned up and in my hand. This is a 70% scale print, so a little smaller than the original, but all the important morphology is clear enough. For one thing, I can finally make sense of the dorsal views of the vertebra.
I have been astonished at how useful a 3D print can be as an aid to thought. The caudals of the Snowmass Haplocanthosaurus are among the smallest sauropod vertebrae I’ve spent a lot of time with, and they’re still heavy enough and fragile enough that I don’t just whip them out and twirl them around in my fingers. But I can do that with the 3D prints, and it really helps ram the morphology home in my brain. There are a thousand subtle things I might not otherwise have noticed if I hadn’t been able to turn those shapes over easily in my hand. Not to mention the other things you can do with prints, like physically sculpt on them without gooping up your fossils (we’re midway through step #8 from that post, BTW).
Anyway, back to Xeno. Mike reminded me that I have seen the actual specimen in person exactly once, very briefly during our 2005 visit to the NHM collections when I was over there for SVPCA. But it wasn’t Xeno yet, and we had other fish to fry, including a lot of pneumatic and possibly-pneumatic stuff for me to see and photograph for my dissertation. So I have to admit that it didn’t register. Being able to handle it now, so much that Mike has written about it snaps into focus. Not that his writing isn’t clear, there’s just a huge gulf between the best written description and holding a thing in your hands.
Why do I have this thing? Partly to educate myself, partly because it’s relevant to a current project, and partly because we may not be done with Xeno. Stay tuned.
Many thanks to Gary Wisser for setting up the print, and to Jeff Macalino for pulling it for me.
Sauropod Vertebra Picture of the Week 
