Can we distinguish taphonomic distortion and (paleo)pathology from normal biological variation?
February 12, 2021
Taylor 2015: Figure 8. Cervical vertebrae 4 (left) and 6 (right) of Giraffatitan brancai lectotype MB.R.2180 (previously HMN SI), in posterior view. Note the dramatically different aspect ratios of their cotyles, indicating that extensive and unpredictable crushing has taken place. Photographs by author.
Here are cervicals 4 and 8 from MB.R.2180, the big mounted Giraffatitan in Berlin. Even though this is one of the better sauropod necks in the world, the vertebrae have enough taphonomic distortion that trying to determine what neutral, uncrushed shape they started from is not easy.
Wedel and Taylor 2013b: Figure 3. The caudal vertebrae of ostriches are highly pneumatic. This mid-caudal vertebra of an ostrich (Struthio camelus), LACM Bj342, is shown in dorsal view (top), anterior, left lateral, and posterior views (middle, left to right), and ventral view (bottom). The vertebra is approximately 5cm wide across the transverse processes. Note the pneumatic foramina on the dorsal, ventral, and lateral sides of the vertebra.
Here’s one of the free caudal vertebrae of an ostrich, Struthio camelus, LACM Ornithology Bj342. It’s a bit asymmetric–the two halves of the neural spine are aimed in slightly different directions, and one transverse process is angled just slightly differently than the other–but the asymmetry is pretty subtle and the rest of the vertebral column looks normal, so I don’t think this rises to the level of pathology. It looks like the kind of minor variation that is present in all kinds of animals, especially large-bodied ones.
This is a dorsal vertebra of a rhea, Rhea americana, LACM Ornithology 97479, in posteroventral view. Ink pen for scale. I took this photo to document the pneumatic foramina and related bone remodeling on the dorsal roof of the neural canal, but I’m showing it here because in technical terms this vert is horked. It’s not subtly asymmetric, it’s grossly so, with virtually every feature–the postzygapophyses, diapophyses, parapophyses, and even the posterior articular surface of the centrum–showing fairly pronounced differences from left to right.
That rhea dorsal looks pretty bad for dry bone from a recently-dead extant animal, but if it was from the Morrison Formation it would be phenomenal. If I found a sauropod vertebra that looked that good, I’d think, “Hey, this thing’s in pretty good shape! Only a little distorted.” The roughed-up surface of the right transverse process might give me pause, and I’d want to take a close look at those postzygs, but most of this asymmetry is consistent with what I’d expect from taphonomic distortion.
Which brings me to my titular question, which I am asking out of genuine ignorance and not in a rhetorical or leading way: can we tell these things apart? And if so, with what degree of confidence? I know there has been a lot of work on 3D retrodeformation over the past decade and a half at least, but I don’t know whether this specific question has been addressed.
Corollary question: up above I wrote, “It looks like the kind of minor variation that is present in all kinds of animals, especially large-bodied ones”. My anecdotal experience is that the vertebrae of large extant animals tend to be more asymmetric than those of small extant animals, but I don’t know if that’s a real biological phenomenon–bone is bone but big animals have larger forces working on their skeletons, and they typically live longer, giving the skeleton more time to respond to those forces–OR if the asymmetry is the same in large and small animals and it’s just easier to see in the big ones.
If either of those questions has been addressed, I’d be grateful for pointers in the comments, and thanks in advance. If one or both have not been addressed, I think they’re interesting but Mike and I have plenty of other things to be getting on with and we’re not planning to work on either one, hence the “Hey, you! Want a project?” tag.
References
- Taylor, Michael P. 2015. Almost all known sauropod necks are incomplete and distorted. PeerJ Preprints 3:e1767. doi:10.7287/peerj.preprints.1418v1
- Wedel, Mathew J., and Michael P. Taylor. 2013. Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus.PLOS ONE 8(10):e78213. 14 pages. doi:10.1371/journal.pone.0078213 [PDF]
In my recent visit to the LACM herpetology collection, I was interested to note that almost every croc, lizard, and snake vertebra I saw had a pair of neurovascular foramina on either side of the centrum, in “pleurocoel” position. You can see these in the baby Tomistoma tail, above. Some vertebrae have a big foramen, some have a small foramen, and some have no visible foramen at all. Somehow I’d never noticed this before.
This is particularly interesting in light of the observation from birds that pneumatic diverticula tend to follow nerves and vessels as they spread through the body. Maybe we find pneumatic features where we do in dinosaurs and pterosaurs because that’s where the blood vessels were going in the babies. Also, these neurovascular foramina in extant reptiles are highly variable in size and often asymmetric – sound familiar?
It should. Caudal pneumaticity in the tail of Giraffatitan MB.R.5000. Dark blue vertebrae are pneumatic on both sides, light blue vertebrae only have fossae on the right side. Wedel and Taylor (2013b: Figure 4).
I am starting to wonder if some of the variability we associate with pneumaticity is just the variability of soft tissue, full stop. Or if pneumaticity is variable because it developmentally follows in the footsteps of the blood vessels, which are themselves inherently variable. That seems like a promising line of inquiry. And also something I should have though of a lot sooner.
UPDATE in 2023: A promising line of inquiry indeed! This spawned a paper in 2021, a conference presentation later in 2021, which will become a paper in time, and the tentacles of this idea — that diverticula following blood vessels has a lot of explanatory power — are wound through a LOT of my current and upcoming projects.
Vertebral orientation: Varanus komodoensis would like a word
September 25, 2018
I am still building up to a big post on vertebral orientation, but in the meantime, check out this caudal vertebra of a Komodo dragon, Varanus komodoensis. This is right lateral view–the vert is strongly procoelous, and the articular ends of the centrum are really tilted relative to the long axis. I find this encouraging, for two reasons. First, it helped me clarify my thinking on how we ought to orient vertebrae, which Mike wrote about here and here. And second, it gives me some hope, because if we can figure out why tilting your articular surfaces makes functional sense in extant critters like monitors, maybe we can apply those lessons to sauropods and other extinct animals.
This is LACM Herpetology specimen 121971. Many thanks again to Neftali Camacho for access and assistance, and to Jessie Atterholt for basically doing all the other jobs while I was faffing about with this Komodo dragon.
Pelican vertebrae are mostly air
July 16, 2014
I was at the Natural History Museum of Los Angeles County yesterday to do some research in the ornithology collection. After lunch I was working on this pelican skeleton and I thought, “Geez, there is just no way to do this thing justice with still photos. I should make a video.” Here it is. You’ll want to see it full-screen–this being my first time out making a video, I didn’t realize that I was holding the phone the wrong way for efficient viewing on other devices.
The specimen is LACM Ornithology 86262. I’m posting this video with the knowledge and kind permission of the ornithology collection staff.
For previous things in this vein, please see:
- There’s almost nothing but nothing there, Brachiosaurus edition
- There’s almost nothing but nothing there, Sauroposeidon edition
If you like it that stuff like this exists, please support your local natural history museum, especially the LACM, which has some really fantastic education and outreach programs.
All I want to do in this post is make people aware that there is a difference between these two things, and occasionally that affects those of us who work in natural history.
In one of his books or essays, Stephen Jay Gould made the point that in natural history we are usually not dealing with whether phenomena are possible or not, but rather trying to determine their frequency. If we find that in a particular population of quail most of the birds eat ants but some avoid them, then we know some things: that quail can tolerate eating ants, that quail are not required to eat ants, and that both strategies can persist in a single population.
This idea has obvious repercussions for paleoart, especially when it comes to “long-tail” behaviors. I dealt with that in this post, and also in the comment thread to this one. But that’s not what I want to talk about today.
Sometimes it is useful to talk about things that never happen, or that have at least never occurred in the sample of things we know of. Obviously how certain you can be in these cases depends on the intensity of sampling and the inherent likelihood of a surprising result, which can be hard to judge. If you argued right now that T. rex lacked feathers because no T. rex specimens have been found with feathers, you’d most likely be wrong; it is almost certainly just a matter of time before someone finds direct evidence of feathers in T. rex, given the number of T. rex specimens waiting to be found and the strength of the indirect evidence (e.g., phylogenetic inference, analogy: ornithomimids are known to be feathered even though most specimens are found without feather impressions). If you argue that sauropods are unique among terrestrial animals in having necks more than five meters long, you’re most likely right; being wrong would imply the existence of some as-yet undiscovered land animal of sauropod size, or with seriously wacky proportions (or both), and our sampling of terrestrial vertebrates is good enough to make that extremely unlikely.
The reason for this post is that sometimes people confuse that last argument, which is about sampling and induction, with the argument from personal incredulity.
For example, in our no-necks-for-sex paper (Taylor et al. 2011), we included this passage:
Sauropoda also had a long evolutionary history, originating about 210 million years ago in the Carnian or Norian Age of the Late Triassic, and persisting until the end-Cretaceous extinction of all non-avian dinosaurs about 65 millions years ago. Thus the ‘necks-for-sex’ hypothesis requires that this clade continued to sexually select for exaggeration of the same organ for nearly 150 million years, a scenario without precedent in tetrapod evolutionary history.
One of the reviewers argued that we couldn’t include that section, because it was just the argument from personal incredulity writ large, like so:
There are no other known cases of X in tetrapod evolutionary history, and therefore we don’t believe that the case in question is the sole exception.
…with the second part of that unstated (by us) but implied. But we disagreed, and argued (successfully) that it was an argument based on sampling, like so:
There are no other known cases of X in tetrapod evolutionary history, and therefore it is unlikely that the case in question is the sole exception.
Now, it is perfectly fair to criticize arguments like that based on the thoroughness of the sampling and the likelihood of exceptions, as discussed above for T. rex feathers. Just don’t mistake arguments like that for arguments from personal incredulity.* On the flip side, if someone makes an argument from personal incredulity, see if the same thing can be restated as an argument about sampling. Maybe they’re correct but just expressing themselves poorly (“I refuse to believe that the moon is made out of cheese”), and maybe they’re wrong and restating things in terms of sampling will help you understand why.
* If you want to get super pedantic about it, they’re both arguments from ignorance. But one of them is at least potentially justifiable by reference to sampling. Absence of evidence is not necessarily evidence of absence, but it may get to be that way as the sampling improves (e.g., there is no evidence of planets closer to the sun than Mercury, and at this point, that is pretty persuasive evidence that no such planets exist).
Parting shot: one thing that has always stuck in my head from Simberloff (1983) is the bit about imagining a large enough universe of possible outcomes. And I’ve always had a perverse fascination with Larry Niven’s “Down in Flames”, in which he pretty much demolished his Known Space universe by assuming that every basic postulate of that universe was false. Neither of these follow directly on from the main point of the post, but they’re not completely unrelated, either. Because I think that they yield a pretty good heuristic for how to do science: imagine what it would take for you to be wrong–imagine a universe in which you are wrong–and then go see if the thing that makes you wrong, whatever it is, can be shown to exist or to work. If not, it doesn’t mean you’re right, but it means you’re maybe less wrong, which, if we get right down to it, is the best that we can hope for.
The photos have nothing to do with the post, they’re just pretty pictures from the LACM to liven things up a little.
References
- Simberloff, D. (1983). Competition theory, hypothesis-testing, and other community ecological buzzwords. The American Naturalist, 122(5), 626-635.
- Taylor, M. P., Hone, D. W., Wedel, M. J., & Naish, D. (2011). The long necks of sauropods did not evolve primarily through sexual selection. Journal of Zoology, 285(2), 150-161.
Night at the Museum: LACM’s Camp Dino
April 8, 2013
Last night London and I spent the night in the Natural History Museum of Los Angeles County (LACM), as part of the Camp Dino overnight adventure. So we got lots of time to roam the exhibit halls when they were–very atypically–almost empty. Above are the museum’s mounted Triceratops–or one of them, anyway–and mounted cast of the Mamenchisaurus hochuanensis holotype, presented in glorious not-stygian-darkness (if you went through the old dino hall, pre-renovation, you know what I mean).
We got there early and had time to roam around the museum grounds in Exposition Park. The darned-near-life-size bronze dinos out front are a minor LA landmark.
The rose garden was already closed, but we walked by anyway, and caught this rainbow in the big fountain.
After we checked in we had a little time to roam the museum on our own. I’ve been meaning to blog about how much I love the renovated dinosaur halls. The bases are cleverly designed to prohibit people touching the skeletons without putting railings or more than minimal glass in the way, and you can walk all the way around the mounted skeletons and look down on them from the mezzanine–none of that People’s Gloriously Efficient Cattle Chute of Compulsory Dinosaur Appreciation business. Signage is discreet and informative, and so are the handful of interactive gizmos. London and I spent a few minutes using a big touch-screen with a slider that controlled continental drift from the Triassic to the present–a nice example of using technology to add value to an exhibit without taking away from the real stuff that’s on display. There are even a few places to sit and just take it all in. That’s pretty much everything I want in a dinosaur hall.
Also, check out the jumbotron on the left in the above photo. It was running a (blessedly) narration-free video on how fossils are found, collected, prepared, mounted, and studied, on about a five-minute loop. Lots of pretty pictures. Including this next one.
There are a couple of levels of perspective distortion going on here, both in the original photo and in my photo of that photo projected on the jumbotron. Still, I feel confident positing that that is one goldurned big ilium. I’m not going to claim it’s the biggest bone I’ve ever seen–that rarely ends well–but sheesh, it’s gotta be pretty freakin’ big. And apparently a brachiosaurid, or close to it. Never mind, it’s almost certainly an upside-down Triceratops skull. Thanks to Adam Yates for the catch. I will now diminish, and go into the West.
Triceratops, Styracosaurus, and Einiosaurus–collect the whole set!
Of course, the centerpiece of the second dinosaur hall–and how great is it that there are two!?–is the T. rex trio: baby, juvenile (out of frame to the right), and subadult. Yes, subadult: the “big” one is not as big as the really big rexes, and from the second floor you can see unfused neural arches in some of the caudal vertebrae (many thanks to Ashley Fragomeni for pointing those out to me on a previous visit).
Awwwww! C’mere, little fella!
Still, this ain’t Vulgar Overstudied Theropod Picture of the Week. Here are some sweet pneumatic diplodocid caudals in the big wall o’ fossils (visible behind Mamenchisaurus in the overhead photo above). The greenish color is legit–in the Dino Lab on the second floor, they’re prepping a bunch of sauropod elements that look like they were carved out of jade.
Sudden violent topic shift, the reason for which will be become clear shortly: London and I have been sculpting weapons of mass predation in our spare time. In some of the photos you may be able to see his necklace, which has a shark tooth he sculpted himself. Here are a couple of allosaur claws I made–more on those another time.
The point is, enthusiasm for DIY fossils is running very high at Casa Wedel, so London’s favorite activity of the evening was molding and casting. Everyone got to make a press mold using a small theropod tooth, a trilobite, or a Velociraptor claw. Most of the kids I overheard opted for the tooth, but London went straight for the claw.
Ready for plaster! Everyone got to pick up their cast at breakfast this morning, with instructions to let them cure until this evening. All went well, so I’ll spare you a photo of this same shape in reverse.
We were split into three tribes of maybe 30-40 people each, and each tribe bedded down in a different hall. The T. rex and Raptor tribes got the North American wildlife halls, but our Triceratops tribe got the African wildlife hall, which as a place to sleep is about 900 times cooler. Someone had already claimed the lions when we got there, so London picked hyenas as our totem animals.
Lights out was at 10:30 PM, and the lights came back on at 7:00 this morning. Breakfast was out from 7:15 to 8:00, and then we had the museum to ourselves until the public came in at 9:30. So I got a lot of uncluttered photos of stuff I don’t usually get to photograph, like this ammonite. Everyone should have one of these.
London’s favorite dino in the museum is Carnotaurus. It’s sufficiently weird that I can respect that choice.
Not that there’s anything wrong with the old standards, especially when they’re presented as cleanly and innovatively as they are here.
Finally, the LACM has a no tripod policy, and if they see you trying to carry one in they will make you take it back to your car. At least during normal business hours. But no one searched my backpack when we went in last night, and I put that sucker to some good use. Including getting my first non-bigfoot picture of the cast Argentinosaurus dorsal. It was a little deja-vu-ey after just spending so much time with the giant Oklahoma Apatosaurus–elements of the two animals really are very comparable in size.
If you’re in the LA area and interested in spending a night at the museum–or at the tar pits!–check out the “Overnight Adventures” page on the museum’s website. Cost is $50 per person for members or $55 for non-members, and worth every penny IMHO. It’s one of those things I wish we’d done years ago.
Sauropod Vertebra Picture of the Week 