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The articulated, almost-complete hand of Hagryphus giganteus. From Zanno and Sampson, 2005.

When I think of oviraptorosaurs – feathered, beaked, omnivorous theropods–my mind immediately jumps to Mongolia’s famous brooding dinosaurs and other forms extracted from Asia’s Cretaceous rock. But these weird dinosaurs were present in North America, too. Among the latest to come to the attention of paleontologists is Hagryphus giganteus–a large oviraptorosaur known from little more than a hand and pieces of foot.

Paleontologists started to report on the oviraptorosaurs of North America’s Late Cretaceous in the 1930s. They just didn’t immediately recognize the dinosaurs for what they were. Scrappy remains of these dinosaurs were attributed to the ostrich-like ornithomimosaurs and Cretaceous birds. It was only in the 80s and 90s that researchers began to untangle the identities of these dinosaurs. Based on specimens found in Canada, Montana, and the Dakotas, there may have been at least three different genera present–Caenagnathus, Chirostenotes, and Elmisaurus–around 75 million years ago. That depends on who you ask, though. Researchers disagree about which genera are valid. The material from these dinosaurs is so fragmentary that it’s difficult to tell just how many different forms we’re looking at.

But Hagryphus, described by paleontologists Lindsay Zanno and Scott Sampson in 2005, was different. Represented by a nearly-complete left hand, part of the left radius, and fragments of the foot, this theropod lived further to the south in the 75-million-year-old swampy environment preserved in Utah’s Grand Staircase-Escalante National Monument. Much like other dinosaurs found in the same formation, and other southern species from roughly contemporaneous deposits, the known remains of Hagryphus are distinct from the equivalent bones known from the northern species. Not only was Hagryphys bigger–Zanno and Sampson estimated that the dinosaur was about 10 feet long, quite large for an oviraptorosaur–but bones in the dinosaur’s hand were much more robust.

Zanno and Sampson considered that the unique nature of Hagryphus might be because the individual was an older specimen of one of the northern oviraptorosaurs. They rejected this hypothesis, arguing the the dinosaur’s distinctive hand proportions were more consistent with being it a different taxon than changes due to growth. If they’re right, this fits the general pattern of Utah’s Kaiparowits Formation in preserving dinosaurs that were related to those found in Montana and Alberta but were unique genera and species.

So how many oviraptorosaurs were there in North America around 75 million years ago? We probably haven’t found traces of all of them, but based on what has been described so far there were probably at least two and as many as four. We need more complete skeletons to be sure.

The same problem affects other small-bodied theropod dinosaurs from the Late Cretaceous. Based on teeth and fragmentary remains, paleontologists used to think that the deinonychosaur Troodon had a range from southern Utah to Alaska. As parts of additional specimens come out of the ground, paleontologists are starting to realize that what seemed to be just one dinosaur is really a collection of different genera or species spread across the latitudes. And regardless of what Hagryphys is, the existence of an oviraptorosaur in Utah greatly extends the range of these dinosaurs during the 75-million-year-old time frame. Exposures between southern Utah and Montana may very well hold additional oviraptorosaur specimens–individuals that will be critical to understanding how these dinosaurs evolved.

This is the latest post in the Dinosaur Alphabet series.

Reference:

Zanno, L., Sampson, S. 2005. A new oviraptorosaur (Theropoda, Maniraptora) from the Late Cretaceous (Campanian) of Utah. Journal of Vertebrate Paleontology. 35:4, 897-904

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

A reconstruction of Deinosuchus at the Natural History Museum of Utah. Photo by the author.

From the time of their origin around 230 million years ago, to the extinction of the non-avian forms 66 million years ago, dinosaurs ruled the Earth. That’s how we like to characterize the Mesozoic menagerie, anyway. We take the long success of the dinosaurs as a sign of their long-lived and terrifying domination, but, despite our belief that they were the most vicious creatures of all time, there were creatures that even the dinosaurs had reason to fear. Chief among them was Deinosuchus – North America’s “terrible crocodile.”

Between 80 and 73 million years ago, when North America was divided in two by the shallow Western Interior Seaway, the marshes and swamps along the coasts were ruled by Deinosuchus. Fossils of this Cretaceous cousin of modern alligators have been found from Mexico to Montana and in east coast states such as North Carolina and Georgia, tracing the margins of the western subcontinent Laramidia and its eastern counterpart, Appalachia. For the most part, paleontologists have found the bony armor, vertebrae, and teeth of Deinosuchus, but pieces of jaw and partial skeletons found in places such as Texas and Utah indicate that this alligatoroid was a giant, growing over thirty feet in length and approaching forty feet among the biggest individuals.

During the heyday of Deinosuchus, adults of the aquatic ambush predator were among the largest carnivores in their ecosystems. The enormous Tyrannosaurus rex was over five million years off, and the tyrannosaurs of the time were not quite so long or bulky. (Teratophoneus, found in southern Utah among strata that also yield Deinosuchus, was about twenty feet long, and Daspletosaurus from Montana grew to be about thirty feet long.) A fully mature Deinosuchus would have outstretched and outweighed the dinosaur competition, and would have undoubtedly been a deadly apex predator in the water habitats it haunted.

The skull of Deinosuchus testifies to its destructive potential. The alligatoroid’s skull was large, broad, and equipped with an array of teeth deployed to pierce and crush. Indeed, even though there were other giant crocodylomorphs of near-equal size during the Mesozoic (such as the narrow-snouted Sarcosuchus), Deinosuchus appears to be unique in having the anatomical necessities to take down hadrosaurs and other unwary dinosaurs at the water’s edge. And, thanks to tooth-damaged fossils, we know that Deinosuchus truly did dine on dinosaurs. Two years ago, Héctor Rivera-Sylva and colleagues described hadrosaur bones bearing tell-tale Deinosuchus toothmarks from Mexico, and similar finds have been reported from Texas. There may be other candidates in museum drawers elsewhere.

Of course, we don’t know whether the bitten bones record hunting or scavenging. Unless the injuries show signs of healing, toothmarks on bones record feeding rather than hunting behavior. The evidence only takes us so far. Adult Deinosuchus were apparently capable of taking down dinosaurs, but, as yet, there’s no direct evidence of such an incident. Indeed, while images of Deinosuchus chomping on dinosaurs fires our imagination, we actually know relatively little about how this alligatoroid fed and what it ate. Probably, like modern alligators, large Deinosuchus were generalists that snagged fish, turtles, and whatever carrion it happened upon. We don’t know for sure. Nevertheless, dinosaurs in the habitat of this monstrous croc would have been wise to carefully approach the water’s edge, looking for teeth and scutes hiding just beneath the surface.

Dilophosaurus, in a restoration based on an impression found at St. George, Utah. Art by Heather Kyoht Luterman, from Milner et al., 2009.

The Early Jurassic is a mysterious time in dinosaur evolution. In North America, at least, paleontologists have uncovered scores of dinosaur tracks from this critical time when dinosaurs had been handed ecological dominance in the wake of a mass extinction, but body fossils are rare. In the orange sandstone that makes up so much of Arches and Canyonlands national parks in Utah, for example, only a handful of skeletons have ever been found. This formation–called the Glen Canyon, Navajo, Nugget or “Nuggaho” depending on who you ask–preserves immense sand dunes that recorded prehistoric footsteps but rarely bone. The recently described sauropodomorph Seitaad, and a group of as-yet-unnamed coelophysoids, are exceptionally rare finds.

Yet, from Connecticut to Arizona, there is one dinosaur that is constantly presented as an icon of dinosaurs circa 190 million years ago. This is Dilophosaurus–the 20-foot-long, double-crested theropod that gained dubious fame thanks to Jurassic Park. (Contrary to the film, there’s no evidence that this carnivore was a “spitter” with a collapsible neck frill.) At sites where Early Jurassic theropod tracks are found in abundance, Dilophosaurus is invoked as a possible trackmaker. But is this really so?

The remains of what would eventually be named Dilophosaurus were discovered in 1942 by Jesse Williams near Tuba City, Arizona. It took another 12 years before paleontologist Samuel Welles mistakenly attributed the bones to a new species of Megalosaurus –“M.” wetherilli–and the name Dilophosaurus itself wasn’t actually coined until 1970. Despite all this shifting around, though, Dilophosaurus wetherilli became a symbol of top Early Jurassic carnivores. Paleontologists had found plenty of Early Jurassic tracks made by a Dilophosaurus-size dinosaur, and now they finally had a body.

Frustratingly, though, we usually don’t know what dinosaur left a particular trace fossil unless the animal literally died in its tracks. While Dilophosaurus is a good fit for many large-size, Early Jurassic tracks, and may very well have left tracks at places such as St. George, Utah’s megatracksite, there’s no way to know for sure. And it seems unlikely that the same species of dinosaur that left tracks in Early Jurassic Utah also made footprints in the mud of what would become the Connecticut Valley. Who knows how many mid-sized theropods might have stalked lakeshores during this time? We don’t know, and the situation is made all the more irksome since the sediments which preserve tracks often don’t contain body fossils. We know these dinosaurs from the bottom of their feet but little else. Until future discoveries fill out the fauna of North America’s Early Jurassic, Dilophosaurus will remain the most familiar and iconic predator of its epoch.

Reference:

Naish, D. 2009. The Great Dinosaur Discoveries. University of California Press: Berkeley. pp. 94-95

Fossil swim tracks indicate that theropods similar to this Megapnosaurus at least occasionally swam in prehistoric lakes and rivers. Art by Dmitry Bogdanov, image from Wikipedia.

Paleontologist R.T. Bird inspected many dinosaur trackways while combing Texas for the perfect set to bring back to the American Museum of Natural History. During several field seasons in the late 1930s, Bird poked around in the Early Cretaceous rock in the vicinity of the Paluxy River for a set of sauropod footprints that would fit nicely behind the museum’s famous “Brontosaurus” mount. Bird eventually got what he was after but not before poring over other intriguing dinosaur traces. One of the most spectacular seemed to be made by a swimming dinosaur.

Known as the Mayan Ranch Trackway, the roughly 113-million-year-old slab is almost entirely made up of front foot impressions. The semicircular imprints were undoubtedly left by one of the long-necked sauropod dinosaurs. But towards the end of the trail, where the dinosaur’s path makes an abrupt turn, there was a single, partial impression of a hind foot.

At the time Bird and his crew uncovered this trackway, sauropods were thought to be amphibious dinosaurs. Other than their immense bulk, what defense would they have had but to trundle into the water, where theropods feared to paddle? Under this framework, Bird thought he knew exactly how the Mayan Ranch Trackway was made. “The big fellow had been peacefully dog-paddling along, with his great body afloat, kicking himself forward by walking on the bottom here in the shallows with his front feet,” Bird wrote in his memoir. The great dinosaur then kicked off with one of its hind feet and turned.

With the exception of well-defended dinosaurs such as the ceratopsids and stegosaurs, many herbivorous dinosaurs were thought to be at least semi-aquatic. There seemed to be only two options for Mesozoic prey species–grow defenses or dive into the water. In time, though, paleontologists realized that the sauropods, hadrosaurs and other herbivores didn’t show any adaptations to swimming. Our understanding of the ecology of these dinosaurs was based on false premises and faulty evidence.

In the case of the Mayan Ranch Trackway, for example, there’s no indication that the sauropod that made the trackway was swimming. A more likely scenario has to do with evolutionary changes among sauropods. While the sauropods that dominated the Late Jurassic of North America–such as Diplodocus, Apatosaurus and Barosaurus–carried much of their weight at the hips and left deeper hindfoot impressions, the center of mass shifted among their successors–the titanosaurs–such that more of the weight was carried by the forelimbs. Hence, in some trackways, the deeper impressions made by the forefeet are more likely to stand out than those made by the hindfeet, especially if some of the top layers of the rock are eroded away to leave only “undertracks.” What seemed to be evidence of swimming sauropods instead owes to anatomy and the characteristics of the mucky substrate the dinosaur was walking on.

As far as I’m aware, no one has yet found definitive evidence of swimming sauropods or hadrosaurs–the two groups previously thought to rely on water for safety. Stranger still, paleontologists have recently uncovered good evidence that theropod dinosaurs weren’t as bothered by water as traditionally believed. In 2006, paleontologists Andrew Milner, Martin Lockley and Jim Kirkland described swim tracks made by Early Jurassic theropods at a site that now resides in St. George, Utah. Such traces weren’t the first of their kind ever discovered, but the tracksite was one of the richest ever found.

Small to medium-sized theropods made the St. George swim tracks–think of dinosaurs similar to Megapnosaurus and Dilophosaurus. Even better, the large number of smaller-size swim tracks hints that whatever dinosaurs made these tracks were moving as a group as they struggled against the current in the lake shallows. The larger dinosaurs, on the other hand, were a bit taller and able to wade where their smaller cousins splashed around.

A different team of researchers announced additional evidence for swimming theropods the following year. Paleontologist Rubén Ezquerra and co-authors described dinosaur swim traces from Early Cretaceous rock near La Rioja, Spain. Based on the details of the track and their direction, the theropod was swimming against a current that pushed the dinosaur diagonally. Along with other theropod swim tracks, the researchers noted, the discovery meant that paleontologists would have to revise their ideas about the kind of habitats theropods lived in and what carnivorous species would do. Theropod dinosaurs were not so hydrophobic, after all.

Does this mean that dinosaurs like Dilophosaurus were adapted to an amphibious lifestyle? Not at all. As Ezquerra and co-authors pointed out, the swimming strokes of these dinosaurs were exaggerated walking motions. The way the dinosaurs moved on land allowed them to be adequate swimmers while crossing rivers or lakes, but, compared with semi-aquatic animals such as crocodiles and otters, no known dinosaur shows traits indicative of a primarily waterlogged existence. (And dinosaurs found in marine sediments don’t count as evidence, as these were washed out to sea prior to burial. I can’t imagine ankylosaurs taking to life among the high seas, in any case.) Some dinosaurs could swim, but that doesn’t mean that they made the water their home. Still, thanks to special prehistoric traces, we can imagine packs of Megapnosaurus fighting to get ashore, and Dilophosaurus strutting into the shallows, aiming to snatch whatever fish were foolish enough to swim into the carnivore’s shadow.

References:

Bird, R.T. (1985). Bones for Barnum Brown, edited by Schreiber, V. Forth Worth: Texas Christian University Press. pp. 160-161

Ezquerra, R., Doublet, S., Costeur, L., Galton, P., Pérez-Lorente, F. (2007). Were non-avian theropod dinosaurs able to swim? Supportive evidence from an Early Cretaceous trackway, Cameros Basin (La Rioja, Spain) Geology, 40 (10), 507-510 DOI: 10.1130/G23452A.1

Milner, A., Lockley, M., Kirkland, J. (2006). A large collection of well-preserved theropod dinosaur swim tracks from the Lower Jurassic Moenave Formation, St. George, Utah. New Mexico Museum of Natural History and Science Bulletin, 37, 315-328