Cutting up dinosaur’s evolutionary tree

For more than 130 years dinosaurs have been classified into two distinct orders, the saurischians and the ornithischians. But as it is common in biological sciences, every theory is true until the opposite is proved. A new study has called into question classical dinosaur classification, destroying and redistributing some of the different dinosaur groups. Even if this new hypothesis isn’t 100% sure yet, in this entry we’ll explain what this dinosaur reordering consists in.

TRADITIONAL DINOSAUR CLASSIFICATION

Since the XIX century, dinosaurs have been divided into two large orders based on their hip anatomy. The order Saurischia (lizard-hipped) includes theropods (carnivorous dinosaurs and current birds) and sauropodomorphs (large, long-necked herbivores); the order Ornithischia (bird-hipped) includes ornithopods (herbivorous and duck-billed dinosaurs), marginocephalians (dinosaurs with horns and hardened skulls) and thyreophorans (armored dinosaurs).

Traditional dinosaur evolutionary tree by Zureks, with the two different hip morphologies at the bottom.

Yet, this classification doesn’t have the last word. Palaeontology is an extremely volatile science, as with each new discovery, you can dismantle everything you knew at that moment, even if it’s a centenary-old hypothesis. This is what has recently happened with dinosaurs.

THE RISE OF A NEW HYPOTHESIS

A new study published in March 2017, has caused the reconsideration of traditional dinosaur classification. Many previous studies assumed the Saurichia/Ornithischia classification as true and so, the used characters and taxons were all focussed on this classification. However, this new study has pioneered in many aspects:

• It includes a larger number of species and taxons (many more than in previous investigations).
• Previous studies gave more importance to basal theropod and sauropodomorph dinosaurs (traditional saurischians), as they were the first dinosaurs to diversify, including few basal ornithischians.
• It has also included many dinosauromorph archosaurs (non-dinosaur taxons).
• Older studies had assumed many ornithischian characters to be symplesiomorphies (ancestral characters of all dinosaurs) and they only focused on a few synapomorphies (characters found in a monophyletic group).

This study has detached from many of the previous assumptions on dinosaur phylogeny and has analysed a large number of species and many characters not included in previous investigations. This has made the resulting evolutionary tree pretty different from the ones obtained before.

RESHAPING THE TREE

Then, how does the dinosaur’s evolutionary tree stand according to this hypothesis? Well, the matter is somewhat complex, even if the different groups are still divided in two orders:

• Order Saurischia which, according to this study, only includes sauropodomorphs and herrerasaurids (a group of carnivorous, non-theropod saurischians).
• The new order Ornithoscelida (bird-limbed) that includes the traditional ornithischians and theropods, which are no longer saurischians.

Keeping this in mind, let’s now see the characteristics that define these two orders.

Saurischians

The order Saurischia is almost the same, except that theropods are no longer part of this group. This order presents the original saurischian hip structure, which the dinosaurs’ ancestors also had. According to this new hypothesis,  herrerasaurids and sauropodomorphs are all included as saurischians.

Herrerasaurids (Herrerasauridae family) were a small group of basal saurischians that evolved towards meat-eating. That’s why for a long time it was thought that they were the sister-taxon of theropods, but it was later seen that they were found among the first saurischians. Even if they were pretty specialized, they were probably displaced by competition with other predators, appearing during the middle Triassic and becoming extinct at the end of it.

Photo by Brian Smith of a Herrerasaurus skeleton and model, from the Field Museum of Natural History of Chicago.

Herrerasaurids occupied a similar ecological niche as theropods. The new hypothesis implies that hypercarnivory (feeding exclusively on meat) evolved independently twice in dinosaurs, which makes some palaeontologist question it. Yet the herrerasaurid and theropod anatomy differed in some aspects, such as the anatomy of their hands (more generalistic in herrerasaurids) and the jaw structure.

The first sauropodomorphs were biped animals just like herrerasaurids, even if they were omnivorous. Yet, sauropodomorphs would end up becoming huge herbivorous quadrupeds with characteristic long necks.

Thecodontosaurus skeleton (by Qilong), a basal sauropodomorph and a reconstruction of Plateosaurus (from Walters, Senter & Robins) a more advanced one. Even if it cannot be appreciated in this image, sauropodomorphs would increase very their size very much during their evolution (Thecodontosaurus 2 metres, Plateosaurus up to 10 metres).

Ornithoscelidans

The new dinosaur order is Ornithoscelida, which groups theropods with ornithischians. This taxon is supported by more than twenty skeletal synapomorphies (derived characters shared by a clade), present both in basal theropods and ornithischians. Some of these characteristics include the presence of a gap between premaxillar and maxillar teeth (diastema) and the fusion of the ends of the tibia and the fibula into a tibiotarsus (even if these characteristics are only found on the most basal species).

Scheme from Baron et al. (2017) of the skulls of two basal ornithoscelidans, Eoraptor (a theropod, top) and Heterodontosaurus (an ornithischian, bottom).

Both theropods and the first ornithischians were bipedal animals. Also, the presence of heterodont teeth in the ancestral members of both groups leads us to think that the first ornithoscelidans were omnivorous, which would later specialise in feeding on meat and on plants (theropods and ornithopods respectively).

Reconstruction of the face of Daemonosaurus, one of the first theropods, by DeadMonkey8984.

A curiosity about the new classification is that accepting Ornithoscelida as a valid taxon, all feathered dinosaurs are put together into one group. Everyone knows that many theropods presented feathers (as they were the ancestors of birds) but, what most people don’t know is that feathers have also been found in some basal ornithischians and in more advanced ones too.

Reconstruction by Tom Parker of Kulindadromeus, a ornithischian which feathers have been proved to be present on most of its body.

KEEP INVESTIGATING

Then, is this hypothesis irrevocable? Well, no of course. Even if it’s pretty tempting to assume that the dinosaur’s natural history has been changed, we cannot say that from now on dinosaurs will be classified this way.

Dinosaur evolutionary tree according to Baron et al. (2017), in which we can see the different clades; Dinosauria (A), Saurischia (B) and Ornithoscelida (C).

Even if this study shows really interesting results about the origin of dinosaurs, we cannot dismiss hundreds of previous studies about this group of animals. We’ll have to remain alert to new articles that step by step will keep unveiling more information about the relationships between these Mesozoic reptiles. And that’s what’s so stimulating about biology, that there’s nothing sure! And that with new investigation techniques and new discoveries, little by little we learn more about the world around us.

Keep your mind open and keep investigating!

REFERENCES

The following sources have been consulted during the elaboration of this entry:

• G. Seeley (1887). On the Classification of the Fossil Animals Commonly Named Dinosauria. Proceedings of the Royal Society of London Vol. 43. Pp 165-171.
• Baron, Norman & Barret (2017). A new hypothesis of dinosaur relationships and early dinosaur evolution. Nature Vol. 543. Pp 501-506.
• Senter & Robins (2015). Resting Orientations of Dinosaur Scapulae and Forelimbs: A Numerical Analysis, with Implications for Reconstructions and Museum Mounts. Plos One.
• Scientific American. Ornithoscelida Rises: A New Family Tree for Dinosaurs.
• Cover image by Charles Knight.

The kingdom of the reptiles: what is a dinosaur?

Dinosaurs (superorder Dinosauria, “terrible reptiles”) are a group of reptiles which dominated all terrestrial ecosystems during the Mesozoic (Secondary Era or “the Age of Reptiles”). Even today, to most people there’s still some confusion over what a dinosaur is and what is not, and the term “dinosaur” is often used to refer to all the reptiles that evolved during the Secondary Era. In this entry I’ll try to give account of some of the different groups of reptiles that appeared during the Mesozoic and I’ll explain the classification of the different dinosaurian groups and some of their adaptations.

MESOZOIC REPTILES: DINOSAURS AND LOTS MORE

The rise of the dinosaurs was possible thanks to a mass extinction phenomenon which occurred 251 million years ago (Permian-Triassic extinction event). That phenomenon annihilated up to 96% of marine species and up to 70% of terrestrial species in that time, leaving lots of empty ecological niches to be inherited by new animal species.

Modified graphic from Rohde & Muller (2005) showing the great massive extinction. The darker zone corresponds to the Mesozoic period.

During the Triassic period (in the early Mesozoic) many different groups of reptiles evolved. One of these groups was the Dinosauria, which at that moment was far from being the dominant group of terrestrial animals. Some other reptilian groups of that time were the terrestrial rauisuchians (clade Rauisuchia) and fully aquatic groups like the sauropterygians (superorder Sauropterygia) and the ichthyopterygians (superorder Ichthyopterygia).

Reconstructions by Dmitry Bogdanov of Prestosuchus (a rauisuchian, top), Nichollsia (a suropterygian, left bottom) and Platypterigius (an ichthyopterygian, right bottom).

A second mass extinction in the late Triassic and the early Jurassic put an end to most of the dominant reptile groups, allowing the yet small dinosaurs to expand and evolve, along with some new groups like the crocodilomorphs (superorder Crocodylomorpha, ancestors of crocodilians), the flying pterosaurs (order Pterosauria).

Reconstructions by Dmitry Bogdanov of Dakosaurus (a crocodilomorph, top) and Scaphognathus (a pterosauria, bottom).

As we can see, dinosaurs are only one of many reptile groups that evolved during the Mesozoic. During the Jurassic period, dinosaurs diversified into many different groups, but they were mostly restricted to terrestrial ecosystems, which they would rule until their practical extinction 65 million years ago at the end of the Cretacic period.

DINOSAUR CLASSIFICATION

The first dinosaurs evolved around 231 million years ago during the mid-Triassic period. They were small in size and were characterized by their limb’s posture, which contrary to most reptiles, grew vertically elevating their body from the ground. That gave them more agility and a more active lifestyle.

Top: Skeleton of Eoraptor, one of the oldest known dinosaurs (Museum of Japan, photo by Kentaro Ohno). Bottom: Representation of the posture common among most reptiles (left) and the posture characteristic of dinosaurs (right).

Then dinosaurs diverged into two different orders: the Saurischia and the Ornithischia. These two groups were distinguished by the structure of their pelvis; saurischians conserved a pelvis more closely similar to that of the other reptiles, while the ornithischians evolved a pelvis superficially similar to that of modern birds.

Representation showing the structure of saurischian hips (left) and ornitischian hips (right). The animals represented are facing left.

ORNITISCHIA: BIRD HIPPED

Evolutionary tree of Ornithischia, modified by Zureks.

Ornithopoda (“bird feet”): Ornithopods were the most diverse group of Ornithischia, characterized by their three-toed feet similar to that of birds. They were herbivores that could combine bipedal and quadruped walking. Among them we can find the Iguanodon, one of the first dinosaurs to be discovered by science.

Iguanodon feet (right) and reconstruction by O. C. Marsh (1896).

Ornithopods acquired many different adaptations; some groups had duck-like bills to feed on aquatic vegetation, others developed specialized hands with a sharp thumb and an opposable little finger to grasp the plants they fed on. Many groups developed bony crests which are thought to be used both for species identification and for communication between members of the same species.

Reconstruction of Parasaurolophus (missing author), an ornithopod which presented a big hollow crest to amplify the sounds it made.

Marginocephalia (“fringed heads”): The so-called marginocephalians were a group of herbivorous dinosaurs related to the ornithopods characterized by a great cranial ossification. These can be divided into two separated groups:

Pachycephalosaurians (suborder Pachycephalosauria, “thick-headed reptiles”) were bipeds which had an extremely thick skull and a series of lateral osteoderms (keratin-covered ossifications) flanking it. It is believed that pachycephalosaurians resolved territorial fights and disputed reproductive rights via head-ramming, similar to goats.

Reconstruction of Pachycephalosaurus by Jordan Mallon.

The other members of the group are the ceratopsians (suborder Ceratopsia, “horned faces”), quadrupeds which presented; neck frills making their skulls look bigger and the “rostral bone”, which formed a beak-shaped structure on the mouth. Lots of species also developed facial horns which could protrude from the cheek-bone, the eyebrow or the neck frill.

Reconstruction of Rubeosaurus by Lukas Panzarin (left) and skull of Triceratops (right), photo by Zachi Evenor.

Thyreophora (“shield bearers”): This basal group of ornithischians was exclusively composed of quadruped herbivores characterized by the presence of heavy osteoderms that constituted their main defence. This group can be divided into:

Stegosaurians (suborder Stegosauria, “roofed reptiles”) were big herbivorous dinosaurs characterized by having two rows of dorsal osteoderms from the neck to the tail, which served as protection and helped them in their thermoregulation. Some species also developed caudal spines called “thagomizers” used as weapons to defend themselves from predators.

Mounted “thagomizer” at Denver Museum of Nature and Science (left) and reconstruction of Stegosaurus by Nobu Tamura (right).

Anchylosaurians (suborder Ankylosauria, “fused reptiles”) developed heavy bony armours that covered most of the body. Some of them, like the Ankylosaurus, developed big bony clubs at the end of the tail to fend off predators.

Reconstruction of Euoplocephalus by Nobu Tamura.

SAURISCHIA: REPTILE HIPPED

Evolutionary tree of Saurischia, modified by Zureks.

Sauropodomorpha (“reptile-shaped feet”): The sauropodomorphs are better known as the “long-necked dinosaurs”. That’s because they adapted to feed on the highest strata of vegetation.

Reconstruction of different sauropodomorphs (left to right): Camarasaurus, Brachiosaurus, Giraffatitan and Euhelopus.

Most species became large quadrupeds, with pillar-like legs similar to those of elephants and long necks to reach the leaves of the highest trees. Later species reached tremendous sizes, like the Amphicoelias which could grow up to 60 metres long.

Theropoda (“beast feet”): This last group is mostly known for two reasons. First of all is that this group includes some taxons of great predators like the Tyrannosauridae and Dromeosauridae families. The second reason is that theropods are the only dinosaurian group that includes living species, because modern birds are included in the suborder Theropoda.

Skeleton of Allosaurus from american museum collections (1915).

All theropods are bipedal and most of the Mesozoic species were carnivorous, with sharp replaceable teeth adapted to predation. Theropods present a saurischian pelvis but later on, birds evolved a hip structure more similar to that found in ornitischian dinosaurs.

Reconstruction by Davide Bonnadonna of the different clades that led to the aparition of birds (left to right): Neotheropoda, Tetanurae, Coelurosauria, Paraves and finally the Archaeopteryx, believed to be the first bird species that ever existed.

Some species had feathers to help thermoregulation. Birds from these groups evolved at the end of the Jurassic period.

REFERENCES

The following sources have been consulted in the elaboration of this entry:

• Cover photo: Gerhard Boeggemann.
• http://web.archive.org/web/20110612180650/http://www.hmnh.org/library/diplodocus/holland1910.html
• http://www.smithsonianmag.com/science-nature/tracking-the-origin-of-dinosaurs-70638045/?no-ist
• http://www.geol.umd.edu/~tholtz/G104/104Syl.html
• http://palaeo.gly.bris.ac.uk/palaeofiles/fossilgroups/dinosauria/index.html

If you enjoyed this article, please share it on social networks tospread it. The aim of the blog, after all, is to spread science and reachas many people as possible.

Licencia Creative Commons Atribución-NoComercial-CompartirIgual 4.0 Internacional.