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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

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.

A reconstruction of Patagonykus, one of South America’s alvarezsaurs. Image from Wikipedia.

If there’s one group of dinosaurs that needs better PR, it’s alvarezsaurs. They’re among the strangest dinosaurs to have ever evolved, yet outside of dinosaur die-hards, few people have ever heard of them. They’re not one of those classic forms–the sauropods, tyrannosaurs, stegosaurs, or ceratopsids–that have been cherished for the past century. Paleontologists only recently began to uncover their bones. Alvarezsaurus itself was named in 1991, but it and its close relatives didn’t quite get swept up in the same wave of dinomania as their other Mesozoic cousins.

Alvarezsaurs weren’t big, toothy, or menacing. That’s part of makes them so special. Alvarezsaurus, Mononykus and their relatives from Cretaceous Asia, South America and North America were small dinosaurs–these feathered dinos ranged from the size of a pigeon to about the size of a turkey. In fact, these dinosaurs were so avian in nature that there was once a debate about whether alvarezsaurs were non-avian dinosaurs or birds that had lost the ability to fly. Since those early debates, numerous studies have confirmed that they were non-avian dinosaurs that were closely related to the strange therizinosaurs and ostrich-like ornithomimosaurs.

But the strangest thing of all is the mystery of what alvarezsaurs ate.

Despite being short, alvarezsaur arms weren’t wimpy. Not at all. Alvarezsaur forelimbs were very stout and included one robust finger tipped in a big claw. (Among these dinosaurs, the total number and development of the fingers varied, but they’re connected by having one finger that was bigger than the others.) In contrast, these dinos often had a reduced number of very small teeth. Paleontologists thought they saw a connection between these traits and a life feeding on social insects. Mammals such as pangolins and ant-eaters also have stout, heavy-clawed arms and are toothless–a functional pairing that goes with a life of tearing into ant and termite nests to slurp up the scurrying insects in their nests.

Could alvarezsaurs have done the same? So far, it’s the most popular hypothesis for their bizarre nature. In a 2005 paper, paleontologist Phil Senter proposed that Mononykus would have been capable of the kind of scratch-digging needed to rip open social insect nests. Then, in 2008, Nicholas Longrich and Philip Currie described the alvarezsaur Albertonykus in deposits that also contained traces of Cretaceous termites. Alvarezsaurs seemed to have the right equipment and live at the right time to be social insect predators.

But we don’t really know. No one has published any direct evidence that Albertonykus or any other alvarezsaur ate ants or termites. The hypothesis is certainly a reasonable one, but we still need a test of the idea. Fossil feces may eventually hold the answer.

If paleontologists eventually uncover dinosaur dung of appropriate size that contains ants or termites and comes from a habitat shared by alvarezsaurs, that discovery would strengthen the ant-eating hypothesis. A cololite would be even better. While coprolites are petrified feces that have already been excreted, cololites are fossil poop preserved inside the prehistoric creature’s body prior to expulsion. If paleontologists found an alvarezsaur with a cololite containing termites, that would be direct evidence that these dinosaurs truly did snarf down hordes of insects. For now, though, we can only hope that some lucky fossil hunter makes such a discovery.

Sinocalliopteryx feeds on the dromaeosaurid Sinornithosaurus (left) and the early bird Confuciusornis (right). Art by Cheung Chungtat, from Xing et al., 2012.

Earlier this week, I got into a snit over the blinkered assertion that feathery dinosaurs are lame. I argued the opposite point–as I wrote at the time “Feathered dinosaurs are awesome. Deal with it.” How fortunate that a new paper this week offers proof of fuzzy dinosaur superiority. The evidence comes in the form of gut contents found within predatory dinosaurs that stalked Cretaceous China around 125 million years ago.

The carnivores in question are a pair of Sinocalliopteryx. These dinosaurs were close cousins of the much earlier Compsognathus, albeit quite a bit larger. While Compsognathus was turkey-size, about three feet long, Sinocalliopteryx grew to be about eight feet long. And this big predator was fluffy. The original description of the dinosaur mentioned the vestiges of simplified dinofuzz around the body of Sinocalliopteryx, and this makes sense given the dinosaur’s relationships. While considerably bigger than its close relatives, Sinocalliopteryx was a compsognathid–a group of theropod dinosaurs that also includes fuzzy forms such as Sinosauropteryx and Juravenator. Big or small, the compsognathids were hunters wrapped in wispy plumage.

And the initial description of Sinocalliopteryx mentioned something else. The skeleton that formed the basis of the original paper contained the leg of an unidentified dromaeosaurid dinosaur in its gut contents. Even though dromaeosaurids have long been cherished as sickle-clawed uber-predators, Sinocalliopteryx had clearly eaten the drumstick of one of the smaller feathered predators. Since then, paleontologists have identified a second Sinocalliopteryx with gut contents, and the two dinosaurs form the basis of a new PLoS One study by University of Alberta paleontologist Lida Xing and colleagues.

Looking back at the first Sinocalliopteryx, Xing and colleagues identified the victim as Sinosauropteryx. The second Sinocalliopteryx specimen had a different menu before it perished–its stomach contains the remains of two Confuciusornis, an archaic bird, and bones from an unidentified ornithischian dinosaur. But these gut contents invoke an aggravating mystery. Did these Sinocalliopteryx hunt their dinosaurian prey, or did they scavenge their meals?

This isn’t the first time paleontologists have puzzled over the meaning of predatory dinosaur gut contents. Earlier this year, Dave Hone and collaborators investigated a pterosaur bone found inside a Velociraptor, and last year Jingmai O’Connor and colleagues described a Microraptor with the remains of a bird in its gut (just to pick two examples of many). Frustratingly, though, it’s difficult to say how the dinosaurs obtained the meat. In the case of the Velociraptor, the researchers could not rule out hunting even though scavenging seemed the more likely option. Likewise, even though O’Connor and co-authors suggested their Microraptor hunted birds in the trees, the non-avian dinosaur could have just as easily scavenged a dead bird that fell to the forest floor. Gut contents tell us about what dinosaurs consumed, but they almost never provide direct evidence of how carnivores obtained flesh and bone to eat.

In the case of Sinocalliopteryx, the PLoS One study concludes that the dinosaur may have been skilled at catching live avian prey. The fact that one Sinocalliopteryx fed on two Confuciusornis in quick succession could mean that the large dinosaur was adept at nabbing early birds. “[T]he evidence of bird predation in Sinocalliopteryx,” Xing and colleagues conclude, “suggests that it was a highly capable stealth hunter.” Then again, the same researchers also note that their scenario “is speculative.” While it may seem improbable, the Sinocalliopteryx in question could have scavenged one or both of those birds, as well as the non-avian dinosaur remains in its stomach. We just don’t know. Like many predators, Sinocalliopteryx most likely hunted live prey and took advantage of carrion. Frustratingly, these fossil gut contents can’t tell us what happened in each case. Sinocalliopteryx may very well have been a skilled bird-slayer. Or perhaps not. The fact is that we don’t know for sure.

Perplexing feeding habits aside, there’s something else about the gut contents of Sinocalliopteryx that can give us a closer look at the dinosaur’s biology. In the dinosaur that ate the two birds and the ornithischian, the bone of the ornithischian dinosaur was corroded by stomach acid. The more delicate bird bones, by contrast, had not been so damaged. This means that the Sinocalliopteryx ate the ornithischian first, followed by one bird and, later, another. More than that, the acid damage indicates that at least some dinosaurs had highly-acidic foreguts where bone was broken down–comparable, but not exactly like, the stomachs of crocodilians and perhaps some bone-eating birds like the bearded vulture.

All of which is to say that Sinocalliopteryx is a great example of a fluffy dinosaur you wouldn’t want to mess with. Even if we can’t discern the backstory of each meaty morsel, the variety of prey in the Sinocalliopteryx stomachs shows that this dinosaur wasn’t a picky eater and may have even been a quick hunter that specializing in snapping up other feathery dinosaurs. For our fuzzy mammalian predecessors, hiding the Cretaceous forests, this would have been one scary dinosaur.

Reference:

Xing L, Bell PR, Persons WS IV, Ji S, Miyashita T, et al. (2012) Abdominal Contents from Two Large Early Cretaceous Compsognathids (Dinosauria: Theropoda) Demonstrate Feeding on Confuciusornithids and Dromaeosaurids. PLoS ONE 7(8): e44012. doi:10.1371/journal.pone.0044012

A reconstructed Acrocanthosaurus at the North Carolina Museum of Natural Sciences. Photo by the author.

Compared to mounted dinosaur skeletons, fossil footprints might seem like mundane objects. They only record one small part of a fantastic creature, and it is harder to envision a whole dinosaur from the ground up than the wrap flesh around a skeletal frame. But we should not forget that dinosaur footprints are fossilized behavior—stone snapshots of an animal’s life. And sometimes, trackways record dramatic moments in dinosaur lives.

In 1938, American Museum of Natural History paleontologist Roland T. Bird traveled to Glen Rose, Texas to investigate rumors of huge dinosaur tracks found in the vicinity of the Paluxy River. Bird found them in abundance, but one site was especially intriguing. Set in 113-million-year-old rock were the footprints of a huge sauropod dinosaur—and it seemed that the long-necked giant was followed. The large, three-toed footprints of a predatory dinosaur, probably the ridge-backed Acrocanthosaurus or a similar theropod, paralleled and eventually converged on the footsteps of the sauropod. And at the point of overlap, the predator seemed to skip a step—a little hop that Bird took to mean that the carnivore had sunk its teeth into the herbivore and was lifted out of its tracks a short distance.

Bird excavated the trackway in 1940. About half of the long trail went to the AMNH and can now be seen stretching out behind the museum’s Apatosaurus mount, despite the fact that Apatosaurus lived millions of years before the tracks were made. The other portion is housed at the Texas Memorial Museum in Austin. Bird’s hypothesis about how the tracks were made has inspired exhibits at other museums, such as the Maryland Science Center and the North Carolina Museum of Natural Sciences. Yet not everyone agrees about what the tracks represent. Do they record an Acrocanthosaurus attack as it happened? Or could the trackway simply be a fortuitous association of tracks from dinosaurs that walked the same ground at different times?

Artist David Thomas and paleontologist James Farlow went back to Bird’s notes and the trackway evidence to reconstruct what might have transpired. The association between the sauropod and theropod tracks seemed too tight to just be coincidence. The predatory dinosaur very closely followed the pathway of the larger herbivore, both moving along a broad left curve. Near the end of the excavated area, both the theropod and sauropod turned abruptly to the right. If the two dinosaurs had passed at different times, then we’d expect that the sauropod or theropod would have continued on in the same trajectory and crossed another set of tracks preserved nearby. Based on the fully reconstructed image, the sauropod and theropod were interacting with each other.

And there’s something else. Just before the enigmatic double-right-footprints made by the theropod, there is a drag mark made by the sauropod’s right hind foot. This might be where the titan was attacked and faltered, or maybe the sauropod threw its weight to avoid being bitten. Frustratingly, we can’t know for sure. And the missing left theropod footprint isn’t a clear sign of an attack, either—all we know is that there’s a missing track right where the animals were in close proximity.

Whether or not the Paluxy River Trackway records a successful Acrocanthosaurus assault is uncertain. But the tight connection between the theropod and sauropod tracks suggests that the carnivore at least stalked the herbivore, and perhaps even took a swipe at it. Specimens like this test our ability to draw brief moments in time from stone. The task is made all the more complicated by the gradual loss of information contained within the rock. While they look sturdy, trackways are actually fragile fossils, and the half of the trackway at the Texas Memorial Museum has significantly deteriorated since it was put on display. The museum is trying to raise a million dollars to properly conserve and house this historically and scientifically significant fossil. If you wish to learn more about their campaign, you can find more information here.