I have long intended to write a paper entitled Why Elephants Are So Small, as a companion piece to Why Giraffes Have Short Necks (Taylor and Wedel 2013). I’ve often discussed this project with Matt, usually under the acronym WEASS, and its substance has come up in the previous post, and especially Mickey Mortimer’s comment:

I think it would be interesting to read a study on that — the order in which various factors restrict body size without transformative adaptations. Similarly, what the differences would be for an aquatic animal like a whale.

That is exactly what the WEASS project was supposed to consist of: a list of many candidate limitations on how big animals can get, some rough attempt to quantify their Big-O behaviour, some discussion of which factors seems to limit the sizes of modern terrestrial animals, and how dinosaurs (especially sauropods) worked around those limitations.

(Whales are different. I have in my mind a half-formed notion for a third paper, completing the trilogy, with a title along the lines of Why Whales Are Dirty Cheaters.)

What are those candidate limitations? Off the top of my head:


  • Bone strength
  • Cartilage strength
  • Cartilage thickness
  • Muscle strength
  • Nerve length and conduction time


  • Blood pressure: column height and capillary length
  • Lung capacity
  • Tracheal dead space
  • Digestive efficiency
  • Metabolic overheating

Those are just some of the physical limits. There is anecdotal evidence that elephants are not very close to their mechanical limits in their usual behaviour: they could get bigger, and still work mechanically. (Follow the link at the start of this paragraph. You will thank me.)

There are plenty of other factors that potentially limit organism size, including:


  • Feeding rate
  • Ability to navigate dense environments
  • Predator avoidance with limited athleticism
  • Difficulties in mating


  • Territory requirement
  • Time taken to reach reproductive maturity
  • Reproductive rate
  • Birth size
  • Lack of selection pressure: when there are no predators bigger than a lion, why would elephants need to evolve larger size?

I’m sure I am missing loads. Help me out!

I am haunted by something Matt wrote a while back when we were discussing this — talking about how alien sauropods are, and how easily we slip into assuming mammal-like paradigms.

We are badly hampered by the fact that all of the 250kg+ land animals are mammals. We only get to see one way of being big, and it’s obviously not the best way of being big. Our perceptions of how hard it is to be big are shaped by the animals that are bad at it.

So having written this blog post, I am wondering whether it’s time to breathe life back into this project, started in 2009 and repeatedly abandoned.

For this forthcoming Barosaurus paper, we would like to include an establishing photo of the AMNH Barosaurus mount. There are two strong candidate photos which we’ve used before in an SVPCA talk, but since this is a formal publication we need to be more careful about copyright. Here are the photos, which are the property of their respective rightsholders:

This one is hard to find at all, at least using Google’s reverse image search. Whereas this one …

… is sprinkled all over the Internet, but (in all the cases I’ve seen so far) without attribution.

Does anyone have the necessary skills to track down who the rightsholder is for either of these? Thank you!

Matt and I are writing a paper about Barosaurus cervicals (yes, again). Regular readers will recall that the best Barosaurus cervical material we have ever seen was in a prep lab for Western Paleo Labs. We have some pretty good photos, such as this one:

Barosaurus cervical vertebra lying on its right side in anterodorsal view (i.e. with dorsal to the left), showing the distinctive shape of the prezygapophyseal rami.

The problem is that this specimen was privately owned at the time we saw it, and so far as we know it still is. So according to all standard procedures, we should consider it unavailable to science until such time as it is deposited in an accredited museum. (I was pretty sure the SVP has an explicit policy to this effect, but I couldn’t find it on the site. Can anyone?)

So what should we do? All the possible courses of action seem unfortunate.

1. We could go ahead and include photos, drawings and descriptions of these vertebrae in the paper — but that would violate community norms by building an argument on observations that cannot be in general be replicated by other researchers. (For all we know, these vertebrae are now decorating Nicolas Cage’s pool room.)

2. We could omit these vertebrae from the paper, but use the information we gained from examining them in formulating our diagnostic criteria for Barosaurus cervicals — but this would also not really be replicable, plus it would have that horrible “we know something that you don’t” quality.

3. We could act as though these vertebrae do not exist, or as though we had never seen them, writing the paper based only on our observations of inferior material and of the good AMNH 6341 that is not accessible for study or photography — but that would make our characterisation of Barosaurus cervical morphology less helpful than it could be.

4. We could refrain from publishing on Barosaurus cervicals at all until such time as these vertebrae, or similarly well-preserved ones, are available to study at accredited institutions — but that would simply deprive the world of an interesting and exciting study.

Is there a fifth path that we have not seen? And if not, which of these four is the least objectionable?

I was looking more closely at the turkey skeleton from my recent post, and zeroed in on the last two dorsal (= thoracic) vertebrae. They articulate very well with each other and with the first vertebra of the sacrum, with the centra and zygapophyses both locking in so that there can only have been very little if any movement between them in life. Here they are, in right lateral view:

Last two dorsal (= thoracic) vertebrae of a mature domestic turkey Meleagris gallopavo domesticus, in right lateral view.

Before we move on, it’s worth clicking through to the full-size version of this image and wondering at both the quality of modern phone cameras (a Pixel 3a in this case) and the variety of textures on these little bones. There is smooth, finished bone on the sides of the neural spines; very fine pits and bumps on the zygapophyseal facets where the thin layer of hyaline cartilage attached; rougher texture in the parapophyseal facets where thicker cartilage attached; and very rough texture on the ends of the transverse processes, where there was relatively thick cartilage.

And there is, unsurprisingly in a bird, pneumaticity everywhere. In the more anterior vertebra alone (to the right) the photo shows pneumatic openings (from bottom to top) low on the centrum (below the parapophysis), high on the centrum (below the lateral process),  in the hollow between the lateral process, the posyzyg and the centrum, on the lateral surface of the prezygapophyseal ramus, and on the rear surface of the lateral process. There are others that are obscured in this photo, including on top of the lateral process where it meets the neural spine. Here they are, pointed out for you (with the hidden one shown translucently):

Last two dorsal (= thoracic) vertebrae of a mature domestic turkey Meleagris gallopavo domesticus, in right lateral view. Pneumatic openings on penultimate vertebra highlighted with red lines; obscured opening above lateral process shown as translucent.

OK, that was the B-movie. Now to the main feature. The next photo shows the same two vertebrae, folded away from each other so that we see the anterior face of the posterior vertebra (on the left) and the posterior face of the anterior vertebra (on the right).

Last two dorsal (= thoracic) vertebrae of a mature domestic turkey Meleagris gallopavo domesticus. Left: last dorsal vertebra in anterior view; right: penultimate dorsal vertebra in posterior view.

Again, do click through to see the exquisite detail, especially the complex of pneumatic features on the anterior face of the neural spine of the last dorsal (on the left) and on the posterior face of the left lateral process of the penultimate dorsal (on the right).

And … in the articular facets of the centra?

Seriously, what the heck is going on here? It doesn’t make sense  that there would be pneumatic openings in articular surfaces, because by definition something else (in this case the adjacent vertebra) is abutted hard up against then, so there is no way for a diverticulum to get in. For the same reason, you don’t get vascular foramina in articular surfaces because there is no way to get an artery in there. And there is no hint in these vertebrae of channels along either articular surface that diverticula or arteries could  possibly have laid in.

And yet, there those big openings are. What are they?

I discussed this with Matt, in case it’s Well Known Phenomenon that I’d somehow not heard about but it seems it is not. What we know for sure is that these openings are present, and that they are not mechanical damage inflicted during preparation. So what are they?

What else even is there for them to be? What penetrates bone apart from diverticula and blood vessels? Nerves follow the blood vessels, so it can’t be nerves in the absence of blood vessels.

By the way, there are similar but smaller openings in the posterior face of the last dorsal (the one on the right in the photo), but none anywhere else along the postcervical column: not on the anterior surface of the penultimate dorsal, not on the front or back of the sacrum, and not in any of the other dorsals.

One possibility we considered is that the vertebrae were locked together in life and that a pneumatic space inside the centrum of the last dorsal worked right through into the penultimate one. But that doesn’t work: the openings are not aligned. Also, those in the penultimate dorsal are definitely blind (i.e. they do not connect to deeper internal air-spaces) and those in the last dorsal probably are, too.

We do not know what is going on here.

Help us! Is this kind of thing common in turkeys? Have people seen it in other taxa? Do we know what it is?

In mammals — certainly the most-studied vertebrates — regional differentiation of the vertebral column is distinct and easy to spot. But things aren’t so simple with sauropods. We all know that the neck of any tetrapod is made up of cervical vertebrae, and that the trunk is made up of dorsal vertebrae (subdivided into thoracic and lumbar vertebrae in the case of mammals). But how do we tell whether a given verebra is a posterior cervical or an anterior dorsal?

Here two vertabrae: a dorsal vertebra (D3) and a cervical vertebra (C13) from CM 84, the holotype of Diplodocus carnegii, modified from Hatcher (1901: plates III and VII):

It’s easy to tell these apart, even when as here we have only lateral-view images: the dorsal vertebra is tall, its centrum is short, its neural spine is anteroposteriorly compressed and its parapophysis is up on the dorsal half of the centrum; but the cervical vertebra is relatively low, its centrum is elongated, its neural spine is roughly triangular and its parapophysis hangs down well below the centrum (and has a cervical rib fused to it and the diapophysis).

But things get trickier in the shoulder region because, in sauropods at least, the transition through the last few cervicals to the first few dorsals is gradual — the vertebrae become shorter, taller and broader — and tends to have no very obvious break point. In this respect, they differ from mammals, in which the regional differentiation of the spinal column is more abrupt. (Although even here, things may not be as simple as generally assumed: for example, Gunji and Endo (2016) argued that the 1st thoracic vertebra of the giraffe behaves functionally like an 8th cervical.)

So here are those two vertebrae in context: the sequence D3 D2 D1 C15 C14 C13 in CM 84, the holotype of Diplodocus carnegii, modified from Hatcher (1901: plates III and VII):

Given that the leftmost is obviously a dorsal and the rightmost obviously a cervical, where would you place the break-point?

The most usual definition seems to be that the first dorsal vertebra is the first one that has a free rib, i.e. one not fused to the vertebra: in the illustration above, you can see that the three cervicals on the right all have their cervical ribs fused to their diapophyses and parapophyses, and the three dorsals on the left do not. This definition of the cervical/dorsal distinction seems to be widely assumed, but it is rarely explicitly asserted. (Does anyone know of a paper that lays it out for sauropods, or for dinosaurs more generally?)

But wait!

Hatcher (1903:8) — the same dude — in his Haplocanthosaurus monograph, writes:

The First Dorsal (Plate I., Fig. 1). […] That the vertebra now under consideration was a dorsal is conclusively shown not by the presence of tubercular and capitular rib facets showing that it supported on either side a free rib, for there are in our collections of sauropods, skeletons of other dinosaurs fully adult but, with the posterior cervical, bearing free cervical ribs articulating by both tubercular and capitular facets as do the ribs of the dorsal region. The character in this vertebra distinguishing it as a dorsal is the broadly expanded external border of the anterior branch of the horizontal lamina [i.e. what we would now call the centroprezygapophyseal lamina]. This element has been this modified in this and the succeeding dorsal, no doubt, as is known to be the case in Diplodocus to give greater surface for the attachment of the powerful muscles necessary for the support of the scapula.

Hatcher’s illustrations show this feature, though they don’t make it particularly obvious: here are the last two cervicals and the first dorsal, modified from Hatcher (1903:plate I), with the facet in question highlighted in pink: right lateral view at the top, then anterior, and finally posterior view at the bottom. (The facet is only visible in lateral and anterior views):

Taken at face value, Hatcher’s words here seem to imply that he considers the torso to begin where the scapula first lies alongside the vertebral column. Yet if you go back to the Diplodocus transition earlier in this post, a similar scapular facet is not apparent in the vertebra that he designated D1, and seems to be present only in D2.

Is this scapular-orientation based definition a widespread usage? Can anyone point me to other papers that use it?

Wilson (2002:226) mentions a genetic definition of the cervical/dorsal distinction

Vertebral segment identity may be controlled by a single Hox gene. The cervicodorsal transition in many tetrapods, for instance, appears to be defined by the expression boundary of the Hoxc-6 gene.

But this of course is no use in the case of extinct animals such as sauropods.

So what’s going on here? In 1964, United States Supreme Court Justice Potter Stewart, in describing his threshold test for obscenity, famously said “I shall not today attempt further to define the kinds of material I understand to be embraced within that shorthand description, and perhaps I could never succeed in intelligibly doing so. But I know it when I see it.” Is that all we have for the definition of what makes a vertebra cervicals as opposed to dorsal? We know it when we see it?

Help me out, folks! What should the test for cervical-vs-dorsal be?


  • Gunji, Mego, and Hideki Endo. 2016. Functional cervicothoracic boundary modified by anatomical shifts in the neck of giraffes. Royal Society Open Science 3:150604. doi:10.1098/rsos.150604
  • Hatcher, Jonathan B. 1901. Diplodocus (Marsh): its osteology, taxonomy and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63 and plates I-XIII.
  • Hatcher, J. B. 1903b. 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 and plates I-VI.
  • Wilson, Jeffrey A. 2002. Sauropod dinosaur phylogeny: critique and cladistic analysis. Zoological Journal of the Linnean Society 136:217-276.

I have several small ordered sequences of data, each of about five to ten elements. For each of them, I want to calculate a metric which captures how much they vary along the sequence. I don’t want standard deviation, or anything like it, because that would consider the sequences 1 5 2 7 4 and 1 2 4 5 7 equally variable, whereas for my purposes the first of these is much more variable.

Here is a matric that I think does what I want, and will allow me to compare different sequences for variability-along-the-sequence.

For the n-1 pairs along the sequence of n elements, I take the difference (absolute value, so always positive) between elements i and i+1. Then I average all those differences. Then I divide the result by the average of the values themselves, to normalise for magnitude.

Some example calculations:

  • For the sequence 1 5 2 7 4, the differences are 4 3 5 3, for a total of 15 and an average of 3.75. The average of the values is 1+5+2+7+4 = 19/5 = 3.8, which gives me a metric of 3.75/3.8 = 0.987.
  • For the sequence 1 2 4 5 7, the differences are 1 2 1 2, for a total of 6 and an average of 1.5. The average of the values is again 3.8, which gives me a metric of 1.5/3.8 = 0.395.
  • So the first sequence is 0.987/0.395 = 2.5 times as sequentially variable as the second sequence.
  • And for the sequence 10 20 40 50 70 (which is the same as the previous one, but all values ten times greater), the differences are 10 20 10 20, for a total of 60 and an average of 15. The average of the values is 38, which gives me a metric of 15/38 = 0.395, the same as before — which is as it should be.

And now, my question! Does this metric, or something similar, already exist? If so, what is it called? Or if I should be using something else instead, what is it?

(It happens that my sequences are the aspect ratios of the cotyles of consecutive vertebrae, but that’s not important: whatever metric we land on should work for any sequences.)

Taylor 2015: Figure 8. Cervical vertebrae 4 (left) and 6 (right) of Giraffatitan brancai lectotype MB.R.2180 (previously HMN SI), in posterior view. Note the dramatically different aspect ratios of their cotyles, indicating that extensive and unpredictable crushing has taken place. Photographs by author.

A couple of months ago, I asked for your help in compiling a list of all known complete sauropods necks. This has gone really well, and I want to thank everyone who chipped in, and all the various authors I have contacted for details as a result.

My next step is to take the raw data in the Google spreadsheet that I have been maintaining, and write it up as prose for the paper that I am shortly going to resubmit, having first done so back in 2015. And I thought it would make sense to draft that section here on SV-POW!, so I can get any further feedback before I finalize it for the manuscript.

Young and Zhao (1972:figure 3). Mamenchisaurus hochuanensis holotype CCG V 20401 as is occurred in the field.

So here goes: any additional comments at this stage will be welcome!

Unambiguously complete necks are known from published accounts of only a few sauropod specimens. In chronological order of description, the following specimens were found with their necks complete and articulated, and have been adequately described:

  • CM 11338, a referred specimen of Camarasaurus lentus described by Gilmore (1925). This is a juvenile specimen, and thus does not fully represent the adult morphology. (McIntosh et al. 1996:76 claim that this specimen is the holotype, but this is not correct: YPM 1910 is the holotype — see below.)
  • CM 3018, the holotype of Apatosaurus louisae, described by Gilmore (1936). The neck was separated from the torso but articulated from C1–C15, though the last three cervicals were badly crushed: see below for details.
  • CCG V 20401, the Mamenchisaurus hochuanensis holotype, described by Young and Zhao (1972). Each vertebra is broken in half at mid-length, with the posterior part of each adhering to the anterior part of the its successor; and all the vertebrae are badly crushed in an oblique plane.
  • ZDM T5402, a Shunosaurus lii referred specimen, described in Chinese by Zhang (1988), with English figure captions. Figure 22 depicts the atlas. Unlike the holotype T5401, this specimen is mature.
  • BYU 9047, the Cathetosaurus lewisi holotype, described by Jensen (1988). (Jensen incorrectly gives the specimen number as BYU 974.) This specimen was redescribed, and the species referred to Camarasaurus, by McIntosh et al. (1996). Although all 12 cervicals are present, “10–12, particularly 12, have suffered such severe damage that it is impossible to restore them” (McIntosh et al. 1996:76).
  • MACN-N 15, the holotype of Amargasaurus cazaui MACN-N 15, described by Salgado and Bonaparte (1991) who desribed “22 presacral vertebrae articulated with each other and attached to the skull and sacrum, relatively complete” (Salgado & Bonaparte 1991:335, translated.
  • ZDM 0083, the holotype of Mamenchisaurus youngi, described in Chinese by Ouyang and Ye (2002) with English figure captions. Figure 14 depicts the atlas and axis.
  • MUCPv-323, the holotype of Futalognkosaurus dukei, initially described by Calvo et al. 2007a and redescribed by Calvo et al. 2007b. The neck was found in two articulated sections which fit together without needing additional vertebrae in between (Jorge O. Calvo, pers. comm., 2021).
  • SSV12001, the holotype of Xinjiangtitan shanshanesis, described by Zhang et al. (2018). The original description of this specimen by Wu et al. 2013 included only the last two cervicals, which were the only ones that had been excavated at that time.

A few additional specimens are known to have complete and articulated necks, but have not yet been described:

  • USNM 13786, a referred subadult specimen of Camarasaurus lentus recently mounted at the Smithsonian. The specimen “was almost completely buried before the sinews had allowed the bones to separate” (letter from Earl Douglass to William J. Holland, 22 August 1918), and photographs kindly supplied by Andrew Moore show that the atlas was preserved.
  • MNBH TIG3, the holotype of Jobaria tiguidensis. Sereno et al. (1999:1343) assert that this species has 12 cervicals in all and say “One articulated neck was preserved in a fully dorsiflexed, C-shaped posture”. Sereno (pers. comm., 2021) confirms that the articulated neck is MNBH TIG3
  • SMA 002, referred to Camarasaurus sp. Tschopp et al. (2016), in a description of its feet, say that this specimen “lacks only the vomers, the splenial bones, the distal end of the tail, and one terminal phalanx of the right pes. The bones are preserved in three dimensions and in almost perfect articulation”.
  • MAU-Pv-LI-595, the “La Invernada” Titanosaur. Filippi et al. (2016) give a very brief account in an abstract. Filippi, pers. comm, 2021) says that the entire preserved specimen was articulated.
  • MAU-Pv-AC-01, an unnamed titanosaur mentioned in abstracts by Calvo et al. (1997) and Coria and Salgado (1999). The specimen was found in perfect articulation from skull down to the last caudal vertebrae (Rodolfo Coria, pers. comm., 2021).

The first cervical (the atlas) in sauropods is very different in form from the other vertebrae, and small and fragile. Consequently it is easily lost. Some further specimens have necks that are complete and articulated from C2 (the axis) backwards:

  • MB.R.4886, the holotype of Dicraeosaurus hansemanni, described by Janensch (1929), has a neck that complete and well preserved from C2 to C12 (the last cervical). Janensch referred to this as “specimen m” and writes “It was found articulated from the 19th caudal vertebra to the 9th cervical vertebra inclusive. The proximal part of the neck from the 8th cervical vertebra up to the axis was bent ventrally and lay at right angles to the distal part of the neck.” (Janensch 1929:41).
  • PMU 233, the holotype of Euhelopus zdanskyi, described by Wiman (1929) as “exemplar a” and redescribed by Wilson and Upchurch (2009).
  • ZDM T5401, the subadult holoype of Shunosaurus lii, described in Chinese by Zhang et al 1984. The quarry map (Zhang et al. 1984:figure 1) suggests that the atlas is missing.
  • MCT 1487-R, informally known as “DGM Series A”, described by Powell (2003). Gomani (2005:9) summarises as “12 cervical vertebrae, except the atlas, preserved in articulation with three proximal dorsal vertebrae”.
  • GCP-CV-4229, the holotype of Spinophorosaurus nigerensis, described by Remes et al. (2009). The specimen was found in very good condition and well articulated from C2 to C13, the last cervical. The atlas seems to be missing (Remes, pers. comm., 2021.

One other sauropod is complete from the first cervical, but probably not to the last:

  • MOZ-Pv1232, the holotype of Lavocatisaurus agrioensis, described by Canudo et al. (2018). This is complete from C1-C11. Canudo’s guess is that this is complete neck (Canudo, pers. comm, 2021), but the specimen doesn’t demand that conclusion and no known eusauropod has fewer than 12 cervicals.

Other sauropod specimens have necks that are complete and articulated from further back in the cervical sequence:

  • YPM 1910, Camarasaurus lentus, a mounted specimen described by Lull (1930). The neck is complete from C2 or C3, Lull was uncertain which.
  • SMA 0004, Kaatedocus siberi, described by Tschopp and Mateus (2012). Cervicals 3-14 are preserved.
  • AODF 888 (informally “Judy”), probably referrable to Diamantinasaurus, briefly described by Poropat et al. (2019). Preserved from C3 or maybe C4. “One posterior cervical (XIII or XIV) found several metres from articulated series, but appears to slot nicely into the gap between the articulated cervical series and the unprepared thoracic section, which might include at least one additional cervical (XIV or XV)” (Poropat, pers. comm. 2021).

Several necks are probably nearly complete, but it is not possible to knew due to their not being found in articulation:

  • CM 84, the holotype of Diplodocus carnegii, described by Hatcher (1901). C2–C15 are preserved, though not all in articulation; C11 may be an intrusion: see below for details.
  • ZDM T5701, the holotype of Omeisaurus tianfuensis, described by He et al. (1988). The neck was not articulated (He et al. 1988:figure 1), and was missing “two elements or so” (He et al. 1988:120).
  • QJGPM 1001, the holotype of Qijianglong guokr, described by Xing et al. (2015). On page 8, the authors say “The axis to the 11th cervical vertebra were fully articulated in the quarry. The atlas intercentrum and the 12th–17th cervical vertebrae were closely associated with the series.”
  • MNBH TIG9, a referred specimen of Jobaria tiguidensis. Wilson (2012:103) writes that this specimen “includes a partially articulated series of 19 vertebrae starting from the axis and extending through the mid-dorsal vertebrae.”
  • MNBH TIG6, another referred specimen of Jobaria tiguidensus, which has not been mentioned in the literature. Sereno (pers. comm., 2021) says that it is “a subadult partial skeleton with excellent neck” and that “the sequence was articulated from C2–11. Most of the ribs were attached as well.”

At the time of writing, the Paleobiology Database (https://paleobiodb.org/) lists more than 270 sauropod species. The nine unambigously complete and articulated necks therefore represent only one in 30 known sauropod species.

Note. The Jobaria tiguidensis individuals previously had specimen numbers beginning MNN, but the Musee National du Niger changed its name to Musée National Boubou Hama and the specimen numbers have changed with it.

Today marks the one-month anniversary of my and Matt’s paper in Qeios about why vertebral pneumaticity in sauropods is so variable. (Taylor and Wedel 2021). We were intrigued to publish on this new platform that supports post-publication peer-review, partly just to see what happened.

Taylor and Wedel (2021: figure 3). Brontosaurus excelsus holotype YPM 1980, caudal vertebrae 7 and 8 in right lateral view. Caudal 7, like most of the sequence, has a single vascular foramen on the right side of its centrum, but caudal 8 has two; others, including caudal 1, have none.

So what has happened? Well, as I write this, the paper has been viewed 842 times, downloaded a healthy 739 times, and acquired an altmetric score 21, based rather incestuously on two SV-POW! blog-posts, 14 tweets and a single Mendeley reader.

What hasn’t happened is even a single comment on the paper. Nothing that could be remotely construed as a post-publication peer-review. And therefore no progress towards our being able to count this as a peer-reviewed publication rather than a preprint — which is how I am currently classifying it in my publications list.

This, despite our having actively solicited reviews both here on SV-POW!, in the original blog-post, and in a Facebook post by Matt. (Ironically, the former got seven comments and the latter got 20, but the actual paper none.)

I’m not here to complain; I’m here to try to understand.

On one level, of course, this is easy to understand: writing a more-than-trivial comment on a scholarly article is work, and it garners very little of the kind of credit academics care about. Reputation on the Qeios site is nice, in a that-and-two-bucks-will-buy-me-a-coffee kind of way, but it’s not going to make a difference to people’s CVs when they apply for jobs and grants — not even in the way that “Reviewed for JVP” might. I completely understand why already overworked researchers don’t elect to invest a significant chunk of time in voluntarily writing a reasoned critique of someone else’s work when they could be putting that time into their own projects. It’s why so very few PLOS articles have comments.

On the other hand, isn’t this what we always do when we write a solicited peer-review for a regular journal?

So as I grope my way through this half-understood brave new world that we’re creating together, I am starting to come to the conclusion that — with some delightful exceptions — peer-review is generally only going to happen when it’s explicitly solicited by a handling editor, or someone with an analogous role. No-one’s to blame for this: it’s just reality that people need a degree of moral coercion to devote that kind of effort to other people’s project. (I’m the same; I’ve left almost no comments on PLOS articles.)

Am I right? Am I unduly pessimistic? Is there some other reason why this paper is not attracting comments when the Barosaurus preprint did? Teach me.



Early in my 2015 preprint on the incompleteness of sauropod necks, I wrote “Unambiguously complete necks are known from published account of only six species of sauropod, two of which are species of the same genus”, and listed them.

Taylor 2015: Figure 3. W. H. Reed’s diagram of Quarry C near Camp Carnegie on Sheep Creek, in Albany County, Wyoming. The coloured bones belong to CM 84, the holotype of Diplodocus carnegii; other bones belong to other individuals, chiefly of Brontosaurus, Camarasaurus and Stegosaurus. Modified (cropped and coloured) from Hatcher (1901: plate I). Cervical vertebrae are purple (and greatly simplified in outline), dorsals are red, the sacrum is orange, caudals are yellow, limb girdle elements are blue, and limb bones are green.

Haha, stupid me! I had hugely under-counted. With thanks to the three peer-reviewers of the submitted manuscript and to SV-POW! commenters, I have revised this list, in preparation for forthcoming resubmission. The table as it stands currently consists of 24 candidates, not all of them very solid. Of these, 15 were found in articulation, the others mostly not — though we don’t know for sure in all cases. Not all of the necks have been properly described, and not all of the ones that have been described have been named. And other questions hang over some of them, very briefly summarised in notes.

Here is the list, sorted by date of description. If I got the Google-docs permissions right, you should be able to see it but not edit it. (If you can edit, please don’t! And let me know.)

Please let me know if you find any mistakes, or if you think I have missed anything. Everyone who contributes will get a mention in the acknowledgements.


We’ve noted many times over the years how inconsistent pneumatic features are in sauropod vertebra. Fossae and formamina vary between individuals of the same species, and along the spinal column, and even between the sides of individual vertebrae. Here’s an example that we touched on in Wedel and Taylor (2013), but which is seen in all its glory here:

Taylor and Wedel (2021: Figure 5). Giraffatitan brancai tail MB.R.5000, part of the mounted skeleton at the Museum für Naturkunde Berlin. Caudal vertebrae 24–26 in left lateral view. While caudal 26 has no pneumatic features, caudal 25 has two distinct pneumatic fossae, likely excavated around two distinct vascular foramina carrying an artery and a vein. Caudal 24 is more shallowly excavated than 25, but may also exhibit two separate fossae.

But bone is usually the least variable material in the vertebrate body. Muscles vary more, nerves more again, and blood vessels most of all. So why are the vertebrae of sauropods so much more variable than other bones?

Our new paper, published today (Taylor and Wedel 2021) proposes an answer! Please read it for the details, but here’s the summary:

  • Early in ontogenly, the blood supply to vertebrae comes from arteries that initially served the spinal cord, penetrating the bone of the neural canal.
  • Later in ontegeny, additional arteries penetrate the centra, leaving vascular foramina (small holes carrying blood vessels).
  • This hand-off does not always run to completion, due to the variability of blood vessels.
  • In extant birds, when pneumatic diverticula enter the bone they do so via vascular foramina, alongside blood vessels.
  • The same was probaby true in sauropods.
  • So in vertebrae that got all their blood supply from vascular foramina in the neural canal, diverticula were unable to enter the centra from the outside.
  • So those centra were never pneumatized from the outside, and no externally visible pneumatic cavities were formed.

Somehow that pretty straightforward argument ended up running to eleven pages. I guess that’s what you get when you reference your thoughts thoroughly, illustrate them in detail, and discuss the implications. But the heart of the paper is that little bullet-list.

Taylor and Wedel (2021: Figure 6). Domestic duck Anas platyrhynchos, dorsal vertebrae 2–7 in left lateral view. Note that the two anteriormost vertebrae (D2 and D3) each have a shallow pneumatic fossa penetrated by numerous small foramina.

(What is the relevance of these duck dorsals? You will need to read the discussion in the paper to find out!)

Our choice of publication venue

The world moves fast. It’s strange to think that only eleven years ago my Brachiosaurus revision (Taylor 2009) was in the Journal of Vertebrate Palaeontology, a journal that now feels very retro. Since then, Matt and I have both published several times in PeerJ, which we love. More recently, we’ve been posting preprints of our papers — and indeed I have three papers stalled in peer-review revisions that are all available as preprints (two Taylor and Wedels and a single sole-authored one). But this time we’re pushing on even further into the Shiny Digital Future.

We’ve published at Qeios. (It’s pronounced “chaos”, but the site doesn’t tell you that; I discovered it on Twitter.) If you’ve not heard of it — I was only very vaguely aware of it myself until this evening — it runs on the same model as the better known F1000 Research, with this very important difference: it’s free. Also, it looks rather slicker.

That model is: publish first, then filter. This is the opposite of the traditional scholarly publishing flow where you filter first — by peer reviewers erecting a series of obstacles to getting your work out — and only after negotiating that course to do get to see your work published. At Qeios, you go right ahead and publish: it’s available right off the bat, but clearly marked as awaiting peer-review:

And then it undergoes review. Who reviews it? Anyone! Ideally, of course, people with some expertise in the relevant fields. We can then post any number of revised versions in response to the reviews — each revision having its own DOI and being fixed and permanent.

How will this work out? We don’t know. It is, in part, an experiment. What will make it work — what will impute credibility to our paper — is good, solid reviews. So if you have any relevant expertise, we do invite you to get over there and write a review.

And finally …

Matt noted that I first sent him the link to the Qeios site at 7:44 pm my time. I think that was the first time he’d heard of it. He and I had plenty of back and forth on where to publish this paper before I pushed on and did it at Qeios. And I tweeted that our paper was available for review at 8:44 — one hour exactly after Matt learned that the venue existed. Now here we are at 12:04 my time, three hours and 20 minutes later, and it’s already been viewed 126 times and downloaded 60 times. I think that’s pretty awesome.


  • 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. [PDF]
  • Taylor, Michael P., and Mathew J. Wedel. 2021. Why is vertebral pneumaticity in sauropod dinosaurs so variable? Qeios 1G6J3Q. doi: 10.32388/1G6J3Q [PDF]
  • Wedel, Mathew J., and Michael P. Taylor 2013b. Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus. PLOS ONE 8(10):e78213. 14 pages. doi: 10.1371/journal.pone.0078213 [PDF]