The Diversity of Neosauropods, or Pity Poor Camarasaurus

Articulated skeleton of juvenile Camarasaurus lentus in the Carnegie Museum of Natural History, photographed by Daderot.

Let me just get the obvious out of the way first: sauropods were huge. Mind-bendingly huge. In some cases, big enough to reduce a human to a sticky puddle under foot and not even break their stride. For close to 150 million years, they were the largest land animals anywhere in the world, and no other terrestrial animal at any time has come even close to rivalling their largest representatives in size. Being around for so long, it should also be no surprise that they were diverse: a large number of sauropod genera have been named, representing a wide variety of forms. Nevertheless, most people's idea of sauropods is encompassed within just four genera from the late Jurassic of North America: Diplodocus, Apatosaurus, Brachiosaurus and Camarasaurus.

These four genera all belong to the clade Neosauropoda, which has been defined as the smallest clade containing the genera Diplodocus and Saltasaurus (a late Cretaceous South American genus). Upchurch et al. (2004) diagnosed the neosauropods by a number of cranial features, together with a reduction in the fourth hind toe (part of a general trend towards toe reduction in sauropods as their feet became more columnar—see this article by Darren Naish for more on the subject). However, Upchurch et al. were writing before the recognition of the Turiasauria, a European clade that is probably the sister group of neosauropods (Royo-Torres et al. 2006), and I don't know how that clade would affect the synapomorphy distribution*. The neosauropods quickly became the dominant sauropod group after their appearance in the middle Jurassic, and the only non-neosauropod sauropods to make it into the early Cretaceous were the aforementioned turiasaurs and possibly Jobaria, an African genus that varying analyses place either just inside or just outside the Neosauropoda.

*There has been an annoying tendency in recent years for many papers featuring phylogenetic analyses to present us with the character data matrix and the final trees from the analysis, but not do anything to map character changes onto the tree. The only way to find that out would be by transcribing the entire matrix and re-running the analysis yourself.

Mounted skeleton of Amargasaurus cazaui in the Melbourne Museum.

The famed North American genera include representatives of each of the two main lineages within the Neosauropoda. Diplodocus and Apatosaurus belong the Diplodocoidea, and Brachiosaurus and Camarasaurus belong to the Macronaria. Diagnostic features of the diplodocoids according the Upchurch et al. (2004) include restriction of teeth to the front of the jaw and a subrectangular snout. This last feature reaches an extreme in the middle Cretaceous Nigersaurus, which was another of those animals that serves to remind us that, if God did indeed create all of nature, then he was taking the piss. Ludicrous diplodocoids also include the late Jurassic South American Brachytrachelopan, which took one look at the graceful, elongate necks of all the other sauropods and decided that it simply couldn't be having with all that.

Reconstruction of Saltasaurus loricatus, by Lady of Hats.

The name of the other lineage, Macronaria, means 'big nostrils', and one of the notable features of this clade is, indeed, a great expansion in the size of the nares, the opening for the nostrils in the skull (though whether the size of the nares directly corresponds to the size of the actual nostrils is, I suppose, another question). As well as Brachiosaurus and Camarasaurus, this clade includes the Titanosauria, a very successful group that included the last surviving sauropods, but whose significance was overlooked for many years because they had the poor judgement not to achieve their main diversity in North America. At least some titanosaurs, such as Saltasaurus pictured above, sported a skin reinforced by nodules of bone.

Which brings us to Camarasaurus. For some reason, of the 'Big Four' genera, this is the one that gets the least love. While the other three have each in their turn enjoyed roles as stars of stage and screen, I'm not aware of a single film in which Camarasaurus has even been given a name-drop*. In John Sibbick's illustration of Brachiosaurus and Camarasaurus together (scroll down a bit at the link) in Norman's (1985) The Illustrated Encyclopedia of Dinosaurs, Brachiosaurus marches confidently towards the front bearing a goofy leer, while Camarasaurus is forced to sulk towards the back. It's all blatantly unfair. It's not as if Camarasaurus is rare: in fact, Camarasaurus may just be the best known of all sauropods, represented in the Morrison Formation by a whole whack of remains, including what is perhaps the single most beautiful sauropod specimen ever found (the one with which I opened this post). It was no slouch in the size department, either: its maximum length of about 23 metres is similar to that of Apatosaurus, despite it having proportionally shorter appendages than the latter. Nor does it lack distinctiveness: the short, bulldog-like face of Camarasaurus instantly stands out in any neosauropod line-up. So after all this time, doesn't Camarasaurus deserve to be given the spotlight?

*Though it is a pity that the name 'Camarasaurus' won out in the priority stakes over its competitor 'Morosaurus', which to those of us from a New Zealand background suggests a dinosaur made out of chocolate.

REFERENCES

Norman, D. 1985. The Illustrated Encyclopedia of Dinosaurs. Salamander Books: London.

Royo-Torres, R., A. Cobos & L. Alcalá. 2006. A giant European dinosaur and a new sauropod clade. Science 314: 1925-1927.

Upchurch, P., P. M. Barrett & P. Dodson. 2004. Sauropoda. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria, 2nd ed., pp. 259-322. University of California Press: Berkeley.