In a comment on the last post, on the mass of Dreadnoughtus, Asier Larramendi wrote:

The body mass should be considerably lower because the reconstructed column don’t match with published vertebrae centra lengths. 3D reconstruction also leaves too much space between vertebrae. The reconstruction body trunk is probably 15-20% longer than it really was. Check the supplementary material: http://www.nature.com/srep/2014/140904/srep06196/extref/srep06196-s1.pdf

So I did. The table of measurements in the supplementary material is admirably complete. For all of the available dorsal vertebrae except D9, which I suppose must have been too poorly preserved to measure the difference, Lacovara et al. list both the total centrum length and the centrum length minus the anterior condyle. Centrum length minus the condyle is what in my disseration I referred to as “functional length”, since it’s the length that the vertebra actually contributes to the articulated series, assuming that the condyle of one vertebra sticks out about as far as the cotyle is recessed on the next vertebra. Here are total lengths/functional lengths/differences for the seven preserved dorsals, in mm:

  • D4 – 400/305/95
  • D5 – 470/320/150
  • D6 – 200/180/20
  • D7 – 300/260/40
  • D8 – 350/270/80
  • D9 – 410/ – / -
  • D10 – 330/225/105

The average difference between functional length and total length is 82 mm. If we apply that to D9 to estimate it’s functional length, we get 330mm. The summed functional lengths of the seven preserved vertebrae are then 1890 mm. What about the missing D1-D3? Since the charge is that Lacovara et al. (2014) restored Dreadnoughtus with a too-long torso, we should be as generous as possible in estimating the lengths of the missing dorsals. In Malawisaurus the centrum lengths of D1-D3 are all less than or equal to that of D4, which is the longest vertebra in the series (Gomani 2005: table 3), so it seems simplest here to assign D1-D3 functional lengths of 320 mm. That brings the total functional length of the dorsal vertebral column to 2850 mm, or 2.85 m.

At this point on my first pass, I was thinking that Lacovara et al. (2014) were in trouble. In the skeletal reconstruction that I used for the GDI work in the last post, I measured the length of the dorsal vertebral column as 149 pixels. Divided by 36 px/m gives a summed dorsal length of 4.1 m. That’s more than 40% longer than the summed functional lengths of the vertebrae calculated above (4.1/2.85 = 1.44). Had Lacovara et al. really blown it that badly?

Before we can rule on that, we have to estimate how much cartilage separated the dorsal vertebrae. This is a subject of more than passing interest here at SV-POW! Towers–the only applicable data I know of are the measurements of intervertebral spacing in two juvenile apatosaurs that Mike and I reported in our cartilage paper last year (Taylor and Wedel 2013: table 3, and see this post). We found that the invertebral cartilage thickness equaled 15-24% of the length of the centra.* For the estimated 2.85-meter dorsal column of Dreadnoughtus, that means 43-68 cm of cartilage (4.3-6.8 cm of cartilage per joint), for an in vivo dorsal column length of 3.28-3.53 meters. That’s still about 15-20% shorter than the 4.1 meters I measured from the skeletal recon–and, I must note, exactly what Asier stated in his comment. All my noodling has accomplished is to verify that his presumably off-the-cuff estimate was spot on. But is that a big deal?

Visually, a 20% shorter torso makes a small but noticeable difference. Check out the original reconstruction (top) with the 20%-shorter-torso version (bottom):

Dreadnoughtus shortened torso comparison - Lacovara et al 2014 fig 2

FWIW, the bottom version looks a lot more plausible to my eye–I hadn’t realized quite how weiner-dog-y the original recon is until I saw it next to the shortened version.

In terms of body mass, the difference is major. You’ll recall that I estimated the torso volume of Dreadnoughtus at 32 cubic meters. Lopping off 20% means losing 6.4 cubic meters–about the same volume as a big bull elephant, or all four of Dreadnoughtus‘s limbs put together. Even assuming a low whole-body density of 0.7 g/cm^3, that’s 4.5 metric tons off the estimated mass. So a ~30-ton Dreadnoughtus is looking more plausible by the minute.

For more on how torso length can affect the visual appearance and estimated mass of an animal, see this post and Taylor (2009).

* I asked Mike to do a review pass on this post before I published, and regarding the intervertebral spacing derived from the juvenile apatosaurs, he wrote:

That 15-24% is for juveniles. For the cervicals of adult Sauroposeidon we got about 5%. Why the differences? Three reasons might be relevant: 1, taxonomic difference between Sauroposeidon and Apatosaurus; 2, serial difference between neck and torso; 3, ontogenetic difference between juvenile and adult. By applying the juvenile Apatosaurus dorsal measurement directly to the adult Dreadnoughtus dorsals, you’re implicitly assuming that the adult/juvenile axis is irrelevant (which seems unlikely to me), that the taxonomic axis is (I guess) unknowable, and that the cervical/dorsal distinction is the only one that matter.

That’s a solid point, and it deserves a post of its own, which I’m already working on. For now, it seems intuitively obvious to me that we got a low percentage on Sauroposeidon simply because the vertebrae are so long. If the length-to-diameter ratio was 2.5 instead of 5, we’d have gotten 10%, unless cartilage thickness scales with centrum length, which seems unlikely. For a dorsal with EI of 1.5, cartilage thickness would then be 20%, which is about what I figured above.

Now, admittedly that is arm-waving, not science (and really just a wordy restatement of his point #2). The obvious thing to do is take all of our data and see if intervertebral spacing is more closely correlated with centrum length or centrum diameter. Now that it’s occurred to me, it seems very silly not to have done that in the actual paper. And I will do that very thing in an upcoming post. For now I’ll just note three things:

  1. As you can see from figure 15 in our cartilage paper, in the opisthocoelous anterior dorsals of CM 3390, the condyle of the posterior vertebra is firmly engaged in the cotyle of the anterior one, and if anything the two vertebrae look jammed together, not drifted apart. But the intervertebral spacing as a fraction of centrum length is still huge (20+4%) because the centra are so short.
  2. Transferring these numbers to Dreadnoughtus only results in 4.3-6.8 cm of cartilage between adjacent vertebrae, which does not seem unreasonable for a 30- or 40-ton animal with dorsal centra averaging 35 cm in diameter. If you asked me off the cuff what I thought a reasonable intervertebral spacing was for such a large animal, I would have said 3 or 4 inches (7.5 to 10 cm), so the numbers I got through cross-scaling are actually lower than what I would have guessed.
  3. Finally, if I’ve overestimated the intervertebral spacing, then the actual torso length of Dreadnoughtus was even shorter than that illustrated above, and the volumetric mass estimate would be smaller still. So in going with relatively thick cartilage, I’m being as generous as possible to the Lacovara et al. (2014) skeletal reconstruction (and indirectly to their super-high allometry-derived mass estimate), which I think is only fair.

References

 

Readers with long memories might recall that, nearly two years ago, we published annotated skeletal reconstructions of Camarasaurus and of Tyrannosaurus, with all the bones labelled. At the time, I said that I’d like to do an ornithischian, too.

Well, here it is at last, based on Marsh’s (1891) classic reconstruction of Triceratops:

Marsh1891--Restoration-of-Triceratops--plate-XV

Click through for the full-sized version (2076 by 864 pixels), which — like the other two — you are welcome to print out and hang on your wall as a handy reference, or to use in teaching. (Marsh’s original is out of copyright; I hereby make my modified version available under the CC By 3.0 licence.)

I recently reread Dubach (1981), “Quantitative analysis of the respiratory system of the house sparrow, budgerigar and violet-eared hummingbird”, and realized that she reported both body masses and volumes in her Table 1. For each of the three species, here are the sample sizes, mean total body masses, and mean total body volumes, along with mean densities I calculated from those values.

  • House sparrow, Passer domesticus, n = 16, mass = 23.56 g, volume = 34.05 mL, density = 0.692 g/mL
  • Budgerigar, Melopsittacus undulatus, n = 19, mass = 38.16 g, volume = 46.08 mL, density = 0.828 g/mL
  • Sparkling violetear,* Colibri coruscans, n = 12, mass = 7.28 g, volume = 9.29 mL, density = 0.784 g/mL

* This is the species examined by Dubach (1981), although not specified in her title; there are four currently-recognized species of violetears. And apparently ‘violetear’ has overtaken ‘violet-eared hummingbird’ as the preferred common name. And as long as we’re technically on a digression,  I’m almost certain those volumes do not include feathers. Every volumetric thing I’ve seen on bird masses assumes plucked birds (read on).

This is pretty darned interesting to me, partly because I’m always interested in how dense animals are, and partly because of how the results compare to other published data on whole-body densities for birds. The other results I am most familiar with are those of Hazlehurst and Rayner (1992) who had this to say:

There are relatively few values for bird density. Welty (1962) cited 0.9 g/mL for a duck, and Alexander (1983) 0.937 g/mL for a domestic goose, but those values may not take account of the air sacs. Paul (1988) noted 0.8 g/mL for unspecified bird(s). To provide more reliable estimates, the density of 25 birds of 12 species was measured by using the volume displacement method. In a dead, plucked bird the air-sac system was reinflated (Saunder and Manton 1979). The average density was 0.73 g/mL, suggesting that the lungs and air sacs occupy some quarter of the body.

That result has cast a long shadow over discussions of sauropod masses, as in this paper and these posts, so it’s nice to see similar results from an independent analysis.  If you’re curious, the weighted mean of the densities calculated from Duchard’s Dubach’s (1981) data is 0.77. I’d love to see the raw data from Hazlehurst and Rayner (1992) to see how much spread they got in their density measurements.  Unfortunately, they did not say which birds they used or give the raw data in the paper (MYDD!), and I have not asked them for it because doing so only just occurred to me as I was writing this post.

There will be more news about hummingbirds here in the hopefully not-too-distant future. Here’s a teaser:

SkeletonFULL

Yes, those are its hyoids wrapped around the back of its head–they go all the way around to just in front of the eyes, as in woodpeckers and other birds that need hyper-long tongue muscles. There are LOADS of other interesting things to talk about here, but it will be faster and more productive if I just go write the paper like I’m supposed to be doing.

Oh, all right, I’ll say a little more. This is a  young adult female Anna’s hummingbird, Calypte anna, who was found by then-fellow-grad-student Chris Clark at a residential address in Berkeley in 2005. She was unable to fly and died of unknown causes just a few minutes after being found. She is now specimen 182041 in the ornithology collection at the Museum of Vertebrate Zoology at Berkeley. Chris Clark and I had her microCTed back in 2005, and that data will finally see the light of day thanks to my current grad student, Chris Michaels, who generated the above model.

This bird’s skull is a hair over an inch long, and she had a body mass of 3.85 grams at the time of her death. For comparison, those little ketchup packets you get at fast-food burger joints each contain 8-9 grams of ketchup, more than twice the mass of this entire bird when it was alive!

References

  • Dubach, M. 1981. Quantitative analysis of the respiratory system of the house sparrow, budgerigar and violet-eared hummingbird. Respiration Physiology 46(1): 43-60.
  • Hazlehurst, G.A., and Rayner, J.M. 1992. Flight characteristics of Triassic and Jurassic Pterosauria: an appraisal based on wing shape. Paleobiology 18(4): 447-463.
Apatosaurus lousiae 1/12 scale skeleton, modelled by Phil Platt, assembled and photographed by Brant Bassam. Image courtesy of BrantWorks.com.

Apatosaurus lousiae 1/12 scale skeleton in left antero-lateral view, modelled by Phil Platt, assembled and photographed by Brant Bassam. Image courtesy of BrantWorks.com.

Today our paper on sauropod neck anatomy is formally published in PeerJ.

There’s not much new to say about the paper, since we posted it to arXiv last year and told the world about it then (post 1, post 2, post 3). Although a lot more attractive in form, this version is almost identical in content, modulo some changes requested by the PeerJ reviewers, and some changes to the figures to make sure every part of every figure was CC BY or otherwise in the public domain. Many thanks to everyone who gave us permission to use their images, especially Scott Hartman, who is rapidly getting to be the go-to person for this sort of thing just by doing good work and being a nice guy.

The big news, of course, is not the paper but the outlet. We’re excited about PeerJ because it promises to be a game-changer, for lots of reasons. Mike has a nice article in the Guardian today about the thing that is getting the most attention, which is the cost to publish. I blogged about it last fall, when I bought the max bling lifetime membership–for about one-tenth of the OA publication fee for a single article from one of the big barrier-based publishers.

Apatosaurus lousiae 1/12 scale skeleton, modelled by Phil Platt, assembled and photographed by Brant Bassam. Image courtesy of BrantWorks.com.

Apatosaurus lousiae 1/12 scale skeleton in left lateral view, modelled by Phil Platt, assembled and photographed by Brant Bassam. Image courtesy of BrantWorks.com.

Then there’s turnaround time: for our paper, a mere 72 days, including both submission day (Dec. 3) and publication day (Feb. 12). My fastest turnaround before this was 73 days for my sauropod nerve paper, but that was from submission to posting of the accepted manuscript, not publication of the final version of record. Prior to that I’d had a couple of papers published within six months of submission, but that was definitely the exception rather than the rule. And sadly, I’ve had several situations now where a paper  languished in peer review for six months.

And that brings me to peer review–the real “peer” in PeerJ. When you sign up a lifetime membership, you agree to review one paper a year for them to keep your membership active. Certainly not a crushing amount of work, especially since I’ve been averaging 5 or 6 reviews a year for much less congenial outlets.

I’ve seen this from both sides now, since I was tapped to review a manuscript for PeerJ back in December. The first thing I liked is that they asked for the review back within 10 days. That’s just about right. I can see a thorough review taking three days (not working straight through, obviously, but taking time to carefully read, digest, look stuff up, and compose the review), and a busy academic maybe needing a week to find that kind of time. If one is too busy to get it done within 10 days, better to just be honest, say that, and decline the review. There is certainly no reason to let reviewers have manuscripts for four to six weeks, let alone the three to four months that was standard when I got into this business.

Apatosaurus lousiae 1/12 scale skeleton in dorsal view, modelled by Phil Platt, assembled and photographed by Brant Bassam. Image courtesy of BrantWorks.com.

Apatosaurus lousiae 1/12 scale skeleton in dorsal view, modelled by Phil Platt, assembled and photographed by Brant Bassam. Image courtesy of BrantWorks.com.

The second thing I liked is that they gave me the option to sign the review (which is almost always implicitly present, whether reviewers take advantage of it or not), and they gave the authors of the manuscript the option to publish my review alongside the paper. I love that. It means that, for the first time ever*, maybe the time and effort I put into the review will not disappear without a trace after I send it off. (It is astonishingly wasteful that we write these detailed technical critiques and then consign them to never be seen by any but a handful of people.) And it had a salutary effect on my reviewing. I always strive to be thoughtful and constructive in my reviews, but the knowledge that this review might be published for the world to see made me a lot more careful, both in what I said and how I said it. Hopefully, the authors I reviewed for will opt to publish my review, so you will be able to judge for yourself whether I succeeded–I’ll keep you posted on that. UPDATE: Hooray! The paper is out, and it’s a beaut, and the authors did publish the review history, which is excellent. The paper is Schachner et al. (2013), “Pulmonary anatomy in the Nile crocodile and the evolution of unidirectional airflow in Archosauria”, the reviews by Pat O’Connor and myself and the author responses and the editor’s letters are all available by clicking the “Peer review history” link on the sidebar, and you should go read all of it right now.

* There are a bare handful of other outlets that publish reviews alongside papers, but I’ve never been tapped to review for them, so this was my first experience with a peer review that might be published.

Naturally Mike and I took the maximum openness option and had our reviews and all the rest of the paper trail published alongside our paper, and I intend to do this every time from here on out. As far as I’m concerned, the benefits of open peer review massively outweigh those from anonymous peer review. There will always be a few jackasses in the world, and if openness itself doesn’t force better behavior out of them, at least they’ll be easier to identify and route around in an open world. Anyway, to see our reviews, expand ‘Author and article information’ at the top of this page, and click the link in the green box that says, “The authors have chosen to make the review history of this article public.”

One happy result of this will manifest in just a few weeks. Bunny-wrangler and sometime elephant-tracker Brian Kraatz and I co-teach a research capstone course for the MS students at WesternU, and one of the things we cover is peer review. Last year I had to dig up a couple of my reviews that were sufficiently old and anonymous that no harm could come from sharing them with the students, but even so, they only got half the story, because I no longer had the manuscripts and couldn’t have shared them if I had. This year I’ll be able to point the students at PeerJ and say, “Go look. There’s the back-and-forth. That’s how we do this. Now you know.”

Science, process and product alike, out in the open, freely available to the world: that’s why I’m proud to be a member of PeerJ.

(And I haven’t even mentioned the preprint server, or all the thought the PeerJ team put into the graphic design of the papers themselves, or how responsive the production team was in helping us get the finished product just right, or….)

Apatosaurus lousiae 1/12 scale skeleton in left postero-lateral view, modelled by Phil Platt, assembled and photographed by Brant Bassam. Image courtesy of BrantWorks.com.

Apatosaurus lousiae 1/12 scale skeleton in left postero-lateral view, modelled by Phil Platt, assembled and photographed by Brant Bassam. Image courtesy of BrantWorks.com.

The pictures in this post have nothing to do with our paper, other than showing off one of the beautiful products of the factors we discuss therein. The images are all borrowed from Brant Bassam’s amazing BrantWorks, which we will definitely be discussing more in the future. Explicit permission to reproduce the images with credit can be found on this page. Thanks, Brant!

UPDATE: Bonus Figure

This special version of Figure 3 from our new paper goes out to Dean, who inspired it with this comment. As Tony Stark said, “It’s like Christmas, only with more…me.” Click to enWedelate.

Matt Wedel (6'2" or 1.88m tall) with various long-necked amniotes for scale.

A selection of Matt Wedels (6’2″ or 1.88m tall) with various long-necked amniotes for scale.

You may remember this:

Rapetosaurus mount at Field Museum

…which I used to make this:

Rapetosaurus skeleton silhouette

…and then this:

Rapetosaurus skeleton silhouette - high neck

The middle image is just the skeleton from the top photo cut out from the background and dropped to black using ‘Levels’ in GIMP, with the chevrons scooted up to close the gap imposed by the mounting bar.

The bottom image is the same thing tweaked a bit to repose the skeleton and get rid of some perspective distortion on the limbs. The limb posture is an attempt to reproduce an elephant step cycle from Muybridge.

That neck is wacky. Maybe not as wrong as Omeisaurus, but pretty darned wrong. As I mentioned in the previous Rapetosaurus skeleton post, the cervicals are taller than the dorsals, which is opposite the condition in every other sauropod I’ve seen. All in all, I find the reposed Rapetosaurus disturbingly horse-like. And oddly slender through the torso, dorsoventrally at least. The dorsal ribs look short in these lateral views because they’re mounted at a very odd, laterally-projecting angle that I think is probably not correct. But the ventral body profile still had to meet the distal ends of the pubes and ischia, which really can’t go anywhere without disarticulating the ilia from the sacrum (and cranking the pubes down would only force the distal ends of the ilia up, even closer to the tail–the animal still had to run its digestive and urogenital pipes through there!). So the torso was deeper than these ribs suggest, but it was still not super-deep. Contrast this with Opisthocoelicaudia, where the pubes stick down past the knees–now that was a tubby sauropod. Then again, Alamosaurus has been reconstructed with a similarly compact torso compared to its limbs–see the sketched-in ventral body profile in the skeletal recon from Lehman and Coulson (2002: figure 11).

I intend to post more photos of the mount, including some close-ups and some from different angles, and talk more about how the animal was shaped in life. And hopefully soon, because history has shown that if I don’t strike while the iron is hot, it might be a while before I get back to it. For example, I originally intended this post to follow the last Rapetosaurus skeleton post by  about a week. So much for that!

Like everything else we post, these images are CC BY, so feel free to take them and use them. If you use them for the basis of anything cool, like a muscle reconstruction or life restoration, let us know and we’ll probably blog it.

In the recent post on OMNH 1670, a dorsal vertebra of a giant Apatosaurus from the Oklahoma panhandle, I half-promised to post the only published figure of this vertebra, from Stovall (1938: fig. 3.3). So here it is:

And in the second comment on that post, I promised a sketch from one of my notebooks, showing how much of the vertebra is reconstructed. Here’s a scan of the relevant page from my notebook. Reconstructed areas of the vert are shaded (confusingly, using strokes going in opposite directions on the spine and centrum, and the dark shaded areas on the front of the transverse processes are pneumatic cavities), and measurements are given in mm.

Next item: is this really a fifth dorsal vertebra?

Apatosaurus louisae CM 3018 D4 and D5, in anterior (top), left lateral, and posterior views, from Gilmore (1936: plate 25).

Here are D4 and D5 of A. louisae CM 3018. They sort of bracket OMNH 1670 in terms of morphology. D4 has a broader spine, and D5 has a narrower one. The spine of D5 lacks the slight racquet-shaped expansion seen in OMNH 1670, but the overall proportions of the spine are more similar. On the other hand, the transverse processes of D4 taper a bit in anterior and posterior view, as in OMNH 1670, and unlike the transverse processes of D5 with their more parallel dorsal and ventral margins. But honestly, neither of these verts is a very good match (and the ones on either side, D3 and D6, are even worse).

Apatosaurus parvus UWGM 15556 (formerly A. excelsus CM 563) D4 (left) and D3 (right) in anterior (top), right lateral, and posterior views, from Gilmore (1936: plate 32).

Here are D3 and D4 of A. parvus UWGM 15556. D3 is clearly a poor match as well–it is really striking how much the vertebral morphology changes through the anterior dorsals in most sauropods, and Apatosaurus is no exception. D3 looks like a dorsal in lateral view, but in anterior or posterior view it could almost pass for a posterior cervical. If I was going to use the term “cervicodorsal”, indicating one of the vertebrae from the neck/trunk transition, I would apply it as far back as D3, but not to D4. That thing is all dorsal.

And it’s a very interesting dorsal from the perspective of identifying OMNH 1670. It has fairly short, tapering transverse processes. The neural spine is a bit shorter and broader, but it has a similar racquet-shaped distal expansion. I’m particularly intrigued by the pneumatic fossae inscribed into the anterior surface of the neural spine–in Gilmore’s plate they make a broken V shapen, like so \ / (or maybe devil eyes). Now, OMNH 1670 doesn’t have devil eyes on its spine, but it does have a couple of somewhat similar pneumatic fossae cut into the spine just below the distal racquet–perhaps a serially modified iteration of the same pair of fossae as in the A. parvus D4. It’s a right sod that D5 from this animal has its spine blown off–but it still has its transverse processes, and they are short and tapering as in OMNH 1670.

Apatosaurus sp. FMNH P25112, dorsal vertebrae 1-10 and sacrals 1 and 2, Riggs (1903: plate 46)

Here are all the dorsals and the first couple sacrals of FMNH P25112, which was originally described as A. excelsus but in the specimen-level analysis of Upchurch et al. 2005) comes out as the sister taxon to the A. ajax/A. parvus/A. excelsus clade. Note the striking similarity of the D5 here with D4 of the A. parvus specimen in Gilmore’s plate (until the careful phylogenetic work up Upchurch et al. 2005, that A. parvus specimen, once CM 563 and now UWGM 15556, was considered to represent A. excelsus as well). But  also notice the striking similarity of D6 to OMNH 1670. It’s not quite a dead ringer–the transverse processes are longer and have weird bent-down “wingtips” (XB-70 Valkyrie, anyone?)–but it’s pretty darned close, especially in the shape of the neural spine.

So what does this all mean? First, that trying to specify the exact serial position of an isolated vertebra is nigh on to impossible, unless it’s something that is one-of-a-kind like an axis. Second, after doing all these comparos I think it’s unlikely that OMNH 1670 is a D4–those are a bit too squat across the board–but it could plausibly be either a D5 or a D6. Third, I’m really happy that it doesn’t seem to match any particular specimen better than all the rest. What I don’t want to happen is for someone to see that this vertebra looks especially like specimen X and therefore decide that it must represent species Y. As I said in the comments of the previous post, what this Oklahoma Apatosaurus material needs is for someone to spend some quality time seeing, measuring, and photographing all of it and then doing a phylogenetic analysis. That sounds like an ambitious master’s thesis or the core of a dissertation, and I hope an OU grad student takes it on someday.

If you were intrigued by my suggestion that the big Oklahoma Apatosaurus rivalled Supersaurus in size, and wanted to see a technical comparison of the two, I am happy to report that Scott Hartman has done the work for you. Here’s one of his beautiful Apatosaurus skeletal reconstructions, scaled to the size of OMNH 1670, next to his Supersaurus silhouette. This is just a small teaser–go check out his post on the subject for a larger version and some interesting (and funny) thoughts on how the two animals compare.

References

  • Gilmore, C.W. 1936. Osteology of Apatosaurus with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11:175-300.
  • Riggs, E.S. 1903. Structure and relationships of opisthocoelian dinosaurs, part I: Apatosaurus Marsh. Field Columbian Museum Publications, Geological Series 2(4): 165–196.
  • Stovall, J.W. 1938. The Morrison of Oklahoma and its dinosaurs. Journal of Geology 46:583-600.

We’re starting the new year with a new feature, in which we answer questions that have come our way. We never had a policy about not answering questions, it’s just that previous ones have tended to arrive in the comments section and have been dealt with there. But suddenly in the last few days I’ve gotten two questions from extrabloggular sources, and rather than hide the replies I thought I’d make them available to all.

One of my cohort at Berkeley texted me the other day with the following questions:

OK, phylobuddy: can you suck the marrow from a chicken bone? If they have hollow bones, where’s the marrow?!? Google is getting me nowhere.

Short answer: yes, one can get marrow from chicken bones, from those bones that contain marrow rather than air. In most fully mature chickens, the pneumatic bones include the braincase, the cervical, dorsal, and most or all synsacral vertebrae, some of the dorsal ribs, the central portion of the sternum, the coracoids, and the humeri (if you’re not a regular and some of these terms are unfamiliar, check out these handy guides [1, 2] to the vertebrate skeleton). That leaves marrow in everything else, although the only bones with large marrow cavities–as opposed to tiny trabecular spaces, which also house marrow–are the radii, ulnae, femora, tibiotarsi, and tarsometatarsi. So if you want to actually see large amounts of chicken marrow, or suck the marrow out of chicken bones, you’re basically stuck with the big distal bones of the wing, the thigh, and the drumstick (tibiotarsus). If you are boiling chicken bones to get stock for soups or stews, might as well throw them all in; even the pneumatic bones will still have bits of adhering meat, cartilage, and ligaments that will give up molecules and flavor to the stock.

The long answer is that the expression “hollow bones” has caused no end of confusion, because there are at least two ways to interpret hollow: filled with air, or not filled with bone (the former is a subset of the latter). If you mean “not filled with bone”, then the bones of almost all amniotes* are hollow, and the spaces inside are occupied by marrow (most commonly) or air. If filled with air, the bones are referred to as pneumatic, and an accessible introduction to them is here.

* At least; I know less about amphibians and fish, although at least one osteoglossomorph (IIRC) pneumatizes its vertebrae from its swim bladder!

The reasons it gets confusing are twofold. First, sometimes authors describe bones as hollow and mean only that they have chambers inside, but later readers see ‘hollow’ and infer ‘pneumatic’. Not all hollow bones are pneumatic; in fact, the vast majority of them are not, including the long bones of your arms and legs. The criteria for inferring pneumaticity from dry bones are more strict, and are explored in this paper and this one. Anyway, this point is just confusion caused by an ambiguous term.

The second case is more interesting, because it involves real unknowns. In the fossil record we can almost always tell if a bone is hollow, sensu lato, but sometimes it is not possible to say for certain whether the hollow space(s) inside were filled with marrow or air. Particularly vexing and intriguing examples include the humerus of Eotyrannus and the iliac chambers of some sauropods, which are discussed in this paper. My guess is that the iliac chambers of sauropods are genuinely pneumatic, because they only occur in sauropods that already have sacral pneumaticity, and we know from broken ilia of more basal sauropods and sauropodomorphs that large marrow-filled chambers are not present in those taxa. Conversely, I suspect that the humerus of Eotyrannus was apneumatic (marrow-filled), given that humeral pneumaticity is otherwise unknown in non-avian theropods, although the pneumatic furcula of Buitreraptor at least shows that the necessary clavicular air sac was present in some.

Next question! This one came to me on Facebook, from ReBecca Hunt-Foster, whom you may know from her awesome Dinochick Blogs. You should also envy her and hubby John Foster for getting the most awesome wedding present of all time: a 1/12 scale skeleton of Apatosaurus sculpted by Phil Platt, which you can read about here. That’s cool enough that I am stealing it for this otherwise picture-challenged post.

ANYWAY, ReBecca wrote on my FB wall today to ask:

Random question: Have you seen many tooth marks on sauro cervical verts? I am debating on whether something I have is a dessication crack or really some tooth marks. Thanks :)

In all the 15 years that I have spent looking at sauropod remains in the bowels of many, many museums, I have never seen a single tooth mark on a sauropod vertebra.

[Update the next day: Er, except for the bitten Apatosaurus tail on display in the AMNH! Many thanks to reptilianmonster and steve cohen for reminding me about this in the comments. I'm going to go hide for a while now.]

Now, that doesn’t mean that they aren’t there. Truth be told, I’ve never looked for them, and my usual mental search pattern for pneumatic traces (large, irregular) would probably exclude tooth scratches (small, linear) as noise. But I’ve certainly never seen any vertebrae with easily recognizable signs of predation or scavenging or with obvious bites removed.

People also sometimes ask me what kinds of healed traumas I’ve seen in pneumatic sauropods bones. That’s easy: apart from vertebral fusions, most of which probably have nothing to do with trauma, I’ve seen zip. Nada. Null set. The wingspan of the average tadpole. I’ve seen some pretty cool pneumatic bones from extant birds that were broken and later healed, including a eagle femur in the UCMP comparative collection that is now shaped like the letter Z, but nothing in sauropods.

I can think of three possible reasons for this, which sort of flow into each other. The first is that apart from the very solid and blocky centra of apneumatic vertebrae, sauropod verts were pretty fragile, and prone to getting distorted and busted up even when they started out intact, and those verts that started out broken just had a tougher time with the taphonomic lottery.

The second is that pneumatic sauropod bones would been nothing to most predators other than a mouthful of relatively dry bone shards, so either carnivores left them alone, or if they were osteovores like T. rex, they ate the shards and whatever is left over is unrecognizable. I have seen, and mostly ignored, plenty of vert-shrapnel in quarries and in collections, and maybe sharper eyes than mine could have discerned evidence of predation from those bits. To me it mostly looked like trampling, hydraulic transport, erosion, and other mundane ways to explode a vertebra.

The third is that in addition to a preservation bias against half-destroyed verts, there is probably also a collection bias against them. I’m probably not the only one would pass up a few shards of excellence to dig out the complete fibula sitting next to them in the quarry, and I love this stuff. That said, we did get a LOT of blasted vert bits out of the Wolf Creek quarry in the Cloverly, so if you want to pore over sauropod shards looking for tooth marks, visit the OMNH.

And, if you do know of tooth marks on sauropod vertebrae, please let us know in the comments. And consider publishing them, given the apparent vacuum of such things.

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