Yes, sauropod neck vertebrae got longer as the animals grew up

November 21, 2016

jvp-fig-12

Fig. 14. Vertebrae of Pleurocoelus and other juvenile sauropods. in right lateral view. A-C. Cervical vertebrae. A. Pleurocoelus nanus (USNM 5678, redrawn fromLull1911b: pl. 15). B. Apatosaurus sp. (OMNH 1251, redrawn from Carpenter &McIntosh 1994: fig. 17.1). C. Camarasaurus sp. (CM 578, redrawn from Carpenter & McIntosh 1994: fig. 17.1). D-G. Dorsal vertebrae. D. Pleurocoelus nanus (USNM 4968, re- drawn from Lull 1911b: pl. 15). E. Eucamerotus foxi (BMNH R2524, redrawn from Blows 1995: fig. 2). F. Dorsal vertebra referred to Pleurocoelus sp. (UMNH VP900, redrawn from DeCourten 1991: fig. 6). G. Apatosaurus sp. (OMNH 1217, redrawn from Carpenter & McIntosh 1994: fig. 17.2). H-I. Sacral vertebrae. H. Pleurocoelus nanus (USNM 4946, redrawn from Lull 1911b: pl. 15). I. Camarasaurus sp. (CM 578, redrawn from Carpenter & McIntosh 1994: fig. 17.2). In general, vertebrae of juvenile sauropods are characterized by large pneumatic fossae, so this feature is not autapomorphic for Pleurocoelus and is not diagnostic at the genus, or even family, level. Scale bars are 10 cm. (Wedel et al. 2000b: fig. 14)

The question of whether sauropod cervicals got longer through ontogeny came up in the comment thread on Mike’s “How horrifying was the neck of Barosaurus?” post, and rather than bury this as a comment, I’m promoting it to a post of its own.

The short answer is, yeah, in most sauropods, and maybe all, the cervical vertebrae did lengthen over ontogeny. This is obvious from looking at the vertebrae of very young (dog-sized) sauropods and comparing them to those of adults. If you want it quantified for two well-known taxa, fortunately that work was published 16 years ago – I ran the numbers for Apatosaurus and Camarasaurus to see if it was plausible for Sauroposeidon to be synonymous with Pleurocoelus, which was a real concern back in the late ’90s (the answer is a resounding ‘no’). From Wedel et al. (2000b: pp. 368-369):

Despite the inadequacies of the type material of Pleurocoelus, and the uncertainties involved with referred material, the genus can be distinguished from Brachiosaurus and Sauroposeidon, even considering ontogenetic variation. The cervical vertebrae of Pleurocoelus are uniformly short, with a maximum EI of only 2.4 in all of the Arundel material (Table 4). For a juvenile cervical of these proportions to develop into an elongate cervical comparable to those of Sauroposeidon, the length of the centrum would have to increase by more than 100% relative to its diameter. Comparisons to taxa whose ontogenetic development can be estimated suggest much more modest increases in length.

Carpenter & McIntosh (1994) described cervical vertebrae from juvenile individuals of Apatosaurus and Camarasaurus. Measurements and proportions of cervical vertebrae from adults and juveniles of each genus are given in Table 4. The vertebrae from juvenile specimens of Apatosaurus have an average EI 2.0. Vertebrae from adult specimens of Apatosaurus excelsus and A. louisae show an average EI of 2.7, with an upper limit of 3.3. If the juvenile vertebrae are typical for Apatosaurus, they suggest that Apatosaurus vertebrae lengthened by 35 to 65% relative to centrum diameter in the course of development.

The vertebrae from juvenile specimens of Camarasaurus have an average EI of 1.8 and a maximum of 2.3. The relatively long-necked Camarasaurus lewisi is represented by a single skeleton, whereas the shorter-necked C. grandis, C. lentus, and C. supremus are each represented by several specimens (McIntosh, Miller, et al. 1996), and it is likely that the juvenile individuals of Camarasaurus belong to one of the latter species. In AMNH 5761, referred to C. supremus, the average EI of the cervical vertebrae is 2.4, with a maximum of 3.5. These ratios represent an increase in length relative to diameter of 30 to 50% over the juvenile Camarasaurus.

If the ontogenetic changes in EI observed in Apatosaurus and Camarasaurus are typical for sauropods, then it is very unlikely that Pleurocoelus could have achieved the distinctive vertebral proportions of either Brachiosaurus or Sauroposeidon.

apatosaurus-cm-555-c6-centrum-and-arch-united

C6 of Apatosaurus CM 555 – despite having an unfused neural arch and cervical ribs, the centrum proportions are about the same as in an adult.

A few things about this:

  1. From what I’ve seen, the elongation of the individual vertebrae over ontogeny seems to be complete by the time sauropods are 1/2 to 2/3 of adult size. I get this from looking at mid-sized subadults like CM 555 and the hordes of similar individuals at BYU, the Museum of Western Colorado, and other places. So to get to the question posed in the comment thread on Mike’s giant Baro post – from what I’ve seen (anecdata), a giant, Supersaurus-class Barosaurus would not necessarily have a proportionally longer neck than AMNH 6341. It might have a proportionally longer neck, I just haven’t seen anything yet that strongly suggests that. More work needed.
  2. Juvenile sauropod cervicals are not only shorter than those of adults, they also have less complex pneumatic morphology. That was the point of the figure at the top of the post. But that very simple generalization is about all we know so far – this is an area that could use a LOT more work.
  3. I’ve complained before about papers mostly being remember for one thing, even if they say many things. This is the canonical example – no-one ever seems to remember the vertebrae-elongating-over-ontogeny stuff from Wedel et al. (2000b). Maybe that’s an argument for breaking up long, kitchen-sink papers into two or more separate publications?

Reference

Wedel, M.J., Cifelli, R.L., and Sanders, R.K. 2000b. Osteology, paleobiology, and relationships of the sauropod dinosaur Sauroposeidon. Acta Palaeontologica Polonica 45:343-388.

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5 Responses to “Yes, sauropod neck vertebrae got longer as the animals grew up”

  1. Anonymous Says:

    “I’ve complained before about papers mostly being remember for one thing, even if they say many things. This is the canonical example – no-one ever seems to remember the vertebrae-elongating-over-ontogeny stuff from Wedel et al. (2000b). Maybe that’s an argument for breaking up long, kitchen-sink papers into two or more separate publications?”

    I’ve had this issue too. My first publication involved the description of previously undescribed material, but no one ever seems to remember the discussion of why this material was important to the evolutionary history of the group, to the point that I have had to point it out to authors when reviewing manuscripts that touch on this subject.

    I ran into the same issue in the last paper I got published. The paper included a discussion of palaeobiology, evolutionary history, and palaeobiological inferences-via-morphology of the group in question, but I just know people are just going to end up remembering it for the new taxa. Of course in this case I could not split the paper up, since the conclusions of one part of the paper depended on the novel observations in the other, and it was unlikely these parts could stand by themselves.

  2. Mike Taylor Says:

    This is a perennial problem. What is the optimal unit of publication? It’s hard enough for authors to make this decision; when reviewers weigh in as well (“You need to add a phylogenetic analysis”, “This discussion is not relevant and should be removed”) it feels like a crap-shoot.

    One of my favourite examples of this is the timing-of-fusions-through-ontogeny section in the long and rather obscure Wedel and Taylor (2013b) paper on neural spine bifurcation in diplodocids — see pages 7 and 8 and Table 1. Understandably lost in a 34-page paper, this information went on to form the backbone of the much more visible subsequent Hone et al. (2015) paper “what, if anything, is an adult dinosaur?”

    I don’t really know what to do about this.

    References

    Hone, David W.E., Andrew A. Farke and Mathew J. Wedel. 2016. Ontogeny and the fossil record: what, if anything, is an adult dinosaur? Biology Letters 12(2):0947. doi:10.1098/rsbl.2015.0947. [PDF]

    Wedel, Mathew J., and Michael P. Taylor. 2013a. Neural spine bifurcation in sauropod dinosaurs of the Morrison Formation: ontogenetic and phylogenetic implications. PalArch’s Journal of Vertebrate Palaeontology 10(1):1-34. [PDF]

  3. AndrewD Says:

    I suppose that you could try, using open source publishing the old trick used by certain chemists in the past. They (he) split the publications up with an overarching title and many(!) sub-papers as e.g :- The affect of Magic Acid* on basicity: Compound A part 1…through compound ZZZ part LXXIIX…and so on.

    *Yes there is such a compound (a 1:1 molar ratio, of fluorosulfuric acid (HSO3F) and antimony pentafluoride (SbF5) if you really want to know)

  4. Jay Says:

    Cervical elongation is especially well documented in basal sauropodomorphs. Massospondylus went from being stocky dog-like critters with short necks as embryos or post-hatchlings (Resisz et al. 2005, 2010), transformed into long-necked gracile animals as sub-adults/adults. There are similar transformations known for abundant specimens of Mussaurus, and less so for Lufengosaurus.


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