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A reconstruction of Irritator. Photo by Kabacchi, image from Wikipedia.

Spinosaurs are often called “fish-eating dinosaurs.” Their long, shallow snouts recall the jaws of crocodiles, and, based on gut contents and fossil geochemistry, it seems that these dinosaurs truly were piscivores. Yet spinosaurs weren’t on a strict fish diet. In 2004, Eric Buffetaut and colleagues described a spinosaur tooth embedded in the fossilized neck vertebrae of an Early Cretaceous pterosaur found in Brazil’s roughly 110-million-year-old Santana Formation. The paleontologists couldn’t say whether the dinosaur caught its prey on the wing or scavenged a fresh carcass, but, based on fossils previously found in the same geologic formation, one spinosaur stood out as the probable culprit–Irritator challengeri.

The spinosaur’s quirky name symbolizes its unconventional back story. As explained in the 1996 description of the dinosaur by David Martill and colleagues, the mostly complete skull of Irritator had been artificially modified by a commercial fossil dealer prior to being purchased and making its way into the collection of Germany’s Stuttgart State Museum of the Natural Sciences. The tip of the snout was made up of bone from elsewhere on the skull, “concealed by blocks of matrix removed from other parts of the specimen and a thick layer of Isopon car body filler.” The fabrication not only deceived the buyers, but was especially difficult to remove from the authentic fossil. Martill and colleagues named the dinosaur Irritator as a tribute to “the feeling the authors felt (understated here) when discovering that the snout had been artificially elongated.”

Martill and collaborators originally proposed that Irritator was a maniraptoran dinosaur–a relative of the feathery deinonychosaurs, oviraptorosaurs, and their kin. That same year, however, paleontologist Andrew Kellner recognized that Irritator was actually a spinosaur–one of the croc-snouted, and often sail-backed, predatory dinosaurs. Kellner also named what he suspected was another spinosaur found in the same geologic formation–”Angaturama limai“–but many researchers suspect that this animal is the same as Irritator, and the so-called “Angaturama” remains may even complete the missing parts of the Irritator skeleton.

But even after Irritator was properly identified, there was still work to be done. Diane Scott undertook the painstaking work of fully cleaning the skull of the encasing matrix, which led to a new description by Hans-Dieter Sues and coauthors in 2002. Irritator is represented by the most complete skull yet known for any spinosaur. Among other new aspects, it was apparent that the back of the skull was significantly deeper among spinosaurs than had previously been thought. And even though Martill and co-authors originally described a prominent crest on the top of the spinosaur’s skull, the fully-prepped fossil showed that this bone did not actually belong to the Irritator skull.

There’s still much we have to learn about spinosaurs. Most of these dinosaurs are only known from bits and pieces. And despite starring in Jurassic Park III, Spinosaurus itself is among the most poorly known dinosaurs of all, and the fragmentary nature of so many of these dinosaurs makes it possible that paleontologists have named too many genera. In their study, Sues and coauthors argue that Suchomimus is really just a different species of Baryonx, and even Irritator might be a distinct species of Spinosaurus. Researchers have only just begun to track the record of these long-snouted dinosaurs, although, hopefully, future finds will not be quite so aggravating as Irritator.

This is the latest post in the Dinosaur Alphabet series.

References:

Buffetaut, E., Martill, D., Escuillie, F. 2004. Pterosaurs as part of a spinosaur diet. Nature. 430: 33

Martill, D., Cruickshank, A., Frey, E., Small, P., Clarke, M. 1996. A new crested maniraptoran dinosaur from the Santana Formation (Lower Cretaceous) of Brazil. Journal of the Geological Society 153: 5-8.

Sues, H., Frey, E., Martill, D., Scott, D. 2002. Irritator challengeri, a spinosaurid (Dinosauria: Theropoda) from the Lower Cretaceous of Brazil. Journal of Vertebrate Paleontology. 22, 3: 535-547

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

A restoration of Gigantspinosaurus. Art by Conty, image from Wikipedia.

Stegosaurus was a weird dinosaur. We’ve known that for well over a century, but, as Darren Naish has often pointed out, Stegosaurus was strange even compared to its Jurassic relatives. The dinosaur’s arrangement of broad, alternating plates is a departure from the arrangements of smaller plates, back spikes and accessory spines seen on many other stegosaurs, including the perplexingly well-armed Gigantspinosaurus sichuanensis.

Ornamented with a double row of short, narrow plates along its back, the roughly 160-million-year-old Gigantspinosaurus generally resembled other stegosaurs from Late Jurassic Asia, such as Tuojiangosaurus. But, as you might be able to guess from the dinosaur’s name, the feature that immediately sets Gigantspinosaurus apart from similar species is a enormous hooked spine that jutted out from behind the shoulder blade. These striking spikes were found close to their life position on the first skeleton of this dinosaur to be found–erroneously attributed to Tuojiangosaurus, before being redescribed as Gigantspinosaurus in 1992–although their exact orientation isn’t entirely clear. Did the shoulder spikes curve straight backward, or were they tiled slightly upwards? And, more significantly, how did such prominent ornaments evolve? No one knows.

As yet, we know relatively little about the natural history of Gigantspinosaurus. The dinosaur has a name, and skin impressions have helped researchers restore what the stegosaur looked like, but many aspects of the spiky herbivore’s biology remain mysterious. In the grand scheme of stegosaur evolution, though, the ornamentation of Gigantspinosaurus has sometimes been taken as evidence that similar forms had shoulder spikes. In addition to paired spikes along its tail, the Late Jurassic stegosaur Kentrosaurus possessed an extra pair of spikes along its side. These were originally placed over the hips, but, due to the discovery of Gigantspinosaurus, some researchers have argued that the spikes truly belong at the shoulders.

Frustratingly, paleontologists have yet to find a Kentrosaurus skeleton with side spikes in place. But the discovery of Gigantspinosaurus doesn’t necessarily mean that its cousin Kentrosaurus had the same arrangement. Among stegosaurs, the two genera were relatively distantly related, and it’s entirely possible that more than one side spike arrangement evolved. As paleontologist Heinrich Mallison has argued, the hips of Kentrosaurus seem to possess areas where the spikes could have articulated, and this arrangement would be consistent with the dinosaur’s ornamentation pattern–small plates at the front give way to spikes along the stegosaur’s back and tail. Indeed, the side spikes on Kentrosaurus more closely resemble the same structures along the dinosaur’s back and tail and the shoulder spike of Gigantspinosaurus. If Kentrosaurus had plates up front and serially homologous spikes along the back, then why shouldn’t the hip spikes remain a reasonable hypothesis? Together, Gigantspinosaurus and Kentrosaurus might represent different alternatives in the stegosaur armory.

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

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