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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

Archaeopteryx had a wing that was different from that of modern birds, and, as seen here, might have been a glider more than a powered flyer. Art by Carl Buell, courtesy of Nicholas Longrich.

How did feathered dinosaurs take to the air? Paleontologists have been investigating and debating this essential aspect of avian evolution for over a century. Indeed, there have been almost as many ideas as they have been experts, envisioning scenarios of dinosaurs gliding through trees, theropods trapping insects with their feathery wings and even aquatic Iguanodon flapping primitive flippers as flight precursors (I didn’t say that all the ideas were good ones). The biomechanical abilities of bird ancestors and their natural history has always been at the center of the debate, and a new Current Biology paper adds more fuel to the long-running discussion.

At present, hypotheses for the origin of avian flight typically fall into one of two categories. Either bird ancestors accrued the adaptations necessary for flight on the ground and, through evolutionary happenstance, were eventually able to take off, or small tree-dwelling dinosaurs used their feathery coats to glide between trees and, eventually, flapped their way into a flying lifestyle. There are variations on both themes, but feathers and the characteristic avian flight stroke are at the core of any such scenario. In the case of the new paper, Yale University paleontologist Nicholas Longrich and colleagues draw from the plumage of early bird Archaeopteryx and the troodontid Anchiornis to examine how feathers changed as dinosaurs started to fly.

In modern flying birds, Longrich and coauthors point out, the wing arrangement typically consists of “long, asymmetrical flight feathers overlain by short covert feathers.” This pattern creates a stable airfoil but also lets the flight feathers separate a little during the upstroke of a wing beat, therefore reducing drag. When the paleontologists examined the fossilized wings of Archaeopteryx and Anchiornis, they found different feather arrangements that would have constrained the flight abilities of the Jurassic dinosaurs.

Both prehistoric creatures had long covert feathers layered on top of the flight feathers. Anchiornis, in particular, appeared to have an archaic wing form characterized by layers of short, symmetrical flight feathers and similarly shaped coverts. Archaeopteryx showed more specialization between the flight feathers and the coverts but still did not have a wing just like that of a modern bird. As a result, Longrich and collaborators hypothesize, both arrangements would have stabilized the wing at the cost of increased drag at low speeds, making it especially difficult for Anchiornis and Archaeopteryx to take off. As an alternative, the researchers suggest that these dinosaurs might have been parachuters who jumped into the air from trees, which might hint that “powered flight was preceded by arboreal parachuting and gliding.”

The trick is determining whether Anchiornis and Archaeopteryx actually represent the form of bird ancestors, or whether the dinosaurs, like Microraptor, were independent experiments in flight evolution. At the Society of Vertebrate Paleontology conference in Raleigh, North Carolina last month, flight expert Michael Habib quipped that all that was needed to make dromaeosaurs aerially competent was the addition of feathers. If Habib is right, and I think he is, then there could have been multiple evolutionary experiments in flying, gliding, wing-assisted-incline-running and other such activities. There’s no reason to think that flight evolved only once in a neat, clean march of ever-increasing aerodynamic perfection. Evolution is messy, and who knows how many ultimately failed variations there were among flight-capable dinosaurs?

The three-step Anchiornis-Archaeopteryx-modern bird scenario of wing evolution fits our expectations of what a stepwise evolutionary pattern would look like, but, as the authors of the new paper point out, shifting evolutionary trees currently confound our ability to know what represents the ancestral bird condition and what characterized a more distant branch of the feathered dinosaur family tree. We need more feathery fossils to further investigate and test this hypothesis, as well as additional biomechanical and paleoecological information to determine whether such dinosaurs really took off from trees. We must take great care in distinguishing between what an organism could do and what it actually did, and with so much up in the air, the debate on the origin of flight will undoubtedly continue for decades to come.

Reference:

Longrich, N., Vinther, J., Meng, Q., Li, Q., Russell, A. 2012. Primitive wing feather arrangement in Archaeopteryx lithographica and Anchiornis huxleyi. Current Biology DOI: 10.1016/j.cub.2012.09.052

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

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.

Thanks to a row of huge bony plates, Stegosaurus remains one of the strangest dinosaurs ever found. Photo by the author at the Utah Field House of Natural History in Vernal, Utah.

Undoubtedly familiar to any dinosaur fan, Stegosaurus remains one of the strangest dinosaurs ever discovered. Even among others of its kind, the iconic Jurassic herbivore looks like an oddball. Many other stegosaur species sported long rows of spikes and short plates, but the flashy Stegosaurus had an alternating row of huge bony plates along its back and a relatively modest set of four tail spikes. How could such a strange arrangement of adornments have evolved?

From the arms of tyrannosaurs to the necks of sauropods and the armor of stegosaurs, bizarre dinosaur structures have frequently made paleontologists wonder “What was that for?” There had to be a reason for the deviations in form, and, paleontologists believe, the immediately recognizable plates on the back of Stegosaurus must have had some function. There has been no shortage of hypotheses. Off-the-wall ideas about flying stegosaurs aside, researchers have proposed that the plates along the spine of Stegosaurus protected the dinosaur from attack, were the Jurassic equivalent of solar panels or acted as sexy billboards to attract the attention of potential mates.

Although Stegosaurus certainly had much to fear from the contemporary Morrison Formation predators Allosaurus, Torvosaurus and Ceratosaurus, the dinosaur’s defensive weapons were its tail spikes (called a “thagomizer” by some). If Stegosaurus was anything like its spikier cousin Kentrosaurus, it could swing its tail with deadly force, and a damaged Allosaurus bone suggests that the “roof lizard” did just that. But the keratin-covered plates of Stegosaurus probably didn’t provide the herbivore with much additional protection. The immobile structures jutted upwards, leaving the dinosaur’s flanks exposed to attack. To call the plates “armor” isn’t quite right.

When I was a kid, though, Stegosaurus plates were more often said to help the dinosaur regulate its body temperature. Presuming that Stegosaurus was an ecothermic animal–that is, had a body temperature determined by the surrounding environment–the plates could have helped the dinosaur heat up by turning broadside in the morning and shed heat by turning toward the sun during midday. Using models of plates in wind tunnel experiments, paleontologist James Farlow and colleagues reported in 1976 that the plates could very well have been used to dissipate heat. This doesn’t mean that the plates evolved for that function, though.

In 2010, Farlow and coauthors followed up on the work by comparing the plates of Stegosaurus to the bony armor along the backs of modern crocodylians. While stegosaur plates might have played some passive role in regulating body temperature, they concluded, there was no indication that Stegosaurus plates evolved for that reason, or even were principally used as thermoregulatory equipment. (Not to mention the fact that we now know that dinosaurs were not lizard-like reptiles whose internal physiology was primarily dictated by the temperature outside.) If Stegosaurus plates made any difference in managing body temperature, it was a happy little quirk that rode along with the principal function of the plates.

At present, it appears that the impressive bony fins on the back of Stegosaurus evolved as display structures. A 2005 study by Russell Main and  collaborators, which focused on the microstructure of stegosaur plates, couldn’t find any evidence that the structures were used to radiate heat. Indeed, if stegosaurs truly required such radiators, it’s surprising that Stegosaurus seems unique in its plate arrangement–if plates were really used to regulate body temperature, you’d expect to see the same arrangement in many closely related species. Instead, much like the horns of ceratopsid dinosaurs, the plates and spikes of stegosaurs varied greatly between species. This suggests that visual display was driving the evolution of these structures. Being recognized as a member of a particular species, or displaying an individual’s maturity and vigor during mating season, probably drove the divergence in form among stegosaur ornaments. The question is whether stegosaur plates made any difference in the mating season or they simply served to help species recognize members of their own kind. That debate–over the sexiness of plates, spikes, horns, crests, sails and domes–is just heating up.

References:

Farlow, J., Thompson, C., Rosner, D. 1976. Plates of the dinosaur Stegosaurus: Forced convection heat loss fins? Science. 192,4244: 1123-1125

Farlow, J., Hayashi, S., Tattersall, G. 2010. Internal vascularity of the dermal plates of Stegosaurus (Ornithischia, Thyreophora). Swiss Journal of Geosciences. 103, 2: 173-185

Hayashi, S., Carpenter, K., Watabe, M., McWhinney, L. 2011. Ontogenetic histology of Stegosaurus plates and spikes. Palaeontology. 55, 1: 145-161

Main, R., de Ricqlès, A., Horner, J., Padian, K. 2005. The evolution and function of thyreophoran dinosaur scutes: implications for plate function in stegosaurs. Paleobiology. 31, 2: 291-314

Padian, K., Horner, J. 2010. The evolution of “bizarre structures” in dinosaurs: biomechanics, sexual selection, social selection, or species recognition? Journal of Zoology. 283,1: 3-17