At least a few parts of the dinosaurian family tree are still relatively little studied and are controversial. One of these is the ankylosaurian dinosaurs, though a new paper seeks to delve into this problem and produced a new and comprehensive analysis. Lead author Rick Thompson takes us through it:
In 1842 Richard Owen coined the term “dinosaur”, helping to imbue the world with a fascination for the gigantic ‘terrible lizards’ of the prehistoric world. Owen’s work was based on the fossil remains of three distinct archosaurs; Megalosaurus and Iguanadon have become fixed in our consciousness as archetypal dinosaurs, while the third, Hylaeosaurus, is very much the ‘ugly-duckling’ of the trio. Hylaeosaurus is a member of the Ankylosauria, a group of Middle Jurassic to the Late Cretaceous heavily armoured herbivorous dinosaurs, closely related to the stegosaurs in the clade Thyreophora. The story of Hylaeosaurus somewhat mirrors that of its ankylosaur relatives; fragmentary, difficult to study and lacking the popularity of its theropod, ornithopod and sauropod cousins. The ankylosaurs barely even worked their way into Jurassic Park (apparently Ankylosaurus does appear in Jurassic Park III – if you can stand it…)! The lasting image of the Ankylosauria in the public consciousness is the classic battle between Ankylosaurus and Tyrannosaurus, with the ankylosaur determinedly swinging its tail club towards the leg of its powerful foe. Indeed, it is the presence of the tail club, widely regarded as a defensive weapon that could shatter bone (Arbour et al. 2009), which has helped to raise the profile of this fascinating dinosaur group.
Of course ankylosaurs are more than just a dinosaur with a weaponized tail. The Ankylosauria includes over 50 valid taxa, only a fraction of which possess this idiosyncratic armament. Fragmentary ankylosaur specimens have been recovered from every continent on Earth, though their record in North America and Asia is by far the best. It is their extensive covering of dermal armour, particularly that of the skull, that has proved to be both the blessing and bane of ankylosaur palaeontology. The high level of morphological diversity in the armour is probably responsible for the large number of ankylosaur taxa. The cranial ossifications completely cover the bony sutures of the skull, simultaneously providing a unique pattern to identify new species, while destroying a wealth of cranial characters that could help to resolve the interrelationships of the wider clade. Thus the evolution of the Ankylosauria is very poorly understood, and presents a tough challenge for systematic palaeontologists.
Traditionally the Ankylosauria is split into two families, the Ankylosauridae and Nodosauridae. The Ankylosauridae includes many of the more recognisable taxa including Ankylosaurus and Euoplocephalus. Ankylosaurids are generally identified by their short-broad skulls, which are ornamented with horn-like ossifications above (supraorbital), behind (squamosal) and below (quadratojugal) the eye, along with the presence of a tail club. The Nodosauridae generally have a more elongate skull, with a domed skull roof (parietal region) and boss-like rather than horn-like cranial ossifications. Their trunk armour often incorporates spike-like ossifications along their flanks, though no tail club is present. In recent years, a number of ankylosaurs, such Gastonia and the aptly named Gargoyleosaurus, have been found to possess combinations of these classic traits, thus blurring the distinction between ankylosaurids and nodosaurids. This has led some authors to erect a new group of ankylosaurs, the Polacanthidae, which includes many of these recent discoveries, along with a few older, fragmentary genera like Owen’s Hylaeosaurus.
A mounted skeleton of Gastonia burgei. Note the large shield over the pelvis, a character suggested to unite polacanthid taxa. Photo by Susie Maidment.
The validity of the polacanthid hypothesis has proved hard to test cladistically, as the majority of ankylosaur character sets are either heavily biased towards the skull, or include a limited number of taxa. As many of the potential polacanthid taxa are fragmentary, or predominantly postcranial, the incorporation of these taxa into ankylosaur phylogeny has been problematic. Studies including some polacanthid genera place them either towards the root of the Ankylosauridae, or as a new family (Polacanthidae), sister to the Ankylosauridae. During my Master’s Degree in Biosystematics at Imperial College, London, I was lucky enough to get the opportunity to try to resolve some of these issues in ankylosaur evolution. Paul Barrett and Susie Maidment offered me the chance to revise and update the unpublished PhD thesis of Jolyon Parish on ankylosaur phylogeny. My aim was to expand the taxon sample of Parish’s thesis, as well as modify his extensive character set. In this way I hoped to produce a ‘palaeontological total evidence’ phylogeny of the Ankylosauria, which included all valid taxa regardless of completeness, and a roughly even mix of cranial and postcranial characters.
The results of this project were recently published in the Journal of Systematic Palaeontology, and give some new insights into ankylosaur evolution. Encouragingly our data did support the traditional split between the nodosaurids and ankylosaurids; however, no polacanthid clade was recovered. Instead, all taxa that have been affiliated with the Polacanthidae by various authors (and Ankylosauridae by others) were resolved within the Nodosauridae. In the strict consensus tree (the tree which summarizes all of the information that each of the shortest possible evolutionary trees agree upon) the nodosaurid clade formed a polytomy – that is, we could say nothing about the internal relationships of the group. This poor resolution of the clade was in fact caused by a number of unstable taxa, whose position on the tree is highly variable. Unsurprisingly three of these unstable taxa have previously been attributed to the Polacanthidae. To counteract their influence we produced a derivative strict reduced consensus tree (shown in the figure). This tree prunes unstable taxa from the strict consensus tree, greatly increasing resolution. In this phylogeny it is clear that the ‘polacanthid’ taxa form a basal grade of nodosaurid dinosaurs, while the traditional nodosaurids (clade D) form a polytomy.
This result simultaneously highlights the strengths and weaknesses of the ‘palaeontological total evidence’ approach. By including all taxa (regardless of completeness) and a character set sampled from the whole skeleton, a large amount of missing data was guaranteed. This lowers the resolution of the tree, as seen in the traditional nodosaurid clade. However, it is the inclusion of such a broad sample of characters and taxa that has allowed completely new relationships to be revealed, placing ‘polacanthid’ taxa in the Nodosauridae. The problems have been further confounded by the extensive cranial ossification of the taxa. The variation in such complex ornamentation is exceptionally hard to capture using traditional discrete characters. This makes it very hard to test for presence of the evolutionary signal in the ornamentation. In the future, a study which incorporates extensive measurements of the cranial armour could better capture this variation, and more clearly reveal the relationships of the group. This is particularly important for the Nodosauridae, many of which are primarily distinguished by their cranial ornamentation.
The phylogeny of the Ankylosauridae was much better resolved, and broadly in agreement with existing studies, if the ‘polacanthid’ taxa are discounted. Again in this clade we see that the basal lineages do not conform to the traditional view of the Ankylosauridae, often having elongate skulls. The tip of the tail has not been recovered in many of these taxa, so the presence of a tail club is uncertain. However, our tree is the first cladistic analysis to place an ankylosaur that clearly lacks a tail club within the Ankylosauridae. Zhongyaunsaurus was originally described as a nodosaurid, though this assignment was subsequently corrected by Ken Carpenter (Carpenter et al. 2008). Our phylogeny confirms that Zhongyaunsaurus was indeed an ankylosaurid, and suggests that the most characteristic trait of the Ankylosauria is only present in the most derived ankylosaurids.
Unfortunately our study has done little to help the plight of the earliest known ankylosaur, Hylaeosaurus. Our analysis suggests that it is a member of the Nodosauridae, but its removal from the reduced consensus tree feels like a sad continuation of its rather anonymous history. Never the less, our study has helped to reveal a new perspective on the Ankylosauria. Although the ankylosaurid-nodosaurid dichotomy has been maintained, it appears that the classical characters of these groups only apply to the most derived of their forms. The earliest lineages of each family show greater levels of diversity in their armour and body form. This has made their classification difficult, and without more fossils, or newer forms of character, it is likely to remain so. There is still much work to do in order to understand the evolution of this extraordinary group of animals, but hopefully our study can serve to trigger a new wave of research in the coming years.