Vertebrae of Haplocanthosaurus (A-C) and a giraffe (D-F) illustrating three ways of orienting a vertebra: articular surfaces vertical — or at least the caudal articular surface vertical (A and D), floor of the neural canal horizontal (B and E), and similarity in articulation (C and F). See the paper for details! Taylor and Wedel (2002: fig. 6).
This is a lovely cosmic alignment: right after the 15th anniversary of this blog, Mike and I have our 11th coauthored publication (not counting abstracts and preprints) out today.
This one started back in 2018, with Mike’s post, What does it mean for a vertebra to be “horizontal”? That post and subsequent posts on the same topic (one, two, three) provoked interesting discussions in the comment threads, and convinced us that there was something here worth grappling with. We gave a presentation on the topic at the 1st Palaeontological Virtual Congress that December, which we made available as a preprint, which led to us writing the paper in the open, which led to another preprint (of the paper this time, not the talk).
Orienting vertebrae with the long axis of the centrum held horizontally seems simple enough, but choosing landmarks can be surprisingly complex. Taylor and Wedel (2022 fig. 5).
This project represented some interesting watersheds for us. It was not our first time turning a series of blog posts into a paper — see our 2013 paper on neural spine bifurcation for that — but it was our first time writing a joint paper in the open (Mike had started writing the Archbishop description in the open a few months earlier). It was also the last, or at least the most recent, manuscript that we released as a preprint, although we’ve released some conference presentations as preprints since then. I’m much less interested in preprints than I used to be, for reasons explained in this post, and I think Mike sees them as rather pointless if you’re writing the paper in the open anyway, which is his standard approach these days (Mike, feel free to correct me here or in the comments if I’m mischaracterizing your position).
So, we got it submitted, we got reviews, and then…we sat on them for a while. We have both struggled in the last few years with Getting Things Done, or at least Getting Things Finished (Mike’s account, my own), and this paper suffered from that. Part of the problem is that Mike and have far too many projects going at any one time. At last count, we have about 20 joint projects in various stages of gestation, and about 11 more that we’ve admitted we’re never going to get to (our To Don’t list), and that doesn’t count our collaborations with others (like the dozen or so papers I have planned with Jessie Atterholt). We simply can’t keep so many plates spinning, and we’re both working hard at pruning our project list and saying ‘no’ to new things — or, if we do think of new projects, we try to hand them off to others as quickly and cleanly as possible.
Two different ways of looking at a Haplocanthosaurus tail vertebra. Read on for a couple of recent real-life examples. Taylor and Wedel (2022: fig. 2).
Anyway, Mike got rolling on the revisions a few months ago, and it was accepted for publication sometime in late spring or early summer, I think. Normally it would have been published in days, but the Journal of Paleontological Techniques was moving between websites and servers, and that took a while. But Mike and I were in no tearing rush, and the paper is out today, so all is well.
One of the bits of the paper that I’m most proud of is the description of cheap and easy methods for determining the orientation of the neural canal. For neural canals that are open, either because they were fully prepped or never full of matrix to begin with, there’s the rolled-up-piece-of-paper method, which I believe first appeared on the blog back when I was posting photos of the tail vertebrae of the Brachiosaurus altithorax holotype. For neural canals that aren’t open, Mike came up with the Blu-tack-and-toothpick method, as shown in Figure 12 in the new paper:
A 3d print of NHMUK PV R2095, the holotype of Xenoposeidon, illustrating the toothpick method of determining neural canal orientation. Taylor and Wedel (2022: fig. 12).
I know both methods work because I recently had occasion to use them, studying the Haplocanthosaurus holotypes (see this post). For CM 572, the neural canal of the first caudal vertebra is full of matrix, so I used a variant of the toothpick method. I didn’t actually have Blu-tack or toothpicks, so I cut thin pieces of plastic from the edge of an SVP scale bar and stuck them in bits of kneadable eraser. It worked just fine:
The neural canal of caudal 2 was prepped, so I could use the rolled-up-piece-of-paper method:
(Incidentally, Mike and I refer to our low-tech orientation-visualizers as “neural-canal-inators”, in honor of Dr. Heinz Doofenshmirtz from Phineas and Ferb.)
In the above photos, notice how terribly thin the base of the neural arch is, antero-posteriorly. Both of these vertebrae are in pretty good shape, without much breakage or missing material, and their morphology is broadly consistent with that of other proximal caudals of Haplocanthosaurus, so we can’t write this off as distortion. As weird as it looks, this is just what Haplo proximal caudals were like. And with the neural canals held horizontally, the first two caudals end up oriented like so:
Now, as we pointed out in the paper, the titular question is not about determining the posture of the vertebrae in life, it’s about defining the directions ‘cranial’ and ‘caudal’ for isolated vertebrae — Mike asked the question back when for the holotype (single) dorsal vertebra of Xenoposeidon. But an interesting spin-off for me has been getting confronted with the weirdness of vertebrae whose articular surfaces are nowhere near orthogonal with their neural canals. I tilted those CM 572 Haplo caudals so that their neural canals were horizontal partly because that’s the preferred orientation that Mike and I landed on in the course of this work, but also partly because to me, that’s a more arresting image than the preceding ones with the articular faces held vertically. I’m both freaked out and fascinated, and that seems like a promising combination — there are mysteries here that cry out to be solved.
As usual, we have loads of people to thank. In addition to all those listed in the Acknowledgments of the new paper, I’m grateful to Matt Lamanna and Amy Henrici of the Carnegie Museum of Natural History for letting me play with study the Haplo specimens in their care. Mike and I also owe a huge thanks to the editorial team at the Journal of Paleontological Techniques. We reached out to them a few days ago to ask if it might be possible to get our in-press paper done and out in time for SV-POW!’s anniversary weekend, and they pitched in to make it happen.
What’s next? We weighed the evidence and formulated what the best solution we could think of. Now it’s up to the world to decide if that was a useful contribution. The comment thread is open — let’s find out.
Things to Make and Do, part 31: redneck CT-scan
March 3, 2022
For reasons that would be otiose, at this moment, to rehearse, I recently found myself in need of a hemisected turkey cervical. Happily, I own five skeletonised turkey necks, so it was with me the work of a moment to select a candidate. But now what? How to hemisect it? We have discussed plenty of hemisected things here at SV-POW!, but they tend to have been produced using heavy machinery such as a band saw: something that I singularly lack.
SPOILER: I found a way. Here is a domestic turkey Meleagris gallopavo domesticus, 9th cervical vertebra, hemisected, in right medial view. Read on to discover the extremely high-tech approach that yielded this prize. It’s propped up on some kind of turkey bone to help me get a good medial perspective, I am thinking maybe the pygostyle?
One idea was to use an angle-grinder: not to cut down the midline of the vertebra — it would be much too blunt and powerful for a small, delicate vertebra — but to use as a sanding surface, locking the grinder in place and holding the vertebra up against the spinning plate. That might work well, assuming I could find a way to secure the angle grinder safely, but as it happened my need for a hemisected vertebra came up during a power cut. (Thanks, Storm Eunice!).
So I did it the way the Amish do their vertebral hemisections: by hand, simply by rubbing the vertebra against a sheet of sandpaper:
CT scanning: the Amish method.
This is not as laborious as you might think. I used a single sheet of medium-grade sandpaper, and it took maybe 15–20 minutes. And I just rubbed back and forth while exerting downward pressure. Initially I worked my way only through the prezygapophyseal ramus, which is the part of the turkey cervical that extends the furthest laterally. Once I was satisfied that the plane between eroded prezyg and the intact postzyg was parasagittal, I just kept the vertebra parallel to the sandpaper and kept rubbing. (Sorry I didn’t think to get a photo at this stage.)
One thing that took me by surprise is that there was so very much bone dust. I mean, I am an idiot that this surprised me, since the whole purpose of this exercise was to reduce one half of this vertebra to bone dust. But the lesson to be learned here is to do it on the easily-cleaned bathroom floor — not on the desk next to the computer keyboard and above a carpet. Learn from my mistakes, folks!
Anyway, after some work on the prezyg/postzyg pair, here’s how the vertebra was looking:
You can see straight away that the prezyg ramus, postzyg ramus and parapophyseal ramus are extensively pneumatized, honeycombed with small, irregular air-spaces. In this image it looks like the region of bone between the pre- and postzygs is much more solid, but this is an illusion: what we’re seeing here is a section through the cortical bone of the neural arch, cut parallel to the surface. Let this be a warning not to over-interpret individual slices of CT-scans!
Once we get a little deeper, we see that the whole wall of the neural arch — and indeed the centrum and the neural spine — is honeycombed, just like the zyg rami:
Now we have another area of what I’m going to call Phantom Apneumaticity: the posterior part of the centrum looks like solid bone, apart from a few pneumatic spaces in the posteroventral extremity. Again, this is an illusion.
Here’s the next place I stopped:
Here, the Phantom Apneumaticity is even more striking: seeing just this as a CT slice could easily mislead someone into thinking that almost the whole of the posteroventral part of the centrum is solid bone. But again, it’s just that we’re very close to the surface of the bone, and seeing a slice parallel to that surface.
This last image also shows an important point of technique: there is a low convex ridge running across the phantom apneumatic area from the top of the cotyle to the base of the centrum. This is where I had changed the angle I was holding the vertebra at, so I accidentally sanded the posteroventral part of the vertebra more than the rest. I found that it was important during this process to keep checking the angles, and to adjust: making sure I wasn’t sanding more from the front than the back, or from the top than the bottom, or leaving a ridge like this.
Also in this last photo you can see that I was just beginning to break through into the neural canal: the anterior part of it is now exposed, between the anterior part of the neural spine and the anterior articular surface. At this stage I sighted along from in front to get a sense of how much further I had to go:
Domestic turkey Meleagris gallopavo domesticus, 9th cervical vertebra, most of right side removed, in anterior view with dorsal to the right. Propped up on the coracoid of a different, larger, turkey.
Quite a way, I guess. Here it is rotated and cropped, so you can more easily recognise it:
Domestic turkey Meleagris gallopavo domesticus, 9th cervical vertebra, most of right side removed, in anterior view. You can see that the neural canal is still mostly intact.
More sanding was required. I sanded some more.
You’ve already seen the final result up at the top of the page, but here is a cleaned-up version of that image, oriented according to Definition 3 of Taylor and Wedel (2019):
Domestic turkey Meleagris gallopavo domesticus, 9th cervical vertebra, hemisected, in right medial view.
And if that isn’t beautiful, what is?
The exciting thing is, anyone can make one of these. Matt’s already explained how to extract and clean up bird vertebrae and given you some ideas of what to do with them. Prepare out some turkey vertebrae and get going with the sandpaper!
I leave you with one more image: the hemisected vertebra in anterior view, oriented with dorsal to the top, and mirrored so it makes up a complete vertebra once more. Enjoy!
References
Fossae/foramina in the articular surfaces of turkey vertebrae
February 21, 2022
I was looking more closely at the turkey skeleton from my recent post, and zeroed in on the last two dorsal (= thoracic) vertebrae. They articulate very well with each other and with the first vertebra of the sacrum, with the centra and zygapophyses both locking in so that there can only have been very little if any movement between them in life. Here they are, in right lateral view:
Last two dorsal (= thoracic) vertebrae of a mature domestic turkey Meleagris gallopavo domesticus, in right lateral view.
Before we move on, it’s worth clicking through to the full-size version of this image and wondering at both the quality of modern phone cameras (a Pixel 3a in this case) and the variety of textures on these little bones. There is smooth, finished bone on the sides of the neural spines; very fine pits and bumps on the zygapophyseal facets where the thin layer of hyaline cartilage attached; rougher texture in the parapophyseal facets where thicker cartilage attached; and very rough texture on the ends of the transverse processes, where there was relatively thick cartilage.
And there is, unsurprisingly in a bird, pneumaticity everywhere. In the more anterior vertebra alone (to the right) the photo shows pneumatic openings (from bottom to top) low on the centrum (below the parapophysis), high on the centrum (below the lateral process), in the hollow between the lateral process, the posyzyg and the centrum, on the lateral surface of the prezygapophyseal ramus, and on the rear surface of the lateral process. There are others that are obscured in this photo, including on top of the lateral process where it meets the neural spine. Here they are, pointed out for you (with the hidden one shown translucently):
Last two dorsal (= thoracic) vertebrae of a mature domestic turkey Meleagris gallopavo domesticus, in right lateral view. Pneumatic openings on penultimate vertebra highlighted with red lines; obscured opening above lateral process shown as translucent.
OK, that was the B-movie. Now to the main feature. The next photo shows the same two vertebrae, folded away from each other so that we see the anterior face of the posterior vertebra (on the left) and the posterior face of the anterior vertebra (on the right).
Last two dorsal (= thoracic) vertebrae of a mature domestic turkey Meleagris gallopavo domesticus. Left: last dorsal vertebra in anterior view; right: penultimate dorsal vertebra in posterior view.
Again, do click through to see the exquisite detail, especially the complex of pneumatic features on the anterior face of the neural spine of the last dorsal (on the left) and on the posterior face of the left lateral process of the penultimate dorsal (on the right).
And … in the articular facets of the centra?
Seriously, what the heck is going on here? It doesn’t make sense that there would be pneumatic openings in articular surfaces, because by definition something else (in this case the adjacent vertebra) is abutted hard up against then, so there is no way for a diverticulum to get in. For the same reason, you don’t get vascular foramina in articular surfaces because there is no way to get an artery in there. And there is no hint in these vertebrae of channels along either articular surface that diverticula or arteries could possibly have laid in.
And yet, there those big openings are. What are they?
I discussed this with Matt, in case it’s Well Known Phenomenon that I’d somehow not heard about but it seems it is not. What we know for sure is that these openings are present, and that they are not mechanical damage inflicted during preparation. So what are they?
What else even is there for them to be? What penetrates bone apart from diverticula and blood vessels? Nerves follow the blood vessels, so it can’t be nerves in the absence of blood vessels.
By the way, there are similar but smaller openings in the posterior face of the last dorsal (the one on the right in the photo), but none anywhere else along the postcervical column: not on the anterior surface of the penultimate dorsal, not on the front or back of the sacrum, and not in any of the other dorsals.
One possibility we considered is that the vertebrae were locked together in life and that a pneumatic space inside the centrum of the last dorsal worked right through into the penultimate one. But that doesn’t work: the openings are not aligned. Also, those in the penultimate dorsal are definitely blind (i.e. they do not connect to deeper internal air-spaces) and those in the last dorsal probably are, too.
We do not know what is going on here.
Help us! Is this kind of thing common in turkeys? Have people seen it in other taxa? Do we know what it is?
Turkey skeleton audit
February 16, 2022
Back in at least 2008 — maybe earlier — I kept all the bones from our good-sized Christmas turkey. Of course, it’s missing the head, neck and feet, but otherwise it’s pretty much all there. (I may also have the neck, but if so then it was supplied as a separate item, and prepared separately.) Here is the box of postcervical bones:
Postcervical skeleton of a mature domestic turkey Meleagris gallopavo domesticus, complete except for feet.
As I was transferring them to a better box today, it occurred to me to lay them out and see how much sense I could make of them. Here’s what I did with the bones I was happy about:
Postcervical skeleton of a mature domestic turkey Meleagris gallopavo domesticus, complete except for feet. Some small bones omitted (see below). Laid out roughly as in life, in dorsal view.
I should have put something in the photo to act as a scale-bar, because it’s not apparent from this photo that a turkey is a pretty big thing. From top to bottom of the skeleton as I laid it out here is about 90 cm.
Here are my (in some cases tentative) identifications of the bones:
Postcervical skeleton of a mature domestic turkey Meleagris gallopavo domesticus, complete except for feet. Some small bones omitted (see below). Laid out roughly as in life, in dorsal view. Bones are tentatively labelled. DO NOT USE FOR TUTORIAL PURPOSES.
Are there any obvious mistakes in there? And have I got any of the bones left-right reversed?
Now, here are all the other bones that I was less confident about the positions of:
Postcervical skeleton of a mature domestic turkey Meleagris gallopavo domesticus, complete except for feet. Small bones only.
On the left of course we have the dorsal ribs, but I’ve not been able to arrange them all into near pairs, nor figure out what order I should impose on the pairs that I do have. I’m not even sure how many pairs I should have. On the right are other paired bones whose identity I can’t figure out. I am guessing that the longer ones are probably sternal ribs and that the irregularly shaped ones might be parts of the wrist, but I would welcome corrections and clarifications. Finally, the middle column contains the bones whose idea I have little or no idea about, and which don’t appear to be paired. Any ideas?
By the way, I found this image useful in figuring out the identities of the appendicular bones:
Skeletal reconstruction of a domestic turkey, published in 1808. Source unknown (leave a comment if you can identify it).
And this one useful for the bones of the wing and especially the hand:
Human and bird arm skeletons compared. Source unknown.
More from this skeleton another time!
Turkeys lie
December 18, 2018
We all know what turkeys look like, right?
Turns out that two thirds of that bird is a lie. Here’s a diagram produced for hunters on which part of the turkey to shoot. (It’s all over the Internet, and I can’t trace the original source, but I got it from here):
Bascially, if you fire an arrow at a visible turkey, there’s a 2/3 chance that it’ll pass straight through feathers and completely miss the actual bird.
Now, then: what do we think a theropod looked like in life? Probably not much like what skeleton reconstructions show as the flesh envlope, as for example in Scott Hartman’s Guanlong:
Instead, it might have looked like this:
(Note: this is not in any way a criticism of Scott’s fine work, which is a scientific restoration of the soft tissue, and does not address integument at all.)
And now that pterosaurs have feathers, too(*), we have to assume that they, too, probably had body outlines bearing little resemblance to the flesh-on-bone shapes we’ve been used to seeing.
(*) As Matt pointed out: “I can’t be bothered to write “integumentary structures” when I mean “feathers”. I realize they may be independently derived, but eyes evolved independently like 40 times and we don’t refer to the other 39 instances as “photoreceptive structures”.” (He actually wrote “I can’t be arsed”, but I changed it to “bothered” to make him appear more professional.)
Epipophyses, the forgotten apophyses: not just for sauropods!
February 2, 2015
Matt’s last post contained a nice overview of the occurrence of epipophyses in sauropodomorphs: that is, bony insertion points for epaxial ligaments and muscles above the postzygapophyseal facets. What we’ve not mentioned so far is that these structures are not limited to sauropods. Back when we were preparing one of the earlier drafts of the paper that eventually became Why sauropods had long necks; and why giraffes have short necks (Taylor and Wedel 2013a), I explored their occurrence in related groups. But that section never got written up for the manuscript, and now seems as good a time as any to fix that.
Theropods (including birds)
Most obviously, epipophyses occur in theropods, the sister group of sauropodomorphs.
Taylor and Wedel (2013a: figure 11). Archosaur cervical vertebrae in posterior view, Showing muscle attachment points in phylogenetic context. Blue arrows indicate epaxial muscles attaching to neural spines, red arrows indicate epaxial muscles attaching to epipophyses, and green arrows indicate hypaxial muscles attaching to cervical ribs. While hypaxial musculature anchors consistently on the cervical ribs, the principle epaxial muscle migrate from the neural spine in crocodilians to the epipophyses in non-avial theropods and modern birds, with either or both sets of muscles being significant in sauropods. 1, fifth cervical vertebra of Alligator mississippiensis, MCZ 81457, traced from 3D scans by Leon Claessens, courtesy of MCZ. Epipophyses are absent. 2, eighth cervical vertebra of Giraffatitan brancai paralectotype HMN SII, traced from Janensch (1950, figures 43 and 46). 3, eleventh cervical vertebra of Camarasaurus supremus, reconstruction within AMNH 5761/X, “cervical series I”, modified from Osborn and Mook (1921, plate LXVII). 4, fifth cervical vertebra of the abelisaurid theropod Majungasaurus crenatissimus,UA 8678, traced from O’Connor (2007, figures 8 and 20). 5, seventh cervical vertebra of a turkey, Meleagris gallopavo, traced from photographs by MPT.
In this figure from the 2013 paper, the rightmost images show cervical vertebrae of Majungasaurus (an abelisaurid theropod) and a turkey, both in posterior view. The red arrows indicate epaxial musculature pulling on the epipophyses. They are particularly prominent in Majungasaurus, rising almost a full centrum’s height above the postzygapophyseal facets.
The epipophyses are very prominent in the anterior cervicals of Tyrannosaurus, but much less so in its posterior cervicals — presumably because its flesh-tearing moves involved pulling upwards more strongly on the anterior part of the neck. Here’s a photo of the AMNH mount, from our post T. rex‘s neck is pathetic:
You can see something similar in the neck of Allosaurus, and the trend generally seems to be widespread among theropods.
Ornithischians
Note the very prominent epipophyses protruding above the postzygs in the anterior cervicals of this Heterodontosaurus in the AMNH public gallery:
Cast of AMNH 28471, Heterodontosaurus tucki, collected from the Early Jurassic Voisana, Herschel district, South Africa. Neck in left lateral view.
Here’s the hadrosaur Corythosaurus:
AMNH 5338, Corythosaurus casuarius, from the Campanian of the Red Deer River, Alberta, Canada. Collected by Barnum Brown and P. C. Kaisen, 1914. Cervicals 1-4 in right lateral view.
The prominent vertebra is C2: note that is has both a modest blade-like neural spine and prominent epipophyses — but that already by C3 the epipophyses are gone. Here is that C2 postzyg/epipophyses complex is close-up, clearly showing anteroposteriorly directed striations on the epipophysis, presumably representing the orientation of the attaching ligaments and muscles:
As previous image: close-up of posterior part of C2.
Here’s a close-up of the neck of the boring ornithopod Tenontosaurus, also in the AMNH gallery. (I’m not sure of the specimen number — if anyone can clarify, please leave a comment).
AMNH ?3554, Tenontosaurus tilletti, cervicals 2-4 in right lateral view.
The interesting thing here is that it its axis (C2) seems to lack epipophyses (unlike C3), and to have a tall blade-like neural spine, as seen in mammals. We don’t really see C2 spines this big in other dinosaurs — compare with the much more modest spine in Corythosaurus, above. The texture of this part of the Tenontosaurus specimen looks suspicious, and I wonder whether that neural spine is a fabrication, created back in the day by AMNH staff who were so used to mammals that they “knew” what a C2 should look like? Anyway, the epipophysis above the postzyg of C3 is very distinct and definitely real bone.
Pterosaurs
Things get much more difficult with pterosaurs, because their cervicals are so fragile and easily crushed (like the rest of their skeleton, to be fair). While it’s easy to find nice, well-preserved ornithischian necks on display, you don’t ever really see anything similar for pterosaurs.
As a result, we have to rely on specimen photographs from collections, or more often on interpretive drawings. Even high-resolution photos, such as the one in Frey and Tischlinger (2012: fig 2) tend not to show the kind of detail we need. Usually, the only usable information comes from drawings made by people who have worked on the specimens.
Here, for example, is Rhamphorhynchus, well known as the most difficult pterosaur to spell, in figure 7 from Bonde and Christiansen’s (2003) paper on its axial pneumaticity:
It’s not the main point of the illustration, but you can make out clear epipophyses extending posteriorly past the postzygapophyseal facets in at least C3 and C5 — in C4, the relevant area is obscured by a rib. (Note that the vertebrae are upside down in this illustration, so you need to be looking towards the bottom of the picture.)
I’m pretty sure I’ve seen a better illustration of Rhamphorhynchus epipophyses, but as I get older my memory for Rhamphorhynchus epipophyses is no longer what it used to be and I can’t remember where. Can anyone help?
But also of interest is the azhdarchid pterosaur Phosphatodraco, here illustrated by Pereda Suberbiola et al. (2003):
Pereda Suberbiola et al. (2003: fig. 3). Phosphatodraco mauritanicus gen. et sp. nov, OCP DEK/GE 111, Late Cretaceous (Maastrichtian), Morocco: (a) cervical five in two fragments, ventral and left lateral views; (b) cervical six in ventrolateral view; (c) cervical seven in ventral view; (d) cervical eight in left lateral view; (e) cervical nine in posterior view; (f) cervical six in anterior view. c, centrum; co, condyle; ct, cotyle; hyp, hypapophysis; nc, neural canal; ns, neural spine; poe, postexapophysis; poz, postzygapophysis; prz, prezygapophysis; su, sulcus; tp, transverse process.
The cervicals of Phosphatodraco seem to have no epipophyses. So they were not ubiquitous in pterosaurs.
What does it all mean? This post has become a bit of a monster already so I’ll save the conclusion for another time. Stay tuned for more hot epipophyseal action!
References
- Bonde, Niels and Per Christiansen. 2003. The detailed anatomy of Rhamphorhynchus: axial pneumaticity and its implications. pp 217-232 in: E. Buffetaut and J-M Mazin (eds), Evolution and Palaeobiology of Pterosaurs. Geological Society, London, Special Publications 217. doi:10.1144/GSL.SP.2003.217.01.13
- Frey Eberhard and Helmut Tischlinger. 2012. The Late Jurassic Pterosaur Rhamphorhynchus, a Frequent Victim of the Ganoid Fish Aspidorhynchus? PLoS ONE 7(3):e31945. doi:10.1371/journal.pone.0031945
- Janensch, Werner. 1950. Die Wirbelsaule von Brachiosaurus brancai. Palaeontographica, Supplement 7 3:27-93.
- O’Connor Patrick M. 2007. The postcranial axial skeleton of Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. pp 127-162 in: S. D. Sampson., D. W. Krause (eds), Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. Society of Vertebrate Paleontology Memoir 8.
- Osborn, Henry F., and Charles C. Mook. 1921. Camarasaurus, Amphicoelias and other sauropods of Cope. Memoirs of the American Museum of Natural History, New Series 3:247-387.
- Pereda Suberbiola, Xabier, Nathalie Bardet, Stéphane Jouve, Mohamed Iarochène, Baadi Bouya and Mbarek Amaghzaz. 2003. A new azhdarchid pterosaur from the Late Cretaceous phosphates of Morocco. pp 79-90 in: E. Buffetaut and J-M Mazin (eds), Evolution and Palaeobiology of Pterosaurs. Geological Society, London, Special Publications 217. doi:10.1144/GSL.SP.2003.217.01.08
- Taylor, Michael P., and Mathew J. Wedel. 2013. Why sauropods had long necks; and why giraffes have short necks. PeerJ 1:e36 doi:10.7717/peerj.36
Photography and illustration talk, Part 1: Intro and Stromer
February 6, 2014
Recently I had the opportunity to give a talk on photographing specimens and preparing illustrations in Jim Parham‘s phylogenetics course at Cal State Fullerton. Jim is having each student (1) write a description of a specimen, (2) run a phylogenetic analysis, and (3) do some kind of calibration on their tree. I think that’s rad.
Anyway, it was fun talk and I wanted to put it up for everyone, but Mike had the idea of posting a batch of slides at a time, to hopefully fire some discussion on different aspects of photography and illustration. So if you’re impatient to see the whole thing, blame him! I will post the whole talk at the end of the post series, and you can find all of the talk posts here.
And now, on with the show.
Tutorial 21: how to measure the length of a centrum
March 11, 2013
For a paper that I and Matt are preparing, we needed to measure the centrum length of a bunch of turkey cervicals. That turns out to be harder than you’d think, because of the curious negative curvature of the articular surfaces.
Above is a C7 from a turkey: anterior view on the left; dorsal, left lateral and ventral views in the middle row; and posterior on the right. As you can see from the anterior, dorsal and ventral views, the anterior articular surface[1] is convex dorsoventrally but concave transversely; and as you can see from the lateral view, the posterior face is concave dorsoventrally and convex transversely.
This means you can’t just put calipers around the vertebra. If you approach the vertebra from the top or bottom, then the upper or lower lip of the posterior articular surface will protrude past the centre of the saddle, and give you too long a length. If you approach from the side, the same will happen with the left and right lips of the anterior articular surface.
What are we trying to measure anyway?
But this raises the question of what it is we’re trying to measure. I said “we needed to measure the centrum length of a bunch of turkey cervicals”, but what exactly is centrum length? Why shouldn’t the upper and lower lips of the posterior articular surface count towards it?
What does centrum length mean?
The problem doesn’t only arise with bird cervicals. The same issue arises in measuring more sensible and elegant vertebrae, such as our old friend HMN SII:C8, or MB.R.2181:C8 as we must now learn to call it.
Although the back of the vertebra is nice and simple here — it’s obvious what line we’re measuring to at the back — we have three choices of where the “front” of the vertebra is, and a case can be made for any of them as being “the length of the vertebra”.
The longest measurement (here marked “T” for “total length”) goes to the front of the prezygapophyseal rami. The next one (“C” for “centrum length”) goes to the anteriormost point of the condyle. The distinction is important: as noted recently, the longest vertebra in the world belongs to Sauroposeidon if we use total length, but to Supersaurus if we use centrum length.
But in life, most of the condyle would be buried in the cotyle of the preceding vertebra. So should it count towards the length of the vertebra? If you consider a string of articulated vertebrae, the buried condyles don’t contribute to the overall length of the neck. So Matt and I call the length from the posterior margin of the condyle to the posterior margin of the cotyle the functional length (marked “F” above), which I believe is a new term.
Another way to think of the functional length is the distance from a given point on a vertebra (in this case the posterior margin of the cotyle) to the same point on the adjacent vertebra:
For our current project, Matt and I are interested in how the lengths of individual vertebrae contribute to total neck length, so for our purposes, functional length is definitely what we want.
By the way, Janensch is the only author I know of to have even recognised the importance of functional length. The measurement tables on pages 39 and 44 have columns for “Gesamtlänge des Wirbels ab Vorderende per Präzygapophyse”, “Gesamtlänge der Wirbel-Körpers in 1/2 Höhe” and “Länge der Wirbel-körpers ohne Condylus in 1/2 Höhe” — that is, “Total length of the vertebra from the anterior end of the prezygapophysis”, “Total length of the centrum measured at mid-height” and “Length of the centrum minus condyle at mid-height”. This is typical of his careful and methodical approach. Kudos!
Hey! I thought this was about turkeys
And so it is. Here is the functional length measurement for a turkey cervical:
It’s the shortest anteroposterior distance between the two articular surfaces.
Measuring functional length
Matt and I chatted about this at some length, and I am ashamed to say that we thought through all sorts of complicated solution involving subtracting measurements from known scaffold length and suchlike.
It took us a stupidly long to to arrive at the very obvious solution, which is just to modify the calipers to have a “tooth” that can protrude into the concavity of the anterior articulation between its left and right lips. Easily done with a flat-ended screw and a blob of wood glue:
With the measurements of all the vertebrae in my series, I can now fairly confidently expect that the sum of the individual lengths will come out at about the length of the complete neck.
You know, unless intervertebral cartilage turns out to be important or something.
References
- Janensch, Werner. 1950. Die Wirbelsaule von Brachiosaurus brancai. Palaeontographica (Suppl. 7) 3:27-93.
Footnotes
1. Matt and I are so used to opisthocoelous sauropod presacrals that when we’re talking about vertebrae — any vertebrae — we tend to say “condyle” and “cotyle” for the anterior and posterior articular surfaces, no matter what their morphology. When talking about crocodile cervicals or titanosaur caudals, we’re even likely to say ridiculous things like “the condyle is concave and the cotyle is convex”. Nonsense, of course: condyle means “A rounded prominence at the end of a bone, most often for articulation with another bone.” What we should say is “the condyle is at the back and the cotyle is in front”.
Tutorial 20: how to measure necks using Duplo
January 10, 2013
I’ve measured a few necks in my time, including the neck of a baby giraffe. I can tell you from experience that necks are awkward things to measure, even if they have been conveniently divested of their heads and torsos. They have a tendency to curl up, which impedes attempts to find the straight-line length. Even when you manage to hold them straight, you want them maximally compressed end-to-end rather than stretched out, which is hard to achieve without buckling them out of the straight line. And then you need to measure between perpendiculars in a straight line.
Awkward.
Tonight, I needed to measure the mass and length of seven turkey necks. (Never mind why, all will become clear in time.) And I found a way to do it that works much better than anything I’ve done before.
Here’s the equipment:
You will need:
- Kitchen scales (for weighing the necks)
- Small numbered labels (for the sandwich bags that the necks will go into for the freezer once they’ve been measured)
- Pen and paper to take down the measurements
- Translucent ruler
- Saucepan full of turkey necks
- Slightly less than one half of a birthday cake decorated like a map of Middle-earth [optional]
- A Duplo baseboard (double-sized Lego) and about fifteen 4×2 bricks
Use the bricks to build an L-shaped bracket on the board — about half way back, so that can rest your hand in front of it.
Now you can push the neck into the angle of the bracket. By keeping it pressed firmly against the back wall (yellow in my construction), you can keep it straight. I find the best way to get the neck exactly abutting the left (red) wall is to start with the neck in its natural position, with the anterior and posterior ends curving towards you, then sort of unroll it against the back wall, and finally push the posterior end into place with your little finger (see below). There is a satisfying moment– almost a click — as the back end pops into place and the neck slides along a little to right as necessary to accommodate the added length.
Now use another brick (blue in this photo) as a bracket: slide it along the back wall from right to left until it’s solidly abutting the anteriormost vertebra. If you do this right, there is very little travel: the entire series of vertebrae is lined up and solidly abutted, with bone pushing against the left wall and your new brick. I find there’s less than half a millimeter of variation between the length under gentle-but-firm pressure (which is what I measured) and under the very strongest force you can exert without buckling the neck.
Once you have found the blue brick’s correct position, you need to hold it firmly in place and measure its position relative the the left wall. (It doesn’t matter if you let the neck re-curl at this point, so long as the blue brick doesn’t shift.)
You need a translucent ruler so that you can lay it across the neck and see where blue brick falls under the scale. (My ruler’s zero is, rather annoyingly, 5 mm from the end; so I needed to subtract 5 mm from the lengths I measured.)
Finally, I bagged up each neck in its own sandwich bag, ready for the freezer. Each neck is labelled with a number so that when I take it out for dissection, I will be able to relate the measurements and observations that I make back to these initial measurements.
For the record, here are the measurements:
- Neck 1: 154 g, 179.5 mm.
- Neck 2: 122 g, 151 mm.
- Neck 3: 154 g, 199.5 mm.
- Neck 4: 133 g, 162.5 mm.
- Neck 5: 142 g, 169 mm.
- Neck 6: 80 g, 167 mm.
- Neck 7: 70 g, 169 mm.
As expected, there is some correlation between neck mass and length; but not as much as you might expect. Naively (i.e. assuming isometric similarity) mass should be proportional to length cubed, but there is a lot of scatter about that line. I don’t know whether that is due to individual variation, or merely because the various necks — all of them incomplete — are different sections of the full neck. Hopefully I will be able to confirm or rule out that possibility when I’ve dissected down to naked vertebrae.
Sauropod Vertebra Picture of the Week 
