The “Morphotype 1″ tunnel complex: points marked “a” represent tunnels, and points marked “b” signify vertical shafts. From Colombi et al., 2012.

Dinosaurs never cease to surprise. Even though documentaries and paleoart regularly restore these creatures in lifelike poses, the fact is that ongoing investigations into dinosaur lives have revealed behaviors that we never could have expected from bones alone. Among the most recent finds is that dinosaurs were capable of digging into the ground for shelter. Burrows found in Australia and Montana show that some small, herbivorous dinosaurs dug out cozy little resting places in the cool earth.

But when did dinosaurs develop burrowing behavior? The distinctive trace fossils found so far are Cretaceous in age, over 100 million years after the first dinosaurs evolved. That’s why a new PLoS One paper by paleontologist Carina Colombi caught my eye. In the Triassic rock of Argentina’s Ischigualasto Basin, Columbi and coauthors report, there are large-diameter burrows created by vertebrates that lived approximately 230 million years ago. Archaic dinosaurs such as Eoraptor and Herrerasaurus roamed these habitats–could dinosaurs be responsible for the burrows?

Colombi and colleagues recognized three different burrow forms in the Triassic rock. Two distinct types–differentiated by their diameter and general shape–were “networks of tunnels and shafts” that the authors attributed to vertebrates. The third type showed a different pattern of “straight branches that intersect at oblique angles” created by the burrowing organism and the plant life. The geology and shapes of the burrows indicate that they were created by living organisms. The trick is figuring out what made the distinct tunnel types.

In the case of the first burrow type, Colombi and collaborators propose that the structures were made by small, carnivorous cynodonts–squat, hairy protomammals. In the other two cases, the identities of the burrow makers isn’t clear. The second type included vertical shafts that hint at a vertebrate culprit. Dinosaurs would have been too big, but, Colombi and coauthors suggest, other cynodonts or the bizarre, ancient cousins of crocodiles–such as aetosaurs or protosuchids–could have created the burrows. Unless remains of these animals are found associated with the burrows, it is impossible to be sure. Likewise, the third type of trace might represent the activities of animals that burrowed around plant roots, but there is no clear candidate for the trace-maker.

As far as we know now, Triassic dinosaurs didn’t burrow. Even though they were not giants, they were still too large to have made fossils reported in the new research. Still, I have to wonder if predatory dinosaurs such as Herrerasaurus, or omnivores like Eoraptor, dug poor little cynodonts out of their burrows much like the later deinonychosaurs scratched after hiding mammals. There’s no direct evidence for such interactions, but, if small animals often sheltered from heat and drought in cool tunnels, perhaps predators tried to nab prey resting in their hiding places. One thing is for sure, though: we’ve only just started to dig beyond the surface of Triassic life.

References:

Colombi, C., Fernández, E., Currie, B., Alcober, O., Martínez, R., Correa, G. 2012. Large-Diameter Burrows of the Triassic Ischigualasto Basin, NW Argentina: Paleoecological and Paleoenvironmental Implications. PLoS ONE 7,12: e50662. doi:10.1371/journal.pone.0050662

Did Deinonychus and other “raptors” use their foot claws to restrain prey? Art by Emily Willoughby, image from Wikipedia.

When paleontologist John Ostrom named Deinonychus in 1969, he provided the spark for our long-running fascination with the “raptors.” Similar dinosaurs had been named before–Velociraptor and Dromaeosaurus were named four decades earlier–but the skeleton of Ostrom’s animal preserved a frightening aspect of the dinosaur that had not yet been seen among the earlier finds. The assembled remains of Deinonychus included the dinosaur’s eponymous “terrible claw”–a wicked, recurved weapon held off the ground on the animal’s hyperextendable second toe. Combined with the rest of the dinosaur’s anatomy, Ostrom argued, the frightening claw indicated that Deinonychus must have been a active, athletic predator.

But how did Deinonychus and its similarly-equipped relatives use that awful toe claw? The appendage looks fearsome, but paleontologists have not been able to agree on whether the claw was using for slashing, gripping, pinning, or even climbing prey. Some researchers, such as Phil Manning and collaborators, have even argued that the claws of Velociraptor and related dinosaurs were best suited to scaling tree trunks–a conclusion consistent with the contentious hypothesis that the ancestors of birds were tree-climbing dinosaurs.

Left hind foot of Deinonychus antirrhopus. Image from Wikipedia.

All this assumes that the claws of deinonychosaurs correspond to a special behavior, but can foot claw shapes really give away the habits of dinosaurs? That’s the question posed by a new PLoS One study by zoologist Aleksandra Birn-Jeffery and colleagues.

Based on observations of living animals, researchers have often tied particular claw shapes to certain behaviors–relatively straight, stubby claws likely belong to an animal that runs on the ground, while tree-climbing species have thin claws with small, sharp points. But nature isn’t quite so neat as to have a single, tell-tale claw shape for perchers, ground-runners, climbers, and predators. Even then, researchers don’t always interpret claw shapes the same way–depending on who you ask, the foot claws of the early bird Archaeopteryx either indicate that it was a climber or could only run on the ground.

To parse this problem, Birn-Jeffery and co-authors studied the geometry of the third toe claw–on dinosaurs, the middle toe claw–in 832 specimens of 331 species, together representing different lifestyles of birds, lizards, and extinct dinosaurs. The claw shapes didn’t strictly conform to particular behaviors. In the climber category, for example, the frill-necked lizard has lower claw curvature than expected, and, among predatory birds, the common buzzard, secretary bird, and greater sooty owl has less sharply recurved claws that anticipated for their lifestyle.

When the dinosaur data was dropped into the mix, the deinonychosaurs didn’t seem to fit in any single category. The sickle-clawed carnivores fell into the range shared by climbers, perchers, predators, and ground dwellers–these dinosaurs could be said to be anything from wholly terrestrial runners to perchers. And even though the researchers identified a general claw shape that corresponded to walking on the ground–deeper claws with less curvature–the dinosaurs did not strictly fit into this category alone.

Some dinosaurs, such as Microraptor, had claws that might have been suited to climbing. However, dinosaurs that we might regard as behaviorally similar showed differences–Velociraptor seemed to best fit the ground-dweller category, while the larger Deinonychus seemed to have claws more akin to those of predatory birds. This doesn’t mean that Microraptor was definitely a climber, or that Velociraptor wasn’t a predator. As the authors show, the different behavioral categories are not so easily distinguishable as previously thought, and saying that an animal definitely engaged in a particular behavior because of claw shape alone tempts oversimplification.

No wonder there has been such a range of interpretation about dinosaur foot claws! While the new study focused on the third toe claw rather than the famous, second deinonychosaur toe claw, the point of the analysis still applies. Claw geometry alone is not a reliable indicator of behavior. That’s to be expected–as the authors point out, claws are multi-functional, are are unlikely to represent just one type of behavior or habitat. Birds that use their claws to perch may also use them to kill prey, or birds that primarily live in the trees may also forage on the ground. Claw shape is constrained by different aspects of natural history, and reflect flexibility rather than strict adherence to a particular lifestyle. Deinonychosaur claws definitely hold clues to the natural history of dinosaurs, but drawing out those clues is a difficult, convoluted process.

Reference:

Birn-Jeffery, A., Miller, C., Naish, D., Rayfield, E., Hone, D. 2012. Pedal Claw Curvature in Birds, Lizards and Mesozoic Dinosaurs – Complicated Categories and Compensating for Mass-Specific and Phylogenetic Control. PLoS ONE. 7,12: e50555. doi:10.1371/journal.pone.0050555

The arms of Therizinosaurus–as yet, the rest of the dinosaur is missing. Photo by FunkMonk, image from Wikipedia.

The most famous set of arms in the history of dinosaurs belong to Deinocheirus–eight foot long appendages from a huge ornithomimosaur that roamed Mongolia around 70 million years ago. But the immense ostrich-mimic wasn’t the only giant omnivore of its time, nor the only one made famous by its imposing arms. About 20 years before the discovery of Deinocheirus, a joint Soviet-Mongolian expedition found extremely long, tapering claws and a few other bones from a gigantic reptile. The identity of this animal took decades to untangle.

Paleontologist Evgeny Maleev described the paltry remains in a 1954 paper. Based on rib fragments, a bone from the hand, and three claws, Maleev believed that he was looking a gargantuan turtle. He named the creature Therizinosaurus cheloniformis–roughly, the “turtle-like scythe lizard.”

The animal’s claws played a key role in the identification. No terrestrial animal had such claws, he argued. Such armaments “may have been originally used by the animal for cutting aquatic vegetation or for another function, constrained by movement and acquiring food.” And even though Maleev only had pieces to work with, he proposed that Therizinosaurus was about 15 feet long with claws at least three feet long. This aquatic, apparently armor-less turtle lived in a time of hadrosaurs, tyrannosaurs, and sauropods.

Therizinosaurus wasn’t recognized as a dinosaur until 1970. In that year, paleontologist Anatoly Konstantinovich Rozhdestvensky published a re-evaluation of Maleev’s fossils that found the rib to be from a sauropod dinosaur, but the hand bone and the claws to be from some as-yet-unknown theropod. This recognition only spawned a new mystery–what sort of theropod dinosaur was Therizinosaurus, and what was the creature doing with such fearsome claws?

More complete forelimb and shoulder material described by Rinchen Barsbold in 1976 showed that Therizinosaurus had extraordinarily robust arms–quite a departure from the trend seen in large carnivorous dinosaurs, in which the arms seemed to become smaller as skulls became more heavily-built. At a time when theropod was generally considered to be synonymous with “carnivorous dinosaur”, it’s not surprising that experts speculated that Therizinosaurus was a monstrous predator who used claws, rather than teeth, to slice up the hadrosaurs and sauropods of its time. That’s the way I encountered the dinosaur in the books I read as a kid–a little-known, Cretaceous hadrosaur-shredder.

What researchers didn’t recognize was that Therizinosaurus represented an entirely new variety of theropod dinosaur. More complete skeletons of related forms such as Segnosaurus, Erlikosaurus, Alxasaurus, and Beipiaosaurus revealed the presence of a previously-unknown group of dinosaurs with long necks, beaked mouths, fat bodies, and stout arms tipped with ludicrously-long claws. These were omnivorous or herbivorous dinosaurs, not carnivores, although paleontologists didn’t immediately agree on what lineage they belonged to. Some thought they might be aberrant ornithischians–on the opposite side of the dinosaur family tree from theropods–or strange variations on the sauropod theme. By the mid-90s, however, paleontologists recognized that these truly were theropods, and ones belonging to the maniraptoran group that also encompasses the strange alvarezsaurs, beaked and crested oviraptorosaurs, the sickle-clawed deinonychosaurs, and birds. This group of tubby, feathery dinosaurs became known as the therizinosaurs.

Although Maleev didn’t recognize it when he named Therizinosaurus, he had found one of the most spectacular dinosaurs of all time–a giant, fluffy, omnivorous dinosaur that challenged what we thought we knew about theropods. Still, our image of Theriziniosaurus relies on the skeletons of more complete, closely-related dinosaurs. So far, we only really know what the arms of this dinosaur looked like, and the hindlimb elements described in the 1980s may or may not belong to another creature. We’re still waiting for the true nature of this undoubtedly bizarre dinosaur to come into focus.

References:

Barsbold, R. 1976. New data on Therizinosaurus (Therizinosauridae, Theropoda) [translated]. In Devâtkin, E.V. and N.M. Ânovskaâ (eds.), Paleontologiâ i biostratigrafiâ Mongolii. Trudy, Sovmestnaâ Sovetsko−Mongol’skaâ paleontologičeskaâ kspediciâ, 3: 76–92.

Maleev, E.A. 1954. “New turtle−like reptile in Mongolia [translated].” Priroda, 1954, 3: 106–108.

Zanno, L. 2010. A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria: Maniraptora). Journal of Systematic Palaeontology. 8, 4: 503–543.

A restoration of Nyasasaurus in its Middle Triassic habitat, based on the known bones and comparisons to closely related forms. Art by Mark Witton.

For the past twenty years, Eoraptor has represented the beginning of the Age of Dinosaurs. This controversial little creature–found in the roughly 231-million-year-old rock of Argentina–has often been cited as the earliest known dinosaur. But Eoraptor has either just been stripped of that title, or soon will be. A newly-described fossil found decades ago in Tanzania extends the dawn of the dinosaurs more than 10 million years further back in time.

Named Nyasasaurus parringtoni, the roughly 243-million-year-old fossils represent either the oldest known dinosaur or the closest known relative to the earliest dinosaurs. The find was announced by University of Washington paleontologist Sterling Nesbitt and colleagues in Biology Letters, and I wrote a short news item about the discovery for Nature News. The paper presents a significant find that is also a tribute to the work of Alan Charig–who studied and named the animal, but never formally published a description–but it isn’t just that. The recognition of Nyasasaurus right near the base of the dinosaur family tree adds to a growing body of evidence that the ancestors of dinosaurs proliferated in the wake of a catastrophic mass extinction.

In March of 2010, Nesbitt and a team of collaborators named a leggy, long-necked creature from the same Triassic rock unit in Tanzania they named Asilisaurus kongwe. This creature was a dinosauriform–a member of the group from which the first true dinosaurs emerged–and, even better, appeared to to be the closest known relative to the Dinosauria as a whole. The find hinted that the dinosaur lineage had probably split off from a common ancestor by this time, meaning that the most archaic dinosaurs may have already existed by 243 million years ago. Roughly 249-million-year-old footprints of dinosauriforms found among Poland’s Holy Cross Mountains, described by different researchers later the same year, added evidence that the dinosauriforms were diversifying right from the beginning of the Triassic–not long after the catastrophe that decimated life on earth at the end of the Permian, around 252 million years ago.

Nyasasaurus is another step closer to the first true dinosaurs, and is just as old as Asilisaurus. To find an animal with such distinctive, dinosaur-like traits in the Middle Triassic indicates that dinosaurs already existed, or their ancestral stem was already established. Either way, Eoraptor and kin from South America can no longer be considered as the first dinosaurs, but rather a later radiation of forms. Even though our knowledge of Nyasasaurus is only fragmentary–the dinosaur is represented by a right humerus and a collection of vertebrae from two specimens–the dinosauriform nonetheless marks an additional 12 million years of dinosaur time that paleontologists are only just starting to explore.

Whether or not we ever achieve a more complete view of Nyasasaurus depends on the luck and the caprices of the fossil record. In the new paper, Nesbitt and coauthors point out that the rare, fragmentary nature of the remains found so far reflects that dinosauriforms–and early dinosaurs–were marginal parts of the ecosystems they inhabited. Dinosaurs did not dominate from the very start. They were  relatively meek, small animals that lived in a world ruled by archosaurs more closely related to crocodiles. It was only in the Late Triassic and Early Jurassic, when their archosaurian competition was diminished, that dinosaurs became dominant. That means the earliest dinosaurs and their ancestors are few and far between in the Triassic record.

Still, when I asked Nesbitt what Nyasasaurus might have looked like, he cited other dinosauriforms and early dinosaurs as templates to constrain our expectations. Nyasasaurus may have looked quite like Asilisaurus–a leggy animal with an elongated neck–although Nyasasaurus may have been bipedal. Future finds will test this idea, but the fact remains that paleontologists are closing in on what the very first dinosaurs were like. As paleontologists uncover more early dinosaurs and dinosauriforms, the dividing line between the two disappears–scientists are starting to smooth out the evolutionary transition between the first dinosaurs and their ancestors. What role Nyasasaurus played in that transformation isn’t yet clear, but the creature is a signal that over 10 million years more of uncharted dinosaur history remains in the rock.

References:

Nesbitt, S., Sidor, C., Irmis, R., Angielczyk, K., Smith, R., Tsuji, L. 2010. Ecologically distinct dinosaurian sister group shows early diversification of Ornithodira. Nature 464, 7285: 95–98. doi:10.1038/nature08718

Nesbitt, N., Barrett, P., Werning, S., Sidor, C., Charig, A. 2012. The oldest dinosaur? A middle Triassic dinosauriform from Tanzania. Biology Letters. http://dx.doi.org/10.1098/rsbl.2012/0949

The arms of the huge ornithomimosaur Deinocheirus. How did such herbivorous theropods get to be so big? Photo by Eduard Solà, image from Wikipedia.

When I was first becoming acquainted with dinosaurs in the mid 1980s, “theropod” was synonymous with “carnivorous dinosaur.” Large or small, from Tyrannosaurus to Compsognathus, every theropod I knew of sustained itself on the flesh of other organisms. But it was just about that time that new discoveries and analyses revealed that many theropod dinosaurs were omnivores, or even herbivores. The ostrich-like ornithomimosaurs, beaked oviraptorosaurs and utterly bizarre therizinosaurs, in particular, embodied a switch from an ancestral meat-filled diet to one more reliant of fruit and foliage. Not only that, but these herbivorous theropods grew almost as large as the biggest carnivores–the ornithomimosaur Deinocheirus, the ovriraptorosaur Gigantoraptor and Therizinosaurus were all enormous Cretaceous dinosaurs. But why did these plant-chomping dinosaurs become giants?

In the latest of a spate of papers considering herbivorous theropods, paleontologists Lindsay Zanno and Peter Makovicky paired evolutionary trees with mass estimates derived from femora lengths and a bit of number crunching to see if there was any distinct evolutionary pattern that might explain why Deinocheirus and similar herbivorous theropods grew to such large sizes. Were these Late Cretaceous dinosaurs just the culmination of an evolutionary trend towards ever-larger body size–called Cope’s Rule–or was something else at work?

Zanno and Makovicky didn’t find any sign of directional selection for larger body size. Even though the earliest representatives of the ornithomimosaurs, oviraptorosaurs and therizinosaurs in Asia were much smaller than their Late Cretaceous relatives, the paleontologists point out that this signal has probably been biased by preservation. The 125-million-year-old deposits that contain small members of these groups seem to be skewed towards “mid-sized vertebrates,” the authors point out, and don’t seem to preserve larger dinosaurs that might belong to the same lineages. Indeed, therizinosaurs of about the same age from North America, such as Falcarius, were larger than species in Asia, meaning that herbivorous dinosaurs might have occupied a range of body sizes and evolved larger body sizes at multiple intervals. There was no simple, straight-line trend of bigger and bigger bodies through time.

Nor did a herbivorous lifestyle alone seem to account for gigantism among these dinosaurs. Even though big herbivores gain particular benefits from their size in terms of breaking down tough, low-quality foods more efficiently, Zanno and Makovicky doubt that this relationship drove the evolution of increased body size in the dinosaurs. Instead, they favor “passive processes” that might be tied to ecology and whether these dinosaurs were omnivores more than herbivores. And, as the paleontologists stress, the pattern relies on how complete we think the dinosaur record is. Some ecosystems might be preferentially preserving larger or smaller dinosaurs, which has the potential to skew the big picture. While Zanno and Makovicky ruled out some possibilities, we still don’t really know what accounts for the multiple herbivorous theropod growth spurts.

Post-Script: After four years working with Smithsonian magazine’s wonderful crew, and over 1,000 posts about various aspects of dinosauriana, it’s time for me to move on. I’ll be leaving Dinosaur Tracking next month. Don’t fret, I’ll still be digging into dinosaur science, but I’ll be at a new blog elsewhere on the web (stay tuned for details). I am deeply indebted to my editors Brian Wolly, Sarah Zielinski and, of course, Laura Helmuth (now doing a great job at Slate), as well as the rest of the Smithsonian staff for inviting me to come here and geek out about dinosaurs every day. And many thanks to all of you–the readers and commenters who have helped make this blog a success. You have all made blogging for Dinosaur Tracking an absolute pleasure.

Reference:

Zanno, L., Makovicky, P. 2012. No evidence for directional evolution of body mass in herbivorous theropod dinosaurs. Proceedings of the Royal Society B. 280. doi: 10.1098/rspb.2012.2526

An illustration showing the only known bones from Genyodectes. Art in Woodward, 1901, image from Wikipedia.

Paleontologists are naming new dinosaurs at an astonishing rate. In fact, they’re only just begun to skim the diversity of dinosaurs preserved in the world’s Mesozoic formations–hundreds of unknown dinosaur species are undoubtedly hiding in stone. But even among dinosaurs that have a formalized identity, there are many that we know relatively little about. Among them is Genyodectes serus, a carnivorous dinosaur known from the tip of its fearsome jaws and little else.

Though it’s far from being a household name, Genyodectes holds a significant place in the history of South American paleontology. Aside from a tooth found a few years before, the incomplete fossil snout of a Genyodectes was the first definitive non-avian theropod dinosaur found on the continent. As described by paleontologist A.S. Woodward in 1901, the remains of Genyodectes mostly consisted of pieces from the lower jaw, as well as the premaxillary bones and fragments of the maxillary bones in the upper jaw, all of which sported frighteningly long, curved teeth.

There was never any question that Genyodectes was a theropod dinosaur. All the principally carnivorous dinosaurs that we know of fell among various branches of this group. But what sort of theropod dinosaur was it? During the 20th century, different paleontologists proposed that it was a megalosaurid (then a generalized term for big predatory dinosaurs), a tyrannosaur or, after additional theropod remains started to come out of South America, one of the stubby-armed abelisaurids.

After the specimen was given a fresh cleaning, paleontologist Oliver Rauhut reexamined Genyodectes with an eye towards what the dinosaur was and where it came from. Based on notes and geological details, Rauhut proposed that the dinosaur was found in Cañadón Grande in Argentina’s Chubut province in a Cretaceous deposit that probably dates to around 113 million years old. And, based on the limited remains, Rauhut hypothesized that Genyodectes was a later, southern cousin of North America’s Ceratosaurus. While the only known specimen of Genyodectes was cracked and damaged by erosion, the size and the anatomy of the dinosaur’s teeth most closely resembled that of Ceratosaurus–especially in having extremely long teeth in the maxilla. Given this relationship, we might expect that Genyodectes had some kind of skull ornamentation like the nasal and eye horns of its cousin, but we need more fossils to be sure.

Reference:

Rauhut, O. 2004. Provenance and anatomy of Genyodectes serues, a large-toothed ceratosaur (Dinosauria: Theropods) from Patagonia. Journal of Vertebrate Paleontology. 24, 4: 894-902

This famous Edmontosaurus skeleton was found with intricate traces of skin over much of its body. Image in Osborn, 1916, from Wikipedia.

Last week, I wrote about attempts by paleontologist Phil Bell and colleagues to extract biological secrets from fossilized traces of dinosaur skin. Among the questions the study might help answer is why so many hadrosaurs are found with remnants of their soft tissue intact. Specimens from almost every dinosaur subgroup have been found with some kind of soft tissue preservation, yet, out of all these, the shovel-beaked hadrosaurs of the Late Cretaceous are found with skin impressions and casts most often. Why?

Yale University graduate student Matt Davis has taken a stab at the mystery in an in-press Acta Paleontologica Polonica paper. Previously researchers have proposed that the abundance of hadrosaur skin remnants is attributable to large hadrosaur populations (the more hadrosaurs there were, the more likely their skin might be preserved), the habits of the dinosaurs (perhaps they lived in environments where fine-resolution fossilization was more likely) or some internal factor that made their skin more resilient after burial. to examine these ideas, Davis compiled a database of dinosaur skin traces to see if there was any pattern consistent with these ideas.

According to Davis, the large collection of hadrosaur skin fossils isn’t attributable to their population sizes or to death in a particular kind of environment. The horned ceratopsid dinosaurs–namely Triceratops–were even more numerous on the latest Cretaceous landscape, yet we don’t have as many skin fossils from them. And hadrosaur skin impressions have been found in several different kinds of rock, meaning that the intricate fossilization occurred in multiple types of settings and not just sandy river channels. While Davis doesn’t speculate about what made hadrosaurs so different, he proposes that their skin might have been thicker or otherwise more resistant than that of other dinosaurs. A sturdy hide might have offered the dinosaurs protection from injury in life and survived into the fossil record after death.

Still, I have to wonder if there was something about the behavior or ecology of hadrosaurs that drew them to environments where there was a greater chance of rapid burial (regardless of whether the sediment was sandy, silty or muddy). And the trouble with ceratopsids is that they have historically been head-hunted. Is it possible that we’ve missed a number of ceratopsid skin traces because paleontologists have often collected skulls rather than whole skeletons? The few ceratopsid skin fossils found so far indicate that they, too, had thick hides ornamented with large, scale-like structures. Were such tough-looking dinosaur hides really weaker than they appear, or is something else at play? Hadrosaurs may very well have had extra-sturdy skin, but the trick is testing whether that characteristic really accounts for the many hadrosaur skin patches resting in museum collections.

Reference:

Davis, M. 2012. Census of dinosaur skin reveals lithology may not be the most important factor in increased preservation of hadrosaurid skin. Acta Paleontologica Polonica http://dx.doi.org/10.4202/app.2012.0077

Osborn, H. 1916. Integument of the iguanodon dinosaur Trachodon. Memoirs of the American Museum of Natural History. 1, 2: 33-54

Sternberg, C.M. 1925. Integument of Chasmosaurus belli. The Canadian Field Naturalist. XXXIX, 5: 108-110

Partial bone shafts found in Late Triassic rock in England might represent a sauropodomorph, similar to this Plateosaurus, or an entirely different kind of creature. Photo by FunkMonk, image from Wikipedia.

Dinosaur giants are among the most famous Mesozoic celebrities. Yet the dinosaur growth spurt didn’t start just as soon as Eoraptor and kin evolved. For most of the Triassic, the first act in their story, dinosaurs were small and gracile creatures, with the first relatively large dinosaurs being the sauropodomorphs of the Late Triassic. Even then, Plateosaurus and kin didn’t come close to the truly enormous sizes of their later relatives–such as Diplodocus and Futalognkosaurus. Discerning when dinosaurs started to bulk up is difficult, however, and made all the more complicated by a set of enigmatic bones found in England.

The fossils at the heart of the in-press Acta Palaeontologica Polonica study, as described by University of Cape Town paleontologist Ragna Redelstorff and coauthors, have been known to researchers for a long time. During the mid-19th century, naturalists described at least five large, incomplete shafts found in the Late Triassic rock of southwest England’s Aust Cliff. Two of these fossils were later destroyed, but, drawing from the surviving specimens and illustrations of the lost bones, paleontologist Peter Galton proposed in 2005 that they came from large dinosaurs that lived over 200 million years ago. In particular, two of the bones resembled stegosaur bones, which would have extended the origin of the armored dinosaurs further back than previously thought.

Not everyone agreed with Galton’s proposal. The bone shafts could be from as-yet-unknown sauropods, some paleontologists argued, while other researchers pointed out that the lack of distinctive features on the bones were unidentifiable beyond the level of “tetrapod” (the major group of vertebrates descended from fish with limbs, similar to Tiktaalik). The bones came from big creatures–possibly more than 20 feet long, based on comparisons to other fossils–but the identity of the Aust Cliff animals is unknown.

Since the outside of the bone shafts provide so little information about their identity, Redelstorff and collaborators looked to the microstructure of two specimens for new clues. While the histological evidence appears to show that the sampled bones belonged to the same species, the authors argue, each individual shows different growth strategies. One bone shaft came from a slightly bigger, rapidly growing individual, while the smaller bone represents an older animal that regularly experienced temporary halts in growth (visible as lines called LAGs in the bone). Why this should be so isn’t clear, but Redelstorff and coauthors suggest individual variation, differences between the sexes or ecological factors as possible causes.

But what sort of animals were the Aust Cliff creatures? When the researchers compared their sample with three kinds of dinosaurs–sauropods, archaic sauropodomorphs and stegosaurs–and Triassic croc cousins called pseudosuchians, the pseudosuchians seemed to be the closest match. Indeed, while the researchers concluded that the “Aust Cliff bones simply do not offer a good match with any previously described histologies,” the specimens appeared to share more in common with those of croc-line archosaurs than with dinosaurs.

This isn’t to say that the Aust Cliff animals were definitely large psuedosuchians, like the recently named Smok. As the researchers point out, the specimens contained a type of bone tissue not previously seen in pseudosuchians–either these animals were not pseudosuchians, or these pseudosuchians were a previously unknown histology. And, Redelstorff and collaborators point out, the bones might be attributable to a sauropodomorph named Camelotia that is found in the same deposits. Studying the bone microstructure of Smok and Camelotia for comparison would be a logical next step in efforts to narrow down the identity of the Aust Cliff animals. Until then, this early “experiment” in gigantism–as Redelstorff and colleagues call it–remains an unresolved puzzle.

Still, the study highlights the importance of building a deep database of paleohistological samples. Had the researchers sampled just one bone, they may have come to the conclusion that all bones of that type would exhibit the same life history–either rapid, continuous growth or a stop-and-go pattern, depending on which they studied. Together, the bones show variations in the natural history of what is presumably the same species, which brings up the question of how quirks of environment, biology and natural history are recorded in bone. If we are going to understand the biology of dinosaurs and other prehistoric animals, we need to cut into as many bones as we can to understand how variable and biologically flexible the creatures truly were.

Reference:

Redelstorff , R., Sander, P., Galton, P. 2012. Unique bone histology in partial large bone shafts from Aust Cliff (England, Upper Triassic): an early independent experiment in gigantism. Acta Palaeontologica Polonica  http://dx.doi.org/10.4202/app.2012.0073

The giant sauropod Futalognkosaurus (at left) with some of its Cretaceous neighbors. Art by Maurilio Oliveira.

Which was the biggest dinosaur ever? We don’t know. Even though the size-based superlative draws a great deal of attention, paleontologists have uncovered so many scrappy sauropod skeletons that it’s difficult to tell who was truly the most titanic dinosaur of all. But, among the current spread of candidates, Futalognkosaurus dukei is one of the most complete giant dinosaurs yet found.

Discovered in 2000, and named in 2007 by Universidad Nacional del Comahue paleontologist Jorge Calvo and colleagues, Futalognkosaurus was one of many dinosaurs found in an exceptionally rich, roughly 90-million-year0old deposit in northwest Argentina. From fossil plants to pterosaurs, fish and dinosaurs, the one site entombed vestiges of a vibrant Cretaceous ecosystem. And, on that landscape, no dinosaur was as grand the newly named titanosaur.

Contrary to what you might expect given their skeletal sturdiness, the biggest sauropods are often found as partial skeletons. Our knowledge of Argentinosaurus, Puertasaurus, Supersaurus, Diplodocus hallorum and other giants is frustratingly incomplete, and figuring out how large they truly were relies on estimation from more complete representatives of other species.

The lack of complete tails from these dinosaurs makes the matter even more problematic. Dinosaur tails varied in length from individual to individual, and different subgroups had proportionally longer or shorter tails. In the case of Diplodocus hallorum, for example, a great deal of the dinosaur’s estimated  100-foot-plus length comes from the fact that other Diplodocus species had very long, tapering tails.

We don’t really know how long Futalognkosaurus was because, with the exception of a single vertebra, the dinosaur’s tail is entirely missing. Nevertheless, the sauropod that Calvo and coauthors described is remarkable for encompassing the entire neck, back and associated ribs, and the majority of the hips. Together, these elements represent over half the skeleton and comprise the most complete giant sauropod individual yet known.

Even if skeletal incompleteness keeps us from knowing exactly how big Futalognkosaurus was, the collected bones can leave no doubt that this was a truly enormous dinosaur. Calvo and coauthors estimated that the whole animal stretched between 105 and 112 feet in length, which would put it in the same class as the more famous (and less complete) Argentinosaurus. As the paleontologists at SV-POW! said when they posted images of Futalognkosaurus bones next to Juan Porfiri, who helped describe the dinosaur, there’s no doubt that the sauropod was “darned big.” The challenge is finding and filling in the parts of the dinosaur’s body that have not yet been found. There will undoubtedly be other challengers for the title of biggest dinosaur, but, for now, Futalognkosaurus remains our most detailed representative of the biggest of the big.

References:

Calvo, J., Porfiri, J., González-Riga, B., Kellner, A. 2007. A new Cretaceous terrestrial ecosystem from Gondwana with the description of a new sauropod dinosaur. Anais da Academia Brasileira de Ciências. 79, 3: 529-541

Calvo, J., Porfiri, J., González-Riga, B., Kellner, A. 2007. Anatomy of Futalognkosaurus dukei Calvo, Porfiri, González Riga, & Kellner, 2007 (Dinosauria, Titanosauridae) from the Neuquen Group, Late Cretaceous, Patagonia, Argentina. Arquivos do Museu Nacional 65, 4: 511–526.

Novas, F. 2009. The Age of Dinosaurs in South America. Bloomington: Indiana University Press. pp. 201-202

Only hindlimb elements of Alnashetri are known so far, but, based on the dinosaur’s relationships, the tiny theropod probably looked something like this Alvarezsaurus. Photo by FunkMonk, image from Wikipedia.

Many dinosaurs have gained fame thanks to their gargantuan size. A creature in the form of a dipldodocid or tyrannosaur would be wonderful at any scale, but the fact that Apatosaurus was an 80-foot-long fern-sucker and Tyrannosaurus was a 40-foot carnivore make their skeletal frames all the more spectacular. Even as an adult, long after my first encounter with their bones at the American Museum of Natural History in New York City, I still feel tiny when I look up at what’s left of the great dinosaurs.

But not all non-avian dinosaurs were gigantic. There were 100-foot giants, like the sauropod Argentinosaurus, but there were also pigeon-sized theropods such as the strikingly-colored Anchiornis. Indeed, a significant part of how we know dinosaurs really ruled the earth is because they occupied such a wide range of body sizes–from the breathtakingly large to the diminutive. And, earlier this month, Field Museum of Natural History paleontologist Peter Makovicky and colleagues added a previously unknown tiny dinosaur to the ever-growing roster of Mesozoic species.

Named Alnashetri cerropoliciensis, the small dinosaur is mostly a mystery. All that we know of it, Makovicky and coauthors report, are a set of articulated hindlimbs from a single animal found in the roughly 95-million-year-old rock of La Buitrera, Argentina. (The dinosaur’s genus name, the paper says, means “slender thighs” in a dialect of the Tehuelchan language.) Yet those appendages contain enough clues about the dinosaur’s identity that the researchers were able to figure out that the specimen represented a new species of alvarezsaur–one of the small, possibly ant-eating dinosaurs recognizable by their short, stout arms and long skulls set with tiny teeth. While the paleontologists acknowledge that their Alnashetri specimen might be a juvenile, Makovicky and collaborators estimate that the dinosaur was comparable to its relative Shuvuuia in size–about two feet long.

How Alnashetri resembled other alvarezsaurs, and where it departed in form, will have to wait for more complete specimens. Further research is also needed to narrow down when this dinosaur lived, but for the moment, Alnashetri appears to be the oldest alvarezsaur found in South America. If only we knew more of this dinosaur! As Makovicky and coauthors conclude, “continued fieldwork and future discoveries hopefully will provide more information on the anatomy of Alnashetri and allow a more definitive evaluation of its affinities and its significance for understanding biogeography and evolutionary trends such as body size evolution within alvarezsaurids.” At least the enigma has a name.

Reference:

Makovicky, P., Apesteguía, S., Gianechini, F. 2012. A new coelurosaurian theropod from the La Buitrera fossil locality of Rio Negro, Argentina. Fieldiana Life and Earth Sciences, 5: 90-98

A reconstruction of an Einiosaurus skull in a ceratopsid gallery at the Natural History Museum of Los Angeles. Photo by Maarten Heerlien, image from Wikipedia.

Xenoceratops was a gnarly-looking ceratopsid. There’s no doubt about that. Much like its horned kin, the dinosaur sported a distinctive array of head ornaments from the tip of its nose to the back of its frill. But that’s hardly the entire story behind this newly named dinosaur.

Contrary to many news reports that focused almost entirely on the dinosaur’s appearance, the real importance of Xenoceratops is in its geological and evolutionary context. The dinosaur is the first identifiable ceratopsid from the relatively unexplored Foremost Formation in Canada, and the creature appears to be at the base of a major horned dinosaur subdivision called centrosaurines. While the dinosaur’s name is certainly aesthetically pleasing, Knight Science Journalism Tracker watchdog Charlie Petit rightly pointed out that the ceratopsid isn’t really any more or less fantastic-looking than close cousins such as Styracosaurus, Spinops and Pachyrhinosaurus. The real importance of the dinosaur–a new data point in an ongoing investigation of a little-known part of the Cretaceous–was obscured by a narrowed focus on the dinosaur’s spiky headgear.

Dinosaurs are perpetually struggling to find context in news reports. Indeed, Xenoceratops is just the latest example and not an anomaly. Theropod dinosaurs are often introduced as Tyrannosaurus rex relatives, even when they’re not particularly closely related to the tyrant king, and journalists had such a fun time giggling over calling Kosmoceratops the “horniest dinosaur ever” that the clues the ceratopsid offered about dinosaur evolution in western North America were almost entirely overlooked. Reports on newly discovered dinosaurs usually contain the vital statistics of when the animal lived, where it was found, how large it was and whatever feature strikes our immediate attention, but the tales dinosaurs have to tell about life, death, evolution and extinction are rarely pulled out by journalistic storytellers.

Fossils don’t divulge their stories all at once, though. Paleontologists spend years drawing paleobiological secrets from dinosaur bones–who was related to whom, grand evolutionary patterns and rates of faunal turnover, and how the animals actually lived. These slowly emerging lines of evidence don’t often receive the same degree of attention. The discovery of a new bizarre species immediately garners journalistic attention, but once the dinosaur has been added to the roster, details about the animal’s life are often forgotten unless the creature earns a new superlative or has been found to have some tenuous connection to T. rex.

Rather than just gripe, though, I want to highlight how discovering and naming a dinosaur is only the initial step in paleontology’s effort to reconstruct prehistoric life. Consider Einiosaurus procurvicornis, a dinosaur I’m selecting here for no other reason than I promised a friend that I’d write about the dinosaur soon.

In 1995, paleontologist Scott Sampson named Einiosaurus from remains of multiple individuals strewn through two bonebeds discovered in Montana’s Late Cretaceous Two Medicine Formation. A geologically younger relative of Xenoceratops by about 4 million years, adults of this ceratopsid species are immediately recognizable by a forward-curved nasal horn, a pair of long, straight spikes jutting from the back of the frill and a suite of more subtle cranial ornaments.

Even before Einiosaurus had a name, though, researchers knew that the collected bones of this dinosaur presented a rich fossil database. Five years before Sampson’s paper, paleontologist Raymond Rogers drew on the two ceratopsid bonebeds to argue that multiple individuals of the species had died in prehistoric droughts. Rather than being places where the bodies of solitary animals accumulated over time, Rogers proposed, the rich assemblages recorded mass mortality events which claimed young and old ceratopsids alike.

The bone assemblages and their geological context outline many tragic dinosaur deaths. But clues about dinosaur lives are preserved inside those bones. For her master’s work at Montana State University, paleontologist Julie Reizner examined the bone microstructure of 16 Einiosaurus tibiae from a single bonebed to reconstruct how these dinosaurs grew and outline their population structure.

The research is still awaiting publication in a journal, but according to Reizner’s 2010 thesis and a poster she presented at the annual Society of Vertebrate Paleontology meeting last month, the histological evidence indicates that these horned dinosaurs grew rapidly until about three to five years of age, when their growth significantly slowed. The dinosaurs did not cease growing entirely, but, Reizner hypothesizes, the slowdown might represent the onset of sexual maturity. Additionally, all the dinosaurs in her sample were either juveniles or subadults–there were no infants or adults (or dinosaurs that had reached skeletal maturity and ceased growing). Even among the two groups, there doesn’t seem to be a continuum of sizes but instead a sharper delineation between juveniles and subadults. If this Einiosaurus bonebed really does represent a herd or part of a herd that died at about the same time, the age gap might mean that Einiosaurus had breeding seasons that occurred only during a restricted part of the year, thus creating annual gaps between broods.

Restored soft tissue profile of Einiosaurus, modified from Hieronymus et al., 2009.

Other researchers have drawn from different bony indicators to restore what the faces of Einiosaurus and similar dinosaurs would have looked like. While the underlying ornamental structures are still prominent in ceratopsid skulls, the horns, bosses and spikes would have been covered in tough sheaths. Thus, in 2009, Tobin Hieronymus and colleagues used the relationship between facial integument and bone in living animals to reconstruct the extent of skin and horn on ceratopsids. While the preservation of the Einiosaurus material frustrated their efforts to detect all the skin and horn structures on the skull, Hieronymus and colleagues confirmed that the nasal horn was covered in a tough sheath and that Einiosaurus had large, rounded scales over the eyes. Artists can’t simply stretch skin over the dinosaur’s skull in restorations–the bone itself shows the presence of soft tissue ornamentation that rotted away long ago.

As with most dinosaur species, we still know relatively little about the biology of Einiosaurus. We are limited to what is preserved in the rock, the technologies at our disposal and the state of paleontological theory. All the same, Einiosaurus is much more than a pretty face. The dinosaur was part of a rich, complex Cretaceous ecosystem, and one in a cast of billions in earth’s evolutionary drama. To me, at least, that is the most entrancing aspect of paleontology. We have only barely begun to plumb the depths of dinosaur diversity, and researchers will continue to introduce us to new species at a breakneck pace, but the true wonder and joy of paleontology lies in pursuing questions about the lives of animals we’ll sadly never observe in the flesh.

References:

Hieronymus, T., Witmer, L., Tanke, D., Currie, P. 2009. The facial integument of centrosaurine ceratopsids: Morphological and histological correlates of novel skin structures. The Anatomical Record 292: 1370-1396

Reizner, J. 2010. An ontogenetic series and population histology of the ceratopsid dinosaur Einiosaurus procurvicornis. Montana State University master’s thesis: 1-97

Rogers, R. 1990. Taphonomy of three dinosaur bone beds in the Upper Cretaceous Two Medicine Formation of northwestern Montana: evidence for drought-related mortality. PALAIOS 5 (5): 394–413.

Sampson, S. 1995. Two new horned dinosaurs from the Upper Cretaceous Two Medicine Formation of Montana; with a phylogenetic analysis of the Centrosaurinae (Ornithischia: Ceratopsidae). Journal of Vertebrate Paleontology 15 (4): 743–760.

The reconstructed skull of Eotriceratops. The actual specimen is not complete, but, based on the recovered elements and the dinosaur’s relationships, we know the dinosaur would have looked similar to Triceratops. Photo by Roland Tanglao, image from Wikipedia.

Triceratops is among the most cherished of dinosaurs. Even that might be a bit of an understatement. Fossil fans threw a conniption when they mistakenly believed that paleontologists were taking the classic “three-horned face” away, after all. But where did the charismatic chasmosaurine come from? Triceratops didn’t simply spring from the earth fully formed–the ceratopsid was the descendant of a long tail of evolutionary forerunners. And in 2007, paleontologist Xiao-chun Wu and collaborators described a 68-million-year-old dinosaur that might represent what one of the close ancestors of Triceratops was like–Eotriceratops.

In 2001, while on an expedition to search the Horseshoe Canyon Formation around the Dry Island Buffalo Jump Provincial Park in Alberta, Canada, Glen Guthrie discovered the partial skeleton of a huge ceratopsid dinosaur. This was the first identifiable dinosaur skeleton found in the top quarter of the formation, and, as Wu and coauthors later argued, the bones represented a new species. They called the animal Eotriceratops xerinsularis.

Paleontological devotees know that “eo” translates to “dawn.” The tiny mammal Eohippus was the “dawn horse” (which Victorian anatomist Thomas Henry Huxley famously characterized for the steed of a tiny “Eohomo“), and there are plenty of dawn dinosaurs such as Eoraptor, Eodromaeus, Eobrontosaurus and Eolambia. The prefix is a kind of honorific, used to indicate the hypothesized beginning of a major lineage or significant change. In the case of Eotriceratops, Wu and colleagues found that the dinosaur was the oldest known member of the evolutionary ceratopsid club containing Triceratops, Torosaurus and Nedoceratops (which, depending on who you ask, may or may not be the same dinosaur).

The individual Guthrie found had fallen apart between death and burial. Aside from some vertebrae, ribs and ossified tendons, the scattered specimen was primarily represented by a dis-articulated skull. When reconstructed, though, the head of Eotriceratops stretched almost ten feet long–about a foot longer than the largest-known Triceratops skull. And while different in some characteristics, Eotriceratops had the same three-horned look of its later relatives Triceratops and Torosaurus.

This isn’t to say that Eotriceratops was directly ancestral to Triceratops, Torosaurus, Nedoceratops or whatever combination of the three paleontologists ultimately settle on. Eotriceratops could be the closest relative of Triceratops to the exclusion of Torosaurus, which would support the idea that those later dinosaurs were separate genera.  Then again, Wu and coauthors pointed out that Eotriceratops might be the most basal member of the subgroup, which would make sense given that it was older than the other three genera. In either case, Eotriceratops can give us a rough idea of the Triceratops and Torosaurus prototype, but we lack the resolution to know if Eotriceratops was ancestral to any later dinosaur. Eotriceratops undoubtedly had some significance in the evolution of the last three-horned dinosaurs, but we need many more fossils to know this little-known dinosaur’s role in the story. Every dinosaur paleontologists find comes with a handful of answers and a myriad of new mysteries.

This post is the latest in the Dinosaur Alphabet series.

Reference:

Wu, X., Brinkman, D., Eberth, D., Braman. 2007. A new ceratopsid dinosaur (Ornithischia) from the uppermost Horseshoe Canyon Formation (upper Maastrichtian), Alberta, Canada. Canadian Journal of Earth Sciences 44: 1243-1265

A restoration of Xenoceratops by Danielle Dufault, courtesy David Evans.

It’s a good time to be a ceratopsid fan. Since 2010, paleontologists have introduced us to a slew of previously unknown horned dinosaurs, and new discoveries are continuing to trickle out of field sites and museums. Long-forgotten specimens and unopened plaster jackets, especially, have yielded evidence of ceratopsids that researchers overlooked for decades, and this week Royal Ontario Museum paleontologist David Evans and colleagues have debuted yet another horned dinosaur that was hiding in storage.

The Late Cretaceous exposures of Alberta, Canada’s Belly River Group are rich with ceratopsid fossils. For over a century, paleontologists have been pulling bones of the fantastically ornamented dinosaurs from these badlands. Yet most of the ceratopsids from this area have been found in the Dinosaur Park Formation, and researchers have paid less attention to the older strata of the Oldman and Foremost Formations nearby.

The Foremost Formation, in particular, has received little attention because diagnostic dinosaur remains seem to be rare within its depths, but a few notable specimens have been found in this slice of time. In 1958, paleontologist Wann Langston, Jr. and a crew from what is now the Canadian Museum of Nature pulled fragments of several ceratopsid specimens from 78-million-year-old deposits in the Foremost Formation. Those bones and skeletal scraps sat in collections for years until they caught the eye of Evans and Michael Ryan (the lead author of the new study) as they made the research rounds for their Southern Alberta Dinosaur Project. Although fragmentary, Langston’s fossils were from a new genus of ceratopsid.

Evans, Ryan and Kieran Shepherd have named the dinosaur Xenoceratops foremostensis in their Canadian Journal of Earth Sciences study. The dinosaur’s name–roughly “alien horned face”–isn’t a testament to the ceratopsid’s distinctive array of horns but to the rarity of horned dinosaur fossils within the Foremost Formation. Indeed, despite Danielle Dufault’s gorgeous restoration of the dinosaur, Xenoceratops is presently represented by skull fragments from several individuals. The researchers behind the new paper pieced them together to create a composite image of what this dinosaur must have looked like, and, in turn, discern its relationships.

Based upon the anatomy of one of the dinosaur’s frill bones–the squamosal–Evans and coauthors are confident that Xenoceratops was a centrosaurine dinosaur. This is the ceratopsid subgroup containing other highly decorated genera such as Styracosaurus, Spinops, Centrosaurus and another dinosaur given a new name in the same paper, Coronosaurus (formerly “Centrosaurus” brinkmani). The other ceratopsid subgroup, the chasmosaurines, encompass Triceratops, Torosaurus and other genera more closely related to them than Centrosaurus.

At approximately 78 million years old, Xenoceratops is currently the oldest ceratopsid known from Canada, beating out its cousin Albertaceratops by half a million years. Given the age of Xenoceratops, and the fact that it had long brow horns and a short nasal horn, instead of the long nasal horn-short brow horns combo seen in its later relatives, it isn’t surprising that the dinosaur seems to be at the base of the centrosaurine family tree. This means that Xenoceratops can help paleontologists examine what the early members of this significant ceratopsid group were like and how drastically centrosaurine ornamentation changed. “Xenoceratops has very well developed frill ornamentation comprised of a series of large spikes and hooks, occurring at multiple parietal loci, that foreshadows the great diversity of these structures in other species that occur later in the Campanian,” Evans says, and this indicates that “complex frill ornamentation is older than we may have thought.”

Still, Evans cautions that Xenoceratops is presently a very scrappy dinosaur. We need more fossils to fully reconstruct this dinosaur and confirm its place in the ceratopsid family tree. The dinosaur’s “true significance in terms of ceratopsid origins will only be revealed with further discoveries,” Evans says, particularly between the time of the slightly older Diabloceratops found in southern Utah, and the even more archaic, roughly 90-million-year-old ceratopsian Zuniceratops. “Our record of ceratopsians in this critical part of their family tree is still frustratingly poor,” Evans laments. In fact, paleontologists know relatively little about dinosaur diversity and evolution during the middle part of the Cretaceous–a critical evolutionary time period for ceratopsians, tyrannosaurs and other lineages that came to dominate the Late Cretaceous landscape. If we are ever going to solve the mystery of how ceratopsids evolved, and why they were such garishly adorned dinosaurs, we must search the lost world of the mid-Cretaceous.

References:

Ryan, M., Evans, D., Shepherd, K. 2012. A new ceratopsid from the Foremost Formation (middle Campanian) of Alberta. Canadian Journal of Earth Sciences 49: 1251-1262

The reconstructed skull of Eolambia–based on a partial adult skull and scaled juvenile elements–and a restoration by artist Lukas Panzarin. From McDonald et al., 2012.

Hadrosaurs were not the most charismatic dinosaurs. Some, such as Parasaurolophus and Lambeosaurus, had ornate, hollow crests jutting through their skulls, but, otherwise, these herbivorous dinosaurs seem rather drab next to their contemporaries. They lacked the garish displays of horns and armor seen among lineages such as the ceratopsians and ankylosaurs, and they cannot compete with the celebrity of the feathery carnivores that preyed upon them. Yet in the habitats where they lived, hadrosaurs were among the most common dinosaurs and essential parts of their ecosystems. What would tyrannosaurs do without ample hadrosaurian prey?

While many hadrosaurs might seem visually unremarkable next to their neighbors, the wealth of these dinosaurs that paleontologists have uncovered represent a huge database of paleobiological information waiting to be tapped for new insights into dino biology and evolution.

In order to draw out dinosaur secrets, though, paleontologists need to properly identify, describe and categorize the fossils they find. We need to know who’s who before their stories can come into focus. On that score, paleontologist Andrew McDonald and colleagues have just published a detailed catalog of Eolambia caroljonesa, an archaic hadrosaur that was once abundant in Cretaceous Utah.

Eolambia is not a new dinosaur. Discovered in the roughly 96-million-year-old rock of the Cedar Mountain Formation, this dinosaur was named by paleontologist James Kirkland–a coauthor on the new paper–in 1998. Now there are multiple skeletons from two different localities representing both sub-adult and adult animals, and those specimens form the basis of the full description.

While the new paper is primarily concerned with the details of the dinosaur’s skeleton, including a provisional skull reconstruction accompanied by an excellent restoration by artist Lukas Panzarin, McDonald and coauthors found a new place for Eolambia in the hadrosaur family tree. When Kirkland announced the dinosaur, he named it Eolambia because it seemed to be at the dawn (“eo”) of the crested lambeosaurine lineage of hadrosaurs. But in the new paper McDonald, Kirkland and collaborators found that Eolambia was actually a more archaic animal–a hadrosauroid that falls outside the hadrosaurid lineage containing the crested forms.

Much like its later relatives, Eolambia would have been a common sight on the mid-Cretaceous landscape. The descriptive paper lists eight isolated animals and two bonebeds containing a total of 16 additional individuals. They lived in an assemblage that was right at the transition between the early and late Cretaceous faunas–tyrannosaurs, deinonychosaurs and ceratopsians have been found in the same part of the formation, as well as Jurassic holdouts like sauropods. How this community fit into the grander scheme of dinosaur evolution in North America is still coming together, though. The Early and Middle parts of the Cretaceous are still poorly known, and paleontologists are just getting acquainted with Eolambia, its kin and contemporaries.

References:

McDonald, A., Bird, J., Kirkland, J., Dodson, P. 2012. Osteology of the basal hadrosauroid Eolambia caroljonesa (Dinosauria: Ornithopoda) from the Cedar Mountain Formation of Utah. PLOS One 7, 10: e45712

The upper and lower jaws of Duriavenator, illustrated when they were thought to belong to Megalosaurus, in A History of British Fossil Reptiles Vol. II. Image from Wikipedia.

If you have been following the Dinosaur Alphabet series so far, you may have noticed a pattern among the first four entries. At one time or another, all the dinosaurs I’ve selected so far were thought to be different animals. The horned Agujaceratops was originally named as a species of Chasmosaurus, the distinctive high-spines of Becklespinax gave Richard Owen’s dopey Megalosaurus its hump, the sauropod Cetiosaurus was originally envisioned as a giant crocodile, and the armored Dyoplosaurus was lumped in with its cousin Euoplocephalus before being split back out again as a distinct genus. I didn’t intend this trend, but it struck me when I came across  one of the rejected candidate for yesterday’s entry for the letter D. Had it not shared much of its story with Becklespinax, I would have picked Duriavenator:

Megalosaurus was a mess. Even though this Jurassic carnivore has been a prehistoric icon ever since it was named by William Buckland in 1824, it has been one of the most confounding of all dinosaurs. That’s because generations of researchers attributed dozens of fragments and isolated bones to the dinosaur, creating a monstrous composite of animals from different places and times. Dinosaurs were unfamiliar animals–the name itself only coined in 1842–and 19th-century naturalists didn’t have the kind of geologic resolution their intellectual descendants rely on to properly constrain when particular species lived. Sometimes researchers named too many species on the basis of scrappy, non-overlapping material, and other times they applied the same name ad infinitum to roughly similar fossils.

Eventually, though, it became apparent that Megalosaurus was unstable. No one could say what the dinosaur really looked like or what bones could accurately be attributed to the predator.  The situation was so bad that, in 2008, paleontologist Roger Benson and colleagues stripped the name Megalosaurus from everything save for the fragment of jaw originally used to name the animal. Whether the rest of the fossils really belonged to Megalosaurus remained to be seen, and, as Benson demonstrated later the same year, at least one other theropod had been improperly obscured behind the famous name.

In 1883, anatomist Richard Owen described a partial theropod skull found on Dorset, England, as another piece of Megalosaurus “bucklandi.” The sharp-toothed dinosaur was only represented by parts of the upper and lower jaws, but, given how little was known about Megalosaurus to start with, Owen’s assignment was reasonable. Nearly a century later, paleontologist Michael Waldman proposed that these fossils represented a previously unknown species of the dinosaur he called Megalosaurus hesperis. Other researchers weren’t sure that the bones really belonged to Megalosaurus, but it wasn’t until Benson’s reexamination that the fossils were split out as a different dinosaur. While the dinosaur was a close cousin of Megalosaurus bucklandii, Benson was able to pick out subtle anatomical characteristics that distinguished the fragmentary skull. In Benson’s analysis, what once was Megalosaurus took on a new life as Duriavenator hesperis.

Benson’s reconstruction of Megalosaurus, with known elements in white and reconstructed portions in grey. While Duriavenator was older and anatomically distinct, the dinosaur would have been similar in form to Megalosaurus. From Benson, 2010.

Unfortunately, we don’t know very much at all about Duriavenator. The dinosaur lived about 170 million years ago in Jurassic England and was a large carnivore of comparable size to the 20-foot-plus Megalosaurus, but that’s where the evidence gives out. Perhaps other Duriavenator specimens are resting in museum collections, but until the discovery of a nearly complete skeleton allows paleontologists to connect the jaws to a body, the dinosaur will be an enigma. But here Megalosaurus itself gives us reason to hope. The Duriavenator paper was just part of Benson’s effort to rehabilitate Megalosaurus, and in 2010 he published a refined, revised reconstruction of the dinosaur’s skeleton based on material collected from Stonesfield, Oxfordshire–the locality where the original jaw came from. Perhaps, with a little detective work in the lab and in the field, paleontologists might also be able to fill out the form of Duriavenator and other Middle Jurassic mysteries.

References:

Benson, R., Barrett, P., Powell, H., Norman, D. 2008.  The taxonomic status of Megalosaurus bucklandii (Dinosauria, Theropoda) from the Middle Jurassic of Oxfordshire, UK. Palaeontology, 51, 2: 419-424.

Benson, R. 2008.  A redescription of “Megalosaurus” hesperis (Dinosauria, Theropoda) from the Inferior Oolite (Bajocian, Middle Jurassic) of Dorset, United Kingdom. Zootaxa 1931: 57-67

Benson, R. 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society 158: 882. doi:10.1111/j.1096-3642.2009.00569.x.

Waldman, M. 1974. Megalosaurids from the Bajocian (Middle Jurassic) of Dorset. Palaeontology 17, 2:325-339.