As part of a major spring cleaning operation that we started the first week of January, this week I opened the last two boxes left over from when we moved into our current house. One of them had a bundle of framed art. I knew most of what was in there before I opened the box, but I had somehow completely forgotten about this. I must have gotten it framed in late 90s, and it hung on the walls of our apartments in Norman and Santa Cruz. At some point it went into a box, and I forgot it even existed.

This is the first technical drawing I ever attempted of OMNH 53062, which would later become the holotype of Sauroposeidon. I drew it for my poster at the 1997 SVP meeting in Chicago, and it went on to become Figure 5 in my undergraduate thesis (which is preserved for posterity here). I’d do other, better drawings of the specimen in later years, but this one came first.

I know I’m biased, but that second vertebra in the preserved series, which I interpreted as C6 back when, will probably always be the most gorgeous natural object on the planet in my book. I don’t expect anyone else to feel the same. I worked on that specimen for three years – some of it seeped into my soul, and vice versa. Then again, I don’t care how jaded you are about long vertebrae, that one is still a pretty arresting sight.

For a much more recent take on the appearance of the Sauroposeidon vertebrae, see this post.

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Cervical rib cross-sections from Mamenchisaurus Giraffatitan and Diplodocus Klein et al 2012 fig 1

Klein et al. (2012: fig. 1)

We have good descriptions of the proximal parts of the cervical ribs for lots of sauropods. We also have histological cross-sections of a few, mostly thanks to the work of Nicole Klein and colleagues (Klein et al. 2012, Preuschoft and Klein 2013), although histological cross-sections of ribs were also figured as long ago as 1999, by Dalla Vecchia (1999: figs. 29 and 30), and as recently as this month, by Lacovara et al. (2014: supplementary figure 4).

What we have very, very few of is series of cross-sections that show how the cr0ss-section of a cervical rib changes along its length. There may be more out there (and if I have forgotten any, please remind me!), but at the moment I can only think of three such figures: two in Janensch (1950: figs. 83 and 85), both on Giraffatitan, and one in Klein et al. (2012: fig. 1), with cross-sections from Mamenchisaurus, Giraffatitan, and Diplodocus (shown at the top of the post).

Sauroposeidon cervical rib cross-sections v3

 

Rarer still are images that show cross-sections of overlapped cervical ribs, stacked in situ. You could use the information in Janensch (1950: figs. 83 and 85) to generate the stacked cross-sections, but you wouldn’t know the spacing between the ribs as they were in the ground. I think the image just above, of the cervical rib bundles in the Sauroposeidon holotype, OMNH 53062, may be the first of its kind–again, if you know of any others, please let me know. I took the notes for this figure back in 2004, sitting down with the holotype and some digital calipers to make sure I could scale everything correctly, I just hadn’t ever put it into a presentable form until now. The first C6 section (blue V-shape) is from right at the root where the capitulum and tuberculum meet and the posterior shaft of the rib begins.

It is by now well-understood that the long cervical ribs of sauropods and other dinosaurs are ossified tendons of the long hypaxial neck muscles, specifically the longus colli ventralis and flexor colli lateralis. We argued this back in 200o on comparative anatomical grounds (Wedel et al. 2000b: pp. 378-379), and it has now been demonstrated histologically (Klein et al. 2012, Lacovara et al. 2014). The system of stacked tendons is also found in most birds. Here’s the bundle of stacked tendons in a rhea neck, only slightly fanned out:

Rhea ventral tendons stacked - full

And the same neck, with both the epaxial and hypaxial muscles more fully separated:

Rhea neck muscles fanned - full

What I’d really like is an MRI of a rhea or ostrich neck, showing the stacked tendons and their associated belts of muscle, to compare with the stacked cervical ribs of Sauroposeidon and other sauropods. Anyone know of any?

Incidentally, I think the cervical ribs and cervical rib bundles of sauropods are one line of evidence for sauropod necks having been rather slenderly-muscled. The long, multi-segment muscles like the longus colli ventralis are the outermost components of the muscular envelope that surrounds the vertebrae, as you can see in the rhea dissection photos. In sauropod specimens with articulated cervical ribs, the ribs do not deviate from one another or fan out. Rather, they lie in vertically stacked bundles that run from one capitulum-tuberculum intersection to the next. So the depth of that intersection–the “root” of the cervical rib of any given vertebra–plus the thickness of the ribs stacked underneath it, is pretty much the thickness of the muscular envelope around the neck, or at least around the ventral half. And the cervical ribs are typically pretty close to the vertebral centra–only weirdos like Apatosaurus and Erketu displace them very far ventrally (see Taylor and Wedel 2013a: fig. 7 and this post). So, thin jackets of muscle around proportionally large vertebrae–or, if you like, corn-on-the-cob rather than shish-kebabs.

As for why sauropods have long cervical ribs, Mike and I discussed some possibilities in our 2013 PeerJ paper (Taylor and Wedel 2013a), and Preuschoft and Klein addressed the issue last fall in PLOS ONE (Preuschoft and Klein 2013). My favorite hypothesis is that long tendons allow an animal to shift the bulk of the muscle–and therefore the center of gravity–toward the base of the neck, but that long unossified tendons can be distorted through stretching, which wastes muscular energy. Ossifying those long tendons is like putting bony wheelbarrow handles on each vertebra, allowing the muscles to move the vertebra from a distance without so much wasted energy, and probably with finer positional control.

That’s a nifty hypothesis in need of testing, anyway. In fact, cervical ribs and their associated muscles could stand a lot more attention on both the descriptive and analytical fronts. I know that Liguo Li has some research in the works on different conformations of hypaxial muscles, tendons, and cervical ribs in birds (you know, when she’s not describing bizarre new titanosaurs like Yongjinglong — see Li et al. 2014). If you saw Peter Dodson give their talk at SVP last fall, you probably remember some stunning images of dissected bird necks. As a famous legislator once said, we shall watch her career with great interest.

References

 

My camera had a possibly-fatal accident in the field at the end of the day on Saturday, so I didn’t take any photos on Sunday or Monday. From here on out, you’re either getting my slides, or photos taken by other people.

Powell Museum sauropod humerus

On Sunday we were at the John Wesley Powell River History Museum in Green River, Utah, for the Cretaceous talks. There were some fossils on display downstairs, including mounted skeletons of Falcarius and one or two ornithischians,* and this sauropod humerus from the Cedar Mountain Formation (many thanks to Marc Jones for the photo).

* A ceratopsian and Animantarx, maybe? They were in the same room as the sauropod humerus, so it’s no surprise that I passed them by with barely a glance.

There were loads of great talks in the Cretaceous symposium on Sunday, and I learned a lot, about everything from clam shrimp biostratigraphy to ankylosaur phylogeny to Canadian sauropod trackways. But I can’t show you any slides from those talks, so the rest of this post is the abstact from Darren’s and my talk, illustrated by a few select slides.

Wedel Naish 2014 Sauroposeidon and kin - slide 1 title

Sauroposeidon is a giant titanosauriform from the Early Cretaceous of North America. The holotype is OMNH 53062, a series of four articulated cervical vertebrae from the Antlers Formation (Aptian-Albian) of Oklahoma. According to recent analyses, Paluxysaurus from the Twin Mountain Formation of Texas is the sister taxon of OMNH 53062 and may be a junior synonym of Sauroposeidon. Titanosauriform material from the Cloverly Formation of Wyoming may also pertain to Paluxysaurus/Sauroposeidon. The proposed synonymy is based on referred material of both taxa, however, so it is not as secure as it might be.

Wedel Naish 2014 Sauroposeidon and kin - slide 34 Sauroposeidon characters

Top row, vertebrae of Paluxysaurus. From left to right, the centrum lengths of the vertebrae are 72cm, 65cm, and 45cm. Main image, the largest and most complete vertebra of the holotype of Sauroposeidon. Labels call out features that are present in every Sauroposeidon vertebra where they can be assessed, but consistently absent in Paluxysaurus. Evaluating the proposed synonymy of Paluxysaurus and Sauroposeidon is left as an exercise for the reader.

MIWG.7306 is a cervical vertebra of a large titanosauriform from the Wessex Formation (Barremian) of the Isle of Wight. The specimen shares several derived characters with the holotype of Sauroposeidon: an elongate cervical centrum, expanded lateral pneumatic fossae, and large, plate-like posterior centroparapophyseal laminae. In all of these characters, the morphology of MIWG.7306 is intermediate between Brachiosaurus and Giraffatitan on one hand, and Sauroposeidon on the other. MIWG.7306 also shares several previously unreported features of its internal morphology with Sauroposeidon: reduced lateral chambers (“pleurocoels”), camellate internal structure, ‘inflated’ laminae filled with pneumatic chambers rather than solid bone, and a high Air Space Proportion (ASP). ASPs for Sauroposeidon, MIWG.7306, and other isolated vertebrae from the Wessex Formation are all between 0.74 and 0.89, meaning that air spaces occupied 74-89% of the volume of the vertebrae in life. The vertebrae of these animals were therefore lighter than those of brachiosaurids (ASPs between 0.65 and 0.75) and other sauropods (average ASPs less than 0.65).

Wedel Naish 2014 Sauroposeidon and kin - slide 64 Mannion phylogeny

Check this out: according to at least some versions of the Mannion et al. (2013) tree, Sauroposeidon and Paluxysaurus are part of a global radiation of andesaurids in the Early and middle Cretaceous. Cool!

Sauroposeidon and MIWG.7306 were originally referred to Brachiosauridae. However, most recent phylogenetic analyses find Sauroposeidon to be a basal somphospondyl, whether Paluxysaurus and the Cloverly material are included or not. Given the large number of characters it shares with Sauroposeidon, MIWG.7306 is probably a basal somphospondyl as well. But genuine brachiosaurids also persisted and possibly even radiated in the Early Cretaceous of North America; these include Abydosaurus, Cedarosaurus, Venenosaurus, and possibly an as-yet-undescribed Cloverly form. The vertebrae of Abydosaurus have conservative proportions and solid laminae and the bony floor of the centrum is relatively thick. In these characters, Abydosaurus is more similar to Brachiosaurus and Giraffatitan than to Sauroposeidon or MIWG.7306. So not all Early Cretaceous titanosauriforms were alike, and whatever selective pressures led Sauroposeidon and MIWG.7306 to evolve longer and lighter necks, they didn’t prevent Giraffatitan-like brachiosaurs such as Abydosaurus and Cedarosaurus from persisting well into the Cretaceous.

Wedel Naish 2014 Sauroposeidon and kin - slide 65 Cloverly sauropods

The evolutionary dynamics of sauropods in the North American mid-Mesozoic are still mysterious. In the Morrison Formation, sauropods as a whole are both diverse and abundant, but Camarasaurus and an efflorescence of diplodocoids account for most of that abundance and diversity, and titanosauriforms, represented by Brachiosaurus, are comparatively scarce. During the Early Cretaceous, North American titanosauriforms seem to have radiated, possibly to fill some of the ecospace vacated by the regional extinction of basal macronarians (Camarasaurus) and diplodocoids. However, despite a flood of new discoveries in the past two decades, sauropods still do not seem to have been particularly abundant in the Early Cretaceous of North America, in contrast to sauropod-dominated faunas of the Morrison and of other continents during the Early Cretaceous.

Wedel Naish 2014 Sauroposeidon and kin - slide 66 acknowledgments

That final slide deserves some explanation. On the way back from the field on Saturday–the night before my talk–a group of us stopped at a burger joint in Hanksville. Sharon McMullen got a kid’s meal, and it came in this bag. We took it as a good omen that Sauroposeidon was the first dinosaur listed in the quiz.

For the full program and abstracts from both days of talks, please download the field conference guidebook here.

Order up!

Sauroposeidon OMNH 53062 articulated right lateral composite with giraffe

Sauroposeidon is stitched together from orthographic views of the 3D photogrammetric models rendered in MeshLab. Greyed out bits of the vertebrae are actually missing–I used C8 to patch C7, C7 to patch C6, and so on forward. The cervical ribs as reconstructed here were all recovered and they are in collections, but they’re in several jackets and boxes and therefore not easily photographed.

The meter bars are both one meter as advertised. The giraffe neck is FMNH 34426 (from this post), which is actually 1.7 meters long, but I scaled it up to 2.4 meters to match that of the tallest known giraffe. I think it’s cool that a world-record giraffe neck is roughly as long as two vertebrae from the middle of the neck of Sauroposeidon.

There are loads of little morphological details in the Sauroposeidon vertebrae that are clearer now than they were in our old photographs, but those will be stories for other posts.

Sauroposeidon in 3D

April 18, 2014

Sauroposeidon meet Sauroposeidon

I was in Oklahoma and Texas last week, seeing Sauroposeidon, Paluxysaurus, Astrophocaudia, and Alamosaurus, at the Sam Noble Oklahoma Museum of Natural History, the Fort Worth Museum of Science and History, the Shuler Museum of Paleontology at SMU, and the Perot Museum of Nature and Science, respectively. I have a ton of interesting things from that trip that I could blog about, but unfortunately I have no time. Ten days from now, I’m off to Colorado and Utah for the Mid-Mesozoic conference and field trip, and between now and then I need to finish up my bits on three collaborative papers, get my summer anatomy lectures posted for internal peer review here at WesternU, and–oh yeah–actually write my conference talk. Fun times.

BUT after being subjected to the horror of the Yale Brontosaurus skull, I figured you all deserved a little awesome.

Photographing Sauroposeidon 2014-04-07

So here’s me getting one of 351 photos of the most posterior and largest of the Sauroposeidon jackets (this is not the awesome, BTW, just a stop along the way). This jacket holds what I once inferred to be the back half of C7 and all of C8. Now that Sauroposeidon may be a somphospondyl rather than a brachiosaur, who knows what verts these are–basal somphospondyls have up to 17 cervicals to brachiosaurids’ probable 13 (for a hypothetical view of an even-longer-necked Sauroposeidon, see this probably-prophetic post by Mike). The vertically-mounted skeleton in the background is Cotylorhynchus. Cotylorhynchus got a lot bigger than that–up to maybe 6 meters long and 2 or 3 tons–and was probably the largest land animal that had ever existed back in the Early Permian. Photo by OU grad student Andrew Thomas, whom you’ll be hearing about more here in the future.

I couldn’t crank the model myself on the road, thanks to the pathetic lack of processing power in my 6-year-old laptop (which will be replaced RSN). Andy Farke volunteered to do the photogrammetricizing with Agisoft Photoscan, if only I’d DropBox him the pictures. Here’s a screenshot from MeshLab showing the result:

Sauroposeidon lateral PLY 10 - 6 and 9 blended

And my best taken-from-overhead quasi-lateral photograph:

Sauroposeidon C8 jacket lateral photo 2014-04-07

If you’re curious, the meter stick at the top is actually one meter long, it just has the English measurement side showing. The giant caliper at the bottom is also marked off in inches, and it is open to 36.0 inches (it didn’t go to 1 meter, or I would have used that). You can tell that there is some perspective distortion involved here since 36 inches on the caliper is 1380 pixels, whereas the 39.4-inch meter stick is only 1341 pixels. Man, I hate scale bars. But they make good calibration targets.

Incidentally, after playing around with the model in orthographic mode in MeshLab, the distortions in the photos of the vertebrae themselves just scream at me. Finally, finally, I can escape the tyranny of perspective. Compare the ends of the big wooden beam at the top of the jacket to get a feel for how much the two views differ.

Working on Sauroposeidon again after all this time made me seriously nostalgic. I love that beast. I don’t think I’m exaggerating when I say that those vertebrae are the most gorgeous physical objects in the universe. Also, an appropriately huge thank-you to preparator Kyle Davies (of apatosaur-sculpting fame), collections manager Jen Larsen, and Andrew Thomas again for help with wrassling those verts around, and for sharing their thoughts and advice. Thanks also to curators Rich Cifelli and Nick Czaplewski for their hospitality and for the go-ahead to undertake this work, and to Andy Farke for generating the model.

I’ll have a lot more to say about this stuff in the future. I didn’t go to all this work just for giggles. For a long time I’ve had a hankering to do a paper on the detailed anatomy of Sauroposeidon, based on all of the things that I’ve noticed in the last decade that didn’t make it into any of the early papers. And now there’s the proposed synonymy of Paluxysaurus with Sauroposeidon. And “Angloposeidon” needs some attention–Darren and I have been thinking about writing “Angloposeidon II” for years now. And…well, plenty more.

So, loads more to come, but not for the next few weeks. Eventually I’ll be publishing all of this–the photos, the 3D models, the whole works. Stay tuned.

UPDATE a few days later

Man, I am frazzled, because I forgot to include the moral of the story: if I can do this, you can do this. There are good, free photogrammetry programs out there–Peter Falkingham published a  whole paper on free photogrammetry in 2012, and posted a guide to an even better program, VisualSFM, on Academia.edu. Even Agisoft Photoscan is not prohibitively expensive–under $200 for an educational license. MeshLab is free and has hordes of good free tutorials. For the photography itself, you basically just build a virtual dome of photos around an object. If you need more instructions than that, Heinrich has written a whole series of tutorials. It doesn’t take a fancy camera–I used a point-and-shoot for the Sauroposeidon work shown here (a Canon S100 operating at 6 megapixels, if anyone is curious). What are you waiting for?

Sauroposeidon and friends

February 24, 2014

Sauroposeidon and kin cervicals - DRAFTAs a break from photography posts, here are four pretty big vertebrae that swirl in the same thought-space in my head. All are shown to scale, in right lateral view. These are not the biggest sauropod cervical vertebrae–Supersaurus beats them all, and there are vertebrae of Puertasaurus, Alamosaurus, and Futalognkosaurus that rival the big Sauroposeidon vert, but those are either less well preserved or still awaiting detailed description.

Incidentally, I think BYU 12867 is a C10. The centrum proportions are about right, compared to Giraffatitan, and the neural spine looks good, too, like a geometric transformation of the big Giraffatitan C8. Also, the drawn-in prezyg outline for MIWG.7306 is a little short; the actual prezyg is a monster and would have overhung the condyle by another 10cm or so. I’m pretty sure that we had a composite photograph showing this at one point, but irritatingly none of us can find it at the moment. If it turns up, I’ll update the image.

For a long time I thought Sauroposeidon was a brachiosaurid. Now it seems to be a somphospondyl (D’Emic 2012) or possibly even a basal titanosaur (Mannion et al. 2013), even if we stick just to the holotype. But if it’s not a brachiosaurid, it’s cervical vertebrae are at least coarsely brachiosaur-y in outline.

You  may recall from Naish et al. (2004) that MIWG.7306 shares several derived characters with the holotype vertebrae of Sauroposeidon. Does that mean that Angloposeidon is a somphospondyl or titanosaur as well? I dunno–as always, we need more material–but it’s an interesting possibility.

References

Following on from Matt’s post about the difficulty of photographing big specimens without distortion, I thought I’d have a play with our best Sauroposeidon C8 photo, which I think is this one:

sauroposeidon-c8-alone

(That’s been the basis for classic SV-POW! posts such as Your neck is pathetic and Darren’s new indeterminate Wealden maniraptoran is inadequate.)

I was motivated by Andy Farke’s comment:

Another–and perhaps more important–area where surface models excel is when you can remove colors on the original specimen that wash out relevant details…I bet this is probably the case for the example vertebra of Sauroposeidon. How many fossae and foramina just don’t show up well on the photos above?

Andy was talking about completely colourless 3d surface models, in which the 3d shape allows a render to make shadows that bring out the subtle shapes. But it made me wonder whether we could get anywhere just by washing out the most prevalent colour in the photo.

I started by doing a big, fat Gaussian blur on a duplicate layer — 500 pixels in each direction — and sampling the colour in the middle, to get a rough-and-ready average. (There may be a better way — please shout if you know one.) That average colour was#7e6b2f. I used it to run Colour To Alpha on another duplicate of the original layer, so that we’d be left with only residual colours. Here’s the result:

sauroposeidon-c8-alone-colour-completely-removed

I’m in two minds about this. It may be informative, but it sure is ugly. To compromise, I reinstated the original layer underneath this mostly-transparent one, and turned its opacity down to 75%. Here’s the result — a nice compromise:

sauroposeidon-c8-alone-colour-removed

Of course, there are endless other approaches you can take — that’s the blessing and the curse of image-editing programs like GIMP. For example, here’s what I got doing a simple Colours → Auto → White Balance:

sauroposeidon-c8-alone-whitebalanced

I’m not sure that isn’t the best of the bunch, in terms of informativeness.

I also tried something else — not amazingly successfully, but I think it’s worth seeing. Since the two photos that Matt showed in the previous post were evidently taken from somewhat different angles, I thought I’d have a go at compositing them into a red-cyan anaglyph. Because the variation in camera position is mostly dorsoventral rather than anteroposterior, the vert has to be pointed upwards for the two eyes to see the two versions from different horizontal points. Here’s the best I could do:

c8-anaglyph

I would say this is of some value; but it’s nowhere near as good as, for example, the anaglyph of Cervical S of the Archbishop. I could sit and look at that one all day. The problems with this one arise for three reasons.

First, I had to reduce both parts of the Sauroposeidon anaglyph to monochrome (since one was already in that form), so all colour information was lost.

Second, I had to scale the high-resolution picture to the same size as the lower-resolution one, throwing away more detail.

Finally, and most important, the two photos were not taken with the intention that they should be used to make an anaglyph. To work well, this has to be done with the images taken under the same lighting conditions, at the same distance from the specimen, from perspectives differing by about the distance between the pupils of the viewer, and with the camera-position difference being perfectly in the plane of the specimen. Needless to say, none of these conditions was met in this case, so it’s actually quite impressive that it works as well as it does.

We have a lot of options for illustrating specimens these days. Postage-stamp-sized greyscale photos really don’t cut it any more.

Here are two photos of what I infer to be C8 of OMNH 53062, the holotype of Sauroposeidon. The top one was taken by Mike during our visit to the OMNH in 2007. If you’re a regular you may recognize it from several older posts: 1, 2, 3. The bottom one was taken by Mike Callaghan, the former museum photographer at the OMNH, sometime in 1999 or 2000. I used it in Wedel et al. (2000) and Wedel and Cifelli (2005).

Sauroposeidon OMNH 53062 C8 photos compared

You’ll notice that the two photos are far from identical. In both cases, the photographers were up on ladders, as far above the vertebra as they could get, and there are still significant perspective effects. That’s just a fact of life when you’re taking photos of a vertebra that is 1.4 meters long, from anything lower than a helicopter. In Mike Taylor’s shot, the neural spine looms a little too large; in Mike Callaghan’s shot, the prezygapophysis looks a little too small, probably because it was curving off at the edge of the shot. So neither photograph is “right”; both distort the morphology of the specimen in different ways. Here’s how the two images stack up, with the outlines scaled to the same length:

Sauroposeidon OMNH 53062 C8 outlines compared

When I ran a draft of this post past Mike, he wrote (with permission to post):

I think the current draft misses an important point: the warning. We really can’t trust photos, however carefully taken, and however beautifully composited into TNFs*. You’re welcome to quote me as having said I’d have assumed the two C8s were different vertebrae. For that matter, I bet I could have worked up several taxonomically significant characters to distinguish them. Yikes.

* TNF = Taylor Normal Form, i.e., making multi-view photos like the ones here and here.

So the moral is, photos of big specimens almost always involve some distortion. This is clearly not ideal. But I have a plan for fixing it. I am hoping to get back to the OMNH this spring, and the next time I’m there, I’m going to take photos of this vertebra from a zillion angles and make a 3D model through photogrammetry. Happily, Heinrich Mallison has been producing a very helpful series of tutorials on that very topic over at dinosaurpaleo: 1, 2, 3, 4, with more on the way (I’ll update the links here later). Update: Don’t forget to check out Peter Falkingham’s (2012) paper in PE on making photogrammetric models with free software.

Armed with that model, it should be possible to produce a perspective-free lateral view image of the vertebra, to which all of the previous photos can be compared. I can’t use CT data because this vertebra has never been CTed; it’s too big to fit through a medical CT scanner, and probably too fragile to be packed up and shipped to an industrial CT machine like they used on Sue (not to mention that would require a significant chunk of money, which is probably not worth spending on a problem that can be solved in other ways).

So, photogrammetry to the rescue, or am I just deluding myself? Let me know what you think in the comments.

Finally, I should mention that the idea of superseding photographs with 3D photogrammetric models is not original. I got religion last week while I was having beers with Martin Sander and he was showing me some of the models he’s made. He said that going forward, he was going to forbid his students to illustrate their specimens only with photographs; as far as he was concerned, now that 3D models could be cheaply and easily produced by just about everyone, they should be the new standard. Inspiring stuff–now I must go do likewise.

Some previous posts on Sauroposeidon:

References

Sauroposeidon OMNH 53062 C7-C8 left side

Sauroposeidon holotype OMNH 53062, posterior half of ?C7 and all of ?C8 in left lateral view. Scale bar is in inches.

There’s a lot more Sauroposeidon material these days than there used to be, thanks to the referral by D’Emic and Foreman (2012) of Paluxysaurus and Ostrom’s Cloverly material and the new Cloverly material to my favorite sauropod genus. I’ve seen almost all of this material firsthand, but obviously the specimen I’m most familiar with is the holotype, OMNH 53062. It was the primary thing occupying my mind from the summer of 1996 through the spring of 2000, and it has remained a frequent object of wonder ever since.

The specimen was found lying on its right side in the field, so that side is in better shape, by virtue of having been more deeply buried and thus protected from the ravages of freezing and thawing and other erosional processes. When the jackets were taken out of the ground and prepared, the not-so-well-preserved left sides were prepped first. Then permanent support jackets were made on the left sides, the vertebrae were flipped onto their left sides, the field jackets were removed from their right sides, and the vertebrae were prepped on the right. They’ve been lying in their support jackets, left side down and right side up, ever since. (For more on the taphonomy and recovery of the specimen, see this post and Wedel and Cifelli 2005 [free PDF linked below].)

Now, if I had known what I was doing, I would have photographed the crap out of the left sides before the verts were flipped. But it was my first project and I was learning on the job, and that didn’t occur to me until later.

It also didn’t occur to me that, once flipped, the left sides would be effectively out of reach forever. But the vertebrae are extremely fragile. The bigger verts have cracks running through them, and the jackets flexed noticeably when we took them for CT scanning. I am worried that if we tried to flip the bigger verts today, they might just crumble. Even the surface bone is fragile. I remember once trying to get some dust off one of the verts with a vacuum cleaner hose, and watching in horror as some of the millimeter-thin external bone just flaked off and flew away. That was in the late 1990s, when the verts were still stored in the dusty, drafty WWII-era buildings that had housed the museum collections for ages. Now they’re in what I still think of as the “new” building, which opened in 2000, in a really nice modern collection room with climate and dust control, and I’ve never seen them with any noticeable dust.

Anyway, the left sides are now obscured by their supporting jackets and will remain that way for the foreseeable future. And I don’t have a complete set of photos of the left sides of the verts. But I do have one, of the back half of ?C7 and all of ?C8, and a scan of it appears at the top of this post. It’s a scan of a physical photograph because it was taken in late 1996 or early 1997–no-one I knew had a digital camera, and if you wanted a digital version of a photograph, you shot it on a film camera, had a big print made, and scanned that on a flatbed scanner.

Here’s another version with the vertebrae outlined:

Sauroposeidon OMNH 53062 C7-C8 left side - outlined

When I and everyone else thought that Sauroposeidon was a brachiosaur, I was pretty sure that these were C7 and C8, out of a total of 13 cervicals, just like Giraffatitan. And it still might be so–a future analysis might find that the newly-expanded Sauroposeidon is a brachiosaurid after all, and even if not, Gomani (2005) posited a primitive cervical count of 13 for titanosaurs. If that’s true, then possibly 13 cervicals are primitive for all titanosauriforms, and the increases beyond that–to 17 in Euhelopus and 14-17 in more derived titanosaurs like Futalognkosaurus and Rapetosaurus–were deviations from that primitive pattern.

But.

If Sauroposeidon was a basal somphospondyl, as posited by D’Emic and Foreman (2012) and as found in the phylogenetic analysis of D’Emic (2012), then maybe it was more like Euhelopus than Giraffatitan, and maybe it had more than 13 cervicals. (Note that D’Emic [2012] found Sauroposeidon to be a basal somphospondyl but outside the Euhelopodidae, so even in his analysis, Euhelopus could have gotten its extra cervicals independently of Sauroposeidon.) That’s an interesting prospect, since the 11.5-meter neck estimate for Sauroposeidon I made back in 2000 was based on the conservative assumption of 13 cervicals. If Sauroposeidon had more cervicals, they were probably mid-cervicals (nobody adds more dinky C3s, or stubby cervico-dorsals*–that would be silly), and therefore between 1 and 1.25 meters long. So if the individual represented by OMNH 53062 had 15 cervicals, as Mike hypothetically illustrated in this post, its neck might was probably more like 14 meters long, and if it had 17 cervicals, like Euhelopus and Rapetosaurus, its neck might have topped 16 meters–as long or longer than that of Supersaurus.

Now, I’m not saying that Sauroposeidon had a 16-meter neck. The conservative estimate is still 13 cervicals adding up to 11.5 meters. But the possibility of a longer neck is tantalizing, and can’t be ruled out based on current evidence. As usual, we need more fossils.

Happily, now that Sauroposeidon is known from Oklahoma, Texas, and Wyoming, and is one of the best-represented EKNApods instead of one of the scrappiest, the chances that we’ll find more of it–and recognize it–are looking good. I will keep my fingers firmly crossed–as they have been for the last 17 years.

* Radical pedantry note: of course we have very good evidence of sauropods getting more cervical vertebrae by recruiting dorsals into the cervical series. So, for example, 13 cervicals and 12 dorsals are supposed to be primitive for neosauropods, but diplodocids have 15 and 10, respectively–the obvious inference being that the first two dorsals got cervicalized. So in this narrow meristic sense, sauropods definitely did add cervicodorsals. But my point above is about the morphology of the verts themselves–once diplodocids had those two extra cervicals at the end, the former cervicodorsals were free to become more “cervicalized” in form. So effectively–in terms of the shapes of their necks–diplodocids added mid-cervicals.

References

YPM 5449, a posterior dorsal vertebra of Sauroposeidon, from D’Emic and Foreman (2012:fig. 6A and C).

Another recent paper (part 1 is here) with big implications for my line of work: D’Emic and Foreman (2012), “The beginning of the sauropod dinosaur hiatus in North America: insights from the Lower Cretaceous Cloverly Formation of Wyoming.” In it, the authors sink Paluxysaurus into Sauroposeidon and refer a bunch of Cloverly material to Sauroposeidon as well. So in one fell swoop Sauroposeidon goes from being one of the most poorly represented Early Cretaceous North American sauropods, based on just four vertebrae from a single individual, to one of the best-known, most complete, and most widespread, based on at least seven individuals from Texas, Oklahoma, and Wyoming.

The web of connections among the different sets of material is complex, and involves the Sauroposeidon holotype OMNH 53062 from the Antlers Formation of southeastern Oklahoma, the type and referred material of Paluxysaurus from the Twin Mountains Formation of northern Texas described by Rose (2007), sauropod material from the Cloverly Formation of north-central Wyoming described and illustrated by Ostrom (1970), and UM 20800, a scap and coracoid newly excavated from one of Ostrom’s old quarries.  D’Emic and Foreman argue that (1) the Cloverly material is referable to Sauroposeidon based on the shared derived characters of a juvenile cervical, YPM 5294, and the Sauroposeidon holotype, and (2) Paluxysaurus is not distinguishable from the Cloverly material and in fact shares several autapomorphies with the Cloverly sauropod. Which means that (3) Paluxysaurus is Sauroposeidon.

But that’s not all! All the new material suggests different phylogenetic affinities for Sauroposeidon. Instead of a brachiosaurid, it is now posited to be a basal somphospondyl. That’s not super-surprising; as we noted back in 2000 (Wedel et al. 2000), if Sauroposeidon was a brachiosaurid it had evolved some features in parallel with titanosaurs, most notably the fully camellate internal structure of the cervical vertebrae. And it also makes sense because other basal somphospondyls include Erketu and Qiaowanlong, the cervicals of which are similar to Sauroposeidon in some features. D’Emic and Foreman (2012) cite a forthcoming paper by Mike D’Emic in the Journal of Systematic Paleontology that contains the cladistic analysis backing all this up, but the case based on comparative anatomy is already pretty strong.

If anyone is unconvinced by all of these referrals, please bear in mind that we haven’t heard the whole story yet, quite probably for reasons that are outside of the authors’ control.  I am inclined to be patient because I have been in that situation myself: Wedel (2003a) was intended to stand on the foundation of evidence laid down by Wedel (2003b), but because of the vagaries of publication schedules at two different journals, the interpretive paper beat the descriptive one into press by a couple of months.

Mid-cervical originally described as Paluxysaurus, now referred to Sauroposeidon, from Rose (2007:fig. 10).

Anyway, if anyone wants my opinion as “Mr. Sauroposeidon“, I think the work of D’Emic and Foreman (2012) is solid and the hypothesis that Paluxysaurus is Sauroposeidon is reasonable. So, if I think it’s reasonable now, why didn’t I synonymize the two myself? Partly because I thought there was a pretty good chance the two were not the same, based mostly on FWMSH 93B-10-8 (which I referred to as FWMSH “A” in Wedel 2003b, since I had only seen in on display without a specimen number), which I thought looked a lot more like a titanosaur cervical than a brachiosaur cervical. But of course I thought Sauroposeidon was a brachiosaur until a couple of months ago, and if it ain’t, and if brachiosaurs and basal somphospondyls have similar cervicals, that objection is considerably diminished. And partly because I’ve had other things to be getting on with, and stopping everything else to spend what would realistically be a few months looking into a possible synonymy (that I didn’t strongly suspect) wasn’t feasible in terms of time or geography. So I’m glad that D’Emic and Foreman have done that work, and I’m excited about the new things they’ve uncovered.

And I’m honored to bring you a new life restoration of Sauroposeidon by uber-talented Bob Nicholls, which we think is the first to show Sauroposeidon in its new guise as a basal somphospondyl. Click through for the mega-awesome version.

Same critter, different views. If anyone wants to GDI this baby, you now have everything you need. Many thanks to Bob for permission to post these and the following making-of images. Please visit him at Paleocreations.com to see a ton of awesome stuff, and give him some love–or at least a few thousand “likes”–on Facebook.

This is Bob’s first foray into 3D modeling, but you’d never know from the quality of his virtual sculpt. And let me tell you, that dude works fast. He sent this initial version, showing Sauroposeidon as an attenuated brachiosaur (sorta like this) on August 23, to solicit comments from Mike and me.

I wrote back and let Bob know about the new work of D’Emic and Foreman, and suggested that he could probably be the first to restore Sauroposeidon as a somphospondyl. Mike and I also voiced our opposition to the starvation-thinned neck, and Mike suggested that the forelimb was too lightly muscled and that the ‘fingers’ were probably too prominent. The very next day, this was in our inboxes:

I wrote back:

Whatever Sauroposeidon was, its neck was fairly tall and skinny in cross-section. It looks like the neck on your model sort of tapers smoothly from the front of the body to the head. I think it would be much narrower, side-to-side, along most of its length, and would have a more pronounced shoulder-step where it met the body.
The bottom view is very useful. It shows the forefeet as being about the same size as the hindfeet. AFAIK all or nearly all known sauropod tracks have much bigger hindfeet than forefeet. Certainly that is the case with Brontopodus birdi, the big Early Cretaceous sauropod tracks from Texas that were probably made by Sauroposeidon. The forefeet should be about 75-80% the width of the hindfeet, and only about half a long front-to-back. Even if you don’t quite get to those numbers, shrinking the forefeet a bit and subtly up-sizing the hindfeet would make the model more accurate.
Mike’s commentary was much shorter–and funnier:
I like how freaky it looks. It looks WRONG, but in a good way.
Bob toiled over the weekend and came back with this subtly different, subtly better version:

I had one more change to recommend:

I’m sorry I didn’t suggest this sooner, but it only just now occurred to me. With the referral of Paluxysaurus and the Cloverly material to Sauroposeidon, we now have dorsal vertebrae, and they are loooong, much more similar in proportion to the dorsals of Brachiosaurus altithorax than those of Giraffatitan brancai. So, as much as I like the compact little body on your Sauroposeidon, I think it was probably fairly long in the torso. You probably already have Mike’s Brachiosaurus paper [Taylor 2009] with the skeletal recon showing the long torso–in the absence of an updated skeletal recon for Sauroposeidon, I’d use Mike’s Figure 7 as a guide for reconstructing the general body proportions.

Bob lengthened the torso to produce the final version, which is the first one I showed above. He sent that over on August 29–the delay in getting this post up rests entirely with me.

So. It is still very weird to think of “my” dinosaur as a somphospondyl rather than a brachiosaur. I had 15 years to get used to the latter idea. But suddenly having a lot more material–essentially the whole skeleton, minus some stinkin’ skull bits–is pretty darned exciting, and the badass new life restoration doesn’t hurt, either.

Now, would it be too much to wish for some more Brontomerus?

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