Sauropods held their necks erect … just like rabbits

May 27, 2009

Welcome, one and all, to Taylor, Wedel and Naish (2009), Head and neck posture in sauropod dinosaurs inferred from extant animals.  It’s the first published paper by the SV-POW! team working as a team, published in Acta Palaeontologica Polonica, and freely available for download here.

Far, far back in the uncharted depths of history, silly people like Osborn and Mook (1921:pl. 84), Janensch (1950b: pl. 8) and Paul (1988:fig. 1), who didn’t know any better, used to depict sauropods with their necks held strongly elevated.

The classic reconstruction of Brachiosaurus brancai, from Janensch (1950b: plate VIII)

The classic reconstruction of Brachiosaurus brancai, from Janensch (1950b: plate VIII. (For some reason, WordPress doesn't allow italics in these captions, hence the roman-font taxonomic names.)

All that began to change with Martin’s (1987) short paper in the Mesozoic Terrestrial Ecosystems volume, and was then turned upside-down by Stevens and Parrish’s (1999) seminal paper in Science: two and a half pages that transformed the way the world looked at sauropods.

xxx

The subhorizontally mounted neck of the Rutland Cetiosaurus skeleton at the Leicester City Museum, in right posterolateral view.

Median part of the subhorizontally mounted neck of the Rutland Cetiosaurus skeleton at the Leicester City Museum, left lateral view.  Mike Taylor for scale.

The median part of the subhorizontally mounted neck of the Rutland Cetiosaurus skeleton at the Leicester City Museum, in left lateral view. Mike Taylor for scale.

John Martin looked at the cervical vertebrae of the Rutland specimen of Cetiosaurus oxoniensis, and concluded that the joints between them couldn’t be as flexible as people thought.  He reconstructed that animal’s neck in a low, near-horizontal pose, and with a very narrow range of movement that didn’t allow it to raise its head far above shoulder level.  Stevens and Parrish brought more rigour to this approach by modelling the cervical articulations of two sauropods (Diplodocus carnegii and Apatosaurus lousiae) using a computer program of their own devising, DinoMorph.  And as most SV-POW! regulars will probably know, they got results similar to Martin’s, showing neutral positions for both animals that were well below horizontal, and finding restricted ranges of motion.  (“neutral pose” here means that the vertebra are aligned such that the zygapophyses overlap as much as possible.)

Diplodocus carnegii, DinoMorph computer model , showing neutral neck posture, and limits of flexibility.  From Stevens (2002:fig. 6a).  [Note that Stevens's more recent models show a slightly higher neck due to its leaving the torso at a less steep angle.]

Diplodocus carnegii, DinoMorph computer model , showing neutral neck posture, and limits of dorsal and ventral flexibility. From Stevens (2002:fig. 6a). (Note that Stevens's more recent models show a slightly higher neck due to its leaving the torso at a less steep angle.)

The DinoMorph posture was quickly adopted as orthodox, and got a lot of exposure in the BBC’s classic CGIumentary, Walking With Dinosaurs: episode 2, Time of the Titans, was primarily about Diplodocus, and under Stevens’s consultancy showed them as having obligate low posture throughout the show.

A still from the BBC Walking With Dinosaurs, episode 2, Time of the Titans, showing Diplodocus in a DinoMorph-compliant posture with a low, horizontal neck.  Image copyright the BBC.

A still from Walking With Dinosaurs, episode 2, Time of the Titans, showing Diplodocus in a DinoMorph-compliant posture with a low, horizontal neck. Image copyright the BBC.

The new horizontal-neck orthodoxy was also reinforced by an exhibition at the American Museum of Natural History featuring a physical metal sculpture of a DinoMorph model:

Physical DinoMorph model at the AMNH, with horizontal-neck advocate Kent Stevens.  Photograph by Rick Edwards, AMNH

Physical DinoMorph model at the AMNH, with horizontal-neck advocate Kent Stevens. Photograph by Rick Edwards, AMNH

This brings us pretty much up to date: there’s been very little in the way of published dissent between 1999 and now, and a couple more Stevens and Parrish papers have reinforced their contention.  Upchurch (2000) published a half-page response to the DinoMorph paper, and Andreas Christian has put out a sequence of papers arguing for an erect neck posture in Brachiosaurus brancai on the basis that this best equalises stress along the intervertebral joints (e.g. Christian and Dzemski 2007), but otherwise all dissent from the DinoMorph posture has been limited to unpublished venues: for example, Greg Paul has posted several messages on the Dinosaur Mailing List disputing the low-necked posture, but has yet to put any of his arguments in print.

But enough of this dinosaury stuff.  Let’s look at a nice, cuddly bunny:

wild-rabbit-41946-480px

Now here’s the thing: you wouldn’t guess by looking at it, but that rabbit has a vertical neck.  In fact, it’s more than vertical: it’s so upright that it bends back on itself.  Don’t believe me?  Then take a look at this X-ray of an unrestrained awake rabbit:

Unrestrained awake rabbit, left lateral view, in X-ray, showing vertical neck. From Vidal et al. (1986:fig. 4B)

Unrestrained awake rabbit, left lateral view, in X-ray, showing vertical neck. From Vidal et al. (1986:fig. 4B)

Amazing.

Can it be that rabbits have unusual cervical vertebrae, such that when you articulate them in neutral pose they curve strongly upwards?  No: and to prove it, here is (ahem) Taylor, Wedel and Naish (2009: fig. 1):

Taylor et al. (2009: fig. 1), reverse for easy comparison with the previous two images: skull and cervical skeleton of the Cape hare (Lepus capensis) in neutral pose and in maximal extension

Taylor et al. (2009: fig. 1), reversed for easy comparison with the previous two images: skull and cervical skeleton of the Cape hare (Lepus capensis) in neutral pose and in maximal extension

(Yes, this is a hare rather than a rabbit, but it’s close enough for government work.)  What we found was that it was only possible to get the cervical skeleton anywhere near the habitual life posture by cranking all the proximal cervical joints up as far as they could physically go.  In fact, it seems that some of the joints in the live animal flex more than the dry bones can — presumably due to intervertebral cartilage moving the centra further apart.

And this is fully in accord with the findings of Vidal et al. (1986), who X-rayed a selection of live animals (human, monkey, cat, rabbit, rat, guinea pig, chicken, monitor lizard, frog) and found that the neck is inclined in all but the frog.  Furthermore, in all the mammals and reptiles, they found that:

  • the cervical column is elevated nearly to the vertical during normal functioning;
  • the middle part of the neck is habitually held relatively rigid;
  • the neck is maximally extended at the cervico-dorsal junction and maximally flexed at the cranio-cervical junction; and
  • it is the cranio-cervical and cervico-dorsal junctions that are primarily involved in raising and lowering the head and neck.

(In life, these facts are obscured from view by soft tissue.)

We also looked at unpublished live-alligator X-rays (thanks to Leon Claessens for access to these) and found that even in these ectothermic sprawlers, the neck is habitually elevated above neutral pose.  Published X-rays of turtles and even (slightly) salamanders also showed the same tendency.

So what does this mean for sauropods?  Simply, unless they were different from all extant terrestrial amniotes, they did not habitually hold their necks in neutral position, but raised well above horizontal.  And if they resembled their closest relatives, the birds — and the only other homeothermic and erect-legged group, the mammals — then their necks were strongly inclined.  As in, all the proximal cervicals were habitually cranked into the most erect positions they could attain.  Kind of like this:

Diplodocus carnegii head, neck and anterior torso, right lateral view, articulated in habitual posture as hypothesised by Taylor et al. (2009).  Skull and vertebrae from Hatcher (1901).

Diplodocus carnegii head, neck and anterior torso, right lateral view, articulated in habitual posture as hypothesised by Taylor et al. (2009). Skull and vertebrae from Hatcher (1901).

Which is a looong way form the DinoMorph posture that we were all getting used to but couldn’t learn to love.  What do you know?  Turns out that Osborn and Mook, and Janensch, were right after all.

So that, in a nutshell, is the contention of the first SV-POW! paper: that sauropods held their heads up high.  That’s not to say that they couldn’t bring them lower when they wanted to — of course they could, otherwise they’d have been unable to drink — but we believe the evidence from extant animals says that they spent the bulk of their time with their heads held high.

I leave you with this rather beautiful piece that noted pterosaurophile Mark Witton drew to illustrate our favoured posture.  Enjoy!

Diplodocus herd -- mostly with necks in habitual raised posture, with one individual drinking.  By Mark Witton.

Diplodocus herd -- mostly with necks in habitual raised posture, with one individual drinking. By Mark Witton.

Stay tuned for more on neck posture …

Update

For more cool stuff about the paper, including blog and media coverage and the chance to hear Mike on BBC Radio(!), see our page about the paper on the sidebar.

References

  • Christian, A. and Dzemski, G. 2007. Reconstruction of the cervical skeleton posture of Brachiosaurus brancai Janensch, 1914 by an analysis of the intervertebral stress along the neck and a comparison with the results of different approaches. Fossil Record 10: 38-­49.
  • Janensch, W. 1950b. Die Skelettrekonstruktion von Brachiosaurus brancai. Palaeontographica (Supplement 7): 97-­103.
  • Martin, J. 1987. Mobility and feeding of Cetiosaurus (Saurischia, Sauropoda) ­ why the long neck? In: P.J. Currie and E.H. Koster (eds.), Fourth Sympo- sium on Mesozoic Terrestrial Ecosystems, Short Papers, 154­-159. Box- tree Books, Drumheller, Alberta.
  • Osborn, H.F. and Mook, C.C. 1921. Camarasaurus, Amphicoelias, and other sauropods of Cope. Memoirs of the American Museum of Natural History, new series 3: 246­-387.
  • Paul, G.S. 1988. The brachiosaur giants of the Morrison and Tendaguru with a description of a new subgenus, Giraffatitan, and a comparison of the world’s largest dinosaurs. Hunteria 2 (3): 1­-14.
  • Stevens, K.A. and Parrish, J.M. 1999. Neck posture and feeding habits of two Jurassic sauropod dinosaurs. Science 284: 798­-800. [Free subscription required]
  • Taylor, M.P., Wedel, M.J. and Naish, D. 2009. Head and neck posture in sauropod dinosaurs inferred from extant animals. Acta Palaeontologica Polonica 54(2): 213-220.
  • Upchurch, P. 2000. Neck posture of sauropod dinosaurs. Science 287: 547b.
  • Vidal, P.P., Graf, W., and Berthoz, A. 1986. The orientation of the cervical vertebral column in unrestrained awake animals. Experimental Brain Research 61: 549­-559.

68 Responses to “Sauropods held their necks erect … just like rabbits”

  1. Andy Says:

    Congratulations! Three cheers for actually testing biological assumptions against reality! :-)

  2. Nathan Myers Says:

    Is that all you got? Extant vertebrates? You haven’t heard the last on this.

    I guess this means Elasmosaurus var. swam with its head out of the water… like Nessie!

    I suppose the return of tail-dragging is next — or maybe wallowing?

  3. Andrea Cau Says:

    Congratulations, SVPoWer Rangers!

  4. Daniel Madzia Says:

    “[…] silly people like Ostrom and Mook […]”

    Osborn;o)

    Sounds interesting… The paper is already available for download.

  5. iljajj Says:

    If this means that all of the Carnegie Diplodocuses will be re-mounted in their original early-20thC design, that WOULD be ironic.

  6. Mark Wildman Says:

    Makes sense to me – always has done. I’ve thought for some time now that intervertebral flexion between centra because of cartilage must increase neck mobility. This is an area for more research, not just along the vertebral column but also the leg joints.
    And how thick were these cartilaginous discs? A lot thicker than some suspect I’d wager……

  7. James Robins Says:

    Brilliant……and a filing cabinet full of sauropod pix unusable for a decade once more released into the light……J

  8. Andreas Johansson Says:

    Tangential, but what would be ONP for human necks and heads?

  9. Darren Naish Says:

    Believe it or don’t, when articulated in ONP the human neck is strongly flexed. It’s actually present in people who have suffered severe neck trauma and are no longer able to properly extend the cervico-dorsal junction.


  10. […] out talking about the position of the neck in sauropod dinosaurs. Read about it in more detail at their hompage, or over at Tetrapod […]


  11. Extremely cool stuff.

  12. Sean Craven Says:

    Does this mean that you guys are going to have to start arguing about blood pressure again?

    Oh — and Witton’s art is gorgeous as always. He’s one of the few whose work strikes me as scientific art rather than scientific illustration — and the idea of scanning a pencil sketch in and coloring it directly is one I believe I will swipe in the near future.

  13. Mark Wildman Says:

    I’ve just heard Mike’s radio interview. Priceless! He nearly choked when the interviewer made the obligatory T.rex reference! “We only work on the sauropods…” Get in there Mike – Love it!

  14. Matt Says:

    Congradulations on the sauropod neck paper! The research all of you have done will hopefully make other sauropod researchers rethink their ideas on sauropod neck posture.
    As someone commented before why would an animal have such a long neck and use it for grazing? It makes no sense what so ever.
    The sauropod which others may use to counter argue with your hypothesis is the Mamenchiasaurus, since it has such a long neck some may feel it is too long to held up. However, Greg Paul’s reconstructions show this animal with an upright neck. Overall looks like sauropod specialists have some new information now on which to debate on. I look foward to reading more on this topic.

  15. domenico Says:

    Great thing !!
    Congratulation !

  16. Zach Miller Says:

    Was that “government work” comment a Grim Fandango reference?!? Just say “yes,” and I’ll think you’re awesome forever. Like I said back on Tet. Zoo, wonderful paper, gents.

    That metal DinoMorph sculpture is tres cool.


  17. Bravo, guys! Hooray for data! Even my phone/email is ringing about this paper. WTF? No T. rex questions so far, thankfully…

  18. Nathan Myers Says:

    This does help answer the question as to how all that unchewed greenery got down such a long gullet without graboids. I wonder, though, where the trachea would connect, if it did. At the bottom seems unlikely. Are there any tetrapods with entirely separate gullet and trachea?

  19. Terry Hunt Says:

    Over at Tet Zoo, Dallas beat me to it with a comment to the effect that it makes better energetic sense to hold the neck as vertical as possible, minimising the effort needed to counter leverage effects.

    So I’ll just suggest that you should adopt this –

    – as the SZ-POW! theme tune.

  20. Nima Says:

    CNGRTS, ll thr f y! Ths s sm mprssv rsrch s wll s brth f frsh r. ‘ve bn wtng t s ppr lk ths fr lng tm.

    Stvns nd Mrtn smply strk m s gtstcl nd nscntfc. A cmptr prgrmmr mks 3d mdl – w, nd h cn mk t NY WY H WNTS. nd rsult – h cms t s ctstrphst dtrctr, clmng tht Brchsrs wr NCPBL f hldng thr ppr-thn nck vrtbr vrtcl!

    Bt y gys lkd t lvng nmls fr nck flxn. Ths s th srt f thng tht Stvns should b slppng hs frhd vr.

    Lw shldrd dpldcds D ppr t hv hrzntl ncks, bt thy’r NT prfctly strght s Stvns sggsts – wldn’t g s fr s Htchr’s ngrving, bt thy D crv PWRD twrds th hd. Ths ws cmfrtbl “cntlvr” pstn fr th twn nck tndns (mr s thn strnd, prfctly rlr-strght nck).

    Thgh wth Brchsrs, thnk Pl gt t rght n 1988 – nd nw t’s cnfrmd! Th bns (s w nw knw thnks t th rmntng jb n Brln) d rtclt n n s-crv. nd Knt Stvns clmng tht thy CN’T nd sng crd drwngs f bns lnd p n flr fr msrng prpss, s smply drwning n chldsh gnrnc f th dt. Crtlg – whch s wht rlly rstrcts nck mvmnt – ds nt vn fsslz, s thr s N gd rsn t sspct Brchsrs “cn’t” hld thr ncks crtn wy, shrt f ll-t DSLCTN (nd y’d hv t crv th nck wll pst vrtcl t chv tht!).

  21. Dan Varner Says:

    I would hope that this discussion remain civil, rather than induldging in personal attacks. This problem has been in contention for over one hundred years and I’m willing to bet will go on for a few more. It need not become nasty.

  22. Matt Wedel Says:

    Yes, absolutely. We all realize that our work is not the last word on the subject. All three of us get along fine with Kent Stevens, too.

    Nima, you’ve been disemvoweled. Any more character assassination and I’ll ban you permanently.

  23. ScottE Says:

    Marvellous! (My single drawing of a sauropod is still mostly correct.)

  24. Jaime A. Headden Says:

    I’m all for this paper, great study. But I noticed something missing on the direct testing of head-attitude: Semicircular canals. Is there work to compare the various skulls of sauropods where the inner ear is known to test the position of the horizontal canal, the orientation of this canal relative to head attitude (on Witmer’s work) and the use of the inference of head-neck flexure to the posture of the head?

    Another interesting position that seems peculiar, though: I do not think it is simple or easy to infer a rabbit’s short neck (encased in robust muscle) is comparable to a sauropod’s neck, as it is so long, unless it were also as encased or proportionately encased in muscle. Do these questions bear on the topic of the neck’s attitude?

  25. Nathan Myers Says:

    I’m going to anticipate that the semicircular canal position doesn’t help matters, because the canal goes with the head, and the head angle tells us little or nothing about the neck posture.

  26. Matt Wedel Says:

    I do not think it is simple or easy to infer a rabbit’s short neck (encased in robust muscle) is comparable to a sauropod’s neck, as it is so long, unless it were also as encased or proportionately encased in muscle. Do these questions bear on the topic of the neck’s attitude?

    I would argue not. The pattern of carrying the neck extended and the head flexed is tetrapod-wide. It holds for things with short, muscle-bound necks and for things with long, slender necks. We couched the paper in terms of sauropods because that’s where the battle’s at, but we could have made the same argument using any extant tetrapods as ammo and any extinct tetrapods as targets.

    I’m going to anticipate that the semicircular canal position doesn’t help matters, because the canal goes with the head, and the head angle tells us little or nothing about the neck posture.

    We-e-ell… Something we noticed, and commented on in the paper, is that if you put the HSCCs level in most sauropods, the occipital condyle points down. That means the anterior neck had to be going up, which in light of the Vidal and Graf work implies that the whole neck is going up. If the HSCCs are angled up, as they are in many mammals and birds, the occipital condyle points further down and the neck must be even steeper. If the head was not carried in neutral position but was instead flexed on the neck, as it is in most extant tetrapods, then the neck had to be steeper still. So the HSCCs aren’t completely uninformative. They tell us that if sauropods were anything like extant tetrapods, at least the anterior part of the neck was angled up steeply. That dovetails nicely with the Vidal and Graf data, so we’ve got two lines of evidence for inclined necks in sauropods. Both lines of evidence stand on their own, but–like the Wonder Twins–their powers combine.

  27. Scott Hartman Says:

    Well done good sirs, well done.

    Now, to show where neutral position actually lies in some of these taxa!


  28. […] It’s been a day spent doing publicity for the new SV-POW! paper on sauropod neck posture. […]


  29. […] a few days on neck posture, I thought I’d expand on what Mike said about bunnies in the first post: in most cases, it is awfully hard to tell the angle of the cervical column when looking at a live […]


  30. […] heads high.” For more be sure to check out the summaries of the paper at Tetrapod Zoology and SV-POW!, written by some of the authors of the study. Posted By: Brian Switek — Discoveries | Link […]

  31. John Dziak Says:

    Although I’m not a paleontologist, I have been really enjoying reading about this debate. I liked Dr. Witton’s drawing with the jaws sloping down even though the necks are sloping up — a little bit like horses maybe. I had always imagined that whatever their necks were doing, sauropods’ heads should always be facing forward so they could see where they’re going. But of course I was thinking like a human being, whereas for a creature with eyes on the sides of its head it wouldn’t have to be that way.

  32. Graham King Says:

    Great! Brilliant! Fantastic! Well done you guys!

    What a relief. Sauropods can hold their heads up high, as lords of all they survey, and abandon that slouchy hangdog look. What’s that slapping sound I hear in the heavens? Pioneer illustrators giving each other high-fives on high?!

    Thanks for the free PDF. I read the whole paper right through straight off. It is so commendably readable!

    Some of your G. brancai (thorax-high and thorax-low) varied neck postures do look quite odd, because an animal would surely not keep the rest of its body in same fixed attitude while reaching way up or down? But of course I see that you were illustrating neck range, not real-life whole-body postures, there.

    (I think that craning, stretching, browsing and drinking postures of sauropods may provide interesting scope for further study. And that’s all without even considering their vertebral columns’ scopes for lateral flexion.. or torsion!)

    Once again, well done guys. You will have cheered up a lot of kids once more.. ;-D

  33. Graham King Says:

    Oops, B.brancai. I was thinking G for Giraffatitan, but B. is what your paper has.


  34. […] of Ted Naismith’s, because it amuses me to see the role of Glaurung so obviously played by a Diplodocus.  Enjoy: "Finduilas is Led Past Túrin at the Sack of Nargothrond" by Ted […]


  35. […] Sauropod Vertebra Picture of the Week: Love to look at and study the remains of ancient sauropods? This blog is perfect for you, with regularly posted photos of all kinds of skeletal remains. Recommended posts: “Lies, damned lies, and Clash of the Dinosaurs” and “Sauropods held their necks erect, just like rabbits.” […]


  36. […] The case for horizontal or near-horizontal habitual pose rests on two assumptions.  First, that osteological neutral pose (ONP) was habitually adopted; and second, that we can know what ONP was.  We still feel that both of these assumptions are false.  We can’t know ONP because there is not a single sauropod neck skeleton anywhere in the world consisting of undistorted cervicals — and even if we knew what ONP was, it wouldn’t tell us much about what I am suddenly going to call mechanical neutral pose (MCN), because we don’t know anything about the intervertebral cartilage.  And we know that extant animals do not habitually adopt ONP because we have X-rays that show us how they habitually rest, and we know that they don’t match what you get by articulating bones. […]


  37. […] to convey that Mama had broken its neck.  Of course, that didn’t really work, because the extension at the base of the neck is habitual for most amniotes anyway, but it at least gave us a sense of the direction we wanted to go […]

  38. Adam Baig Says:

    I know this is an older topic of discussion; but may I offer my two cents in this please?
    I have heard that with such long necks, it may be extremely energy-consuming to lift such a long neck way up high on behalf of the heart. So, it may seem feasible to say that sauropod necks and heads were held low to reduce systolic blood pressure on the heart. I disagree.
    Sauropods could not at all keep their long necks and heads more horizontal than vertical because keeping the neck level would also be energy consuming. Try to hold a long rod horizontal for a long period of time. Keeping the neck and head more vertical would allow more normal force in the pectoral region to assist the weight of the long neck and head.
    Also, sauropods (even diplodocids and camarasaurids, not just brachiosaurids) keeping their necks and heads more vertical would allow more surveillance on their environment, for food, potential mates and predators.
    Also, keeping the necks and heads more horizontal would mean being more vulnerable to charging predators, who would have an easier chance of seizing the sauropods’ head and neck.
    Furthermore, why would sauropods have evolved such a long neck for low grazing, when they could just walk to where more grass would be? Giraffes have long necks for higher browsing.
    I would postulate that sauropods must have had a specialized heart (or entire circulatory system) to deal with such blood pumping, regardless if it would be fully divided or semi-divided.

  39. Mike Taylor Says:

    Adam,

    We are in broad agreement with all the points you make here, and it is certainly true that holding an elongate appendage such as a long neck horizontally requires more muscular exertion than holding it vertically.

    That said, the issue of the heart is a real one, and as things stand imposes a real barrier to the habitually-elevated neck hypothesis that we favour. The issue has been extensively discussed in a sequence of publications by Roger Seymour, going back as far as 1976, and not yet satisfactorily rebutted in the literature.

    We hope to change that soonish, but for obvious reasons we can’t say much about it at this stage. (This must be infuriating for Roger, who is probably just itching to smack down our argument, but can’t get started on that project until we actually publish it. I’m sympathetic: having been on the other side of the equation a few times, I know how annoying it can be.)

  40. Adam Baig Says:

    Hi Mike,

    Quick question on this topic.

    In addition to the other things I have said (of which you are all in broad agreement with me upon) on behalf of sauropods being high-neck-positioned browsers, I thought that sauropods had teeth adapted for raking fern and conifer leaves. I would think that their tooth structure is not adapted for grazing, which would be further evidence that sauropods did not have their necks low.

    This is just an observational comment. That’s all.

  41. Mike Taylor Says:

    Sauropod tooth morphology varied enormously. The teeth of Nigersaurus, for example, look very highly adapted to ground-level grazing. It’s an easy mistake to generalise across Sauropoda; but a clade that persisted for 150 million years and encompassed a couple of hundred genera that we know about, in numerous different ecological settings, is going to have encompassed a huge range of feeding styles. Asking “how did sauropods eat?” is like asking “how do artiodactyls eat?”


  42. […] As usual for tetrapods, the neck is extended at the base and flexed at the head. […]


  43. […] neck posture is fine. Easy to say, but since I’m one of the “sauropods held their necks erect” guys, it might need some […]


  44. […] things). Along the way, we’ve shared the authorship of a few papers with other authors (Taylor, Wedel and Naish 2009 on habitual neck posture; Taylor, Wedel and Cifelli 2011 on Brontomerus; and Taylor, Hone, Wedel […]


  45. […] in the cotyle. Whether this posture was attainable in life is debatable; I’ve seen some pretty weird stuff. In any case, we didn’t use this joint for estimating cartilage thickness because we had no […]


  46. […] still held by terms that have nothing to do with sauropods. “Rabbit” can only be due to this post on sauropod neck posture; “Leopard seal” is due to the inclusion of a single sensational (but off-topic) photo […]


  47. […] RUBBISH. What was I thinking, and why did my SV-POW!sketeer co-authors let me choose such an uninformative title? We should of course have gone with a title that says what posture we inferred. The associated blog-post had a much better title: Sauropods held their necks erect … just like rabbits. […]

  48. Mark Sutherland Says:

    While I appreciate the perspective of extant morphology, I think in regards to mass, birds and rabbits are a very poor comparison point and would actually be a point of easy contrast. Also, when you look at the rabbit’s lower cervical structure, where the angular posture is at its utmost, you see highly specialized vertebrates which are not so much apparent in sauropods.

    I see also that Taylor is taking the corollary with birds to a rather extreme degree in his “Quantifying the effect of intervertebral cartilage on neutral posture in the necks of sauropod dinosaurs”: https://peerj.com/articles/712/
    He takes it to the point where his diagrams take the exaggerated argument of CNP to the extant of a bird-analogue in its S-shape which stretches credulity.

    This doesn’t speak overall against an erect neck, I would just caution against suggesting the strength of your argument lies in birds and rabbits.

  49. Mike Taylor Says:

    Well, Mark, perhaps we didn’t explain our point well enough in the blog-post. But it certainly is not “Look, rabbits do this, so sauropods must have”. The point is that all tetrapods habitually hold their necks extended from neutral posture — and we emphasised this by showing that it’s true even of animals such as rabbits that we don’t usually think of as even having a neck. If we wanted to extrapolate sauropod behavior from any single extant animal, then the obvious candidate would be a big bird; but that’s not what we were saying. We were making a much broader, and stronger, point.

    Also, be careful not to conflate our 2009 paper, which was all about behaviour extrapolated from that of extant critters, with my 2014 paper, which is simple geometry. The diagrams that you describe as “exaggerated” certainly look weird — as the paper points out, I do not think they were habitually adopted in life — but geometry tells us that neutral postures when cartilage is taken into account were much, much more elevated than those that omit cartilage. I don’t think there’s any avoiding that conclusion. Or at least, anyone who wants to disagree has the responsibility of showing why that paper is wrong. Just saying “it looks weird” won’t cut it.


  50. […] These folks start to shift in their seats when we put up too many posts in a row on open access or rabbits or…okay, mostly just OA and bunnies. If that’s you – or, heck even if it […]


  51. […] the bones, then the necks of many sauropods had the flexibility to loop in a complete circle.  The brilliant minds behind the SV-POW! team over at svpow.com (always enjoyable to read, but definitely more on the technical side) helped rekindle my love for […]


  52. […] Sauropods held their necks erect … just like rabbits — 27,454 […]


  53. […] something useful, because it’s essentially ubiquitous among birds and many mammals … including rabbits, as long-time readers will […]


  54. […] extant wildlife continued to be a source of enjoyment and inspiration, especially this cottontail. Rabbits, […]


  55. […] couple of decades, it’s that life posture for extinct animals is controversial — and that goes double for sauropod necks. Heck, even the neck posture of extant animals is terribly easy to misunderstand. We really […]


  56. […] conviction the quality of brachiosaurosity. It has a very posable neck that can be placed in a realistic life posture, and there are even hints of scapulae and […]

  57. Don Says:

    Mike Taylor wrote: “…postulate that sauropods must have had a specialized heart (or entire circulatory system) to deal with such blood pumping…”

    Is “dive time” relevant here? The ability to store O2 scales positively with body mass – add in pneumaticity, and the potential for a breath hold time of an hour is not extreme. If they could maintain adequate O2 levels in the brain for say 30 minutes, without pumping, then a dip of the head to heart level would allow a “normal” heart to recharge the brain…

    And:

    If holding a neck level is hard, why not a tail?

  58. Mike Taylor Says:

    Two excellent points here, Don, many thanks!

    On the first, I don’t know how dive time would play in here, but it certainly seems a relevant enough concept for someone to work on. If that’s how sauropod necks worked, I wonder how long the “recharge cycle” with a lowered neck would need to be?

    On the second, my instant thought is that saurpods had very elevated sacra, but nothing analgous in the shoulders. So it might be possible to suspend the tail by ligaments from the top of the sacrum in such a way that the muscularly neutral position was roughly horizontal. Related to this, the sacra of saurpods are wedged in a way that would tend to make the base of the tail leave the torso at a slightly inclined angle, so I wonder whether the very straight tails we’ve been used to seeing are an oversimplification, and the true trajectory was like a gently arc?

    Both of these are ideas that could use a lot of work, which I simply don’t have time to even get started on. If anyone wants to take them on, please be my guest.

  59. Matt Wedel Says:

    Relevant: elephant seals spend most of their lives in apnea (not breathing). When they are at sea, they dive for 30-60 minutes, then surface and breathe for 5-10 minutes, then dive again. They manage this by oxygenating a huge volume of red blood cells, which are then slowly leaked by the spleen during the dives, so that there is enough oxygen reaching the brain and muscles while they’re down.

    Oh, and apparently they basically pass out while they’re on the descent phase of the dive, and wake up when they get to depth, to conserve oxygen.

    The crazy thing is that they don’t breathe regularly while they’re sleeping on the beach. They’ll sleep for 20 minutes at a time, not breathing, then wake up, breath a bit, shuffle around, change position, bark at other seals, and then go back to sleep, and as soon as they are out they stop breathing. So in a given hour on the beach, they might only be awake and breathing for 10 or 15 minutes.

    The arcing tail idea is interesting, and worthy of further investigation.

  60. Mike Taylor Says:

    Well, today I learned that elephant seals are idiots.

    Unfortunately, the mechanism they use for oxygenating while diving is no use to sauropods, because unlike diving animals their problem isn’t getting oxygen into the lungs, it’s getting the oxygenated blood from there (or from the cache in the spleen) to the brain.

  61. Matt Wedel Says:

    Yeah. In general vertebrates are very, very good at evolving sophisticated vascular networks to solve all kinds of problems, and to handle varying pressure requirements on the fly. I didn’t bring up the elephant seal as the solution for sauropods, just as an example of that (for lack of a better term) “problem solving ability”. ‘Evolvability’ or ‘adaptability’ might be better terms for what I’m grasping at.

    Anyway, I’m not super-worried about how sauropods oxygenated their brains. For us getting blood to the brain is a huge deal, it’s typically 15% of our cardiac output. For sauropods it was probably more like 1% or less. The most obvious solution for sauropods would be to have a dilation of the carotid or vertebral arteries near the head, which could be recharged with blood when the head was lowered, and then keep a trickle of oxygenated blood flowing to the brain when the head was raised–assuming that there was any difficulty with just pumping blood straight up to the head like everything else does, and I’m far from convinced that was actually a problem.

    I’m sure we’ll never get any direct evidence of whatever weird thing sauropods actually did, but to assume that their vascular systems weren’t well-adapted to handle periodically high and varying pressures, when other vertebrates are so versatile, is loony. Basically it’s special pleading that sauropods sucked.

  62. Don Says:

    Thanks for the response, guys.

    “…sauropods would be to have a dilation of the carotid or vertebral arteries near the head, which could be recharged with blood when the head was lowered, and then keep a trickle of oxygenated blood flowing to the brain when the head was raised…”

    Indeed! Feeding motions would keep it sloshing around nicely. All the heart would need do is provide enough back pressure to prevent a siphon…

    If I have it right, head size and neck diameter both scale negatively with neck length in sauropods (?) – this strikes me as at least mild support for vertical posture, if true – I see no need to reduce brain size (or taper the fluid column) in the horizontal flow case.

  63. Mike Taylor Says:

    Don, I actually don’t know how head size and neck diameter scale with neck length in sauropods, so you’ve asked a good question here. Do you remember where you picked up that impression? Maybe there was a paper that I missed; or perhaps this is a fruitful study that someone could do.

    But I think that if it’s shown to be true, that would make sense either for horizontal or more vertical neck postures, so it wouldn’t really help to resolve that issue either way.

  64. Don Says:

    ” Do you remember where you picked up that impression?”

    No. A minimal search is not helpful. That said, I am reasonably comfortable with it as a personal observation. If brain size is taken as a proxy for “head size”, then the relative brain size of the long necks generally is very low.

    Giraffes have the lowest relative brain size of any mammal I have tested them against (cow, elephant, dog, human), by wide margins.

    Sauropods have the smallest relative brain size of anything, ever, of course – lower than T rex and even stegosaurus.

    Matt stated that he was not concerned with how sauropods got blood to their brains, because their brain needs were a small percentage of their “cardiac output”.

    I reverse that, and posit/conjecture/speculate or maybe observe that their brains are small because they are very hard to get blood to, and selection has reduced them accordingly – which implies vertical posture, given that getting blood to the brain is not an issue in the horizontal posture case.

    Googly stuff – a giraffe brain size of 750 g and a body of 1000 kg yields a ratio of .00075, which would leave a 100 kg man with a 75 g brain – assuming the brain of a 15000 kg titanosaurid weighed 40 g, then our 100 kg guy would have a 266 mg brain, lol. Seems so right sometimes…

  65. Mike Taylor Says:

    Hi, Don. The measurement you should probablybe using using here is encephalization quotient (EQ). See https://en.wikipedia.org/wiki/Encephalization_quotient

    Very roughly it compares brain mass with body mass raised to an allometric constant typically in the range 2/3–3/4. On this basis, we would expect sauropods to have proportionally smaller brains than smaller animals do. We’d need to compare endocast volumes with the brain masses predicted by EQ for animals of the appropriate mass.

  66. Don Says:

    Witmer cast Ampelosaurus – iirc, 39 – 40 cc was the result.

    Pretending the brain was made of water, and filled the case, gave me an upper bound brain/body ratio of around 2 E-6 at .040/15000 kg.

    That is the only sauropod data a minimal search turned up… But rest easy, sauropods are so far below the expectation of mass^3/4 it is almost funny.


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