Last Wednesday, May 9, Brian Engh and I bombed out to Utah for a few days of paleo adventures. Here are some highlights from our trip.

We started at a Triassic tracksite on Thursday. But I’m not going to post any pictures of the tracks – those will be coming to a Brian Engh joint near you in the future. Instead, I’m going to talk about this little male collared lizard whose territory included the tracksite. He was fearless – didn’t want to run off and leave us yahoos wandering around his patch of desert unsupervised. Brian tickled his chin at one point.

Getting this close to him is how I got shots like this one:

Click through to the big version, it’s worth it.

One more shot of a couple of cool desert dwellers. I was so fixated on the lizard that I didn’t realize until later that Brian was in the frame, taking a much-needed hydration break.

On Friday we had a temporary breaking of the fellowship. I went to Fruita, Colorado, to visit the Dinosaur Journey museum. You’ve seen photos from DJ here before, from the 2014 Mid-Mesozoic Field Conference and the 2016 Sauropocalypse. Here’s an apatosaur pubis with some obvious bite marks on the distal end. This is on display next to a similarly-bitten ischium, which is shown in the MMFC14 post linked above.

Here’s a big apatosaur cervical, in antero-ventral view, with a dorsal rib draped over its left side. The cervical ribs are not fused in this specimen, so it was probably still growing. Here’s a labeled version:

The short centrum and nearly-vertical transverse processes indicate that this is a pretty posterior cervical, possibly a C13 or thereabouts. This specimen was over the fence in the exhibit area and I couldn’t throw a scale bar at it, but I’d describe it as “honkin'”. Like most of the apatosaur material at DJ, this vert is from the Mygatt-Moore Quarry.

Of course the real reason I was at Dinosaur Journey was to see the Snowmass Haplocanthosaurus that John Foster and I described back in 2014. You may remember that its caudal vertebrae have wacky neural canals. You may also have noticed a recent uptick in the number of posts around here about wacky neural canals. The game is afoot.

But as cool as they were, the Triassic tracks, the collared lizard, and even the Snowmass Haplo were only targets of opportunity. Brian and I had gone to Utah for this:

That photo was taken by Paige Wiren of Salt Lake City, on the day that she discovered that bone eroding out of a riverbank, just as you see it.

Here’s Paige with the element, which proved to be the left femur of an apatosaurine sauropod. It’s face down in these photos, so we’re looking at the medial side. The articular head is missing from the proximal end – it should be facing toward Paige’s right knee in the above photo – but other than that and a few negligible nicks and dings, the femur was complete and in really good shape.

Paige did the right thing when she found the femur: she contacted a paleontologist. Specifically, she asked a friend, who in turn put her in touch with Carrie Levitt-Bussian, the paleontology Collections Manager at the Natural History Museum of Utah. Based on Paige’s photos and maps, Carrie was able to identify the element as a dinosaur femur, probably sauropod, within the territory of the BLM Hanksville Field Office. John Foster, the Director of the Museum of Moab, has a permit to legally collect vertebrate fossils from that area, and he works on sauropods, so Carrie put Paige in touch with John and with ReBecca Hunt-Foster, the district paleontologist for the BLM’s Canyon Country District in Utah.

Now, I know there’s a lot of heated rhetoric surrounding the Bureau of Land Management, but whatever your political bent, remember this: those are our public lands. Therefore the fossils out there are the collective property of all of us, and we should all be upset if they get poached or vandalized. Yes, that is a big problem – the Brontomerus type quarry was partially poached before the bones we have now were recovered, and vandalism at public fossil sites in Utah made the national news while we were out there.

So that’s what we went to do: salvage this bone for science and education before it could be lost to erosion or asshats. Brian and I were out there to assist John, ReBecca, and Paige, who got to see her find come out of the ground and even got her hands dirty making the plaster jacket. Brian and John headed out to the site Friday morning and met up with Paige there, and ReBecca and I caravanned out later in the day, after I got back from Fruita.

But I’m getting ahead of myself a bit. We didn’t have to jacket the whole thing. It had naturally broken into three pieces, with thin clay infills at the breaks. So we just slid the proximal and middle thirds away as we uncovered them, and hit any loose-looking pieces with consolidant. The distal third was in more questionable shape, so we did make a partial jacket to hold it together.

We also got to camp out in gorgeous country, with spectacular (and welcome) clouds during the day and incredible starry skies at night.

We floated the femur out of the site using the Fosters’ canoe at the end of the day on Saturday, and loaded up to head back to Moab on Sunday. At one point the road was empty and the sky was not, so I stood on the center line and took some photos. This one is looking ahead, toward I-70 and Green River.

And this one is looking behind, back toward Hanksville.

Here are John and Brian with the femur chunks in one of the back rooms of the Museum of Moab. The femur looks oddly small here, but assembled it was 155 cm (5’1″) long and would have been 160 (5’3″) or more with the proximal head. Smaller than CM 3018 and most of the big mounted apatosaurs, but nothing to sneeze at.

What happens to it next? It will be cleaned, prepped, and reassembled by the volunteers and exhibit staff at the Museum of Moab, and eventually it will go on public display. Thousands of people will get to see and learn from this specimen because Paige Wiren made the right call. Go thou and do likewise.

That was the end of the road for the femur (for now), but not for Brian and me. We had business in Cedar City and St. George, so we hit the road Sunday afternoon. Waves of rainclouds were rolling east across Utah while we were rolling west, with breaks for sunlight in between. I miiiight have had to swerve a couple of times when all the scenery distracted me from driving, and I definitely made an obnoxious number of stops to take pictures.

I don’t remember which scenic overlook this was, but it was a pretty darned good view. This is another one that will reward embiggening – check out those mesas marching off into the distance.

In Cedar City we were guests of Andrew R.C. Milner, Site Paleontologist and Curator at the St. George Dinosaur Discovery Site at Johnson Farm (SGDS). We spent most of Monday at SGDS, getting our minds comprehensively blown by the amazing trace and body fossils on display. It was my first time visiting that museum, but it sure as heck won’t be the last.

I didn’t take nearly enough photos in St. George – too busy helping Brian do some filming for a future project – but I did get this gem. This is a Eubrontes track, from a Dilophosaurus-sized theropod. This is a positive track, a cast of the dinosaur’s foot made by sandy sediment that filled the natural mold formed when the dino stepped into mud. The high clay content of the mud recorded the morphology of the foot in fine detail, including the bumps of individual scales on the foot pads. The vertical streaks were cut into the side of the track by similar scales as the animal’s foot pushed into the mud.

The full story of the Johnson Farm tracks and trackmakers is beautifully told in the book Tracks in Deep Time: The St. George Dinosaur Discovery Site at Johnson Farm, by Jerry Harris and Andrew Milner. I hadn’t read it before, so I picked up a copy in the gift shop and I’ve been devouring it. As a professional scientist, educator, and book author myself, I’m jealous of what Jerry and Andrew produced – both the text and the abundant full-color illustrations are wonderfully clear, and the book is well-produced and very affordable.

From St. George we hit the road home, and rolled into Claremont just before midnight on Monday. It was a whirlwind tour – 1800 miles, three museums, and two fossil sites in six days – and my brain is still fizzing with all of the things we got to see and do.

One of the many pros of having a professional artist as a friend is that minimal hospitality, like letting him crash on my couch, is sometimes rewarded with original art. Brian was already gone when I got up Tuesday morning, but this was waiting for me on the dining room table. (Want your own? Help Brian make more monsters here.)

I owe plenty of thanks myself: to the Foster and Milner families for their near-maximal hospitality, to Julia McHugh of Dinosaur Journey for assistance in collections, to Diana Azevedo, Jalessa Spor, Jerry Harris, and the rest of the SGDS staff for being such gracious hosts, to Brian for being such a great friend and traveling companion, and most of all to Paige Wiren for finding the apato femur and helping us save it for science. You’re all top-notch human beings and I hope our paths cross again soon.

Norwescon 41 Guests of Honor: Ken Liu, Galen Dara, and, er, me. Mike would like to remind you that you can get your own ‘Kylo Stabbed First’ t-shirt here.

The week before last I was fortunate to be the Science Guest of Honor at Norwescon 41 in Seattle (as threatened back when). I had a fantastic time. I got to give talks on binocular stargazing and the sizes of the largest sauropods and whales (ahem), participate on panels on alien biology and creature drawing, and meet a ton of cool people, including my fellow Guests of Honor, multiple-award-winning author Ken Liu and multiple-award-winning artist Galen Dara, both of whom turned out to be humble, easygoing, regular folks (if frighteningly talented).

I also had a lot of great conversations with folks who were attending the con, which is exactly what I wanted. One of the most interesting was a hallway conversation with a fellow DM named Shawn Connor. He had a great question for me, which I liked so much I wanted to answer it here on the blog. Here’s his question, copied with permission from a follow-up email:

I run tabletop RPGs, and in my current game one of the characters is a caveman type who naturally grew up hunting dinosaurs. As one does. His weapon is a dinosaur bone, customized and used as a club. I have attached the picture that he came up with [below]. Now understanding the picture is obviously not of a real dinosaur bone – it’s probably a chicken bone or a cow bone or something – let’s assume for the sake of this exercise that it is and that it is four feet long stem to stern. Given that, two questions: discounting the extra bling attached how heavy would such a bone be, and what kind of dinosaur could it have come from?

I’m going to answer those questions out of order. Advance warning: this will be a loooong post that will go down several rabbit holes that are likely of more intense interest to me, personally, than to anyone else on the planet. Read on at your own risk.

Whose femur is in the image?

First, Shawn is correct in noting that the femur in the image provided by his player is not a dinosaur femur. The prominent trochanters and spherical head offset on a narrow neck clearly make it a mammal femur, and if it’s four feet long, it could only have come from an elephant or an indricothere. Or a giant humanoid, I suppose, which is what the anatomy of the bone in the image most closely resembles. (It also appears to be foreshortened to make the distal end look bigger, or deliberately distorted to enhance the clubby-ness.)

Mounted elephant at the Museum of Osteology in Oklahoma City, with Tyler Hunt for scale.

But let’s play along and assume it’s from a non-human mammal. How big? Back in 2016 I was fortunate to get to measure most of the mounted large mammal skeletons at the Museum of Osteology in Oklahoma City, along with Tyler Hunt, then a University of Oklahoma undergrad and now finishing up his MS thesis under my mentor, Rich Cifelli.* The mounted elephant at the Museum of Osteology has a shoulder height of 254 cm (8 ft, 4 in) and a femur length of 102 cm (3 ft, 4 in). Assuming isometric scaling, a world record elephant with a shoulder height of 366 cm (12 ft) would have a femur length of 147 cm (4 ft, 10 in). So a four-foot (122 cm) femur would belong to an elephant roughly in the middle of that range, about ten feet (3 m) tall at the shoulder. That’s the size of the big bull elephant mounted at the Field Museum in Chicago.

The big mounted bull elephant at the Field Museum is 10 feet tall at the shoulder and weighed 6 tons in life. Note Mike for scale on the lower right. He and the elephant are about equidistant from the camera, so he should make a roughly accurate scale bar. Photo from our visit in 2005!

* Two further notes: first, I have roughly a zillion awesome photos from that 2016 visit to the Museum of Osteology, both of the specimens and of Tyler and me measuring them – not having posted them yet is one of the things I was whingeing about in the post that kicked off our return-to-weekly-posting thing this year. And second, I owe a belated and public thanks to the folks at the Museum of Osteology for accommodating Tyler and me. They helped us with ladders and so on and basically gave us free rein to play with collect data from their mounted skeletons, which was incredibly generous and helpful, and fortunately reflects the pro-research and pro-researcher attitude of most museums.

Which dinos had four-foot femora?

As for what kind of dinosaur a four-foot femur could have come from, we can rapidly narrow it down to a handful of clades: sauropods, ornithopods, theropods, and stegosaurs.

  • Sauropods. The longest complete femora of Patagotitan are 238 cm (7 ft, 10 in; Carballido et al. 2017), and an incomplete femur of Argentinosaurus has an estimated complete length of 250 cm (8 ft, 2 in; Mazzetta et al. 2004). So a four-foot femur would not be from a particularly large sauropod – something about elephant-sized, as you might expect from the elephant comparison above. Our old friend Haplocanthosaurus will fit the bill, as we’ll see in a bit.
  • Ornithopods. Femora of 172 cm (5 ft, 8 in) are known for the hadrosaurs Shantungosaurus (Hone et al. 2014) and Huaxiaosaurus (Zhao and Li 2009), and Zhao et al. (2007) reported a 170 cm (5 ft, 7 in) femur for Zhuchengosaurus (Huaxiaosaurus and Zhuchengosaurus may be junior synonyms of Shantungosaurus). But those are all monsters, well over 10 metric tons in estimated mass. So a four-foot femur would be from a large but not insanely large hadrosaur.

Mmmmmm…suffering. OM NOM NOM NOM!!

  • Theropods. Among the largest theropods, the holotype of Giganotosaurus has a femur length of 143 cm (4 ft, 8 in; Coria and Salgado 1995), and ‘Sue’ the T. rex (a.k.a. FMNH PR2081) has a right femur 132 cm long (4 ft, 4 in; Brochu 2003). So a four-foot femur from a theropod would definitely be from one of the monsters. The femur of Saurophaganax was 113.5 cm long (Chure 1995), just under four feet, which I only note as an excuse to use the above photo, which I adore.
  • Stegosaurs. I don’t know the longest femur that has been recovered from a stegosaur, but getting in the ballpark is easy. NHMUK PV R36730 has a femur 87 cm long, and the whole animal was approximately 6 m long (Maidment et al. 2015). Partial bits and bobs of the largest stegosaurs suggest animals about 9 m long, implying a femur length of about 130 cm (4 ft, 3 in), or just over the line.

I think that’s it. I don’t know of any ceratopsians or ankylosaurs with femora long enough to qualify – I assume someone will let me know in the comments if I’ve forgotten any.

How much would a four-foot femur weigh?

There are a couple of ways to get to the answer here. One is to use Graphic Double Integration, which is explained in this post.

Limb bones are not solid – in terrestrial tetrapods there is virtually always a marrow cavity of some sort, and in marine tetrapods the limb bones tend to be cancellous all the way through. Estimating the mass of a limb bone is a lot like estimating the mass of a pneumatic bone: figure out the cross-sectional areas of the cortex and marrow cavity (or air space if the bone is pneumatic), multiply by the length of the element to get volumes, and multiply those volumes by the density of the materials to get masses. I piled up all the relevant numbers and formulas in Tutorial 24, a move that has frequently made me grateful to my former self (instead of cussing his lazy ass, which is my more usual attitude toward Past Matt).

Currey and Alexander (1985: fig. 1)

Sauropod limb bones are pretty darned dense, with extremely thick cortices and smallish marrow spaces that are not actually hollow (tubular) but are instead filled with trabecular bone. My gut feeling is that even a four-foot sauropod femur would be almost too heavy to lift, let alone wield as a club, so in the coming calculations I will err in the direction of underestimating the mass, to give our hypothetical caveman the best possible chance of realizing his dream.

Some of the proportionally thinnest cortices I’ve seen in sauropod limb bones are those of the macronarian Haestasaurus becklesii NHMUK R1870, which Mike conveniently showed in cross-section in this post. I could look up the actual dimensions of the bones (in Upchurch et al 2015: table 1 – they passed the MYDD test, as expected), but for these calculations I don’t need them. All I need are relative areas, for which pixels are good enough.

First, I took Mike’s photo into GIMP and drew two diameters across each bone, one maximum diameter and a second at right angles. Then I drew tick marks about where I think the boundaries lie between the cortex and the trabecular marrow cavity. Next, I used those lines as guides to determine the outer diameters (D) and inner diameters (d) in pixels, as noted in the image.

For the radius, on the left, the mean diameters are D = 891 and d = 648. I could divide those by 2 to get radii and then plug them into the formula for the area of a circle, etc., but there’s an easier way still. For a tubular bone, the proportional area of the inner circle or ellipse is equal to k^2, where k = r/R. Or d/D. (See Wedel 2005 and Tutorial 24 for the derivation of that.) For the Haestasaurus radius (the bone, not the geometric dimension), d/D = 0.727, and that number squared is 0.529. So the marrow cavity occupies 53% of the cross-sectional area, and the cortex occupies the other 47%.

For the ulna, on the right, the mean diameters are D = 896 and d = 606, d/D = 0.676, and that number squared is 0.457. So in this element, the marrow cavity occupies 46% of the cross-sectional area, and the cortex occupies the other 54%.

(For this quick-and-dirty calculation, I am going to ignore the fact that limb bones are more complex than tubes and that their cross-sectional properties change along their lengths – what I am doing here is closer to Fermi estimation than to anything I would publish. And we’ll ground-truth it before the end anyway.)

Left: rat humerus, right: mole humerus. The mole humerus spits upon my simple geometric models, with extreme prejudice. From this post.

You can see from the photo (the Haestasaurus photo, not the mole photo) that neither bone has a completely hollow marrow cavity – both marrow cavities are filled with trabecular bone. By cutting out good-looking chunks in GIMP and thresholding them, I estimate that these trabecular areas are about 30% bone and 70% marrow (actual marrow space with no bone tissue) by cross-sectional area. According to Currey and Alexader (1985: 455), the specific gravities of fatty marrow and bone tissue are 0.93 and 2.1, respectively. The density of the trabecular area is then (0.3*2.1)+(0.7*0.93) =  1.28 kg/L, or about one quarter more dense than water.

But that’s just the trabecular area, which accounts for about one half of the cross-sectional area of each bone. The other half is cortex, which is probably close to 2.1 kg/L throughout. The estimated whole-element densities are then:

Radius: (0.53*1.28)+(0.47*2.1) = 1.67 kg/L

Ulna: (0.46*1.28)+(0.54*2.1) = 1.72 kg/L

Do those numbers pass the sniff test? Well, any skeletal elements that are composed of bone tissue (SG = 2.1) and marrow (SG = 0.93) are constrained to have densities somewhere between those extremes (some animals beat this by building parts of their skeletons out of [bone tissue + air] instead of [bone tissue + marrow]). We know that sauropod limb bones tend to have thick cortices and small marrow cavities, and that the marrow cavities are themselves a combination of trabecular bone and actual marrow space, so we’d expect the overall density to be closer to the 2.1 kg/L end of the scale than the 0.93 kg/L end. And our rough estimates of ~1.7 kg/L fall about where we’d expect.

Femur of Haplocanthosaurus priscus, CM 572, modified from Hatcher (1903: fig. 14).

To convert to masses, we need to know volumes. We can use Haplocanthosaurus here – the femur of the holotype of H. priscus, CM 572, is 1275 mm long (Hatcher 1903), which is just a hair over four feet (4 ft, 2.2 in to be exact). The midshaft width is 207 mm, and the proximal and distal max widths are 353 and 309 mm, respectively. I could do a for-real GDI, but I’m lazy and approximate numbers are good enough here. Just eyeballing it, the width of the femur is about the same over most of its length, so I’m guessing the average width is about 23 cm. The average width:length ratio for the femora of non-titanosaur sauropods is 3:2 (Wilson and Carrano 1999: table 1), which would give an anteroposterior diameter of about 15 cm and an average diameter over the whole length of 19 cm. The volume would then be the average cross-section area, 3.14*9.5*9.5, multiplied by the length, 128 cm, or 36,273 cm^3, or 36.3 L. Multiplied by the ~1.7 kg/L density we estimated above, that gives an estimated mass of 62 kg, or about 137 lbs. A femur that was exactly four feet long would be a little lighter – 86.6% as massive, to be exact, or 53.4 kg (118 lbs).

I know that the PCs in RPGs are supposed to be heroes, but that seems a little extreme.

But wait! Bones dry out and they lose mass as they do so. Lawes and Gilbert (1859) reported that the dry weight of bones of healthy sheep and cattle was only 74% of the wet mass. Cows and sheep have thinner bone cortices than sauropods or elephants, but it doesn’t seem unreasonable that a dry sauropod femur might only weigh 80% as much as a fresh one. That gets us down to 43 kg – about 95 lbs – which is still well beyond what anyone is probably going to be wielding, even if they’re Conan the Cimmerian.

Picture is unrelated.

I mentioned at the top of this section that there are a couple of ways to get here. The second way is to simply see what actual elephant femora weigh, and then scale up to dinosaur size. According to Tefera (2012: table 1), a 110-cm elephant femur has a mass of 21.5 kg (47 lbs). I reckon that’s a dry mass, since the femur in question had sat in a shed for 50 years before being weighed (Tefera 2012: p. 17). Assuming isometry, a four-foot (122 cm) elephant femur would have a dry mass of 29.4 kg (65 lbs). That’s a lot lighter than the estimated mass of the sauropod femur – can we explain the discrepancy?

 

Femora of a horse, a cow, and an elephant (from left to right in each set), from Tefera (2012: plate 1).

I think so. Elephant femora are more slender than Haplocanthosaurus femora. Tefera (2012) reported a circumference of 44 cm for a 110-cm elephant femur. Scaling up from 110 cm to 122 cm would increase that femur circumference to 49 cm, implying a mean diameter of 15.6 cm, compared to 19 cm for the Haplo femur. That might not seem like a big difference, but it means a cross-sectional area only 2/3 as great, and hence a volume about 2/3 that of a sauropod femur of the same length. And that lines up almost eerily well with our estimated masses of 29 and 43 kg (ratio 2:3) for the four-foot elephant and sauropod femora.

A Better Weapon?

Could our hypothetical caveman do better by choosing a different dinosaur’s femur? Doubtful – the femora of ‘Sue’ are roughly the same length as the Haplo femur mentioned above, and have similar cross-sectional dimensions. Hadrosaur and stegosaur femora don’t look any better. Even if the theropod femur was somewhat lighter because of thinner cortices, how are you going to effectively grip and wield something 15-19 cm in diameter?

I note that the largest axes and sledgehammers sold by Forestry Suppliers, Inc., are about 3 feet long. Could we get our large-animal-femur-based-clubs into the realm of believability by shrinking them to 3 feet instead of 4? Possibly – 0.75 to the third power is 0.42. That brings the elephant femur club down to 12.3 kg (27 lbs) and the sauropod femur club down to 18 kg (40 lbs), only 2-3 times the mass of the largest commonly-available sledgehammers. I sure as heck wouldn’t want to lug such a thing around, much less swing it, but I can just about imagine a mighty hero doing so.

Yes, there were longer historical weapons. Among swing-able weapons (as opposed to spears, etc.), Scottish claymores could be more than four feet long, but crucially they were quite light compared to the clubs we’ve been discussing, maxing out under 3 kg, at least according to Wikipedia.

T. rex FMNH PR2081 right fibula in lateral (top) and medial (bottom) views. Scale is 30 cm. From Brochu (2003: fig. 97).

If one is looking for a good dinosaur bone to wield as a club, may I suggest the fibula of a large theropod? The right (non-pathologic) fibula of ‘Sue’ is 103 cm long (3 ft, 4.5 in), has a max shaft diameter just under 3 inches – so it could plausibly be held by (large) human hands, and it probably massed something like 8-9 kg (17-20 lbs) in life, based on some quick-and-dirty calculations like those I did above. The proximal end is even expanded like the head of a war club. The length and mass are both in the realm of possibility for large, fit, non-supernaturally-boosted humans. Half-orc barbarians will love them.

And that’s my ‘expert’ recommendation as a dice-slinging paleontologist. Thanks for reading – you have Conan-level stamina if you got this far – and thanks to Shawn for letting me use his question to freewheel on some of my favorite geeky topics.

References

I choose Haplocanthosaurus

November 18, 2016

snowmass-haplocanthosaurus-caudals

Oh man, 2016, you are really working on my nerves.

Sometimes it’s a positive balm to hold a piece of an animal dead and gone for 145 million years, or stare at a thousand vertical feet of sandstone, and know that we are all ants.

These lovelies here intrigue me deeply. They’re three of the caudal vertebrae recovered from the Snowmass Haplocanthosaurus that John Foster and I described a couple of years ago. Pretty sure I’ll have more to say about them in the future. For now it’s enough that they’ve come across such a vast gulf of time and given this stressed-out primate a little perspective.

Reference

Foster, J.R., and Wedel, M.J. 2014. Haplocanthosaurus (Saurischia: Sauropoda) from the lower Morrison Formation (Upper Jurassic) near Snowmass, Colorado. Volumina Jurassica 12(2): 197–210. DOI: 10.5604/17313708 .1130144

Wedel 2005 Morrison sauropod cervicals 1 - Diplodocus

When I was back in Oklahoma in March, I met with Anne Weil to see some of the new Apatosaurus material she’s getting out of her Homestead Quarry. It’s nice material, but that’s a post for another day. Anne said something that really resonated with me, which was, “I love it when you guys post about vertebral morphology, because it helps me learn this stuff.” Okay, Anne, message received. This will begin to make things right.

Wedel 2005 Morrison sauropod cervicals 2 - Barosaurus and centra shapes

I spent a week at BYU back in 2005, collecting data for my dissertation. One of the first things I had to do was teach myself how to identify the vertebrae of different sauropods, because BYU has just about all of the common Morrison taxa. These are the notes I made back then.

Wedel 2005 Morrison sauropod cervicals 3 - Brachiosaurus and Apatosaurus

I always planned to do something with them – clean them up, get them into a more usable form. There are a lot of scribbly asides that are probably hard for others to read, and it would be more useful if I put the easily confused taxa next to each other – Barosaurus next to Brachiosaurus, for example. And I didn’t go into serial changes at all.

Wedel 2005 Morrison sauropod cervicals 4 - Camarasaurus and Haplocanthosaurus

Still, hopefully someone will find these useful. If there are things I missed or got wrong, the comment thread is open. And if you want all four spreads in one convenient package, here’s a PDF: Wedel 2005 notes on Morrison sauropod cervicals

Mike and I leave for the Sauropocalypse tomorrow. I’m hoping to post at least a few pretty pictures from the road, as I did for the Mid-Mesozoic Field Conference two years ago. Stand by…

Foster and Wedel 2014 fig 3 - dorsals

Fig. 3. MWC 8028, Haplocanthosaurus dorsal vertebrae. A. Lateral view of dorsal centrum with bottom edge of lateral pneumatic fossa preserved. B. Dorsal view of same centrum as in A, showing the median septum between the paired lateral fossae. C. Lateral view of dorsal centrum with smaller segment of the lateral pneumatic fossa margin preserved. D. Dorsal view of same centrum as in C, again showing the median septum and paired lateral fossae. E. Lateral view of dorsal centrum with partial pleurocoel preserved. F. Cross-sectional (posterior) view of same dorsal as in E. G. Dorsal neural spines in lateral (top) and anterior or posterior (center, bottom) views. Scale bars = 10 cm.

Right on the heels of Aquilops last week, my paper with John Foster on the new specimen of Haplocanthosaurus from Snowmass, Colorado, was just published in Volumina Jurassica. I’ll have more to say about it later, but right now I am up against a deadline on a big project and I need to go work on that. I’m only popping up here to note two quick things.

First, if you’re not familiar with Volumina Jurassica – and I wasn’t, before this project – it’s a free-to-access* journal that publishes papers on all aspects of the Jurassic. The current issue is specifically dedicated to the Jurassic formations of the American West. There’s a lot of interesting stuff in there, but of special interest to SV-POW! readers will be the paper by Cary Woodruff and John Foster on the legendary and possibly apocryphal Amphicoelias fragillimus.

* But not truly open access since the journal claims to retain exclusive rights to distribute the papers. That seems like a quaint affectation now that the internet is here, but whatever – at least they let anyone download the PDF for free, which is primarily what I care about.

Foster and Wedel 2014 fig 4 - sacrum

Fig. 4. Sacra of Haplocanthosaurus.  A. MWC 8028, sacrum in right lateral view. B. MWC 8028, close-up of S4 and S5 centra highlighting pneumatic fossae. C. MWC 8028 with divisions between the vertebrae overlaid. D. CM 879, sacrum in right lateral view with divisions between the vertebrae overlaid. E. CM 572 in right lateral view, after Hatcher (1903c: plate 4). B–E are not shown at the same scale, scale bar for A = 20 cm. Note that the neural arches in CM 572 were restored during preparation, and the sacral neural spines as shown here are probably lower than they would have been in life.

Second, the figure resolution in the PDF of the Haplocanthosaurus paper is not stellar, so as is the case with almost all of our papers, the full-color, high-resolution figures are available at the paper’s page on the sidebar.

Gotta run.

For our previous posts on Haplocanthosaurus, go here; for those on Amphicoelias, including Mike’s very popular, “How big was Amphicoelias fragillimus? I mean, really?”, go here.

References

No time for anything new, so here’s a post built from parts of other, older posts.

The fourth sacral centrum of Haplocanthosaurus CM 879, in left and right lateral view. This is part of the original color version of Wedel (2009: figure 8), from this page. (Yes, I know I need to get around to posting the full-color versions of those figures. It’s on my To Do list.)

Note the big invasive fossa on the right side of the centrum. The left side is waisted (narrower at the middle than the ends) like most vertebrae of most animals, but has no distinct fossa on lateral face of the centrum. What’s up with that? Here’s an explanation from an old post (about another sauropod) that still fits:

Now, this asymmetry is also weird, but it’s expected weirdness. Pneumaticity seems to just be inherently variable, whether we’re talking about human sinuses or the facial air sacs of whales or the vertebrae of chickens. It appears that the form of pneumatic features is entirely determined by local tissue interactions, with little or no genetic control of the specific form. Think of it this way: genes prescribe certain developmental events, and those events bring tissues into contact–such as pneumatic epithelium and bone. The morphology of the bone arises out of that interaction, and each interaction of bone and pneumatic epithelium has the potential to produce something new. In this case, the diverticula on the left side of the vertebral column come from the lungs or air sacs on the left, and those on the right side come from the lungs or airs sacs on the right, so it’s really two sets of diverticula contacting the bone independently. The wonder, then, is not that pneumatic bones are so variable, but that we see any regularities at all.

Matt and I have been looking in more detail at indications of maturity in sauropod skeletons, as we prepare the submission of the paper arising from our response to Woodruff and Fowler (2012) [part 1, part 2, part 3, part 4, part 5, part 6].  Here is an oddity.

Sacra of Haplocanthosaurus. Top, H. utterbacki holotype CM 879 in right lateral view, from Hatcher (1903:fig. 15). Bottom, H. priscus holotype CM 572 in left lateral view (reversed), from Hatcher (1903:pl. IV, part 3). To the same scale.

H. priscus is the type species of Haplocanthosaurus; H. utterbacki is the second species, named by Hatcher in the 1903 monograph that described the original material in detail.  As previously noted, the type species is based on adult material, and the referred specimen on subadult material.  This is shown by their different stages of neurocentral fusion, and corroborated by the size of the specimens as indicated in the composite illustration above.

There is a lot of fusion going on in the sacra of dinosaurs:

  1. sacral neural arches fused to their centra
  2. consecutive sacral centra fused together
  3. consecutive sacral neural spines fused together
  4. sacral lateral processes fused to ilia

As we would expect, the less mature of the two Haplocanthosaurus individuals is less fused in most respects: none of the centra were fused either to each other or their respective neural arches, and the ilium was not fused to any of the lateral processes, whereas in the adult all neural arches are fused to their centra, the five sacral centra are all fused together, and the ilium is fused to the lateral processes.

How strange, then, that the consecutive neural spines are more fused in the juvenile!  Not only are spines 1, 2 and 3 fused along their entire dorsolateral length, as in the adult, but spine 4 is similarly fused.  And more: the neurapophysis of spine 5 is fused to that of 4, even though the spines are not fused more ventrally.

What does this mean?  Hatcher (1903:27-28) took it as indicative of species-level separation.  After briefly noting that the posterior dorsal centra of H. utterbacki are more opisthocoelous than those of H. priscus, and speculating that the adult of the referred species was probably larger than that of the type, he continued:

But the most distinctive character is to be found in the sacrum which, in the present species, has the five neural spines normally coössified.  The first four are cocoössified throughout their entire length, forming a long bony plate.  The union between the fourth and fifth is limited to the extremities while medially [sic, presumably meaning half way up the spines] they are separated by an elongated foramen.  In H. priscus only the spines of the three anterior sacrals are coössified, those of the first and second [sic, presumably intending fourth and fifth] sacrals remaining free.  This difference exists notwithstanding that the type of the present species was scarcely adult, the sacral centra being neither coössified with one another nor with their neural arches. By some this character might be considered as of generic importance although I prefer to consider it as of only specific value since in all other parts of the skeleton preserved, there are no distinguishing characters which could be considered as of generic value.

At present, however, the synonymy of H. utterbacki with the type species, proposed by McIntosh and Williams (1988:22), seems to be universally accepted.  If they truly belong to the same taxon then the only realistic possibility is that we are seeing individual variation in the timing of fusion.  That certainly seems to have been the opinion of McIntosh and Williams (1988:14), writing about the sacrum of their own specimen, the H. delfsi holotype CMNH 10380:

As in CM 572 the short to moderately long spines of sacrals one through three are firmly united throughout, and those of sacrals four and five are firmly united to midheight. In CM 572 spines four and five are free, but this is probably an individual character because in the even younger CM 879 all five spines are united.

All of which means: we need to be really careful when drawing conclusions about taxonomy or ontogeny from individual observations of skeletal fusion.

Bonus Pneumaticity Observation: In the image at top, you’ll see that the centrum of sacral 4 in CM 879 has a couple of pneumatic fossae. For more than you probably wanted to know about those specific holes in that specific bone, see this post and the linked paper.

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.
  • McIntosh, J.S., and Williams, M. E. 1988. A new species of sauropod dinosaur, Haplocanthosaurus delfsi sp. nov., form the Upper Jurassic Morrison Fm. of Colorado. Kirtlandia 43:3-26.
  • 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.

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 Haplocanthosaurusmuch 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:

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.

Broadly speaking, pneumatic sauropod vertebrae come in two flavors. In more primitive, camerate vertebrae, modeled here by Haplocanthosaurus, the centrum is a round-ended I-beam and the neural arch is composed of intersecting flat plates of bone called laminae (lam above; fos = fossa, nc = neural canal, ncs = neurocentral suture; Ye Olde Tyme vert pic from Hatcher 1903).

In more derived, camellate vertebrae, the centrum and neural arch are both honeycombed with many small air spaces. This inflated-looking morphology is very similar to that seen in birds, like the turkey we recently discussed. The fossae and foramina on the outside tend to be smaller and more numerous than in camerate vertebrae, as shown here in a titanosauriform axis from India (Figure 3 from Wilson and Mohabey 2006). The green arrows show that the fossae visible on the external surface are excavations or depressions into the honeycombed internal structure of the bone.

External fossae on bones can house many different soft tissues, including muscles, pads of fat or cartilage, and pneumatic diverticula (O’Connor 2006). Pneumatic fossae are often strongly lipped and internally subdivided and may contain pneumatic foramina, which makes them easier to diagnose (but they may also be simple, smooth, and “blind”, which makes them harder to diagnose as pneumatic). But in all of these cases we are usually talking about the same thing: a visible excavation into a corpus of bony tissue, which may have marrow spaces inside if it is apneumatic, or air spaces inside if it is pneumatic (the corpus of bone, not the dent). That’s probably how most of us think about fossae, and it would hardly need to be explained…except that sometimes, something much weirder happens.

Consider this cervical of Brachiosaurus (this is BYU 12866, from Dry Mesa, Colorado). Brachiosaurus and Giraffatitan have an in-between form of vertebral architecture that my colleagues and I have called semicamellate (Wedel et al. 2000); the centrum does have large simple chambers (camerae), but smaller, thin-walled camellae are also variably present, especially along the midline of the vertebra and in the ends of the centrum. As in Haplocanthosaurus, the neural arch is composed of intersecting plates of bone; unlike Haplocanthosaurus, these laminae are not flat or smooth but are instead highly sculpted with lots of small fossae. Janensch (1950) called these “Aussenkaverne”, or accessory outside cavities, because and they are smaller and more variable than the large fossae and foramina that invade the centrum.

And that’s the weird thing. As the red arrows in the above image show, the “Aussenkaverne” are not excavations or depressions into anything, except the space on the other side of the lamina (which in life would have been occupied by another diverticulum). The neural arches of Brachiosaurus and Giraffatitan are not excavated by fossae, they’re embossed, like corporate business cards and fancy napkins.

What’s up with that!? We tend to think of pneumaticity as reducing the mass of the affected elements, but the shortest distance between two vertebral landmarks is a smooth lamina. These embossed laminae actually require slightly more bony material than smooth ones would.

As you can see above, the outer edges of the laminae are thick but the bone everywhere else is very thin. Maybe, like the median septa in pneumatic sauropod vertebrae, the thin bone everywhere except the edges of the laminae was just not loaded very much or very often, and was therefore free to get pushed around by the diverticula on either side, in the sense of being continually and quasi-randomly remodeled into shapes that don’t strike us as being very mechanically efficient. But also like the median septa, the thin parts of the laminae are only rarely perforated (but it does happen), for possible (read: arm-wavy) reasons discussed in the recent FEA post. And maybe the amount of extra bone involved in making embossed laminae versus smooth ones was negligible even by the very light standards of sauropod vertebrae.

Another question: since these thin sheets of bone were sandwiched in between two sets of diverticula, why are the “unfossae” always embossed into them, in the medial or inferior direction? Why don’t any of them pop out laterally or dorsally, looking like domes or bubbles instead of holes, like Mount Fist-of-God from Larry Niven’s Ringworld? Did the developmental program get accustomed to making fossae that went down and into a corpus of bone, and just kept on with business as usual even when there was no corpus of bone to excavate into? I’m only half joking.

I don’t have good answers for any of these questions. I scanned this vert a decade ago and I only noticed how weird the “unfossae” were a few months ago. I’m putting all this here because “Hey, look at this weird thing that I can only wave my arms about” is not a great basis for a peer-reviewed paper, and because I’d like your thoughts on what might be going on.

In Other News

The Discovery Channel’s Clash of the Dinosaurs premiered last night. I would have given you a heads up, except that I didn’t get one myself. I only discovered it was on because of a Facebook posting (thanks, folks!).

COTD is intended to be the replacement, a decade on, for Walking With Dinosaurs. I’m happy to report that one of the featured critters is Sauroposeidon. I grabbed a couple of frames from the clips posted here.

I haven’t seen the series yet, because I don’t have cable. But I’m hoping to catch it at a friend’s place next Sunday night, Dec. 13, when the entire series will be shown again. With any luck, I’ll have more news next week.

Finally, I got to do an interview at Paw-Talk, a forum for all things animal. I’m very happy with how it turned out, so thanks to Ava for making it happen. While you’re over there, have a look around, there’s plenty of good stuff. Brian Switek, whom you hopefully know from this and this, is a contributor; check out his latest here.

References

Bifid Brachiosaurs, Batman!

September 6, 2009

These are the days of miracle and wonder, especially for all you right-minded people out there who are lovers of fine brachiosaurs.  I heard yesterday evening about a new paper in Proceedings of the Royal Society B: You and Li’s (2009, duh) description of a new brachiosaur, the first one known from the Cretaceous of Asia: Qiaowanlong kangxii. Best of all, it’s based primarily on vertebral material:

You and Li (2009:fig. 2)  Cervical vertebrae of Qiaowanlong kangxii holotype FRDC GJ 07-14.

You and Li (2009:fig. 2) Cervical vertebrae of Qiaowanlong kangxii holotype FRDC GJ 07-14. (a) Photograph and (b) interpretative line drawing of C4-C7 in left lateral view; (c) a distal portion of a cervical rib; C9 in (d) cranial, (e) left lateral, (f) caudal, (g) right lateral, (h) dorsal and (i) ventral views. di, diapophysis; f1-f5, fossa 1-fossa 5; pa, parapophysis; poz, postzygapophysis; prz, prezygapophysis; sp, neural spine. Scale bars, 10 cm.

Brachiosaur aficionados will be gazing slack-jawed at parts d, f and h of this figure (the anterior, posterior and dorsal views of C9), which clearly show that the neural spines of the new taxon are bifid (i.e. have two peaks side by side and a trough between them) — just like the cervical neural spines of flagellicaudatans (diplodocids and dicraeosaurs) and camarasaurs.  And mamenchisaurs.  And some titanosaurs.  And Erketu.  Finding this feature yet again — apparently independently evolved in brachiosaurs — makes it about the most plastic character in the matrix.  Very exciting.

That is, it’s exciting if this really is a brachiosaurid.  Now as it happens, Matt was one of the reviewers for this paper (and by the way did an amazingly professional job of not telling me about it until it came out, the git).  He’s told me in email that he’s satisfied that Qiaowanlong really is a brachiosaur, and I hesitate to question that identification given that (A) unlike the authors I’ve never seen the material, and (B) unlike Matt, I’ve spent most of my brachiosaur-presacral quality time with dorsals rather than cervicals.  But, with that caveat, I’m not sure that a compelling case has yet been made for a brachiosaurian identity.

The authors cite three characters in support of a brachiosaurid identity:

  • The most persuasive is the deeply excavated cervical neural spines.
  • Next is a transition in neural spine height: this is quite abrupt in “Brachiosaurusbrancai between cervicals 6 and 7, and also in Sauroposeidon — presumably also between C6 and C7, but that can’t be known for sure, since it’s only the assumption that this is the case that led to the identification of the four preserved Sauroposeidon cervicals as C5-C8 in the first place.  In Qiaowanlong, this transition is “much less pronounced”, with spines increasing in height by only 25% rather then 100% in the other taxa — and occurs between C8 and C9.  All in all, not really very similar to the condition in “B.” brancai.
  • The final character supporting the brachiosaurid identity of Qiaowanlong is the absence of an anterior centrodiapophyseal lamina.  As the authors point out, though, this lamina does exist in “B.” brancai and is absent only in Sauroposeidon; so if this is evidence of anything, it’s a synapomorphy of a clade uniting Qiaowanlong and Sauroposeidon to the absence of other brachiosaurs — something that seems very unlikely given the proportions of the vertebrae.

Putting it all together, there seems to be only one convincing brachiosaur character cited; and that stands against several non-brachiosaur characters, most obviously the bifurcation of the neural spine and the low Elongation Index (centrum length divided by cotyle height) but also by a few other characters that are not discussed in the paper.  For example, Matt has previously noted that in brachiosaur cervicals, the diapophyses are more anteriorly positioned than the parapophyses whereas in diplodocids the opposite is the case: as shown in fig 2(b) above, C6 at least of Qiaowanlong resembles diplodocids in this respect.

To try to get more of a handle on this, I put together a comparative figure of the 8th and 9th cervicals of various sauropods:

8th/9th cervicals vertebrae of various sauropods, scaled to the same centrum length.  "Brachiosaurus" brancai, Sauroposeidon; Qiaowanlong, Diplodocus; Haplocanthosaurus, Camarasaurus

8th/9th cervicals vertebrae of various sauropods, scaled to the same centrum length. From top to bottom and left to right: "Brachiosaurus" brancai, Sauroposeidon; Qiaowanlong, Diplodocus; Haplocanthosaurus, Camarasaurus. Six sauropod vertebrae for the price of one!

Based on overall proportions, I don’t find it intuitively obvious that the Qiaowanlong (middle row, left) more closely resembles the brachiosaurs (top row) than it does the other three.

What does all this mean?  Probably nothing: most likely there are further reasons for the brachiosaurid identification of the new taxon, and lack of space prevented their explanation and illustration.  We can hope that the authors, having placed an initial brief description in Proc. B, will follow it up with a more comprehensive description and analysis in a journal that does not impose such tight length restrictions.  But for now at least, my feeling is that the case for a bifid brachiosaur has yet to be made.

Moving on … Qiaowanlong is also represented by some nice appendicular material: the entire right side of the pelvis (ilium, ischium and pubis).  The ilium certainly looks brachiosaury, so that is another bit of support for the systematic hypothesis, but the proportions of the pelvic bones are very odd:

Right pelvis of "Brachiosaurus" brancai (left), based on composite of Janensch's (1961) figures, and Qiaowanlong (from You and Li 2009: fig. 3a).  Scaled to same ilium length.

Right pelvis of "Brachiosaurus" brancai (left), based on composite of Janensch's (1961) figures, and Qiaowanlong (from You and Li 2009: fig. 3a). Scaled to same ilium length.

You and Li (2009) describe their pelvis as having a “much reduced ischium”, but as is apparent by comparison with the pelvis of “Brachiosaurusbrancai, the ischium is in reasonable proportion to the ilium, and the oddity is more that the pubis is enormous.  So much so that it makes me feel a little ill looking at it, and it makes me wonder how certain it is that all three of these bones are from the same individual — sadly, the paper doesn’t discuss the association of the material.

[Not to flog a dead horse, but this kind of omission shows once more the perils of publishing new taxa in general-interest journals such as Proc. B that impose draconian length limits.  This paper just creeps onto page 7, and I simply don’t believe that it’s possible to do anything like justice to the description of a new taxon in that little space, especially when there is also geography, geology, phylogeny and discussion to be got through.  I don’t want to go all This Is How To Do It, but I can’t help remembering that Darren and I took 18 pages, nearly three times as long, to describe the single partial vertebra that is Xenoposeidon (Taylor and Naish 2007), and it’s not as though that paper wastes a lot of words.  To give You and Li credit, they did squeeze in photos of a representative vertebra from all six cardinal directions, which is great; but only at the cost of the photos being too tiny to be much use.  Please, folks: send your new taxon descriptions to a proper descriptive journal, not to a tabloid!  </hobbyhorse>]

Back on the Dinosaur Mailing List, B tH asked how big Qiaowanlong was.  According to the BBC, the authors say that “the dinosaur would have been a relatively small sauropod about 12m long, 3m high, and weighing perhaps 10 tonnes”.  Can we confirm that?  Well, the excellently comprehensive table of measurements in the paper gives centrum lengths, not counting the condyle, totalling 267 cm for the seven vertebrae C5-C11.  Janensch (1950a:44) gave measurements for the corresponding vertebrae of “Brachiosaurusbrancai HMN SII totalling 577 cm, which is more than twice as long.  If Qiaowanlong was 267/577 = 0.46 times as long as HMN SII, which Janensch (1950b:102) gave as 22.46 m, then it would have been 10.4 m long; it’s not obvious how the authors got the larger figure of 12 m unless they had reason to think the neck was proportionally shorter than in HMN SII.  If Qiaowanlong was isometrically similar to HMN SII, then it was 0.46^3 = 0.99 0.099 times as heavy.  Using my own in-press mass of 23337 kg for HMN SII, this would make Qiaowanlong only 2312 kg in mass — pretty pathetic for a sauropod.

That’s it for now.  I’d be the first to admit that there’s an awful lot of speculation in this post based on relatively little published information.  Hopefully You Hai-Lu will drop by and comment — I’ll be letting him know that I’ve posted this.

References

  • Janensch, Werner.  1950.  Die Wirbelsaule von Brachiosaurus brancai. Palaeontographica (Suppl. 7) 3: 27-93.
    Janensch, Werner.  1950.  Die Skelettrekonstruktion von Brachiosaurus brancai.  Palaeontographica (Suppl. 7) 3: 97-103.
    Janensch, Werner.  1961.  Die Gliedmaszen und Gliedmaszengurtel der Sauropoden der Tendaguru-Schichten.  Palaeontographica, suppl. 7 (1), teil 3, lief. 4: 177-235.
    Taylor, Michael P. and Darren Naish.  2007.  An unusual new neosauropod dinosaur from the Lower Cretaceous Hastings Beds Group of East Sussex, England.  Palaeontology 50 (6): 1547-1564.  doi: 10.1111/j.1475-4983.2007.00728.x
    You, Hai-Lu, and Li, Da-Qing.  2009.  The first well-preserved Early Cretaceous brachiosaurid dinosaur in Asia.  Proceedings of the Royal Society B: Biological Sciences.  doi: 10.1098/rspb.2009.1278.
  • Janensch, Werner.  1950.  Die Wirbelsaule von Brachiosaurus brancai. Palaeontographica (Suppl. 7) 3: 27-93.
  • Janensch, Werner.  1950.  Die Skelettrekonstruktion von Brachiosaurus brancai.  Palaeontographica (Suppl. 7) 3: 97-103.
  • Janensch, Werner.  1961.  Die Gliedmaszen und Gliedmaszengurtel der Sauropoden der Tendaguru-Schichten.  Palaeontographica, suppl. 7 (1), teil 3, lief. 4: 177-235.
  • Taylor, Michael P. and Darren Naish.  2007.  An unusual new neosauropod dinosaur from the Lower Cretaceous Hastings Beds Group of East Sussex, England.  Palaeontology 50 (6): 1547-1564.  doi: 10.1111/j.1475-4983.2007.00728.x
  • You, Hai-Lu, and Li, Da-Qing.  2009.  The first well-preserved Early Cretaceous brachiosaurid dinosaur in Asia.  Proceedings of the Royal Society B: Biological Sciences.  doi: 10.1098/rspb.2009.1278.

And finally … two announcements!

Traumador the Tyrannosaur has asked us to point out that over on ART Evolved (the palaeo-art blog), the next big art gallery is to be sauropod themed.  Details are on the site, so get over there and submit your sauropod art!

And Matt and I will shortly be teaming up with Andy Farke, the open-source paleontologist, on a new project where we plan to actually do some of this Shiny Digital Future that we keep on talking about.  Andy will be announcing the details on Tuesday 8th September.  Mark the date well!  For now, I shall say no more …