Anatomical features of the neural canal in birds and other dinosaurs. A. MWC 9698, a mid caudal vertebra of Apatosaurus in posterodorsal view. Arrows highlight probable vascular foramina in the ventral floor of the neural canal. B. LACM 97479, a dorsal vertebra of Rhea americana in left anterolateral view. Arrows highlight pneumatic foramina inside the neural canal. C. A hemisected partial synsacrum of a chicken, Gallus domesticus, obtained from a grocery store. Anterior is to the right. The bracket shows the extent of the dorsal recess for the glycogen body, which only spans four vertebrae. Arrows highlight the transverse grooves in the roof of the neural canal for the lumbosacral organ. D. Sagittal (left) and transverse (right) CT slices through the sacrum of a juvenile ostrich, Struthio camelus. The bracket shows the extent of the lumbosacral expansion of the spinal cord. Indentations in the roof of the neural canal house the lumbosacral organ. In contrast to the chicken, the ostrich has a small glycogen body that does not leave a distinct osteological trace. Yellow arrows show the longitudinal troughs in the ventral floor of the neural canal that house the ventral eminences of the spinal cord. Wedel et al. (2021: fig. 4).

This is the second in a series of posts on our new paper about the expanded neural canals in the tail vertebrae of the Snowmass Haplocanthosaurus. I’m not going to talk much about Haplo in this post, though. Instead, I’m going to talk about chickens, and about how you can see a lot of interesting spinal anatomy in a living dinosaur for about two bucks.

You know by now that Academia Letters publishes peer reviews, which is one of the things that drew me to this fairly new journal. More on that in a later post, but in the meantime, the peer reviews for the Haplo paper are on the right sidebar here. I confess, I had a total forehead-slap moment when I read the opening lines of Niels Bonde’s review: 

This paper is interesting, and should be published and discussed by others with interest in dinosaur-bird relations. However, as these publications are also meant for the general public, I would recommend that 2 – 3 illustrations were added of the features mentioned for birds under nos. 3 – 6, because the general public (and many paleontologists) have no ideas about these structures, and what they look like.

The original submission only had figures 1 and 2. And this request is totally fair! If you are going to discuss six alternative hypotheses for some mysterious anatomical structure, it’s just responsible reporting to illustrate those things. That goes double if, as Niels Bonde noted, the anatomy in question is unfamiliar to a lot of people, even many paleontologists. Huxley’s quote after first reading Darwin’s Origin of Species flashed through my head: “How extremely stupid not to have thought of that.”

Slide 21 of my 2014 SVPCA talk on supramedullary diverticula in birds and other dinosaurs, illustrating pneumatic foramina in the roof, walls, and floor of the neural canal.

At the time I read that review, I already had images illustrating five of the six hypotheses. A juvenile ostrich synsacrum that Jessie Atterholt and I had CT scanned gave us three of them all by itself: the lumbosacral expansion of the spinal cord to run the hindlimbs, as in all limbed tetrapods and in some fish with sensitive fins; the transverse channels in the dorsal wall of the neural canal to accommodate the lumbosacral balance organ; and the paired troughs in the floor of the neural canal that house the ventral eminences of the spinal cord (Figure 4D in the image at the top of this post). I had good photos of pneumatic foramina in the walls and floor of the neural canal in a dorsal vertebra of a rhea from my 2014 SVPCA talk (Figure 4B), and some photos of small foramina, presumably for blood vessels rather than air spaces, in the floor of the neural canal in a caudal vertebra of Apatosaurus (Figure 4A).

What I did not have is a photo illustrating the fairly abrupt, dome-shaped space in the sacral neural canal that houses the glycogen body of birds. I mean, I had published images, but I didn’t want to wrestle with trying to get image reproduction rights, or with redrawing the images. Instead, I went to the grocery store to buy some chicken.

I don’t know how universally true this is, but IME in the US when you buy a quartered chicken, the vertebrae are usually nicely hemisected by the band saw that separated the left and right halves of the animals. So you can see the neural canal in both the dorsal and sacral parts of the vertebral column. Here are the hemisected dorsal vertebrae in the breast quarter from a sectioned rotisserie chicken:

That’s just how it came to lie on my plate, but it’s not in anatomical position. Let’s flip it over to sit upright:

And label it:

I could and probably should do a whole post just unpacking this image, but I have other fish to fry today, so I’ll just note a couple of things in passing. The big interspinous ligament is the same one you can see in transverse section in the ostrich dissection photos in this post and this one. Also, the intervertebral joints heading toward the neck, on the left of the image, have much thicker intervertebral cartilage than the more posterior dorsals. That’s because the posterior ones were destined to fuse into a notarium. You can see a diagram and a photograph of a chicken notarium in figures 4 and 5, respectively, here. And finally, the big takeaway here is that the neural canal is normal, just a cylindrical tube to hold the spinal cord.

The thigh quarter usually has the pelvis and the hemisectioned synsacrum attached. Here’s a lateral view of the left half of the pelvis and synsacrum:

And the same thing labeled:

And now flipped around so we can see it in medial view:

And now that image labeled:

And, hey, there are three of our alternative hypotheses on display: the long (many vertebral segments) lumbosacral expansion of the spinal cord, which is reflected in a gradually expanded neural canal in the synsacrum; the shorter, higher dome-shaped recess for the glycogen body; and finally the transverse spaces for the lumbosacral balance organ.

As a refresher, there’s nothing terribly special about the lumbosacral expansion of the spinal cord — you have one, labeled as the ‘lumbar enlargement’ in the above diagram. Where the spinal cord has adjacent limbs to run, it has more neurons, so it gets fatter, so the neural canal gets fatter to accommodate it. The cord itself doesn’t look very expanded in the chicken photo above, but that chicken has been roasted rotisserie-style, and a lot of lipids probably cooked out of the cord during that process. What’s more important is that the neural canal is subtly but unmistakably expanded, over the span of many vertebrae.

The lumbosacral spinal cord of a 3-week-old chick in dorsal view. The big egg-shaped mass in the middle is the glycogen body. Watterson (1949: plate 1).

That’s in contrast to the recess for the glycogen body, which is colored in blue in the chicken photo. Glycogen bodies, like the egg-shaped one in the young chicken in the image immediately above, tend not to go on for many vertebral segments. Instead they balloon up and subside over the space of just 4 or 5 vertebrae, so they leave a different skeletal trace than other soft tissues.

Finally, there are the transverse spaces for the lumbosacral balance organ, which I discussed in this post. Those are the things that look like caterpillar legs sticking up from the sacral endocasts in the above figure from Necker (2006). In life, the spaces are occupied by loops of meningeal membranes, through which cerebrospinal fluid can slosh around, which in turn puts pressure on mechanoreceptive cells at the edge of the spinal cord and gives birds a balance organ in addition to the ones in their heads. In the photo of the cooked chicken, the delicate meninges have mostly fallen apart, leaving behind the empty spaces that they once occupied.

I really liked that chicken synsacrum, and I wanted to use it as part of Figure 4 of the new paper, but it needed a little cleaning, so I simmered it for a couple of hours on low heat (as one does). And it promptly fell apart. At least in the US, most of the chickens that make it to table are quite young and skeletally immature. That particular bird’s synsacrum wasn’t syn-anything, it was just a train of unfused vertebrae that fell apart at the earliest opportunity. I had anticipated that might be an issue, so I’d gotten a lot of chicken, including a whole rotisserie chicken and four thigh quarters from the deli counter at the local supermarket. Happily this fried chicken thigh quarter had a pretty good neural canal:

And it cleaned up nicely:

And with a little cropping, color-tuning, and labeling, it was ready for prime time:

I didn’t label them in the published version, for want of space and a desire not to muddy the waters any further, but the jet-black blobs I have colored in the lower part of that image are the exit holes that let the spinal nerves out of the neural canal so they could go serve the hindlimbs, pelvic viscera, and tail. We have them, too.

At my local grocery store, a fried chicken thigh costs about $1.65 if you get it standalone, or you can buy in bulk and save. You get to eat the chicken, and everything else I’ve done here required only water, heat, soap, and a little time. The point is that if I can do this, you can do this, and if you do, you’ll get to see some really cool anatomy. I almost added, “which most people haven’t seen”, but given how much chicken we eat as a society these days, probably most people’s eyes have fallen on the medial surface of a cooked chicken thigh quarter at one time or another. Better to say, “which most people haven’t noticed”. But now you can. Go have fun. 

Way back in January of 2019, I finished up “Things to Make and Do, Part 25b” with this line: “I have one more thing for you to look for in your bird vertebrae, and that will be the subject of the next installment in this series. Stay tuned!” Here we are, 2.3 years later, and I’ve finally made good. So if there’s a promised post you’ve been waiting for, stick around, we may get to it yet.


A. Recovered skeletal elements of Haplocanthosaurus specimen MWC 8028. B. Caudal vertebra 3 in right lateral view. C. The same vertebra in posterior view. Lines show the location of sections for D and E. D. Midsagittal CT slice. The arrow indicates the ventral expansion of the neural canal into the centrum. E. Horizontal CT slice at the level of the neural arch pedicles, with anterior toward the top. Arrows indicate the lateral expansions of the neural canal into the pedicles. B-E are shown at the same scale. Wedel et al. (2021: fig. 1).

New paper out today:

Wedel, Mathew; Atterholt, Jessie; Dooley, Jr., Alton C.; Farooq, Saad; Macalino, Jeff; Nalley, Thierra K.; Wisser, Gary; and Yasmer, John. 2021. Expanded neural canals in the caudal vertebrae of a specimen of Haplocanthosaurus. Academia Letters, Article 911, 10pp. DOI: 10.20935/AL911 (link)

The paper is new, but the findings aren’t, particularly. They’re essentially identical to what we reported in our 1st Paleo Virtual Conference slide deck and preprint, and in the “Tiny Titan” exhibit at the Western Science Center, just finally out in a peer-reviewed journal, with better figures. The paper is open access and free to the world, and it’s short, about 1600 words, so this recap will be short, too.

A. Photograph of a 3D-printed model of the first three caudal vertebrae of Haplocanthosaurus specimen MWC 8028, including endocasts of the neural canal (yellow) and intervertebral joints (blue), in right lateral view, and with the neural canal horizontal. B. Diagram of the same vertebrae in midsagittal section, emphasizing the volumes of the neural canal (yellow) and intervertebral joint spaces (blue). Anterior is to the right. Wedel et al. (2021: fig. 2).

John Foster and I described Museum of Western Colorado (MWC) specimen 8028, a partial skeleton of Haplocanthosaurus from Snowmass, Colorado, in late 2014. One weird thing about that specimen (although not the only weird thing) is that the neural canals of the tail vertebrae are bizarrely expanded. In most vertebrae of most critters, the neural canal is a cylindrical tunnel, but in these vertebrae the neural canals are more like spherical vacuities.

John and I didn’t know what to make of that back in 2014. But a few years later I started working with Jessie Atterholt on bird anatomy, which led me to do a little project on the whole freaking zoo of weird stuff that birds and other dinosaurs do with their neural canals, which led to the 1PVC presentation, which led to this. 

Caudal vertebra 3 of Haplocanthosaurus specimen MWC 8028 in left posterolateral (A), posterior (B), and right posterolateral (C) views, with close-ups (D and E). In A and B, a paintbrush is inserted into one of the lateral recesses, showing that the neural canal is wider internally than at either end. Wedel et al. (2021: fig. 3).

Of course there will be more posts and more yapping, as signaled by the ‘Part 1’ in the post title. Although I am extremely satisfied with the streamlined, 1600-word missile of information and reasoning that just dropped, there are parts that I want to unpack, that haven’t been unpacked before. But the paper launched at midnight-thirty, Pacific Daylight Time, I’m up way too late finishing this first post, and I reckon the rest will keep for a few hours at least.

Anatomical features of the neural canal in birds and other dinosaurs. A. MWC 9698, a mid caudal vertebra of Apatosaurus in posterodorsal view. Arrows highlight probable vascular foramina in the ventral floor of the neural canal. B. LACM 97479, a dorsal vertebra of Rhea americana in left anterolateral view. Arrows highlight pneumatic foramina inside the neural canal. C. A hemisected partial synsacrum of a chicken, Gallus domesticus, obtained from a grocery store. Anterior is to the right. The bracket shows the extent of the dorsal recess for the glycogen body, which only spans four vertebrae. Arrows highlight the transverse grooves in the roof of the neural canal for the lumbosacral organ. D. Sagittal (left) and transverse (right) CT slices through the sacrum of a juvenile ostrich, Struthio camelus. The bracket shows the extent of the lumbosacral expansion of the spinal cord. Indentations in the roof of the neural canal house the lumbosacral organ. In contrast to the chicken, the ostrich has a small glycogen body that does not leave a distinct osteological trace. Yellow arrows show the longitudinal troughs in the ventral floor of the neural canal that house the ventral eminences of the spinal cord. Wedel et al. (2021: fig. 4).

I have a ton of people to thank. John Foster, obviously, for initiating the line of research that led here. Julia McHugh for access to the MWC collections, and for being an excellent sounding board regarding the Morrison Formation, sauropod dinosaurs, and crafting ambitious but tractable research projects. Anne Weil for helping me be methodical in thinking through the logic of the paper, and Mike Taylor for helping me get it polished. Niels Bonde, Steven Jasinski, and David Martill for constructive reviews, which were published alongside the paper. We couldn’t take all of their suggestions because of space limitations, but figures 3 and 4 were born because they asked for them, and that’s not a small thing. Vicki and London Wedel for putting up with me at various points in this project, especially in the last few days as I’ve been going bonkers correcting page proofs. And finally, because I’m the one writing this blog post, my coauthors: Jessie Atterholt, Alton Dooley, Saad Farooq, Jeff Macalino, Thierra Nalley, Gary Wisser, and John Yasmer, for their contributions and for their patience during the unusually long gestation of this very short paper.

More to say about all that in the future. For now, yay, new paper. Have fun with it. Here’s the link again.


This is the Jurassic World Legacy Collection Brachiosaurus. I think it might be an exclusive at Target stores here in the US. It turns up on other sites, like Amazon and eBay, but usually from 3rd-party sellers and with a healthy up-charge. Retails for 50 bucks. I got mine for Christmas from Vicki and London. Here’s the link to if you want to check it out (we get no kickbacks from this).

I thought it would be cool to leverage this thing at outreach events to talk about the new Brachiosaurus humerus that Brian Engh found last year, which a team of us got out of the ground and safely into a museum last October (full story here). But I needed a Brachiosaurus humerus, so I made one, and in this post I’ll show you how to do the same, for next to no money.

Depending on what base you start with and what materials you use, you could build a scale model of a Brachiosaurus humerus at any size. I wanted one that would match the JWLC Brach, so I started by taking some measurements of that. Here’s what I got:


  • Head: 45mm
  • Neck: 455mm (x 20 = 9.1m = 29’10”)
  • Torso: 320mm
  • Tail: 320mm
  • Total: 1140 (x 20 = 22.8m = 74’10”)


  • Max head height: 705mm (x 20 = 14.1m = 46’3″)
  • Withers height: 360mm (x 20 = 7.2m = 23’7″)

The neck length, total length, and head height are pretty close to the mounted Giraffatitan in Berlin. The withers are a little high, as is the bottom of the animal’s belly. I suspect that the limbs on the model are oversized by about 10%. Nevertheless, the numbers say this thing is roughly 1/20 scale.

The largest humeri of Brachiosaurus and Giraffatitan are 213cm, which is about 3mm shy of 7 feet. So a 1/20 scale humerus should be 106.5mm, or 4.2 inches, or four-and-a-quarter if you want a nice, round number.

Incidentally, Chris Pratt is 6’2″ (74 inches), and the Owen Grady action figure is 3.75″, which is 1/20 of 6’3″. So the action figure, the Brachiosaurus toy highly detailed scientific model, and a ~4.2″ humerus model will all be more or less in scale with each other.

I used a chicken humerus for my base. The vast majority of chickens in the US are slaughtered at 5 months, so they don’t get nearly big enough for their humeri to be useful for this project. Fortunately, there’s a pub in downtown Claremont, Heroes & Legends, that has giant mutant chicken hot wings, so I went there and collected chicken bones in the guise of a date. The photo above shows three right humeri (on the left) and one left humerus (on the right) after simmering and an overnight degreasing in a pot of soapy water. I used the same bone clean-up methods as in this post.

What should you do if you don’t have access to giant mutant chicken wings? My method of Brachio-mimicry involves some sculpting, so any reasonably straight bone that bells out a bit at the ends would work. You could use a drumstick in a pinch. Here are my humeri whitening in a tub of 3% hydrogen peroxide from the dollar store down the street.

Brachiosaurid humeri vary somewhat but they all have certain features in common. Here’s the right humerus of Vouivria, modified from Mannion et al. (2017: fig. 19) to show the features of interest to brachiosaur humerus-sculptors. The arrows on the far left point to a couple of corners, one where the deltopectoral crest (dpc in the figure) meets the proximal articular surface, and the other where the articular surface meets the long sweeping curve of the medial border of the humeral shaft.

Here’s a more printer-friendly version of the same diagram. Why did I use Vouivria for this instead of one of the humeri of Brachiosaurus itself? Mostly because it’s a complete humerus for which a nice multi-view was available. Runner-up in this category would have to go to the humerus of Pelorosaurus conybeari figured by Upchurch et al. (2015: fig. 18) in the Haestasaurus paper–here’s a direct link to that figure.

I knew that I’d be doing some sculpting, and I wanted a scale template to work off of, so I made these outlines from the Giraffatitan humerus figured by Janensch (1950) and reproduced by Mike in this post (middle two), and from the aforementioned Pelorosaurus conybeari humerus shown by Mike in this post (outer two). I scaled this diagram so that when printed to fill an 8.5×11 piece of printer paper, the humerus outlines would all be 4.25″–the same nice-round-number 1/20 scale target found above. Here’s a PDF version: Giraffatitan and Pelorosaurus humeri outlines for print.

Here’s the largest of my giant mutant chicken humeri, compared to the outlines. The chicken humerus isn’t bad, but it’s too short for 1/20 scale, the angles of the proximal and distal ends are almost opposite what they should be, and the deltopectoral crest is aimed out antero-laterally instead of facing straight anteriorly. Modification will be required!

Here’s my method for lengthing the humerus: I cut the midshaft of another humerus out, and swapped it in to the middle of the prospective Brachiosaurus model humerus.

To my immense irritation, I failed to get a photo of the lengthened humerus before I started sculpting on it. In the first wave of sculpting, I built up the proximal end and the deltopectoral crest, but missed some key features. On the right, I glued the proximal and distal ends of the donor humerus together; I might make this into a Haestasaurus humerus in the future.

I should mention my tools and materials. I have a Dremel but it wasn’t charged the evening I sat down to do this, so I made all the humerus cuts with a small, cheap hacksaw. I used superglue (cyanoacrylate or CA) for quick joins, and white glue (polyvinyl acetate or PVA) to patch holes, and I put gobs of PVA into the humeral shafts before sealing them up. For additive sculpting I used spackling compound, same stuff you use to patch holes in walls and ceilings, and for reductive sculpting I used sandpaper. I got most of this stuff from the dollar store.

Here we are after a second round of sculpting. The proximal end has its corners now, and the distal end is more accurately belled out, maybe even a bit too wide. It’s not a perfect replica of either the Giraffatitan or Pelorosaurus humeri, but it got sufficiently into the brachiosaurid humerus morphospace for my taste. A more patient or dedicated sculptor could probably make recognizable humeri for each brachiosaurid taxon or even specimen. I deliberately left it a bit rough in hopes that it would read as timeworn, fractured, and restored when painted and mounted. Again, a real sculptor could make some hay here by putting in fake cracks and so on.

The cheap spackling compound I picked up did not harden as much as some other I have used in the past. I had planned on sealing anyway before I painted, and for porous materials a quick, cheap sealant is white glue mixed with water. Here that coat of diluted PVA is drying, and I’m holding up a spare chicken humerus to show how far the model humerus has come.

Before painting, I drilled into the distal end with a handheld electric drill, and used a bamboo barbeque skewer as a mounting rod and handle. I hit it with a couple of coats of gray primer, then a couple of coats of black primer the next day. I could have gotten fancier with highlights and washes and so on, but I was scrambling to get this done for a public outreach event, in an already busy week.

And here’s the finished-for-now product. A couple of gold-finished cardboard gift boxes from my spare box storage gave their lids to make a temporary pedestal. When I get a version of this model that I’m really happy with, either by hacking further on this one or starting from scratch on a second, I’d love to get a wooden or stone trophy base with a little engraved plaque that looks like a proper museum exhibit, and replace the bamboo skewer with a brass rod. But for now, I’m pretty happy with this.

The idea of making dinosaurs out of chicken bones isn’t original with me. I was inspired by the wonderful books Make Your Own Dinosaur Out of Chicken Bones and T-Rex To Go, both by Chris McGowan. Used copies of both books can be had online for next to nothing, and I highly recommend them both.

If this post helps you in making your own model Brachiosaurus humerus, I’d love to see the results. Please let me know about your model in the comments, and happy building!


  • Janensch, Werner. 1950. Die Wirbelsaule von Brachiosaurus brancai. Palaeontographica (Suppl. 7) 3: 27-93.
  • Mannion PD, Allain R, Moine O. (2017The earliest known titanosauriform sauropod dinosaur and the evolution of BrachiosauridaePeerJ 5:e3217
  • Upchurch, Paul, Philip D. Mannion and Micahel P Taylor. 2015. The Anatomy and Phylogenetic Relationships of “Pelorosaurus” becklesii (Neosauropoda, Macronaria) from the Early Cretaceous of England. PLoS ONE 10(6):e0125819. doi:10.1371/journal.pone.0125819

In short, no. I discussed this a bit in the first post of the Clash of the Dinosaurs saga, but it deserves a more thorough unpacking, so we can put this dumb idea to bed once and for all.

As Marco brought up in the comments on the previous post, glycogen bodies are probably to blame for the idea that some dinosaurs had a second brain to run their back ends. The glycogen body is broadly speaking an expansion of the spinal cord, even though it is made up of glial cells rather than neurons — simply put, help-and-support cells, not sensory, motor, or integration cells. When the spinal cord is expanded, the neural canal is expanded to accommodate it; as usual, the nervous system comes first and the skeleton forms around it. This creates a cavity in the sacrum that is detectable in fossils.

avian lumbosacral specializations - glycogen body

Giffin (1991) reviewed all of the evidence surrounding endosacral enlargements in dinosaurs (primarily sauropods and stegosaurs) and concluded that the explanation that best fit the observations was a glycogen body like that of birds. I agree 100%. The endosacral cavities of sauropods and stegosaurs (1) expand dorsally, instead of in some other direction, and (2) expand and contract over just a handful of vertebrae, instead of being more spread out. Of the many weird specializations of the spinal cord in birds, the glycogen body is the only one that produces that specific signal.

If any part of the nervous system of birds and other dinosaurs might be described as a ‘second brain’, it wouldn’t be the glycogen body, it would be the lumbosacral expansion of the spinal cord, which really is made up of neurons that help run the hindlimbs and tail (more on that in this previous post). But there’s nothing special about that, it’s present in all four-limbed vertebrates, including ourselves. Interestingly, that bulk of extra neural tissue in the sacral region of birds was referred to as a sort of ‘second brain’ by Streeter way back in 1904, in reference to the ostrich, but it’s clear that he meant that as an analogy, not that it’s literally a second brain.

So to sum up, a gradual expansion of the spinal cord to help run the hindlimbs and tail IS present in dinosaurs — and birds, and cows, and frogs, and us. But if that qualifies as a ‘second brain’, then we also have a ‘third brain’ farther up the spinal cord to run our forelimbs: the cervical enlargement, as shown in the above figure. These spinal expansions aren’t actual brains by any stretch and referring to them as such is confusing and counterproductive.

The sharp expansion of the neural canal over just a few vertebrae in birds does not house a ‘second brain’ or even an expansion of the neural tissue of the spinal cord. It contains the glycogen body, which is not made of neurons and has no brain-like activity. The sacral cavities of non-avian dinosaurs replicate precisely the qualities associated with the glycogen bodies of birds, and there’s no reason to expect that they contained anything else. That we don’t know yet what glycogen bodies do, even in commercially important species like chickens, may make that an unsatisfying answer, but it’s what we have for now.

The next installment will be way weirder (edit: it was, and is!). Stay tuned!


  • Giffin, E.B.,1991. Endosacral enlargements in dinosaurs. Modern Geology 16: 101-112.
  • Streeter, G.L. 1904. The structure of the spinal cord of the ostrich. American J. Anatomy 3(1): 1-27.

I planned to post this last spring but I never got around to it. I think I have a mental block about discussing the glycogen body. Partly because I’ve been burned by it before, partly because no-one knows what it does and that’s unsatsifying, partly because I didn’t want to plow through all the new literature on it (despite which, the function remains unknown).

Then I decided, screw it, I’ll let the slides speak for themselves, and the actual text of the post can just be navel-gazing and whingeing. Which you are “enjoying” right now.

So, there’s the glycogen body. It balloons out between the dorsal halves of the spinal cord, it’s made of glial cells (neuron support cells) that are packed with glycogen, and nobody knows why it’s there. On the graph of easy-to-find and frustrating-to-study it is really pushing the envelope.

Update: the role of the glycogen body in the ‘second brain’ myth is covered in the next post.

Previous entries in the “Bird neural canals are weird” series:

Here are some stubbornly-not-updated references for the images I used in the slides:

  • Huber, J.F. 1936. Nerve roots and nuclear groups in the spinal cord of the pigeon. Journal of Comparative Neurology 65(1): 43-91.
  • Streeter, G.L. 1904. The structure of the spinal cord of the ostrich. American Journal of Anatomy 3(1):1-27.
  • Watterson, R.L. 1949. Development of the glycogen body of the chick spinal cord. I. Normal morphogenesis, vasculogenesis and anatomical relationships. Journal of Morphology 85(2): 337-389.

Dorsal vertebra of a rhea from the LACM ornithology collection. Note the pneumatic foramina in the lateral wall of the neural canal.

If you’ve been here for very long you know I have a bit of a neural canal fixation. Some of this is related to pneumaticity, some of it is related to my interest in the nervous systems of animals, and some of it is pure curiosity about an anatomical region that seems to receive very little attention in proportion to its weirdness – especially in birds.

Human thoracic vertebrae in midsagittal section showing vertebral venous plexus. Gray (1918, image 579), available from

The neural canals of mammals are pretty boring. The canal is occupied by the spinal cord and its supporting layers of meninges, and the rest of the volume is padded out by adipose tissue and blood vessels, notably an extra-dural venous plexus. Aaand that’s about it, as far as I know. (If there are weird things inside mammalian neural canals that I’ve missed, please let me know in the comments – I’m a collector.)

But not so in birds, which have a whole festival of weird stuff going on inside their neural canals. Let’s start with pneumaticity, just to get it out of the way. Many birds have supramedullary diverticula inside their neural canals, and these can leave osteological traces, such as pneumatic foramina, in the walls of the neural canal. That’s cool but it’s a pretty well-known system – see Muller (1908) on the pigeon, Cover (1953) on the turkey, and these previous posts – and I want to get on to other, even stranger things.

The lumbosacral spinal cord of a 3-week-old chick in dorsal view. The big egg-shaped mass in the middle is the glycogen body. Watterson (1949: plate 1).

The spinal cords of birds have several gross morphological specializations not seen in mammals, as do their meninges, and most of these apomorphic structures can also leave diagnostic traces on the inner walls of the neural canal. In fact, birds have so many weird things going on with their spinal cords – at least five different things in the lumbosacral region alone – that I spent a week back in January just sorting them out. To crystalize that body of knowledge while I had it all loaded in RAM, I made a little slideshow for myself, and I’ll use screenshots of those slides to illustrate the morphologies I want to discuss. We’ll cover the vanilla stuff in the next post, and the really weird stuff in subsequent posts.

Stay tuned!


Illustration talk slide 51

Here’s a working version of that link.

Illustration talk slide 52

Illustration talk slide 53

Working link.

Illustration talk slide 54

Illustration talk slide 55

Illustration talk slide 56

Illustration talk slide 57

Working links:

The rest of this series.


  • Powell, Jaime E.  2003.  Revision of South American Titanosaurid dinosaurs: palaeobiological, palaeobiogeographical and phylogenetic aspects.  Records of the Queen Victoria Museum 111: 1-94.

This  is the fifth in a series of posts reviewing the Apatosaurus maquette from Sideshow Collectibles. Other posts in the series are:

There are really only a couple of interesting points to discuss for posture: the neck and the feet.

The neck posture is fine. Easy to say, but since I’m one of the “sauropods held their necks erect” guys, it might need some unpacking.

On one hand, animals really do use stereotyped postures, especially for the neck and head (Vidal et al. 1986, Graf et al. 1995, van der Leeuw et al. 2001). The leading hypothesis about why animals do this is that the number of joints and muscle slips involved in the craniocervical system permits an almost limitless array of possible postures, and that having a handful of stereotyped postures cuts down on the amount of neural processing required to keep everything going. That doesn’t mean that animals only use stereotyped postures, just that they do so most of the time, when there’s no need to deviate.

This might work something like the central pattern generators in your nervous system. When you’re walking down the sidewalk thinking about other things or talking with a friend, a lot of the control of your walk cycle is handled by your spinal cord, not your brain. Your brain is providing a direction and a speed, but the individual muscles are being controlled from the spinal cord. Key quote from the Wikipedia article: “As early as 1911, it was recognized, by the experiments of T. Graham Brown, that the basic pattern of stepping can be produced by the spinal cord without the need of descending commands from the cortex.”

But then you see a puddle or some dog doo and have to place your foot just so, and your brain takes over for a bit to coordinate that complex, ad hoc action. After the special circumstance is past, you go back to thinking about whatever and your spinal cord is back in charge of putting one foot in front of the other. This is the biological basis of the proverbial chicken running around with its head cut off: thanks to the spinal cord, the chicken can still run, but without a brain it doesn’t have anywhere to go (I have witnessed this, by the way–one of the numerous benefits to the future biologist of growing up on a farm).

Similarly, if the craniocervical system has a handful of regular postures–alert, feeding, drinking, locomoting, and so on–it lightens the load on the brain, which doesn’t have to figure out how to fire every muscle slip inserting on every cervical vertebra and on the skull to orient the head just so in three-dimensional space. That doesn’t mean that the brain doesn’t occasionally step in and do that, just like it takes over for the spinal cord when you place your feet carefully. But it doesn’t have to do it all the time.

van der Leeuw et al. (2001) took this a step further and showed that birds not only hold their heads and necks in stereotyped postures, they move between stereotyped postures in very predictable ways, and those movement patterns differ among clades (fig. 7 from that paper is above). There is a lot of stuff worth thinking about in that paper, and I highly recommend it, along with Vidal et al. (1986) and Graf et al. (1995), to anyone who is interested in how animals hold their heads and necks, and why.

So, on one hand, its wrong to argue that stereotyped postures are meaningless. But it’s also wrong to infer that animals only use stereotyped postures–a point we were careful to make in Taylor et al. (2009). And it’s especially wrong to infer that paleoartists only show animals doing familiar, usual things–I wrote the last post partly so I could make that point in this one.

For example, I think it would be a mistake to look at Brian Engh’s inflatable Sauroposeidon duo and infer that he accepts a raised alert neck posture for sauropods. He might or might not–the point is that the sauropods in the picture aren’t doing alert, they’re doing “I’m going to make myself maximally impressive so I can save myself the wear and tear of kicking this guy’s arse”. The only way the posture part of that painting can be inaccurate is if you think Sauroposeidon was physically incapable of raising its neck that high, even briefly (the inflatable throat sacs and vibrant colors obviously involve another level of speculation).

Similarly, the Sideshow Apatosaurus has its neck in the near-horizontal pose that is more or less standard for depictions of diplodocids (at least prior to 2009, and not without periodic dissenters). But it doesn’t come with a certificate that says that it is in an alert posture or that it couldn’t raise its neck higher–and even if it did, we would be free to ignore it. Would it have been cool to see a more erect-necked apatosaur? Sure, but that’s not a new idea, either, and there are other restorations out there that do that, and in putting this apatosaur in any one particular pose the artists were forced to exclude an almost limitless array of alternatives, and they had to do something. (Also, more practically, a more erect neck would have meant a larger box and heftier shipping charges.)

So the neck posture is fine. Cool, even, in that the slight ribbing along the neck created by the big cervical ribs (previously discussed here) gives you a sense of how the posture is achieved. Visible anatomy is fun to look at, which I suspect is one of the drivers behind shrink-wrapped dinosaur syndrome–even though it’s usually incorrect, and this maquette doesn’t suffer from it anyway.

Next item: the famous–or perhaps infamous–flipped-back forefoot. I have no idea who first introduced this in skeletal reconstructions and life restorations of sauropods, but it was certainly popularized by Greg Paul. It’s a pretty straightforward idea: elephants do this, why not sauropods?

Turns out there are good reasons to suspect that sauropods couldn’t do this–and also good reasons to think that they could. This already got some air-time in the comments thread on the previous review post, and I’m going to start here by just copying and pasting the relevant bits from that discussion, so you can see four sauropod paleobiologists politely disagreeing about it. I interspersed the images where they’re appropriate, not because there were any in the original thread.

Mike Taylor: the GSP-compliant strong flexion of the wrist always look wrong to me. Yes, I know elephants do this — see Muybridge’s sequence [above] — but as John H. keeps reminding us all, sauropods were not elephants, and one might think that in a clade optimsied for size above all else, wrist flexibility would not be retained without a very good reason.

Adam Yates: Yes I agree with Mike here, the Paulian, elephant-mimicking hyperflexion of the wrist is something that bugs me. Sauropod wrist elements are rather simple flat structures that show no special adaptation to achieve this degree of flexion. [Lourina sauropod right manus below, borrowed from here.]

Heinrich Mallison: Hm, I am not too sure what I think of wrist flexion. Sure it looks odd, but if you think it through the very reasons elephant have it is likely true in sauropods. And given the huge amount of cartilage mossing on the bones AND the missing (thus shape unknown) carpals I can well imagine that sauropods were capable of large excursions in the wrist.

 Mike: What are those reasons?

Heinrich: Mike, long humeri, very straight posture – try getting up from resting with weak flexion at the wrist. Or clearing an obstacle when walking. I can’t say too much, since this afternoon this has become a paper-to-be.

Mike: OK, Heinrich, but the Muybridge photos (and many others, including one on John H.’s homepage) show that elephants habitually flex the wrist in normal locomotion, not just when gwetting up from resting or when avoiding obstacles. Why?

The interesting thing here is that this is evidence of how flawed our (or maybe just my) intuition is: looking at an elephant skeleton, I don’t think I would ever have guessed that it would walk that way. (That said, the sauropod wrist skeleton does look much less flexible than that of the elephant.)

Matt: (why elephants flex their wrists) Possibly for simple energetics. If the limb is not to hit the ground during the swing phase, it has to be shortened relative to the stance limb. So it has to be bent. Bending the limb at the more proximal joints means lifting more weight against gravity. Flexing the wrist more might be a way to flex the elbow less.

(sauropod wrists look less flexible) Right, but from the texture of the ends of the bones we already suspect that sauropods had thicker articular cartilage caps than do mammals. And remember the Dread Olecranon of Kentrosaurus (i.e., Mallison 2010:fig. 3).

Mike: No doubt, but that doesn’t change the fact that elephant wrists have about half a dozen more discrete segments.

Matt: Most of which are very tightly bound together. The major flexion happens between the radius and ulna, on one hand, and the carpal block on the other, just as in humans. Elephants may have more mobile wrists than sauropods did–although that is far from demonstrated–but if so, it’s nothing to do with the number of bony elements. [Loxodonta skeleton below from Wikipedia, discovered here, arrow added by me.]

(Aside: check out the hump-backed profile of the Asian Elephas skeleton shown previously with the sway-backed profile of the African Loxodonta just above–even though the thoracic vertebrae have similar, gentle dorsal arches in both mounts. I remember learning about this from the wonderful How to Draw Animals, by Jack Hamm, when I was about 10. That book has loads of great mammal anatomy, and is happily still in print.)

And that’s as far as the discussion has gotten. The Dread Olecranon of Kentrosaurus is something Heinrich pointed out in the second of his excellent Plateosaurus papers (Mallison 2010: fig. 3).

Heinrich’s thoughts on articular cartilage in dinosaurs are well worth reading, so once again I’m going to quote extensively (Mallison 2010: p. 439):

Cartilaginous tissues are rarely preserved on fossils, so the thickness of cartilage caps in dinosaurs is unclear. Often, it is claimed that even large dinosaurs had only thin layers of articular cartilage, as seen in extant large mammals, because layers proportional to extant birds would have been too thick to be effectively supplied with nutrients from the synovial fluid. This argument is fallacious, because it assumes that a thick cartilage cap on a dinosaur long bone would have the same internal composition as the thin cap on a mammalian long bone. Mammals have a thin layer of hyaline cartilage only, but in birds the structure is more complex, with the hyaline cartilage underlain by thicker fibrous cartilage pervaded by numerous blood vessels (Graf et al. 1993: 114, fig. 2), so that nutrient transport is effected through blood vessels, not diffusion. This tissue can be scaled up to a thickness of several centimeters without problems.

An impressive example for the size of cartilaginous structures in dinosaurs is the olecranon process in the stegosaur Kentrosaurus aethiopicus Hennig, 1915. In the original description a left ulna (MB.R.4800.33, field number St 461) is figured (Hennig 1915: fig. 5) that shows a large proximal process. However, other ulnae of the same species lack this process, and are thus far less distinct from other dinosaurian ulnae (Fig. 3B, C). The process on MB.R.4800.33 and other parts of its surface have a surface texture that can also be found on other bones of the same individual, and may indicate some form of hyperostosis or another condition that leads to ossification of cartilaginous tissues. Fig. 3B–D compares MB.R.4800.33 and two other ulnae of K. aethiopicus from the IFGT skeletal mount. It is immediately obvious that the normally not fossilized cartilaginous process has a significant influence on the ability to hyperextend the elbow, because it forms a stop to extension. Similarly large cartilaginous structures may have been present on a plethora of bones in any number of dinosaur taxa, so that range of motion analyses like the one presented here are at best cautious approximations.

One of the crucial points to take away from all of this is that thick cartilage caps did not only expand or only limit the ranges of motions of different joints. The mistake is to think that soft tissues always do one or the other. The big olecranon in Kentrosaurus probably limited the ROM of the elbow, by banging into the humerus in extension. In contrast, thick articular cartilage at the wrist probably expanded the ROM and may have allowed the strong wrist flexion that some artists have restored for sauropods. I’m not arguing that it must have done so, just that I don’t think we can rule out the possibility that it may have. And so the flipped-back wrist in the Sideshow Apatosaurus does not bother me–but not everyone is convinced. Welcome to science!

I can’t finish without quoting a comment Mike left on Matt Bonnan’s blog a little over a year ago:

Ever since I saw Jensen’s (1987) paper about how mammals are so much better than dinosaurs because their limb-bones articulate properly, I’ve been fuming on and off about this — the notion that the clearly unfinished ends we see are what was operating in life. No.

This is a pretty fair summary of Jensen’s position. Of course, thanks to Heinrich, now we know why dinosaurs had such crap distal limb articulations: they weren’t mammals (part 1part 2part 3).

Finally, interest in articular cartilage is booming right now, as Mike blogged about here. In addition to the Dread Olecranon of Kentrosaurus, see the Dread Elbow Condyle-Thingy of Alligator from Casey Holliday’s 2001 SVP talk, and of course the culmination of that project in Holliday et al. (2010), and, for a more optimistic take on inferring the shapes of articular surfaces from bare bones, read Bonnan et al. (2010).

Next time: texture and color.


Necks lie, redux

September 1, 2011

In a recent post I showed photos of the trachea in a rhea, running not along the ventral surface of the neck but along the right side. I promised to show that this is not uncommon, that the trachea and esophagus of birds are usually free to slide around under the skin and are not constrained to like along the ventral midline of the neck, as they usually are in mammals. Here goes.

Here’s figure 5 from van der Leeuw et al. (2001): a lateral x-ray of a duck, reaching up just a bit with its head and neck, possibly to get a bite or just look around. Click through for the unlabeled version.

There’s a LOT of stuff going on in this image:

  • As promised, the trachea (blue lines) is taking a very different path to the head than the vertebrae and skeletal muscles.
  • As usual for tetrapods, the neck is extended at the base in the caudal half and flexed at the head in the cranial half.
  • The epaxial (dorsal) muscles at the base of the neck are not tied down to the vertebral column so they are free to bowstring across the U-bend at the base of the neck (black arrow)–this was the point of the figure in the original paper. Although the gross outline of the neck also deviates from the vertebral column on the ventral side near the head, this is caused by the trachea and gullet approaching the pharynx, not because the hypaxial muscles are bowstringed across the curve.
  • As the post title intimates, this neck lies: the cervical vertebrae are significantly more extended than one would expect based on the external appearance of the neck alone. The red line shows the angle of the most strongly retroverted vertebra, which I measure at 48.5 degrees from vertical (41.5 degrees above horizontal)–slightly closer to horizontal than to vertical! We have seen this before, in most mammals and in a couple of small birds (see this post); here we see it even in a reasonably large, long-necked bird.
  • Worse, the gross outline of the neck–what one can see from the outside–lines up with nothing on the inside: the trachea is less curved and the vertebral column is more curved.

Same points again, this time in a chicken in an alert posture (Vidal et al. 1986: fig. 7). Here the most strongly retroverted cervical is 36 degrees from vertical (54 degrees above horizontal).

What’s all this got to do with sauropods?

First, it shows that even in animals with long, slender necks, it’s not enough to show a photo or painting of an extant animal and make assertions about what the cervicals are doing (necks lie, again). It’s even less defensible to make the dual assertions that (a) the gross outline of the neck shows the path of the cervicals and (b) the cervicals are in ONP, all based on a photo or painting of a living animal. The first point can only be established by radiography, and the second by manipulation of the skeleton, either physically or digitally. It may seem like I’m tilting at windmills here, but we’ve seen these very assertions made in conference talks. As always, we’ll follow where the evidence leads, but not until we see some actual evidence.

Second,  I am increasingly haunted by the idea that we are all waaay too influenced, even (maybe especially) subconsciously, by big mammals when we think about sauropods and their necks. Big mammals–like, say, horses and giraffes–have:

  • only 7 cervical vertebrae;
  • lots of big muscles that attach to the thorax and the head and cross the cervical column without attaching to it much or at all;
  • presacral neural spines that max out, height-wise, over the shoulders, creating withers;
  • alert neck postures that are elevated (like all tetrapods) but often short of vertical, with the vertebrae often held more-or-less straight through the middle section of the neck (camels are an obvious exception here).

In contrast, birds have:

  • many cervical vertebrae, from a 12  or so up to 27 or 28;
  • almost no muscles that span from thorax to skull;
  • presacral neural spines that rise monotonically to the synsacrum (except–maybe–in Giraffatitan);
  • alert neck postures that are S-shaped, with the craniocervical joint over or just slightly in front of the cervicodorsal junction.

Which group sauropods had more in common with is left as an exercise for the reader.


  • van der Leeuw, A.H.J., Bout, R.G., and Zweers, G.A. 2001. Evolutionary morphology of the neck system in ratites, fowl, and waterfowl. Netherlands Journal of Zoology 51(2):243-262.
  • Vidal, P.P., Graf, W., and Berthoz, A. 1986. The orientation of the cervical vertebral column in unrestrained awake animals. Experimental Brain Research 61: 549­-559.