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 Bartleby.com.

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!

References

In the first installment in this series (link), we looked at a couple of weird sauropod vertebrae with neurocentral joints that were situated either entirely dorsal or ventral to the neural canals. This post has more examples of what I am calling “offset” neurocentral synchondroses.

I decided it made more sense to refer to the synchondrosis as being offset, instead of referring to the neural canal as offset. Because the neural canal in all of these vertebrae is right where it pretty much always is, just dorsal to the articular surfaces of the centrum. In an adult, fused vertebra, there’d be no sign that anything unusual had ever happened. So I think it makes more sense to talk about the neurocentral joint having migrated dorsally or ventrally relative to the canal, rather than vice versa. If you know differently, or if these weirdos have been addressed before elsewhere and I’ve just missed it, please let me know in the comments!

Here’s a plate from Marsh (1896) showing caudal vertebrae of Camarasaurus (“Morosaurus” in O.C. Marsh parlance), which echo the Alamosaurus caudal from the first post in having the neurocentral joint almost entirely ventral to the neural canal. The neural arch here doesn’t just arch over the canal dorsally, it also cuts under it ventrally, at least in part.

This plate is also nice because it shows the relationships among the arch, centrum, and caudal ribs before they fuse. Here’s the caption, from Marsh (1896):

Here’s the preceding plate, Plate 33, with illustrations of an unfused Camarasaurus sacrum.

And its caption:

This plate not only shows how the sacral ribs fuse to the arch and spine medially, and to each other laterally (forming the sacrocostal yoke), it also shows a last sacral that is very similar to the aforementioned caudals in the position of the neurocentral joint. But interestingly that neurocentral joint offset only seems to be present in the last caudal sacral – the lower figure shows widely-separated neurocentral joint surfaces in the more anterior centra, indicating that the neural arches (not figured in this dorsal view) did not wrap around the neural canal to approach the midline. (I think we’re looking at S2 through S5 here, and missing a dorso-sacral.)

So now I’m freaked out, wondering if this neural arch wrap-around in the caudals is common to most sauropods and I just haven’t looked at enough juvenile caudals to have spotted it before. As always, feel free to ablate my ignorance in the comments, particularly if you know of more published examples. I’m a collector.

The neural canal of that last sacral also has a very interesting cross-sectional shape, like a numeral 8. I have some thoughts on that, but they’ll keep for a future post in this series.

Here’s a dorsal vertebra of Camarasaurus in anterior view (from Ostrom & McIntosh 1966, modified by Wilson & Sereno 1998). It is one of the most disturbing things I have ever seen in a sauropod. It makes my skin crawl.

Here’s why: the centrum and the thing we habitually call the ‘neural arch’ aren’t fully fused, and as this modified version makes clear, the ‘neural arch’ is neither neural nor an arch. Instead of being bounded ventrally by the centrum and dorsally and laterally by the neural arch, the neural canal lies entirely below the synchondrosis between the not-really-an-arch and the centrum.

Why?! WHY WOULD YOU DO THAT, CAMARASAURUS? This is not ‘Nam. This is basic vertebral architecture. There are rules.

Look at c6 of Apatosaurus CM 555 here, behaving as all good vertebrae ought to. Neural arch be archin’, as the kids say.

And if you are seeking solace in the thought that maybe the artist just drew that Cam dorsal incorrectly, forget it. I’ve been to Yale and examined the original specimen. I’ve seen things, man!

Camarasaurus isn’t the only pervert around here. Check this out:

Unfused neural arch of a caudal vertebra of a juvenile Alamosaurus from Big Bend. And I mean, this is a neural arch. This may be the most neural of all neural arches, in that it contains the entire neural canal. It’s more of a neural…ring, I guess. That’s right, this Alamosaurus caudal is batting for the opposite team from the Cam dorsal above. And it’s a team that neither you nor I play on, because we have well-behaved normal-ass vertebrae with neural arches that actually arch, and then stop, like God and Richard Owen intended.

Scientifically, my question about these vertebrae is: well, that is, I mean to say, what!? I think they have damaged me in some fundamental way.

If you have anything more intelligent to add (or even less intelligent – consider the gauntlet thrown down!), the comment thread is open.

References

  • Ostrom, John H., and John S. McIntosh. 1966. Marsh’s Dinosaurs. Yale University Press, New Haven and London. 388 pages including 65 absurdly beautiful plates.
  • Wilson, J. A. and Paul C. Sereno. 1998. Early evolution and higher-level phylogeny of sauropod dinosaurs. Society of Vertebrate Paleontology, Memoir 5: 1-68.