An Apatosaurus worthy of All Yesterdays
December 11, 2012
A few months ago, Matt and Darren saw a picture someone had done of an Apatosaurus with huge neck-flaps. Since they, they’ve tried to find it again but without success. Then, happily, I stumbled across it in this All Yesterdays review, so here it is:
Unfortunately, I can’t tell you much about it. I know it’s the work of Emiliano Troco, but I’ve not been able to find his web-site, nor a description of the piece, nor a version in a decent resolution. So all we have to go on at the moment is this thumbnail. If you know more, please leave a comment!
Is it credible? Who’s to say? The one thing we know for certain about Apatosaurus is that it had truly crazy cervical vertebrae, unlike those of any other animal. In our recent arXiv paper, we wrote:
It is difficult to see the benefit in Apatosaurus excelsus of cervical ribs held so far below the centrum – an arrangement that seems to make little sense from any mechanical perspective, and may have to be written off as an inexplicable consequence of sexual selection or species recognition.
It certainly seems to have been doing something weird with its neck. It’s no obvious why big flaps like these would require honking great cervicals ribs to hang down from, but maybe it was swinging them around or something?
[We've featured bizarrely ornamented sauropods here before, notably Brian Engh's pouch-throated Sauroposeidon.]
2000 AD’s flagrantly plagiarised Brontosaurus
October 18, 2012
Another blast from the past:
Like the recent Compsognathus, this is a card from the “Flesh” card-game that was printed across several progs (issues) of the comic 2000 AD in 1977. This one is from the back cover of Prog 10. (Click through the picture for the whole back cover.)
What’s interesting about this one is how very flagrant a rip-off it is of Rudolph Zallinger’s 1960 painting of Brontosaurus being attacked by Allosaurus:
I know this painting best from Dinosaurs and other Prehistoric Reptiles, a 1966 book that I had as a boy, and which I believe is the same thing as the Giant Golden Book of Dinosaurs. Here is a high-resolution scan of my copy of that book, pages 24-25. (Click through for 5472 by 3669 version.)
And while I’m here, I may as well throw in my scan of the “Brachiosaurus” (i.e. Giraffatitan)on pages 20-21. (Click through for 5431 by 3162 version.)
I will leave it to others to point out which other classic piece of sauropod art this one plagiarises.
AMNH mounted Apatosaurus with Taylor for scale
September 8, 2012
We’re off to Oxford next week for SVPCA, so things may be quiet around here for a few days. Catch you on the flip side.
Apatosaurus sacra of the National Science Museum, Tokyo
July 19, 2012
In a comment on the previous post, Steve P. asked whether “Apatosaurus” minimus might not be a Apatosaurus specimen after all — particularly, an Apatosaurus ajax individual resembling NSMT-PV 20375, the one in the National Science Museum, Tokyo, that Upchurch et al. (2005) so lavishly monographed.
Initially, I dismissed this idea out of hand, because the “Apatosaurus” minimus sacrum-pelvis complex is so very different to that of the “Brontosaurus” illustrated by Hatcher (1903: fig. 4), as seen in an earlier post. But on going back to the Upchurch et al. monograph I realised that their sacrum-ilium complex is very different from Hatcher’s. Here it is, cleaned up from scans and re-composed in the same format as the Camarasaurus and “Apatosaurus” minimus from last time, for easy comparison.
Sacrum and fused ilia of Apatosaurus ajax NSMT-PV 20375. Top row: dorsal view with anterior to left. Middle row, left to right: anterior, right lateral (reversed), posterior. Bottom row: ventral view with anterior to left. Modified from Upchurch et al. (2005: plate 4 and text-figure 9).
Here’s Hatcher’s “Brontosaurus” illustration (from his plate 4) again:
I’m not sure what to make of this. The Tokyo Apatosaurus seems to be intermediate in some respects between Hatcher’s specimen and “Apatosaurus” minimus.
One important difference is that the neural spines are much taller in Hatcher’s illustration than in the Tokyo Apatosaurus. Could that be ontogenetic? (IIRC the Tokyo individual is subadult). Or are they in fact different species? Or is it just individual variation?
I don’t know. Anyone?
References
- Hatcher, J.B. 1903. Osteology of Haplocanthosaurus with description of a new species, and remarks on the probable habits of the Sauropoda and the age and origin of the Atlantosaurus beds; additional remarks on Diplodocus. Memoirs of the Carnegie Museum 2:1-75.
- Upchurch, Paul, Yukimitsu Tomida, and Paul M. Barrett. 2005. A new specimen of Apatosaurus ajax (Sauropoda: Diplodocidae) from the Morrison Formation (Upper Jurassic) of Wyoming, USA. National Science Museum Monographs No.26. Tokyo.
Upchurch et al. (2005)
How fat was Brontosaurus? Well, not this fat, anyway!
July 16, 2012
This is the Brontosaurus that I grew up with:
It’s by Kenyon Shannon, found on page 14 of The How And Why Wonder Book of Dinosaurs (Geis 1960). I call it Brontosaurus rather than Apatosaurus because this outdated rendition is forever tied to the outdated name in my mind.
It would be terribly easy to pick holes in this representation — the completely wrong head, the bizarre croc-like eyes, the incorrect attitude of the head, the tubular shape and much too thin profile of the neck, the full set of manual unguals and so on. But the part that’s most shocking to me is just how darned fat it is.
Even given that Apatosaurus was among the most robustly build of sauropods — certainly the most robust diplodocoid — how wrong is that super-fat forearm? Here, for comparison, is an Apatosaurus humerus:
Right humerus of Apatosaurus ajax NSMT-PV 20375 in (left to right) anterior, medial, lateral and posterior views, and (top to bottom) proximal and distal views. From Upchurch er al. (2005:fig. 5)
And here is the lower arm:
Right forearm of Apatosaurus ajax NSMT-PV 20375. A: “anterior” (more like anterolateral) view with ulna to left, radius to right; B: “posterior” (more like posteomedial) view with radius to left, ulna to right; C; proximal view; D: distal view.
Put them together and you can get some impression of the proportions of the complete forelimb:
In short: it’s a solidly build animal; but the Shannon depiction that we started with is not merely solid, it’s what we refer to as “a lardy bloater”. Just say no.
References
- Geis, Darlene. 1960. The How and Why Wonder Book of Dinosaurs. Price Stern Sloan, Los Angeles. 48 pages. ISBN: 0-8431-4250-2.
- Upchurch, Paul, Yukimitsu Tomida, and Paul M. Barrett. 2005. A new specimen of Apatosaurus ajax (Sauropoda: Diplodocidae) from the Morrison Formation (Upper Jurassic) of Wyoming, USA. National Science Museum Monographs No. 26. Tokyo. ISSN 1342-9574.
I don’t have time to write about this properly, but a few people have asked me about the new Sellers et al. (2012) paper on measuring the masses of extinct animals — in particular, the Berlin Giraffatitan — by having a CAD program generate minimal complex hulls around various body regions. Rather than write something new about it, I’m going to publish the comments that I sent Ed Yong for his Discover piece on the new technique:
Hi, Ed, good to hear from you. Yes, it’s a good paper: a useful new technique that has some useful properties, most importantly that it requires no irreproducible judgements on the part of the person using it, and that it’s ground-truthed on solid data from extant animals.
It’s a reassuring sanity-check to find that my (2009) mass estimate falls well within their method’s 95% confidence interval, and is in fact within 0.6% of their best estimate.
There are a couple of problems with this study, which I hope will be addressed in followups. The authors are honest enough to touch on all of these problems themselves, though! They are:
1. All the extant animals used to determine the fudge factor are mammals, which means they are not necessarily completely relevant to dinosaurs. In particular I would very much like to have seen regression lines and correlation coefficients for this method for birds and crocodilians, both of which are much more closely related to Giraffatitan.
2. Much depends on the reconstruction of the torso, particular the position of the ribs, which is very difficult to do well and confidently with dinosaurs. In my volumetric analysis (Taylor 2009:803) I found that the torso accounts for 70% of total body volume in Giraffatitan, so rib orientation will make a big difference to overall mass. Sauropod ribs that are well preserved and undistorted along their whole length are extremely rare.
3. Use of a single density value for the whole animal, while appropriate for mammals, really isn’t for brachiosaurs, in which the very long neck likely had a density no more than half that of the legs. I’m not sure what can be done about this, though, since any attempt to correct for density variation involves subjective guesswork. Then again, so do all guesses at overall body density in dinosaurs.
Issue 1 bothers me most, because the convex hulls of limb segments in mammals will be proportionally much larger than in sauropods, due to the complex shapes of mammalian long-bone ends. I worry that using mammals as a baseline will underestimate sauropod leg mass.
Still, even with these caveats, it’s a good exposition of an important new method which I expect to see widely adopted.
Hope that’s helpful.
In short: good work, widely applicable, and probably the best mass-estimation technique we now have available for complete and near-complete skeletons. It would be good to see it applied to (say) the Yale, AMNH and CM apatosaurs.
Composite illustration from Sellers et al.’s press release. Top left: bear skeleton from the Oxford University Natural History Museum, presumably Ursus maritimus: original skeleton, derived point cloud and convex hulls (also used as Sellers et al. 2012:fig. 1). Top right: shedloads of awesome. Bottom: complex hulls around body segments of Giraffatitan.
References
The giant Oklahoma Apatosaurus: OMNH 1670 redux
April 30, 2012
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.
The giant Oklahoma Apatosaurus: OMNH 1670
April 25, 2012
Left: the Queen of England, 163 cm. Middle, the Oklahoma apatosaur dorsal, 135 cm. Right, classic “big Apatosaurus” dorsal, 106 cm. To scale.
Something I’ve always intended to do but never gotten around to is posting on some of the immense Apatosaurus elements from the Oklahoma panhandle. Here’s one of the most impressive, OMNH 1670, an isolated dorsal. Notice that the tip of the neural spine is ever-so-shallowly bifurcated, which in Apatosaurus indicates a D4, D5, or D6. The low parapophyses and fat transverse processes are similar to D4, but Apatosaurus D4s usually have somewhat broader spines, so I’m guessing this thing is a D5. These things vary and I could easily be off by a position in either direction.
Next to it is D5 of CM 3018, the holotype specimen of Apatosaurus louisae (from Gilmore 1936: plate 25), which has served as the basis for many of the published mass estimates of the genus Apatosaurus. OMNH 1670 is 135 cm tall, compared to 106 cm for D5 of CM 3018. If the rest of the animal scaled the same way, it would have been 1.27^3 = 2 times as massive. Mass estimates for CM 3018 are all over the map, from about 18 tons up to roughly twice that, so the big Oklahoma Apatosaurus was probably in Supersaurus territory, mass-wise, and may have rivaled some of the big titanosaurs (Update: see the two giant diplodocids square off in a cool follow-up post by Supersaurus wrangler Scott Hartman). Here’s a fun rainy-day activity: take any skeletal reconstruction of Apatosaurus, clone it in Photoshop or GIMP, scale it up by 27%, and park it next to the original. It looks a lot bigger. So I’m continually surprised that Apatosaurus is so rarely mentioned in the various roundups of giant sauropods, both in the technical literature and in popular articles online. This vertebra was figured by Stovall (1938)–if I get inspired, I’ll dig up that figure and post it another day (hey, look, I did!).
Fun fact: in Apatosaurus the tallest (most posterior) dorsals are 1.3-1.5 times as tall as D5 (Gilmore 1936: 201). So D10 from this individual was probably between 1.7 and 2 meters tall–not quite in Amphicoelias fragillimus territory but getting closer than I’ll bet most people suspected.
NB: if you try to use the scale bar lying on the centrum of OMNH 1670 to check my numbers, you will get a wonky answer. The problem is that the vertebra is so large that it is almost impossible to get far enough back from it (above it, in this case, since it is lying on a padded pallet) to get a shot free from distortion due to parallax. For this shot, the pallet with the vert was on the floor, and I was standing on top of the tallest ladder in the OMNH collections, leaning out over the vert to get centered over the prezygapophyses, and shooting straight down–in other words, I had done everything possible to minimize the visual distortion. But it still crept in. Anyway, trust the measurements, which I–and presumably Gilmore–made with a good old reliable tape measure.
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.
- Stovall, J.W. 1938. The Morrison of Oklahoma and its dinosaurs. Journal of Geology 46:583-600.
Neural spine bifurcation in sauropods, Part 6: more reasons why Haplocanthosaurus is not a juvenile of a known diplodocid
April 15, 2012
Last time, we saw why Haplocanthosaurus couldn’t be a juvenile of Apatosaurus or Diplodocus, based on osteology alone. But there’s more:
Ontogenetic status of Haplocanthosaurus
Here is where is gets really surreal. Woodruff and Fowler (2012) blithely assume that Haplocanthosaurus is a juvenile of something, but the type specimen of the type species — H. priscus CM 572 — is an adult. As Hatcher (1903:3) explains:
The type No. 572 of the present genus consists of the two posterior cervicals, ten dorsals, five sacrals, nineteen caudals, both ilia, ischia and pubes, two chevrons, a femur and a nearly complete series of ribs, all in an excellent state of preservation and pertaining to an individual fully adult as is shown by the coössified neural spines and centra.
So far as I can see, Woodruff and Fowler are confused because the second species that Hatcher describes, H. utterbacki, is based on the subadult specimen CM 879. Where possible in the previous post, I have used illustrations of the adult H. priscus, so that the comparisons are of adult with adult. The exceptions are the two anterior cervicals and the first dorsal, which are known only from H. utterbacki. And sure enough, if you look closely at the illustrations, you can see that in these vertebrae and only these vertebrae, Hatcher had the neurocentral junction illustrated — because it wasn’t yet fused.
Haplocanthosaurus posterior, mid and anterior cervical vertebrae, C14, C9 and C4, in right lateral view. C14 of adult H. priscus (from Hatcher 1903:plate I); C9 and C4 of H. utterbacki (from plate II). Red ellipses highlight neurocentral sutures.
As it happens, the difference in ontogenetic status between these two specimens is nicely illustrated by Wedel (2009), although he was only in it for the pneumaticity:
Neurocentral fusion in Haplocanthosaurus. A, B. Posterior cervical vertebra C?12 of sub-adult H. utterbacki holotype CM 879: A, X-ray in right lateral view; B, coronal CT slice showing separate ossificaton of centrum and neural arch. C, D. Mid-dorsal vertebra D6 of adult H. priscus holotype CM 572: X-rays in (A) right lateral and (B) anterior view, showing fully fused neural arch. Wedel (2009:fig. 6)
So H. utterbacki CM 879 certainly is an immature form of something; and that something is Haplocanthosaurus, most likely H. priscus. (The characters which Hatcher used to separate the two species are not particularly convincing.)
With that out the way, we can move on to …
Phylogenetic analysis
A simple way to evaluate the parsimony or otherwise of a synonymy is to use a phylogenetic analysis. In their abstract, Woodruff and Fowler claim that “On the basis of shallow bifurcation of its cervical and dorsal neural spines, the small diplodocid Suuwassea is more parsimoniously interpreted as an immature specimen of an already recognized diplodocid taxon”. Without getting into the subject of Suuwassea again — Matt pretty much wrapped that up in part 4 — the point here is that the word “parsimony” has a particular meaning in studies of evolution: it refers to minimising the number of character-state changes. And we have tools for measuring those.
So let’s use parsimony to evaluate the hypothesis that Haplocanthosaurus is one of the previously known diplodocids. Pretending for the moment that Haplocanthosaurus really was known only from subadults, how many additional steps would we need to account for if ontogeny were to change its position to make it group with one of the diplodocids?
You don’t need to be a cladistics wizard to do this. (Which is handy, since I am not one.) Here’s the method:
- Start with an existing matrix, add constraints, re-run it, and see how the tree-length changes. Since I am familiar with it, I started with the matrix from my 2009 paper on brachiosaurs.
- Re-run the matrix to verify that you get the same result as in the published paper based on it. This gives you confidence that you’re running it right. In this case, I got a minimum tree length of 791 steps, just as in Taylor (2009).
- Add extra instructions to the run-script defining and imposing constraints. Note that you do not have to mess with the characters, taxa or codings to do this.
- Run the matrix again, with the constraint in place, and see how the tree-length changes.
- Repeat as needed with other constraints to evaluate other phylogenetric hypotheses.
(This is how we produced the part of the Brontomerus paper (Taylor et al. 2011:89) where we said “One further step is sufficient to place Brontomerus as a brachiosaurid, a basal (non−camarasauromorph) macronarian, a basal (non−diplodocid) diplodocoid or even a non−neosauropod. Three further steps are required for Brontomerus to be recovered as a saltasaurid, specifically an opisthocoelicaudiine”. And that’s why we weren’t at all dogmatic about its position.)
Anyway, going through this exercise with Haplocanthosaurus constrained in turn to be the sister taxon to Apatosaurus, Diplodocus, etc., yielded the following results:
- (no constraint) – 791 steps
- Apatosaurus — 817 (26 extra)
- Diplodocus — 825 (34 extra)
- Barosaurus — 815 (24 extra)
- Camarasaurus — 793 (2 extra)
- Brachiosaurus — 797 (6 extra)
(I threw in the other well-known Morrisson-Formation sauropods Camarasaurus and Brachiosaurus, even though Woodruff and Fowler don’t mention them, just because it was easy to do and I was interested to see what would happen. And when I say Brachiosaurus, I mean B. altithorax, not Giraffatitan.)
I hope you’re as shocked as I am to see that for Haplocanthosaurus to emerge as the sister taxon of any diplodocid needs a minimum of 24 additional steps — or an incredible 34 for it to be sister to Diplodocus. In other words, the hypothesis is grossly unparsimonious. Of course, that doesn’t in itself mean that it’s false: but it does render it an extraordinary claim, which means that it needs extraordinary evidence. And while “the simple spines of Haplocanthosaurus might bifurcate when it grows up” is extraordinary evidence, it’s not in the way that Carl Sagan meant it.
In short, running this simple exercise — it took me about a hour, mostly to remember how to do constraints in PAUP* — would have given Woodruff and Fowler pause for thought before dragging Haplocanthosaurus into their paper.
Oh, and it’s ironic that placing Haplo as sister to Brachiosaurus requires only a quarter as many steps as the closest diplodocid, and as sister to Camarasaurus requires only two steps. If you really want to synonymise Haplocanthosaurus, Camarasaurus is the place to start. (But don’t get excited, it’s not Camarasaurus either. It’s Haplocanthosaurus.)
[By the way, anyone who'd like to replicate this experiment for themselves is welcome: all the files are available on my web-site. You only really need the .nex file, which you can feed to PAUP*, but I threw in the log-file, the generated tree files and the summary file, too. Extra Credit: run this same exercise to evaluate the parsimony of Suuwassea as a subadult of one of these other genera. Report back here when you're done to earn SV-POW! points.]
Conclusion
It’s a truism that we stand on the shoulders of giants. In the case of sauropod studies, those giants are people like J. B. Hatcher, Charles Gilmore, Osborn and Mook and — bringing it up to date — John McIntosh, Paul Upchurch, Jeff Wilson and Jerry Harris. When Hatcher described Haplocanthosaurus as a new genus rather than a subadult Diplodocus, he wasn’t naive. He recognised the effects of ontogeny, and he was aware that one of his two specimens was adult and the other subadult. He was also probably more familiar with Diplodocus osteology than anyone else has ever been before or since, having written the definitive monograph on that animal just two years previously (Hatcher 1901).
By the same token, people like Upchurch and Wilson have done us all a huge favour by making the hard yards in sauropod phylogenetics. If we’re going to go challenging the standard consensus phylogeny, it’s just good sense to go back to their work (or the more recent work of others, such as Whitlock 2011), re-run the analyses with our pet hypotheses encoded as constraints, and see what they tell us.
So in the end, my point is this: let’s not waste our giants. Let’s take the time to get up on their shoulders and survey the landscape from up there, rather than staying down at ground level and seeing how high we can jump from a standing start.
The rest of the series
Links to all of the posts in this series:
- Part 1: what we knew a month ago
- Part 2: why serial position matters
- Part 3: the evidence from ontogenetic series
- Part 4: is Suuwassea a juvenile of a known diplodocid?
- Part 5: is Haplocanthosaurus a juvenile of a known diplodocid?
- Part 6: more reasons why Haplocanthosaurus is not a juvenile of a known diplodocid
and the post that started it all:
References
- Hatcher, J.B. 1901. Diplodocus (Marsh): its osteology, taxonomy, and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63.
- Hatcher, J.B. 1903. Osteology of Haplocanthosaurus with description of a new species, and remarks on the probable habits of the Sauropoda and the age and origin of the Atlantosaurus beds; additional remarks on Diplodocus. Memoirs of the Carnegie Museum 2:1-75.
- Taylor, M.P. 2009. A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensch 1914). Journal of Vertebrate Paleontology 29(3):787-806.
- Taylor, M.P., Wedel, M.J. and Cifelli, R.L. 2011. A new sauropod dinosaur from the Lower Cretaceous Cedar Mountain Formation, Utah, USA. Acta Palaeontologica Polonica 56(1):75-98. doi:10.4202/app.2010.0073
- Wedel, M.J. 2009. Evidence for bird-like air sacs in saurischian dinosaurs. Journal of Experimental Zoology 311A:611-628.
- Whitlock, J.A. 2011. A phylogenetic analysis of Diplodocoidea (Saurischia: Sauropoda). Zoological Journal of the Linnean Society 161(4):872-915. doi: 10.1111/j.1096-3642.2010.00665.x
- Woodruff, D.C, and Fowler, D.W. 2012. Ontogenetic influence on neural spine bifurcation in Diplodocoidea (Dinosauria: Sauropoda): a critical phylogenetic character. Journal of Morphology, online ahead of print.
Neural spine bifurcation in sauropods, Part 5: is Haplocanthosaurus a juvenile of a known diplodocid?
April 14, 2012
Introduction
Last time around, Matt walked through a lot of the detailed cervical morphology of Suuwassea and known diplodocids to show that, contra the suggestion of Woodruff and Fowler (2012), Suuwassea is distinct and can’t be explained away as an ontogenomorph of a previously known genus.
Although Suuwassea is singled out for special treatment in this paper, other genera do not escape unscathed. From the Conclusions section on page 9:
Just as particularly large diplodocid specimens (e.g., Seismosaurus; Gillette, 1991) have been more recently recognized as large and potentially older individuals of already recognized taxa (Diplodocus; Lucas et al., 2006; Lovelace et al., 2007), taxa defined on small specimens (such as Suuwassea, but also potentially Barosaurus, Haplocanthosaurus, and ‘‘Brontodiplodocus’’), might represent immature forms of Diplodocus or Apatosaurus.
I have to admit I more or less fell out of my chair when I saw the suggestion that poor old Haplocanthosaurus might be Diplodocus or Apatosaurus. I think this idea comes from a misstatement in the very first sentence of the abstract:
Within Diplodocoidea (Dinosauria: Sauropoda), phylogenetic position of the three subclades Rebbachisauridae, Dicraeosauridae, and Diplodocidae is strongly influenced by a relatively small number of characters.
As a statement of fact, this is simply the opposite of the truth: in all the major phylogenetic analyses, the arrangement of subclades with Diplodocoidea is the most stable part of the tree, supported by more characters than all the other clades.
For example, in the analysis of Upchurch et al. (2004) in The Dinosauria II, fig. 13.18 shows that the nodes with the highest bootstrap percentages are Diplodocinae (96%), Dicraeosauridae (95%) and Diplodocidae (93%).
Or consider the analysis of Wilson (2002). While it’s getting on a bit, it still scores highly by being the most explicit published sauropod analysis, with comprehensive lists of apomorphies. Table 12 lists the decay indexes for the 24 nodes in the strict consensus tree. Apart from the three very basal nodes separating sauropods from their outgroups, the two highest-scoring clades are Diplodocidae and Diplodocinae (DI=7), followed by four clades all with DI=5 of which two are Dicraeosauridae and Flagellicaudata (which Wilson just called “Dicraeosauridae + Diplodocidae” as it had not yet been named). (It’s well worth reading Wilson’s Appendix 3 to see the synapomorphies supporting these nodes in the MPTs: he lists 14 separating Diplodocimorpha from the node it shares with Haplocanthosaurus, 18 separating Flagellicaudata from the node it shares with Rebbachisauridae, 16 separating Diplodocidae from the node it shares with Dicareosauridae, and seven separating Diplodocinae from the node it shares with Apatosaurus).*
* Why are the lists of apomorphies longer than the decay indexes? Because they list the apomorphies as they occur in the specific topology of the consensus tree. Nodes within that tree can be made to collapse without wiping out all the apomorphies by rejuggling other parts of the tree to move character-state transitions around. So although (for example) 26 characters separate Flagellicaudata from Rebbachisauridae (18 + 8 synapomorphies respectively) you can rejuggle the whole tree to break the monophyly of Flagellicaudata while making the entire tree only five steps longer.
Anyway, for whatever reason, Woodruff and Fowler felt that the stability of the diplodocoid clades was in question, and this presumably influenced their hypothesis that Haplocanthosaurus could be easily moved down into one of the diplodocid genera.
Next time we’ll be considering the implications for the tree. But today, let’s take a moment to do this the old-fashioned way, by looking at …
Osteology
Pelvis
Hatcher (1903), ever helpful, included a comparative plate in his monograph which should help us to evaluate the idea that Haplo is a known diplodocid:
Pelves of diplodocids and Haplocanthosaurus. 1. Pelvis of Brontosaurus excelsus (No. 568); 2. Pelvis of Diplodocus carnegii (No. 94); 3. Pelvis of Haplocanthosaurus priscus (No. 572). All seen from left side. 1, 2, 3, 4, 5 indicate neural spines of respective sacral vertebra. Presumably to scale. Direct from Hatcher (1903:plate IV).
Based on this, the pelvis of Haplocanthosaurus differs from those of the diplodocids in having a proportionally lower ilium, in the absence of the laterally facing rugosity on the posterodorsal margin of the ilium, in the very small distal expansion of the pubis and in the almost non-existent distal expansion of the ischium. These are all characters of the limb-girdle elements, which do not change greatly through ontogeny in sauropods.
But the evidence from the sacral vertebrae is just as significant: the neural spines in the sacral area are less than half as tall as in the diplodocids — and this in an animal whose dorsal neural spines are conspicuously tall. The spines are also more anteroposteriorly elongate and plate-like. What’s more, sacral spines 1, 2 and 3 have fused into a single plate in Haplocanthosaurus, while the spine of S1 remains well separated from 2 and 3 in the diplodocids. So the ontogenetic hypothesis would have to say that the spine of S1 unfuses through ontogeny. Which is not something I’ve heard of happening in any sauropod, or indeed any animal.
So the pelvis and sacrum seem distinct. But Woodruff and Fowler’s (2012) notion of ontogenetic synonymy is built on the idea that the differences in the cervical and dorsal vertebrae are ontogenetic. So let’s take a look at them.
Cervical vertebrae
Posterior, mid and anterior cervical vertebrae, in right lateral view, of (top to bottom), Haplocanthosaurus, Apatosaurus louisae CM 3018 (from Gilmore 1936:plate XXIV, reversed for ease of comparison) and Diplodocus carnegii CM 84 (from Hatcher 1901:plate III), scaled to roughly the same size. For the diplodocids, we illustrate C13, C9 and C4. For Haplocanthosaurus, we illustrate C14 of H. priscus (from Hatcher 1903:plate I) and C9 and C4 of H. utterbacki (from plate II).
It should be immediately apparent that the Haplocanthosaurus cervicals have less extensive pneumatic features than those of the diplodocids, but that is one feature which we know does vary ontogenetically. There are other differences: for example, the cervical ribs in Haplocanthosaurus are level with the bottom centrum rather than hanging below. Still, if you kind of squint a bit, you could probably persuade yourself that the Haplocanthus vertebrae look like possible juveniles of Diplodocus.
Unless you look at them from behind:
Posterior cervical vertebrae C15 and C14, in posterior view, of (top to bottom), Haplocanthosaurus priscus CM 572 (from Hatcher 1903:plate I), Apatosaurus louisae CM 3018 (from Gilmore 1936:plate XXIV) and Diplodocus carnegii CM 84 (from Hatcher 1901:plate III), scaled to the same centrum-to-neural-spine height.
(Unfortunately, these are the only Haplocanthosaurus cervical vertebrae that Hatcher had illustrated in posterior view, so we can’t compare more anterior ones.)
From this perspective, we can immediately significant differences:
- First, that unsplit spine. Yes, we know that Woodruff and Fowler (2012) have argued that it could be ontogenetic, but these are vertebrae from the most deeply bifurcated region of a diplodocid neck, in a decent sized animal, and there is nothing that so much as hints at bifurcation.
- That whacking great ligament scar running right down the back (and also the front, not pictured) of the neural spine. There is nothing like this in any diplodocid — neither on the metapophyses nor running though the trough. And remember, scars like these tend to become more prominent through ontogeny.
- The neural arch (i.e. the region between the postzygapophyses and the centrum) is taller in Haplocanthosaurus — much taller in the case of C15.
- The plates running out to the diapophyses are less dorsoventrally expanded in Haplocanthosaurus.
- The centrum is smaller as a proportion of total height — especially, much smaller than in Diplodocus.
- The parapophyses extend directly laterally rather than ventrolaterally (hence the position of the cervical ribs level with the bottom of the centrum).
So it doesn’t look good for the juvenile-diplodocid hypothesis. But let’s take a look at the …
Dorsal vertebrae
Posterior, mid and anterior dorsal vertebrae, in right lateral view, of (top to bottom), Haplocanthosaurus, Apatosaurus louisae CM 3018 (from Gilmore 1936:plate XXV, reversed for ease of comparison) and Diplodocus carnegii CM 84 (from Hatcher 1901:plate VII), scaled to roughly the same size. For the diplodocids, we illustrate D9, D5 and D2. For Haplocanthosaurus, which has four more dorsals, we illustrate D13 and D7 of H. priscus (from Hatcher 1903:plate I) and D2 of H. utterbacki (from plate II).
Here we see that Haplocanthosaurus has dorsolaterally inclined diapophyses (which we’ll see more clearly in a minute), a prominent spinodiapohyseal lamina in posterior dorsals, and no infraparapophyseal lamination. Also, the dorsal vertebrae have reached their full height by the middle of the series (in fact the last nine dorsals are startlingly similar in proportions), whereas in diplodocids, total height continues to increase posteriorly.
Now let’s see those vertebrae in posterior view:
Posterior, mid and anterior dorsal vertebrae, in posterior view, of (top to bottom), Haplocanthosaurus priscus CM 572 (From Hatcher 1903:plate I), Apatosaurus louisae CM 3018 (from Gilmore 1936:plate XXV) and Diplodocus carnegii CM 84 (from Hatcher 1901:plate VII), scaled to the same height of the mid dorsal. For the diplodocids, we illustrate D9, D5 and D1. For Haplocanthosaurus, which has four more dorsals, we illustrate D13, D6 and D1.
Here is where it all falls apart. The Haplocanthosaurus dorsals differ from those of the diplodocids in almost every respect:
- Of course we have the non-bifid spine in again, in the anterior dorsal, but let’s not keep flogging that dead horse.
- In the mid and posterior dorsals, the neurapophysis is rounded in posterior view rather than square.
- In the posterior dorsal, the neural spine has laterally directed triangular processes near the top.
- All three Haplocanthosaurus neural spines have broad, rugose ligament scars, whereas those of the diplodocids have narrow postspinal laminae.
- The neural spines (measured from the diapophyses upwards) are much shorter than in the diplodocids; but
- The neural arches (measured from the centrum up to the diapophyses) are much taller.
- The diapophyses have distinct club-like rugosities at their tips.
- the diapophyses of the mid and posterior dorsals are inclined strongly upwards
- The hyposphenes of mid and posterior dorsals have very long centropostzygapophyseal laminae curving up in a gentle arch.
- The centra are smaller than those of Apatosaurus, and much smaller than those of Diplodocus.
(By the way, it’s interesting how very different the D5s of Apatosaurus and Diplodocus are. Since both are from uncontroversially adult specimens, bifurcation was evidently very different between these genera.)
So based on the vertebrae alone, the case of Haplocanthosaurus as an immature form of Diplodocus or Apatosaurus is blown right out of the water. And this is without even looking at the appendicular material — for example, the scapula and coracoid illustrated by Hatcher (1903:figs 17-19), which are so completely different from those of diplodocids.
But there’s more. Tune in next time for the rest.
The rest of the series
Links to all of the posts in this series:
- Part 1: what we knew a month ago
- Part 2: why serial position matters
- Part 3: the evidence from ontogenetic series
- Part 4: is Suuwassea a juvenile of a known diplodocid?
- Part 5: is Haplocanthosaurus a juvenile of a known diplodocid?
- Part 6: more reasons why Haplocanthosaurus is not a juvenile of a known diplodocid
and the post that started it all:
References
- Gillette, D.D. 1991. Seismosaurus halli, gen. et sp. nov., a new sauropod dinosaur from the Morrison Formation (Upper Jurassic/Lower Cretaceous) of New Mexico, USA. Journal of Vertebrate Paleontology 11(4):417-433.
- Gilmore, C.W. 1936. Osteology of Apatosaurus with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11:175-300.
- Hatcher, J.B. 1901. Diplodocus (Marsh): its osteology, taxonomy, and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63.
- Hatcher, J.B. 1903. Osteology of Haplocanthosaurus with description of a new species, and remarks on the probable habits of the Sauropoda and the age and origin of the Atlantosaurus beds; additional remarks on Diplodocus. Memoirs of the Carnegie Museum 2:1-75.
- Lovelace, D.M., Hartman, S.A., Wahl, W.R. 2008. Morphology of a specimen of Supersaurus (Dinosauria, Sauropoda) from the Morrison Formation of Wyoming, and a re-evaluation of diplodocid phylogeny. Arquivos do Museu Nacional, Rio de Janeiro 65(4):527-544.
- Lucas, S.G., Spielmann, J.A., Rinehart, L.F., Heckert, A.B., Herne, M.C., Hunt, A.P., Foster, J.R., Sullivan, R.M. 2006, Taxonomic status of Seismosaurus hallorum, a Late Jurassic sauropod dinosaur from New Mexico. New Mexico Museum of Natural History and Science Bulletin 36:149-162.
- Upchurch, P. Barrett, P.M., Dodson, P. 2004. Sauropoda. pp. 259-322 in D.B. Weishampel, P. Dodson and H. Osmólska (eds.), The Dinosauria, 2nd edition. University of California Press, Berkeley and Los Angeles. 861 pp.
- Wilson, J.A. 2002. Sauropod dinosaur phylogeny: critique and cladistic analysis. Zoological Journal of the Linnean Society 136:217-276.
- Woodruff, D.C, and Fowler, D.W. 2012. Ontogenetic influence on neural spine bifurcation in Diplodocoidea (Dinosauria: Sauropoda): a critical phylogenetic character. Journal of Morphology, online ahead of print.
Special bonus illustrations
I composited the cervical and dorsal series above into the following compound illustrations. As always, click through for full resolution.
Lateral view:
Posterior, mid and anterior dorsal vertebrae and cervical vertebrae, in right lateral view, of (top to bottom), Haplocanthosaurus, Apatosaurus louisae CM 3018 (from Gilmore 1936:plates XXIV and XXV, reversed for ease of comparison) and Diplodocus carnegii CM 84 (from Hatcher 1901:plates III and VII), scaled to roughly the same size. For the diplodocids, we illustrate D9, D5, D2, C13, C9 and C4. For Haplocanthosaurus, which has four more dorsals, we illustrate D13, D7 and C14 of H. priscus (from Hatcher 1903:plate I) and D2, C9 and C4 of H. utterbacki (from plate II).
Posterior view:
Posterior, mid and anterior dorsal vertebrae and posterior cervical vertebrae C15 and C14, in posterior view, of (top to bottom), Haplocanthosaurus priscus CM 572 (From Hatcher 1903:plate I), Apatosaurus louisae CM 3018 (from Gilmore 1936:plates XXIV and XXV) and Diplodocus carnegii CM 84 (from Hatcher 1901:plates III and VII), scaled to the same height of the mid dorsal. For the diplodocids, we illustrate D9, D5 and D1. For Haplocanthosaurus, which has four more dorsals, we illustrate D13, D6 and D1.
