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…

In my recent preprint on the incompleteness and distortion of sauropod neck specimens, I discuss three well-known sauropod specimens in detail, and show that they are not as well known as we think they are. One of them is the Giraffatitan brancai lectotype MB.R.2181 (more widely known by its older designation HMN SII), the specimen that provides the bulk of the mighty mounted skeleton in Berlin.

Giraffatitan c8 epipophyses

That photo is from this post, which is why it’s disfigured by red arrows pointing at its epipophyses. But the vertebra in question — the eighth cervical of MB.R.2181 — is a very old friend: in fact, it was the subject of the first ever SV-POW! post, back in 2007.

In the reprint, to help make the point that this specimen was found extremely disarticulated, I reproduce Heinrich (1999:figure 16), which is Wolf-Dieter Heinrich’s redrawing of Janensch’s original sketch map of Quarry S, made in 1909 or 1910. Here it is again:

Taylor 2015: Figure 5. Quarry map of Tendaguru Site S, Tanzania, showing incomplete and jumbled skeletons of Giraffatitan brancai specimens MB.R.2180 (the lectotype, formerly HMN SI) and MB.R.2181 (the paralectotype, formerly HMN SII). Anatomical identifications of SII are underlined. Elements of SI could not be identified with certainty. From Heinrich (1999: figure 16), redrawn from an original field sketch by Werner Janensch.

Taylor 2015: Figure 5. Quarry map of Tendaguru Site S, Tanzania, showing incomplete and jumbled skeletons of Giraffatitan brancai specimens MB.R.2180 (the lectotype, formerly HMN SI) and MB.R.2181 (the paralectotype, formerly HMN SII). Anatomical identifications of SII are underlined. Elements of SI could not be identified with certainty. From Heinrich (1999: figure 16), redrawn from an original field sketch by Werner Janensch.

For the preprint, as for this blog-post (and indeed the previous one), I just went right ahead and included it. But the formal version of the paper (assuming it passes peer-review) will by very explicitly under a CC By licence, so the right thing to do is get formal permission to include it under those terms. So I’ve been trying to get that permission.

What a stupid, stupid waste of time.

Heinrich’s paper appeared in the somewhat cumbersomely titled Mitteilungen aus dem Museum fur Naturkunde in Berlin, Geowissenschaftliche Reihe, published as a subscription journal by Wiley. Happily, that journal is now open access, published by Pensoft as The Fossil Record. So I wrote to the Fossil Record editors to request permission. They wrote back, saying:

We are not the right persons for your question. The Wiley Company holds the copyright and should therefore be asked. Unfortunately, I do not know who is the correct person.

I didn’t know who to ask, either, so I tweeted a question, and copyright guru Charles Oppenheim suggested that I email permissions@wiley.com. I did, only to get the following automated reply:

Dear Customer,

Thank you for your enquiry.

We are currently experiencing a large volume of email traffic and will deal with your request within the next 15 working days.

We are pleased to advise that permission for the majority of our journal content, and for an increasing number of book publications, may be cleared more quickly by using the RightsLink service via Wiley’s websites http://onlinelibrary.wiley.com and www.wiley.com.

Within the next fifteen working days? That is, in the next three weeks? How can it possibly take that long? Are they engraving their response on a corundum block?

So, OK, let’s follow the automated suggestion and try RightsLink. I went to the Wiley Online Library, and searched for journals whose names contain “naturkunde”. Only one comes up, and it’s not the right one. So Wiley doesn’t admit the existence of the journal.

Despite this, Google finds the article easily enough with a simple title search. From the article’s page, I can just click on the “Request Permissions”  link on the right, and …

rightslink-fail

Well, there’s lots to enjoy here, isn’t there? First, and most important, it doesn’t actually work: “Permission to reproduce this content cannot be granted via the RightsLink service.” Then there’s that cute little registered-trademark symbol “®” on the name RightsLink, because it’s important to remind me not to accidentally set up my own rights-management service with the same name. In the same vein, there’s the “Copyright © 2015 Copyright Clearance Center, Inc. All Rights Reserved” notice at the bottom — copyright not on the content that I want to reuse, but on the RightsLink popup itself. (Which I guess means I am in violation for including the screenshot above.) Oh, and there’s the misrendering of “Museum für Naturkunde” as “Museum für Naturkunde”.

All of this gets me precisely nowhere. As far as I can tell, my only recourse now is to wait three weeks for Wiley to get in touch with me, and hope that they turn out to be in favour of science.

sadness_____by_aoao2-d430zrm

It’s Sunday afternoon. I could be watching Ireland play France in the Rugby World Cup. I could be out at Staverton, seeing (and hearing) the world’s last flying Avro Vulcan overfly Gloucester Airport for the last time. I could be watching Return of the Jedi with the boys, in preparation for the forthcoming Episode VII. Instead, here I am, wrestling with copyright.

How absolutely pointless. What a terrible waste of my life.

Is this what we want researchers to be spending their time on?

Promoting the Progress of Science and useful Arts, indeed.

Update (13 October 2015): a happy outcome (this time)

I was delighted, on logging in this morning, to find I had email from RIGHTS-and-LICENCES@wiley-vch.de with the subject “Permission to reproduce Heinrich (1999:fig. 16) under CC By licence” — a full thirteen working days earlier than expected. They were apologetic and helpful. Here is key part of what they said:

We are of course happy to handle your request directly from our office – please find the requested permission here:
We hereby grant permission for the requested use expected that due credit is given to the original source.
If material appears within our work with credit to another source, authorisation from that source must be obtained.
Credit must include the following components:
– Journals: Author(s) Name(s): Title of the Article. Name of the Journal. Publication  year. Volume. Page(s). Copyright Wiley-VCH Verlag GmbH & Co. KGaA. Reproduced with permission.

So this is excellent. I would of course have included all those elements in the attribution anyway, with the exception that it might not have occurred to me to state who the copyright holder is. But there is no reason to object to that.

So, two cheers for Wiley on this occasion. I had to waste some time, but at least none of it was due to deliberate obstructiveness, and most importantly they are happy for their figure to be reproduced under CC By.

References

  • Heinrich, Wolf-Dieter. 1999. The taphonomy of dinosaurs from the Upper Jurassic of Tendaguru, Tanzania (East Africa), based on field sketches of the German Tendaguru expedition (1909-1913). Mitteilungen aus dem Museum fur Naturkunde in Berlin, Geowissenschaftliche Reihe 2:25-61.

Here I am at SVPCA in 2015. I am haunted by the fact that ten years ago at SVPCA 2005, I gave a talk about the NHM’s Tendaguru brachiosaurid, NHMUK R5937. And the description is still not done and submitted a full decade later. Even though it’s objectively one of the most beautiful specimens in the world:

dorsals-ab-composite

So here is my pledge to the world:

By this time next year (i.e. the start of SVPCA 2016 in Liverpool), I will have written and submitted this description. If I fail, I give you all permission — no, I beg you — to mock me mercilessly. Leave mocking comments on this blog, yes; but more than that, those of you at SVPCA are invited to spend the week pointing contemptuously at me and saying “Ha!”

Let’s hope it doesn’t come to that.

Update (6 September): see also.

A couple of months ago, Darren (the silent partner in the SV-POW! organisation) tweeted this photo …

felch-quarry-brachiosaur-skull

… describing it as “Skull of the Morrison Formation Brachiosaurus at Denver Museum of Nature & Science”.

Well.

As Darren knows well (but didn’t have have space to explain in the tweet), it’s not quite as simple as that. What follows is adapted from Taylor 2009:789.

In 1883, a large sauropod skull (81 cm in length) was found in Felch Quarry 1, Garden Park, Colorado. It was shipped to O. C. Marsh in Yale that year and an illustration of the skull was used in his second attempt at reconstructing the skeleton of Brontosaurus (Marsh, 1891: plate 16).

Marsh's second attempt at reconstructing the skeleton of Brontosaurus, based primary on the holotype YPM 1980, using a skull based on the Felch Quarry specimen. From Marsh (1891:plate XVI)

Marsh’s second attempt at reconstructing the skeleton of Brontosaurus, based primary on the holotype YPM 1980, using a skull based on the Felch Quarry specimen. From Marsh (1891:plate XVI)

And here’s that skull in close-up:

Marsh1891-plateXVI-Apatosaurus-skull-UNREVERSED

This is often described as a “Camarasaurus-type” skull, but it’s not, really. It’s too long and low, and not stupid and ugly enough, to be Camarasaurus.

As we described in a previous post, this skull was also apparently the inspiration for the horrible, horrible sculpted skull that was originally used on the mounted Brontosaurus. (And let me reiterate my praise of the Yale museum for displaying this important historic object in their gallery instead of hiding it away.)

Anyway, the Felch Quarry skull was subsequently transferred to the National Museum of Natural History, where it was accessioned as USNM 5730. McIntosh and Berman (1975:195-198) recognized that whatever the skull was, it wasn’t Brontosaurus, but chickened out a bit by describing it as being “of the general Camarasaurus type” (p. 196). But McIntosh subsequently identified the skull tentatively as Brachiosaurus (Carpenter and Tidwell, 1998:70) and it was later described by Carpenter and Tidwell (1998), who considered it intermediate between the skulls of Camarasaurus and Giraffatitan, and referred it to Brachiosaurus sp.

The skull may be that of Brachiosaurus altithorax, but this is currently impossible to test due to the lack of comparable parts. Near this skull was a 99 cm cervical vertebra, probably of Brachiosaurus, but this was destroyed during attempts to collect it (McIntosh and Berman, 1975:196). Shame there are no photos.

References

  • Carpenter, Kenneth, and Virginia Tidwell. 1998. Preliminary description of a Brachiosaurus skull from Felch Quarry 1, Garden Park, Colorado. Modern Geology 23:69-84.
  • Marsh, Othniel Charles. 1891. Restoration of Triceratops. American Journal of Science, Series 3, 41:339-342.
  • McIntosh, John S., and David S. Berman. 1975. Description of the palate and lower jaw of the sauropod dinosaur Diplodocus (Reptilia: Saurischia) with remarks on the nature of the skull of Apatosaurus. Journal of Paleontology 49:187-199.
  • Taylor, Michael P. 2009. A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensch 1914). Journal of Vertebrate Paleontology 29(3):787-806.

Look on my works, ye mighty, and despair!

DSCN0476

[Giraffatitan brancai paralectotype MB.R.2181 (formerly HMN S II), mounted skeleton in left anteroventrolateral view. Presacral vertebrae sculpted, skull scaled and 3d-printed from specimen T1. Round the decay of that colossal wreck, boundless and bare, the lone and level sands stretch far away.]

I’ve been taking a long-overdue look at some of the recently-described giant sauropods from China, trying to sort out just how big they were. Not a new pursuit for me, just one I hadn’t been back to in a while. Also, I’m not trying to debunk anything about this animal – as far as I know, there was no bunk to begin with – I’m just trying to get a handle on how big it might have been, for my own obscure purposes.

‘Huanghetitan’ ruyangensis was named by Lu et al. (2007) on the basis of a sacrum, the first 10 caudal vertebrae, some dorsal ribs and haemal arches, and a partial ischium. The holotype is 41HIII-0001 in the Henan Geological Museum. Lu et al. (2007) referred the new animal to the genus Huanghetitan, which was already known from the type species H. liujiaxiaensis (You et al., 2006). However, Mannion et al. (2013) found that the two species are not sister taxa and therefore ‘H.’ ruyangensis probably belongs to another genus, which has yet to be erected. Hence my use of scare quotes around the genus name.

Huanghetitan ruyangensis sacrum comparison

Here’s the sacrum of ‘H.’ ruyangensis from Lu et al. (2007: fig. 2). The original small scale bar is supposed to be 10cm. You know how I feel about scale bars (or maybe you don’t, in which case read this and this), but in this case the scale seems pretty legit based on limited measurements that are also given in the paper. I comped in the sacrum of Brachiosaurus altithorax FMNH P25107 from this post (many thanks to Phil Mannion for the photos!), and scaled it according to the max width across the second pair of sacral ribs, which Riggs (1904: p. 236) gives as 105 cm. The sacrum of ‘H.’ ruyangensis is a little bigger, but not vastly bigger. ‘H.’ ruyangensis had six sacrals to Brachiosaurus‘s five, so extra length is mostly illusory, whereas the extra width is mostly legit.

According to Lu et al. (2007), the anterior face of the first caudal vertebra in ‘H.’ ruyangensis measures 26.9 cm tall by 32 cm wide, and the centrum is 18.2 cm long. The same measurements in Brachiosaurus are 28 x 33 cm for the anterior face and 16 cm for the centrum length. It’s basically a tie.

What about the big rib? Lu et al. (2007) show a complete dorsal rib of ‘H.’ ruyangensis that is 293 cm long. That’s nothing to sniff at – the longest rib of Brachiosaurus, and the cause for the specific name altithorax (‘tall-bodied’), measures 274.5 cm, so the ‘H.’ ruyangensis rib is about 7% longer. But it’s not the longest rib known for any sauropod. As far as I know, that honor goes to a Supersaurus dorsal rib measuring 305 cm (Lovelace et al., 2008). The biggest Supersaurus caudal also blows away the caudals of both ‘H.’ ruyangensis and Brachiosaurus, with a anterior face 39 cm tall by 46 cm wide. But then diplodocids were all about that bass, so there’s not much point in comparing tail size with a titanosauriform if you’re trying to get a handle on overall body size. Still, the 35-40 ton Supersaurus shows that you can have 3-meter ribs without being anywhere near Argentinosaurus territory, mass-wise.

So what’s the verdict? ‘H.’ ruyangensis was a little bigger than the holotype of Brachiosaurus altithorax, but only by a few percent. It might have been about the same size as the XV2 specimen of Giraffatitan brancai. Or, who knows, it could have had completely different proportions and massed considerably more (or less). But on the current evidence, it doesn’t seem to have been one of the biggest sauropods of all time. I hope we get some more of it one of these days.

References

 

Introduction and Background

2005-09-27 CM 555 c6 480

An epipophysis in a neural arch of a juvenile Apatosaurus, CM 555. From this post.

I have three goals with this post:

  1. To document the range of variation in epipophyses in the cervical vertebrae of sauropods.
  2. To show that the “finger-like processes” overhanging the cervical postzygapophyses in the newly described Qijianglong are not novel or mysterious structures, just very well developed epipophyses.
  3. Finally, to show that similar long, overhanging epipophyses are present in other mamenchisaurids, although as far as I can tell no-one has noted them previously.

Epipophyses are muscle attachment points dorsal to the postzygapophyses, for the insertion of long, multi-segment epaxial (dorsal) neck muscles in birds and other dinosaurs. I know that they turn up occasionally in non-dinosaurian archosaurs, and possibly in other amniotes, but for the purposes of this post I’m only considering their distribution in sauropods. For some quick background info on epipophyses and the muscles that attach to them, see the second half of this post, and see Wedel and Sanders (2002) and Taylor and Wedel (2013a) for further discussion and more pictures.

OMNH emu vert 480

Before we start with the pictures, a fiddly nomenclatural point: this muscle attachment point dorsal to the postzyg has traded under at least six names to date.

  1. The ‘Owenian’ term, used by virtually all non-avian theropod workers, by Sereno et al. (1999) for Jobaria, and probably by loads of other sauropod workers (including myself, lately) is epipophysis.
  2. Beddard (1898) referred to this feature in birds as the hyperapophysis; this term seems to have fallen completely out of use.
  3. Boas (1929), again referring to birds, called it the processus dorsalis. Zweers et al. (1987: page 138 and table 1) followed this terminology, which is how I learned of it when I was an undergrad at OU.
  4. Baumel and Witmer (1993) called this feature in birds the torus dorsalis (note 125 on page 87), which some authors have informalized to dorsal torus (e.g., Harris 2004: page 1243 and fig. 1). Baumel and Witmer (1993: page 87) note that, “the use of ‘Torus’ is preferable since it avoids confusion with the spinous [dorsal] process of the neural arch”.
  5. In my own early papers (e.g., Wedel et al. 2000b) and blog posts I called this feature the dorsal tubercle, which was my own attempt at an informal term matching ‘processus dorsalis’ or ‘torus dorsalis’. That was unfortunate, since there are already several other anatomical features in vertebrates that go by the same name, including the dorsal-facing bump on the dorsal arch of the atlas in many vertebrates, and a bump on the humerus in birds and some other taxa. In more recent papers (e.g., Taylor and Wedel 2013a) I’ve switched over to ‘epipophysis’.
  6. In the last post, Mike coined the term parapostzygapophysis for this feature in Qijianglong. [Note: he now regrets this.]

As usual, if you know of more terms for this feature, or additional history on the ones listed above, please let us know in the comments.

Now, on to the survey.

Prosauropods

Leonerasaurus_cervical_vertebrae - Pol et al 2011 fig 5

I haven’t seen very many prominent epipophyses in basal sauropodomorphs. Probably the best are these in the near-sauropod Leonerasaurus, which is very sauropod-like in other ways as well. Modifed from Pol et al. (2011: fig. 5).

This combination of photograph and interpretive drawing neatly shows why it’s often difficult to spot epipophyses in photos: unless you can make out the postzygapophyseal facet, which is often located more anteriorly than you might guess, you can’t tell when the epipophysis projects further posteriorly, as in the last of these vertebrae. In this case you can make it out, but only because the interpretive drawing shows the facet much more clearly than the photo.

Basal sauropods

Tazoudasaurus cervical - Allain and Aquesbi 2008 fig 9i-j

The most basal sauropod in which I have seen clear evidence of epipophyses is Tazoudasaurus. They’re not very apparent in lateral view, but in posterior view the epipophyses are clearly visible as bumps in the spinopostzygapophyeal laminae (SPOLs). Modified from Allain and Aquesbi (2008: fig. 9).

Jobaria epipophyes

In addition to Qijianglong, some other basal eusauropods have prominent epipophyses. Probably the best known is Jobaria; Sereno et al. (1999: fig. 3) figured and labeled the epipophysis in one of the cervical vertebrae. The vertebra image in that figure is tiny (nice work, glam-magz!), so here are some sketches of Jobaria mid-cervicals (from two different individuals) that I made back in the day when I was doing the research for Gary Staab’s Jobaria neck sculpture (see Sanders et al. 2000 for our SVP abstract about that project).

Turiasaurus also has prominent, overhanging epipophyses in at least some of its cervical vertebrae. You can just make one out as a tiny spike a few pixels long in Royo-Torres et al. (2006: fig. 1K). I have seen that cervical firsthand and I can confirm that the epipophyses in Turiasaurus are virtually identical to those in Jobaria.

Other mamenchisaurids

It’s not air-tight, but there is suggestive evidence of projecting epipophyses in some other mamenchisaurids besides Qijianglong.

Mamenchisaurus epipophyses - lateral view

If you’re really hardcore, you may remember that back in 2005, Mike got to go up on a lift at the Field Museum of Natural History to get acquainted with a cast skeleton of Mamenchisaurus hochuanensis that was mounted there temporarily. During that adventure he took some photos that seem to show projecting epipophyses in at least two of the mid-cervicals. At least, if they’re not epipophyses, I don’t know what they might be.

Mamenchisaurus epipophyses - medial view

Here they are again in medial view. My only reservation is that these vertebrae were distorted to begin with, and some features of the cast are very difficult to interpret. So, probably epipophyses, but it would be nice to check the original material at some point.

Mamenchisaurus youngi epipophyses

Something similar may be present in some posterior cervical vertebrae of Mamenchisaurus youngi. Here’s Figure 17 from Ouyang and Ye (2002). The “poz” label does not not seem to be pointing to the articular facet of the postzygapophysis, which looks to be a little more anterior and ventral, below the margin of the PODL. If that’s the case, then C15 has long, overhanging epipophyses like those of Jobaria. C16 has a more conservative bump, which is to be expected – the epipophyses typically disappear through the cervico-dorsal transition.

Omeisaurus epipophysis

Finally, here’s a cervical vertebra of Omeisaurus junghsiensis from Young (1939: fig. 2). I don’t want to hang very much on just a few pixels, but my best guess at the extent of the postzygapophyseal articular facet is shown in the interpretation above. If that’s correct, then this specimen of Omeisaurus had really long epipophyses, rivaling those of Qijianglong. Unfortunately that’s impossible to check, because this specimen has been lost (pers. comm. from Dave Hone, cited in Taylor and Wedel 2013).

Diplodocoidea

Haplocanthosaurus epipophyses - Hatcher 1903

Haplocanthosaurus nicely shows that the epipophyses can be large in terms of potential muscle attachment area without projecting beyond the posterior margins of the postzygapophyses. Here is C14 of H. priscus, CM 572, in posterior and lateral views, modified from Hatcher (1903: plate 1).

diplodocid epipophyses

Epipophyses that actually overhang the postzygapophyses are not common in Diplodocidae but they do occasionally occur. Here are prominent, spike-like epipophyses in Diplodocus (upper left, from Hatcher 1901: plate 3), Barosaurus (upper right), Kaatedocus (lower left, Tschopp and Mateus 2012: fig. 10), and Leinkupal (lower right, Gallina et al. 2014: fig. 1).

NIgersaurus cervical - Sereno et al 2007 fig 3

Of course, the champion epiphysis-bearer among diplodocoids is the weird little rebbachisaurid Nigersaurus. Here’s a Nigersaurus mid-cervical, from Sereno et al. (2007: fig. 3). Note that the projecting portions of the epipophysis is roughly as long as the articular surface of the postzygapophysis.

Macronaria

Australodocus epipophysis

The epipophysis in this cervical of Australodocus just barely projects beyond the posterior margin of the postzygapophysis.

Giraffatitan c8 epipophyses

In Giraffatitan, epipophyses are absent or small in anterior cervicals but they are prominent in C6-C8. Here’s a posterolateral view of C8, showing very large epipophyses that are elevated several centimeters above the postzygapophyses. You can also see clearly in this view that the spinopostzygapophyseal lamina (SPOL) and postzygodiapophyseal lamina (PODL) converge at the epipophysis, not the postzygapophysis itself.

Sauroposeidon epipophyses

The holotype of Sauroposeidon, OMNH 53062, is similar to Giraffatitan in that the two anterior cervical vertebrae (possibly C5 and C6) have no visible epipophyses, but epipophyses are prominent in the two more posterior vertebrae (possibly C7 and C8). Click to enlarge – I traced the articular facet of the postzygapophysis in ?C8 to more clearly separate it from the epipophysis. For a high resolution photograph of that same vertebra that clearly shows the postzyg facet and the epipophysis dorsal to it, see this post.

Oddly enough, I’ve never seen prominent epipophyses in a titanosaur. In Malawisaurus, Trigonosaurus, Futalognkosaurus, Rapetosaurus, Alamosaurus, and Saltasaurus, the SPOLs (such as they are – inflated-looking titanosaur cervicals do not have the same crisply-defined laminae seen in most other sauropods) merge into the postzygapophyseal rami and there are no bumps sticking up above or out beyond the articular facets of the postzygs. I don’t know what to make of that, except to note that several of the animals just mentioned have mediolaterally wide, almost balloon-shaped cervical neural spines. In our 2013 PeerJ paper, Mike and I argued that the combination of tall neural spines and tall epipophyses in the cervical vertebrae of sauropods made them functionally intermediate between crocs (huge neural spines, no epipophyses) and birds (small or nearly nonexistent neural spines, big epipophyses). Perhaps most titanosaurs reverted to a more croc-like arrangement with most of the long epaxial neck muscles inserting on the neural spine instead of the postzygapophyseal ramus. I’ve never seen that possibility discussed anywhere, nor the apparent absence of epipophyses in most titanosaurs. As usual, if you know otherwise, please let me know in the comments!

malawisaurus-cervicals

Cervical vertebrae of Malawisaurus from Gomani (2005: fig. 9): not an epipophysis in sight. But check out the spike-like neural spines – these are so wide from side to side that from the front they look like party balloons.

And as long as we’re discussing the phylogenetic distribution of epipophyses, it is interesting that long, overhanging epipophyses are so broadly but sporadically distributed. They turn up in some non-neosauropods (Jobaria, Turiasaurus, Omeisaurus) and some diplodocoids (Nigersaurus, the occasional vertebra in Diplodocus and Leinkupal), but not in all members of either assemblage, and they seem to be absent in Macronaria (although many non-titanosaurs have shorter epipophyses that don’t overhang the postzygs). I strongly suspect that a lot of this is actually individual variation that we’re not perceiving as such because our sample sizes of almost all sauropods are tiny, usually just one individual. Epipophyses are definitely muscle attachment sites in birds and no better hypothesis has been advanced to explain their presence in other archosaurs. Muscle attachment scars are notoriously variable in terms of their relative development and expression among individuals, and it would be odd if epipophyses were somehow exempt from that inherent variability.

It also seems more than likely that ontogeny plays a role: progressive ossification of tendons attached at the epipophyses would have the effect of elongating the preserved projection. And since for some aspects of sauropod vertebral morphology, serial position recapitulates ontogeny (Wedel and Taylor 2013b), it shouldn’t be surprising that we see differences in the prominence of the epipophyses along the neck.

Back to Qijianglong

By now it should be clear that the “finger-like processes” in Qijianglong are indeed epipophyses, and although they are quite long, they aren’t fundamentally different from what we see in many other sauropods. I haven’t gone to the trouble, but one could line up all of the vertebrae figured above in terms of epipophysis size or length, and Qijianglong would sit comfortably at one end with Omeisaurus and Mamenchisaurus, just beyond Nigersaurus and Jobaria.

FIGURE 11. Anterior cervical series of Qijianglong guokr (QJGPM 1001) in left lateral views unless otherwise noted. A, axis; B, cervical vertebra 3; C, cervical vertebra 4; D, cervical vertebrae 5 and 6; E, cervical vertebra 7 and anterior half of cervical vertebra 8 (horizontally inverted; showing right side); F, posterior half of cervical vertebra 8 and cervical vertebra 9; G, cervical vertebra 10; H, cervical vertebra 11; I, close-up of the prezygapophy- sis-postzygapophysis contact between cervical vertebrae 3 and 4 in dorsolateral view, showing finger-like process lateral to postzygapophysis; J, close- up of the postzygapophysis of cervical vertebra 5 in dorsal view, showing finger-like process lateral to postzygapophysis. Arrow with number indicates a character diagnostic to this taxon (number refers to the list of characters in the Diagnosis). All scale bars equal 5 cm. Abbreviations: acdl, anterior centrodiapophyseal lamina; cdf, centrodiapophyseal fossa; plc, pleurocoel; pocdl, postcentrodiapophyseal lamina; poz, postzygapophysis; pozcdf, post- zygapophyseal centrodiapophyseal fossa; pozdl, postzygodiapophyseal lamina; ppoz, finger-like process lateral to postzygapophysis; ppozc, groove for contact with finger-like process; przdl, prezygodiapophyseal lamina; sdf, spinodiapophyseal fossa.

Cervical vertebrae of Qijianglong (Xing et al. 2015: fig. 11)

The strangest thing about the epipophyses in Qijianglong is that they seem to be bent or broken downward in two of the vertebrae (B and H in the figure above). I assume that’s just taphonomic distortion – the cervical shown in H wouldn’t even be able to articulate with the vertebra behind it if the epipophysis really drooped down like that. The epipophyses in Qijianglong seem to mostly manifest as thin spikes of bone (or maybe plates, as shown in B and I), so it’s not surprising that they would get distorted – most of the vertebrae shown above have cervical ribs that are incomplete or missing as well.

One more noodle-y thought about big epipophyses. I wrote in the last section that I’ve never seen them in titanosaurs, possibly because titanosaurs have big neural spines for their epaxial muscles to attach to. Maybe long, overhanging epipophyses are so common in mamenchisaurids because their neural spines are so small and low. Although we tend to think of them as a basal group somewhat removed from the “big show” in sauropod evolution – the neosauropods – mamenchisaurids did a lot of weird stuff. At least in terms of their neck muscles, they may have been the most birdlike of all sauropods. Food for thought.

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