The Ichnology of Jurassic Park

All paleontologists remember their first time. Mine was in 1993, during a hot, steamy summer in Atlanta, Georgia. I had just spent the previous month camping in wilderness areas of Wyoming, so coming back to a big city with all of its urban temptations meant I was weak and susceptible to seeking out unusual sources of pleasure. Although I was not quite ready to be taken for such an exhilarating ride, it was an experience I’d never forget. Afterwards, once I had recovered enough to be able think about it, I wanted to do it again.

I am, of course, talking about seeing the film Jurassic Park on a movie screen. Sure, this movie has been around long enough (20 years) that nearly every paleontologist has also watched it on a TV, computer, or mobile device. But there is something about seeing dinosaurs – which, let’s face it, are most famous for their size – shown big. The initial glimpse of a Brachiosaurus munching on the tops of tall trees, a herd of Paralophosaurus ringing a glistening lake, an ill Triceratops dwarfing its human caretakers, the grand entrance of the Tyrannosaurus: all of these “actors” were meant to be seen large, and fill us with awe. It worked. Plot, acting, and science aside, Jurassic Park was, and probably still is, the best movie made for conveying what it would feel like for us humans, separated by a minimum of 65 million years, to see real, living dinosaurs.

“It’s, it’s a dinosaur.” That pretty much said it all in 1993, and still does. But what traces were being made by this Brachiosaurus as it strolled through its all-you-can-eat salad bar, known to you and me as a “landscape”? Please read on.

In 1993, though, I did not have an appreciation for some of the smaller details included in this film, and how my research specialty of ichnology – the study of traces, like tracks, burrows, and nests – was reflected throughout it. What dinosaur traces were included in the movie, and how were these traces used to serve or advance the plot? I also wondered how 20 years of field experience and scholarly research in ichnology might have changed my perceptions of it since that first viewing.

So with the re-release of Jurassic Park in 3-D last week, I decided it was time to view it from an ichnological perspective and share these thoughts with others. After all, my next book, Dinosaurs Without Bones (Pegasus Press), is about dinosaur trace fossils, and written for a popular audience. Also, in between the movie’s first release and now, I wrote two editions of a college textbook on dinosaurs (Introduction to the Study of Dinosaurs). Thus I went to the theater well justified in watching Jurassic Park once more, to see for myself how dinosaur traces were portrayed in the most well-loved of all dinosaur movies. And oh yes, for the fun.

For the sake of simplicity, I’ve divided these traces into two categories: those viewers could directly observe in the film, and others that could be inferred from the dinosaurs’ behaviors. Wherever possible, I also connect traces shown in the movie to research done on dinosaur trace fossils during the last 20 years, giving a broad sense of how far dinosaur ichnology has progressed since 1993.

(Ichnologist’s note: Even though all of the live dinosaurs in the movie were set in the 1990s, the study of their modern traces still qualifies as neoichnology. In contrast, any reference I make to actual dinosaur trace fossils is paleoichnology. Just so you know.)

Dinosaur Traces in Jurassic Park

Velociraptor hatchling traces. Jurassic Park shows two different but complementary examples of hatchling traces for “Velociraptor.” (I will call this dinosaur Velociraptor throughout this post, but as most dino-philes know, the director, Steven Spielberg, scaled up the Late Cretaceous Velociraptor to maximize its frightfulness. Hence it is actually more like the Early Cretaceous Deinonychus or Utahraptor.)

One is an egg-emergence trace, which is the hole left in an eggshell where a dinosaur broke out of its egg. In this scene, a cooing Velociraptor hatchling pokes its cute little nose out of its egg. (This nose, if everything worked out for it, would some day would be warmed by fresh human viscera.) We first witness this tracemaking in the Jurassic Park laboratory toward the start of the film, the same day most of the protagonists arrive on the island (Isla Nublar). As far as I know, such trace fossils have not been interpreted from the fossil record, or if they have, they have not been referred to as trace fossils: which they should be.

The next day, after dinosaur paleontologist Alan Grant and his two companions – Lex and Tim Murphy – were sufficiently terrified (and enthralled) by various dinosaur encounters out in the park, they come across a Velociraptor nest. The nest has about 15-20 broken eggs, and the fracture patterns of the eggshells provide clear evidence of hatching. But these traces also had tiny, two-toed tracks leading away from them. The tracks, with two toes studded by digital pads, were typical for deinonychosaurs. However, unlike nearly every theropod track I’ve seen, these tracks lacked claw marks at their ends. (Tsk, tsk, says this nitpicking ichnologist.)

Baby-Velociraptor-Traces-JPAw, look at the cute little Velociraptor tracks and hatched eggs. Don’t these traces just make you want to say, “Who’s the cutest little predator in the world?” Still from Jurassic Park (1993), taken from www.anyclip.com.

Even though these tracks were flashed on the screen for only a few seconds, what’s really cool is how they convey three important pieces of information. One is that the Velociraptor chicks hatched after the torrential rainstorm of the previous night, and thus only mere hours before our wandering heroes saw their traces. Second, the tracks demonstrate that the hatchlings were not altricial, but ready to move and leave the nest immediately, presumably without parental care. Third, Dr. Grant realizes that these successfully fertilized and hatched eggs mean the “female-only” genetic fail-safe plan for the dinosaurs just got disproved. In other words, these traces mattered.

One point about that nest: as far as I could tell from, this Velociraptor mother did not make a rimmed structure to protect the eggs, such as those made by another Late Cretaceous theropod, Troodon, or Late Cretaceous sauropods in Argentina. Instead, the eggs were laid out in the open, like some ground-nesting shorebirds might do. In contrast, the Jurassic Park sequels featured Velociraptor nests that were much more overt as traces, such as a rimmed nest seen in Jurassic Park III.

Troodon-Rim-NestPartially preserved rimmed nest structure of Troodon, a Late Cretaceous theropod that lived in what we now call Montana. The rim has eroded quite a bit since its discovery in the mid-1990s; the Troodon egg clutch was in the area of the foreground before its extraction. (Photograph by Anthony Martin; scale in centimeters.)

Triceratops feces. “That is one big pile of sh*t,” observes Dr. Ian Malcolm, the “chaotician” (mathematician) who supplies both pessimism and comic relief throughout the movie. In this scene, where the main protagonists attend to an under-the-weather Triceratops, two impressive piles of fecal material inspire Dr. Ellie Satler, a paleobotanist, to figure out whether or not the ceratopsian had eaten any toxic plants.

Somehow I suspect this scene was meant as a metaphor for what most paleontologists have to do in their day jobs in order to do any paleontology at all.

Still, when added together, this amount of still-moist waste was far too voluminous to have been from one or two depositional events: I mean, this dinosaur was sick, but not that much. As a result, park personnel must have been responsible for making these dung heaps from several days worthy of Triceratops contributions. (Strictly speaking, then, these heaps were composite traces.) If so, this would have been a rather unenviable job, but maybe they were better paid than Dennis Nedry, the disgruntled computer programmer who later provided ample fodder for Dilophosaurus.

Unfortunately, fossil Triceratops feces (coprolites) are thus far unknown from the geologic record. What is exciting, though, is that several excellent studies have been done by Dr. Karen Chin on Late Cretaceous hadrosaur coprolites. These coprolites, like the fictionalized Triceratops feces, contain lots of plant material, telling us much about what these hadrosaurs ate 75 million years ago. They also tell us what ate the feces or grazed on them, which were dung beetles and snails, respectively. (Indeed, I now wonder if Isla Nubar had a severe shortage of dung beetles, which might explain how those Triceratops feces got piled higher and deeper.)

Two-Medicine-CoproliteDinosaur coprolite – probably from a large hadrosaur, such as Maisaura – and filled with wood fragments, accompanied by special bonus trace fossils: dung beetle burrows! Specimen from the Two Medicine Formation (Late Cretaceous, Montana) as part of a Museum of the Rockies traveling exhibit at Fernbank Museum of Natural History. (Photograph by Anthony Martin, taken in 2001 and scanned from a 35-mm slide; scale in centimeters.)

• Tyrannosaurus tracks. Probably the most memorable scene in Jurassic Park is the grand entrance of the Tyrannosaurus, whose approach is first detected by “impact tremors” transmitted in cups of water on the dashboard of a jeep. Following this first bout of terror and the arrival of Ellie Sattler and big-game hunter Robert Muldoon, Malcolm, convalescing in the back of a jeep, looks down at a three-toed Tyrannosaurus track. The track, filled with water from the rain, communicates a warning as it vibrates from the footfalls of the approaching giant theropod. This repeats the previous image of the trembling water in the cup, but is made doubly dreadful by happening in a freshly made footprint of the same animal causing the tremors.

So what was by far the most exciting moment in the movie for me, ichnologically speaking? The Tyrannosaurus making a track, in which mud pushes up and out to the sides of its right foot, observed at 2:38 in the following video clip. Just watch:

This was already a great scene for all of its action, suspense, and lawyer eating. But check out that track-making!

Only a few fossil tracks have been attributed to Tyrannosaurus or other tyrannosaurids, mostly for being the right size (really big) and geologic age (Late Cretaceous). One was discovered in New Mexico in 1983, but wasn’t reported in a scientific journal until the year after Jurassic Park came out. More than a decade passed before another was found in Montana in 2007 and reported in 2008. Tragically, both were isolated tracks, and a Tyrannosaurus trackway, with two or more consecutive steps, has not yet been found. If so, it would make for a pretty darned nice find. So please do let the world know if you find one.

Large-Theropod-Track-CretaceousA large and well-preserved three-toed theropod track from the Early Cretaceous Glen Rose Formation of Texas, made about 95 million years ago. Although this track was more likely made by Acrocanthosaurus, rather than Tyrannosaurus rex, you can be assured that this theropod, like all living things, was a distant relative of T. rex. (Photograph taken by Anthony Martin; scale in centimeters.)

• Velociraptor tracks (adults). These tracks, shown only for a few seconds, are outside of the Velociraptor enclosure after the power was shut down. Muldoon, accompanied by Sattler, spots the footprints, and he wordlessly identifies them. (His expression also tells the audience that Sattler and he are going to be in deeper doo-doo than the Triceratops piles.) The twisted and broken bars of the enclosure provide additional traces affirming the conclusion that these ‘raptors are on the loose. All of these traces are shown only minutes before Muldoon utters his meme-inspiring last words, “Clever girl.”

Tracking-Velociraptors-JPUh oh: Velociraptor tracks, and these don’t look like they’re from hatchlings. Good thing Muldoon is a big-game hunter, who’s skilled at tracking and predicting Velociraptor behavior based on their tracks. But too bad his hypothesis was falsified in such an unpleasant way, but I suppose he could have picked kinder reviewers. Still from Jurassic Park (1993), taken from www.anyclip.com.

Deinonychosaur-Track-UtahHere’s what a real deinonychosaur track looks like. This one, from the Early Cretaceous of Utah, is a right foot impression, and is just slightly smaller than the adult tracks depicted in Jurassic Park. Notice the digits are thinner and end with sharp clawmarks, too. (Photograph by Anthony Martin; scale in centimeters.)

• Bioerosion of fossil dinosaur bones by modern dinosaurs. Toward the end of the film, the main human heroes – Grant, Sattler, Murphy, and Murphy (which sounds like a law firm, but we know how T. rex feels about those) – are confronted by a Velociraptor pack in the Jurassic Park visitor center. During their attempts to flee the ‘raptors, both species end up disarticulating and breaking some of the mounted dinosaur skeletons in the atrium of the visitor center. Their actions were thus a form of bioerosion, in which the fractured dinosaur bones are traces of their activities. Alternatively, the bones may have been artificial casts, in which case their breakage would have constituted bioerosion of modern substrates.

This bioerosion is made more complicated when the Tyrannosaurus rex (who everyone agrees is the ultimate protangonist of the movie) enters the atrium and, among other antics, smashes a skeleton of itself with a recently crunched Velociraptor. As a result, the jumbled assemblage of bones at the end is attributable to three separate, interacting tracemakers: four humans (two adult, two juvenile), two Velociraptors (both adults), and one Tyrannosaurus (adult). What should be noted, though, is that if the Velociraptor was already dead when flung by the Tyrannosaurus, then the broken skeleton is a trace of the Tyrannosaurus, not the Velociraptor. In other words, the Velociraptor body was just being used as a tool.

Bioerosion in action, as a result of Velociraptor and human interactions. Also, at 2:45: T. rex smash!

Dinosaur Trace-Making Behaviors in Jurassic Park

• Brachiosaurus tracks, browsing traces. When Drs. Grant and Sattler experience their first jaw-dropping glimpse of a Brachiosaurus, they watch it rear up on its hind feet, and come down hard on front feet. Considering that a Brachiosaurus this size might have weighed at least 30 tonnes, it surely would have left deep tracks in both the front and rear from the increased stresses imparted by these actions. Also, its cropping the tops of trees would have caused some easily visible gaps in the canopies of forests on Isla Nubar.

• Theropod toothmarks. Part of the fun of Jurassic Park was indulging in our worst nightmares about these non-avian theropods frequently sampling human flesh. Assuming that the theropod teeth in each instance penetrated skin and muscle and contacted bone, toothmarks would have included the following: (1) Tyrannosaurus toothmarks on goat, human, and Velociraptor bones; and (2) Velociraptor and Dilophosaurus toothmarks on human bones.

• Triceratops resting trace. When the paleontologists and others visit the ailing Triceratops, it was lying on its right side. I couldn’t help but think that if Triceratops or any other large ceratopsian dinosaur ever reclined like that, and in the right type of sediment, it would have left a gorgeous body impression. This scene also reminded me of bison traces I’ve seen in Yellowstone National Park, in which bisons roll onto their sides for a dust bath, leaving outlines of their bodies. Did dinosaurs – especially the feathered ones – ever take dust baths, and leave similar body impressions? We don’t know yet, but such trace fossils are something to look for.

• Dinosaur stampede. One of the most astonishing computer-generated effects of Jurassic Park, and one that was especially effective in 3-D, was of a dinosaur stampede. In this scene, a flock of Gallimus run toward and around Grant, Murphy, and Murphy, just before the ambush-hunting Tyrannosaurus rex slaughters one of them (the Gallimus, that is). I’ve written about this scene before, connecting it to a dinosaur tracksite in Queensland, Australia that has more than 3,000 tracks preserved. Although the site was originally interpreted as the only evidence of a dinosaur stampede, the tracks were recently reinterpreted as swim tracks. I’ll write about this topic at length in my upcoming book, so for now, I ain’t saying nothing more.

Run away, run away!

• Tyrannosaurus drag mark. After the Tyrannosaurus rex decides that a measly goat was just an appetizer and begins seeking out the nearest available mammals for nomming purposes, at some point it overturns and begins pushing an SUV, which still has Lex and Tim Murphy trapped underneath it. Its flipping the SUV with its head certainly would have left a substantial mark on the muddy ground, a trace that then would have been connected to a semi-circular dragmark (clockwise oriented), and with tracks directly adjacent to these traces. Her tracks also may have been deeper in their fronts because of her head being down as she pushed, reflecting a shift in her weight distribution. However, I should again remind everyone that just like with the dead Velociraptor used for bioerosion by this same T. rex later in the film, the SUV is not making the trace. It is only a tool being used by the tyrannosaur, which is the real tracemaker.

• Tyrannosaurus running trackway – This pulse-quickening chase scene, in which the T. rex pursues a jeep driven by Muldoon and with Malcolm and Sattler as passengers, very likely would have caused a wonderful sequence of tracks. These tracks would have shown increasing stride lengths from a standing start to full-speed run, and each successive track would have registered external and internal structures consistent with this acceleration. Even better, the tracks would have cross-cut the jeep tire-tracks at some points, demonstrating to a later observer that the tyrannosaur was very likely following the jeep. (The demolition of a low-hanging tree branch by the T. rex during the chase also counts as some bioerosion along the way.) Some convincing studies have been done since showing that an adult Tyrannosaurus could not have moved as fast as the one in Jurassic Park, but it still could have caught most running humans. And just to repeat what I said earlier, it’d be really nice for someone to find a T. rex trackway, which would give us more direct evidence of how these massive theropods moved.

• Velociraptor scratch marks and other traces. This time, while watching the harrowing and claustrophobic scene in which a pair of Velociraptors hunt the Murphy siblings in a kitchen, I started thinking about the traces they might have left. Did their claws leave scratch marks on the door handles and kitchen counters? Did they indent the steel counters when they jumped up on these surfaces? A broken window is also shown as a trace of their smashing through glass once they became frustrated with a locked door.

OK, enough of the fanciful ichnology. What about other dinosaurs and their traces? Well, it turns out that Jurassic Park saved the real, living dinosaurs for the very end of the movie. These were five brown pelicans (Pelecanus occidentalis), flying in formation as Grant, Sattler, and their companions leave Isla Nubar in a helicopter. For Grant, this is a poignant moment, as he is likely reflecting on how dinosaurs were still with us today as birds. With that thought, I will say “amen,” and add that dinosaur traces – tracks, nests, feces, bite marks, and all – are still here with us, too, and don’t require special glasses to see them in three dimensions. Thanks for that reminder, Jurassic Park.

Pelican-Tracks-SapeloWant to see some modern dinosaur traces? Here are tracks of a brown pelican, made in wet sand while it was loafing on a beach at low tide on Sapelo Island, Georgia. To see more modern dinosaur traces, just go outside, and you’ll find them wherever birds are found. (Photograph by Anthony Martin; scale in centimeters.)

Further Reading

Chiappe, L.M., Schmitt, J.G., Jackson, F., Dingus, L., and Grellet-Tinner, G. 2004. Nest structure for sauropods: sedimentary criteria for recognition of dinosaur nesting traces. Palaios, 19: 89–95.

Chin, K. 2007. The paleobiological implications of herbivorous dinosaur coprolites from the Upper Cretaceous Two Medicine Formation of Montana: why eat wood? Palaios, 22: 554-566.

Chin, K., and Gill, B.D. 1996. Dinosaurs, dung beetles, and conifers: participants in a Cretaceous food web. Palaios, 11: 280-285.

Elbroch, M., and Marks, E. 2001. Bird Tracks and Sign of North America. Stackpole Books, Mechanicsburg, Pennsylvania.

Erickson, G. M., Van Kirk, S. D., Su, J., Levenston, M. E., Caler, W. E., & Carter, D. R. 1996. Bite force estimation for Tyrannosaurus rex from tooth-marked bones. Nature, 382: 706–708.

Gignac, P.M., Makovicky, P.J., Erickson, G.M., and Walsh, R.P. 2010. A description of Deinonychus antirrhopus bite marks and estimates of bite force using tooth indentation simulations. Journal of Vertebrate Paleontology, 30: 1169-1177.

Hutchinson, J.R., and Garcia, M. 2002. Tyrannosaurus was not a fast runner. Nature, 415: 1018-1021.

Jacobsen, A.R. 1998. Feeding behaviour of carnivorous dinosaurs as determined by tooth marks on dinosaur bones. Historical Biology, 13: 17-26.

Lockley, M.G., and Hunt, A.P. 1994. A track of the giant theropod dinosaur Tyrannosaurus from close to the Cretaceous/Tertiary Boundary, northern New Mexico. Ichnos, 3: 213-218.

Manning, P.L., Ott, C., and Falkingham, P.L. 2008. A probable tyrannosaurid track from the Hell Creek Formation (Upper Cretaceous), Montana, United States. Palaios, 23: 645-647.

Martin, A.J. 2013. Life Traces of the Georgia Coast: Revealing the Unseen Lives of Plants and Animals. Indiana University Press, Bloomington, Indiana: 692 p.

Romilio, A., and Salisbury, S.W. 2011. A reassessment of large theropod dinosaur tracks from the mid-Cretaceous (late Albian–Cenomanian) Winton Formation of Lark Quarry, central-western Queensland, Australia: a case for mistaken identity. Cretaceous Research, 32: 135-142.

Romilio, A., Tucker, R., Salisbury, S. 2013. Reevaluation of the Lake Quarry dinosaur tracksite (late Albian-Cenomanian Winton Formation, central-western Queensland, Australia): no longer a stampede? Journal of Vertebrate Paleontology. 33: 1, 102-120.

Sellers, W.I., and Manning, P.L. (July 2007). Estimating dinosaur maximum running speeds using evolutionary robotics. Proceedings of the Royal Society of London, B, 274: 2711-2716.

Thulborn, R.A., and Wade, M., 1979. Dinosaur stampede in the Cretaceous of Queensland. Lethaia, 12: 275-279.

Varricchio, D.J., Jackson, F. and Trueman, C.N. 1999. A nesting trace with eggs for the Cretaceous theropod dinosaur Troodon formosus. Journal of Vertebrate Paleontology, 19: 91-100.

 

Tracking Wild Turkeys on the Georgia Coast

Of the many traditions associated with the celebration of Thanksgiving in the U.S., the most commonly mentioned one is the ritual consumption of an avian theropod, Meleagris gallopavo, simply known by most people as “turkey.” The majority of turkeys that people will eat this Thursday, and for much of the week afterwards, are domestically raised. Yet these birds are all descended from wild turkeys native to North America. This is in contrast to chickens (Gallus gallus), which are descended from an Asian species, and various European mammals, such as cattle, pigs, sheep, and goats (Bos taurus, Sus scrofa, Ovis aries, and Capra aegagrus, respectively).

Trackway of a wild turkey (Meleagris gallopavo) crossing a coastal dune on Cumberland Island, Georgia. Notice how this one, which was likely a big male (“tom”), was meandering between clumps of vegetation and staying in slightly lower areas, its behavior influenced by the landscape. Scale = 20 cm (8 in). (Photograph by Anthony Martin.)

American schoolchildren are also sometimes taught that one of the founding fathers of the United States, Benjamin Franklin, even suggested that the wild turkey should be elevated to the status of the national bird, in favor of the bald eagle (Haliaeetus leucocephalus). With an admiring (although I suspect somewhat facetious) tone, he said:

He [the turkey] is besides, though a little vain & silly, a Bird of Courage, and would not hesitate to attack a Grenadier of the British Guards who should presume to invade his Farm Yard with a red Coat on.”

There are eight of us, and only one of you. Do you really want to mess with us? (Photograph by Anthony Martin, taken on Cumberland Island, Georgia.)

Unfortunately, because I live in the metropolitan Atlanta area, I never see turkeys other than the dead packaged ones in grocery stores. Nonetheless, one of the ways I experience turkeys as wild, living animals is to visit the Georgia barrier islands, and the best way for me to learn about wild turkey behavior is to track them. This is also great fun for me as a paleontologist, as their tracks remind me of those made by small theropod dinosaurs from the Mesozoic Era. And of course, as most schoolchildren can tell you, birds are dinosaurs, which they will state much more confidently than anything they might know about Benjamin Franklin.

Compared to most birds, turkeys are relatively easy to track. Their footprints are about 9.5-13 cm (3.7-5 in) long and slightly wider than long, with three long but thick, padded toes in front and one shorter one in the back, pointing rearward. In between these digits is a roundish impression, imparted by a metatarsal. This is a trait of an incumbent foot, in which a metatarsal registers behind digit III because the rear part of that toe is raised off the ground. The short toe is digit I, equivalent to our big toe, but not so big in this bird. Despite the reduction of this toe, its presence shows that turkeys probably descended from tree-dwelling species, as this toe was used for grasping branches. Clawmarks normally show on the ends of each toe impression, and when a turkey is walking slowly, it drags the claw on its middle toe (digit III), thus making a nicely defined linear groove.

Wild turkey tracks made while it was walking slowly up a gentle dune slope, dragging the claw on the middle digit of its right foot, making a long groove. Also notice the bounding tracks of a southern toad (traveling lower right –> upper left), cross-cutting the turkey tracks. (Photograph by Anthony Martin, taken on Cumberland Island.)

A normal walking pace (right foot –> left foot, left foot –> right foot) for a turkey is anywhere from 15-40 cm (6-16 in), and its stride (right foot –> right foot, left foot –> left foot) is about twice that, or 30-80 cm (12-32 in), depending on the age and size of the turkey. Their trackways show surprisingly narrow straddles for such wide-bodied birds, only 1.5 times more than track widths. This is because they walk almost as if on a tightrope, with angles between each step approaching 180°; so they still make a diagonal pattern, but nearly define a straight line. However, turkeys meander, stop, or change direction often enough to make things interesting when tracking them. Their flocking behavior also means their tracks commonly overlap with one another or cluster, making it tough to pick out the trackways of individual turkeys. However, in such flocks, the dominant male’s tracks are noticeably larger than those of the females or younger turkeys, so these can be picked out and help with sorting who’s who.

Turkey trackway in which it walked across the wind-rippled surface of a coastal dune on Cumberland Island, meandering while moseying. Same photo scale as before. (Photograph by Anthony Martin.)

An abrupt right turn recorded by a turkey’s tracks. Check out that beautiful metatarsal  impression in the second track from the right, and how the claw dragmark in the thrid track from the right points in the direction of the next track. (Photograph by Anthony Martin.)

One of the more remarkable points about these Georgia barrier-island turkeys, though, is how their tracks belie their stereotyped image as forest-only birds. Although they do spend much of their time in the forest, I’ve tracked turkeys through broad swaths of coastal dunes, and sometimes they will stop just short of primary dunes at the beach. So however difficult it might be to think about these birds as marginal-marine vertebrates, their tracks overlap the same places with ghost-crab burrows and shorebird tracks. Geologists and paleontologists take note: this exemplifies the considerable overlap between terrestrial and marginal-marine tracemakers that can happen in coastal environments. This also happened with dinosaurs that strolled onto tidal flats or otherwise passed through marginal-marine ecosystems.

Turkey tracks heading toward the beach, with the open ocean visible just beyond. Is this close enough to consider turkeys as marginal-marine tracemakers? (Photograph by Anthony Martin.)

Do these turkeys also have an impact on the dunes themselves? Yes, although these effects vary, from trackways disrupting wind ripples to more overt changes to the landscape. For instance, one of the most interesting effects I’ve seen is where they’ve caused small avalanches of sand downslope on dune faces. Interestingly, this same sort of phenomenon was also documented for Early Jurassic dinosaurs that walked across dry sand dunes, which caused grainflows that cascaded downhill with each step onto the sand.

Grainflow structure (arrow), a small avalanche caused by a turkey walking down a dune face. (Photograph by Anthony Martin.)

Close-up of grainflow structure (right) connected to turkey tracks, which become better defined once the turkey reached a more level surface. (Photograph by Anthony Martin, taken on Cumberland Island.)

What other traces do turkeys make? A lot, although I’ve only seen their tracks. Other traces include dust baths, feces, and nests. Dust baths, in which turkeys douse themselves with dry sediment to suffocate skin parasites, must be awesome structures. These are described as 50 cm (20 in) wide, 5-15 (1-3 in) deep, semi-circular depressions, and feather impressions show up in those made in finer-grained sediments. Although such structures would have poor preservation potential in the fossil record, I hold out hope that if paleontologists start looking more at modern examples, they are more likely to find a fossil dust bath, whether in Mesozoic or Cenozoic rocks.

Turkey feces, like most droppings from birds, have white caps on one end, but are unusual in that these can tell you the gender of their depositor. Male turkeys tend to make curled cylinders that are about 1 cm wide and as much as 8 cm long (0.4 X 3 in), whereas females make more globular (not gobbular) droppings that are about 1 cm (0.4 in) wide. These distinctive shapes are a result of their having different digestive systems. Turkeys are herbivores, so their scat normally includes plant material, but don’t be surprised to see insects parts in them, too. Still think about how exciting it would be to find a grouping of same-diameter cylindrical and rounded coprolites in the same Mesozoic deposit, yet filled with the same digested material, hinting at gender differences (sexual dimorphism) in the same species of dinosaur maker.

Turkeys normally make nests on the ground by scratching out slight depressions with their feet, but evidently this is a flexible behavior. On at least one of the Georgia barrier islands (Ossabaw), these birds have been documented as building nests in trees. Although this practice seems very odd for a large, ground-dwelling bird, it is an effective strategy against feral hogs, which tend to eat turkey eggs, as well as eggs of nearly every other species of bird or reptile, for that matter. Just to extend this idea to the geologic past, ground nests are documented for several species of dinosaurs, but tree nests are unknown, let alone whether species of ground-nesting dinosaurs were also capable of nesting in trees.

As everyone should know from their favorite WKRP episode, domestic turkeys can’t fly. But wild turkeys can, and use this ability to get into the branches of live oaks (arrow), high above their predators, or even curious ichnologists. (Photograph by Anthony Martin, taken on Cumberland Island.)

So whether or not you have tryptophan-fueled dreams while dozing later this week, keep in mind not just the evolutionary heritage of your dinosaurian meal, but also what their traces tell us about this history. Moreover, it is an understanding aided by these magnificent and behaviorally complex birds on the Georgia barrier islands. For this alone, we should be thankful.

Paleontologist Barbie, tracking wild turkeys on the Georgia coast to learn more about how these tracemakers can be used as modern analogs for dinosaur behavior and traces, and once again demonstrating why she is the honey badger of paleontologists. (Yes, photograph by me, and taken on Cumberland Island. P.S. Happy Thanksgiving!)

Further Reading

Dickson,J.G. (editor). 1992. Wild Turkeys: Biology and Management. Stackpole Books, Mechanicsburg, Pennsylvania: 463 p.

Elbroch, M., and Marks, E. 2001. Bird Tracks and Sign of North America. Stackpole Books, Mechanicsburg, Pennsylvania: 456 p.

Fletcher, W.O., and Parker, W.A. 1994. Tree nesting by wild turkeys on Ossabaw Island, Georgia. The Wilson Bulletin, 106: 562.

Loope, D.B. 2006. Dry-season tracks in dinosaur-triggered grainflows. Palaios, 21: 132-142.

Deep in the Dinosaur Tracks of Texas

Given the continuing public mania over dinosaurs, and recent important discoveries of yet more exquisite specimens of feathered theropod dinosaurs discovered in countries far away from the U.S. (here and here), it is sometimes easy to forget what has long been known about these animals, and right here in my own “backyard” (globally speaking).

Need to see some of the best dinosaur tracks in the world, and you live in the southeastern U.S.? Guess what: you can seen them in Glen Rose, Texas. Not China, Mongolia, Canada, Utah, or some other far-off land inhabited by strange people with unusual customs, but Texas. Saddle up! (Photograph by Michael Blair, taken in Dinosaur Valley State Park, Texas.)

So on July 22, just to jog my memory a bit, I flew from Atlanta, Georgia to the Dallas-Ft. Worth (Texas) airport, and only a few hours later was gazing upon dinosaur tracks accompanied by the burrows of invertebrate animals, both trace fossils having been made more than 100 million years ago. It was a fitting welcome to Glen Rose, Texas, a place famous for its dinosaur trace fossils since the 1930s, and where dinosaurs were an integral part of its culture long before it was cool, hip, and contemporary elsewhere.

In Glen Rose, Texas, the dinosaur tracks are so abundant, you can choose whether to see these just outside of your hotel room, or go to the hotel jacuzzi and pool. Naturally, I chose both. (Photograph by Anthony Martin, taken in Glen Rose, Texas.)

So just how did I end up in Glen Rose, Texas, looking at Cretaceous dinosaur tracks and invertebrate burrows? I was lucky enough to be there as an invited participant in an expedition sponsored by the National Geographic Society. I say “lucky” because luck was certainly a part of it, a fortuitous connection made through my writing a book about the modern traces of the Georgia coast. James (Jim) Farlow, a paleontologist at Indiana-Purdue University Fort Wayne (IPFW) and an associate editor with Indiana University Press, reviewed the first draft of my book, but he was also in charge of this dinosaur-track expedition to Glen Rose. Evidently he was impressed enough about what I knew about invertebrate burrows (or at least what I wrote about them) that he considered me as a possible member for his team of scientists, field assistants, and teachers on this expedition.

Dr. Jim Farlow, the world expert on the Glen Rose dinosaur tracks, having a reflective moment at Dinosaur Valley State Park near Glen Rose, Texas. What’s with the broom? He and other people in the expedition used these to sweep river sediment out of dinosaur tracks submerged in the river. In 100° F (38° C) temperatures. On the other hand, I just described invertebrate trace fossils, which was more of a job, not work. (Photograph by Anthony Martin, taken in Dinosaur Valley State Park, Texas.)

Thus when Jim asked me last fall if I would be interested in joining them to describe and interpret the Cretaceous invertebrate burrows that occur with the dinosaur tracks there, I jumped at the opportunity. The Glen Rose dinosaur tracksites, most of which crop out in the Paluxy River bed in Dinosaur Valley State Park, are world famous for their quantity and quality, and they connect with an important part of the history of dinosaur studies. Going there, experiencing these tracks for myself, and better understanding their paleoecological and geological context would be of great benefit to me, my students, and of course, you, gentle readers.

Just to back up a bit, and clarify for anyone who doesn’t know why these tracks are so darned important, here’s a brief background. In November 1938, Roland T. Bird, an employee of the American Museum of Natural History and a field assistant to flamboyant paleontologist Barnum Brown (the guy who named Tyrannosaurus rex), saw large, isolated limestone slabs with theropod dinosaur tracks in a Native American trading post in Gallup, New Mexico. Upon inquiring about the origin of these tracks, Bird was told they came from Glen Rose, Texas. So he set out in his Buick for Glen Rose to see for himself whether these tracks were real or not, and whether there were any more to see in the rocks around Glen Rose. The theropod track set in the town bandstand – pictured below – was one of the first sites that greeted him, and Glen Rose locals told him about the tracks in the Paluxy River.

Glen Rose, Texas, the only place in the world where the town bandstand has an Early Cretaceous theropod dinosaur track on display. Wish I could also tell you about all of those little holes in the rock with that track, but I can’t right now. Nonetheless, rumor has it they are burrows made by small, marine invertebrates that lived at the same time as the dinosaurs. (Photograph by Anthony Martin, taken in Glen Rose, Texas.)

Bird had hit the jackpot, the motherlode, the bonanza, the surfeit, the, well, you get the point. Not only did the Paluxy River outcrops contain hundreds of theropod dinosaur tracks – many as continuous trackways – but also the first known evidence of sauropod dinosaur tracks.

A couple of beautifully preserved theropod tracks under shallow water in the Paluxy River. Note that the track at the bottom also has a partial metatarsal (“heel”) impression, and look closely for the digit I (“thumb”) imprint on the right. Scale is about 20 cm (8 in) long. (Photograph by Anthony Martin, taken in Dinosaur Valley State Park, Texas.)

Funny how those “potholes” in the limestone bedrock of the Paluxy River have oblong outlines and form regular alternating patterns, isn’t it? Well, them ain’t no potholes, y’all. They’re sauropod tracks, and were among the hundreds recognized as the first know =n such tracks from the geologic record. (Photograph by Anthony Martin, taken in Dinosaur Valley State Park, Texas.)

The discovery of sauropod tracks was as huge as the tracks. Up until then, sauropods were assumed to have been so large that they could not support their weights on land and spent most of their time in water bodies. These tracks said otherwise, that these sauropods were walking along mudflats along with the theropods. In short, the trace fossil evidence contradicted the assumed story about how these massive animals moved. After all, trace fossils are direct records of animal behavior, and if interpreted correctly, can tell paleontologists more about what an animal was doing on a given day than any amount of shells, bones, and yes, even feathers.

Sauropod tracks from the main tracksite in Dinosaur Valley State Park, Texas. The sauropod was moving away in this view, and the trackway pattern is a typical diagonal-walking one, right-left-right. In parts of this trackway, both the manus (front foot) and pes) rear foot registered, something Bird noticed in 1938, his observation accompanied by more than a little bit of excitement. (Photograph by Anthony Martin, taken in Dinosaur Valley State Park, Texas.)

The details preserved in these sauropod tracks are also astounding. Most sauropod tracks I have seen elsewhere, in Jurassic and Cretaceous rocks of the American West, Europe, and Western Australia, are only evident as large, rounded depressions that you would only know are tracks because they form diagonal-walking patterns. In contrast, the Glen Rose tracks include all five toe and claw impressions on the rear feet (pes) and full outlines of the front feet (manus). The original calcium-carbonate mud in the shoreline environments where the sauropods walked, similar to mudflats I’ve seen in the modern-day Bahamas, is what made this exquisite preservation possible. The mud had to be firm enough to preserve these specific details of the sauropods’ feet, but not so soft that the mud would collapse into the tracks after the sauropods extracted their feet.

Beautifully preserved tracks, manus (top) and pes (bottom). Note the five toe impressions in the pes, which along with its size confirms that these were made by a large sauropod. Meter stick for scale. (Photograph by Anthony Martin, taken in Dinosaur Valley State Park, Texas.)

One sauropod trackway, preserved with a theropod trackway paralleling and intersecting it, was actually quarried out of the river and taken to the American Museum. Once there, its pieces stay disassembled for years, before Bird helped with putting the puzzle pieces back together so that it could be used as part of a display there.

Archival video footage of Roland Bird and his field crew working on the dinosaur tracks in the Paluxy River near Glen Rose, Texas. More about this tracksite and its role in the history of dinosaur paleontology is ably conveyed by Brian Switek here.

Photos at the visitor’s center at Dinosaur Valley State Park, showing the sequence of clearing (left) and extraction (right) of the limestone bed containing the theropod and sauropod dinosaur tracks. (Photographs taken of the photographs, then enhanced, cropped, and placed side-by-side by Anthony Martin.)

A lasting trace today of Roland Bird and his field helpers from the 1940s, in which they took out a sauropod and theropod trackway from this place and transported it to New York City. (Photograph by Anthony Martin, taken in Dinosaur Valley State Park, Texas.)

Other than some of the best-preserved Early Cretaceous dinosaur tracks in the world, one other claim to fame for the Glen Rose area, and not such a proud one, is its attraction to evolution deniers, a few charlatans who used the tracks to promote what might be mildly termed as cockamamie ideas. You see, Glen Rose is also the site of the infamous “man tracks.” These tracks are preservational variants of theropod tracks that – through a combination of the theropods sinking into mud more than 100 million years ago and present-day erosion of the tracks in the Paluxy River – prompted some people to claim these were the tracks of biblical giants who were also contemporaries of the dinosaurs. (Perhaps this is as good of a time as any to start humming the theme music for The Flintstones.)

Rare documentary footage of humans and dinosaurs interacting with one another during the Early Cretaceous Period, or the Late Jurassic Period. Whatever. Note the inclusion of other seemingly anachronistic mammals, too, such as the saber-toothed felid Smilodon. Perhaps this footage could be included in the curriculum of some U.S. public schools, providing a formidable counter to the views of 75 Nobel laureate scientists. Then we’ll let the kids decide which is right.

I will not waste any further electrons or other forms of energy by continuing to flog this already thoroughly discredited notion, but instead will direct anyone interested to a thorough accounting of this debacle to some actual scholarship here, summarizing original research by Glen Kuban and others in the 1980s through now that have laid to rest such spurious notions. Speaking of Mr. Kuban, I was delighted to meet him for the first time during while in Glen Rose (we had corresponded a few times years ago). I was even more gratified to spend a few hours in the field with him, discussing the genuinely spectacular trace fossils there in Dinosaur Valley State Park with these directly in front of us. Again, I’m a lucky guy.

The expedition was scheduled in Glen Rose for three weeks during late July through early August, but with so many commitments for this summer, I could only carve out a week for myself there, from July 22-29. Fortunately, this was enough time for me to accomplish what was needed to do, while also having fun getting to know the rest of the expedition crew – teachers, artists, videographers, laborers – and enjoying wonderful discussions (and debates) with colleagues in the field. The people of Glen Rose were also exceedingly welcoming and accommodating to us: we felt like rock stars (get it – “rock”?), and were feted by local folks three nights in a row during the week I was there. Many thanks to these Glen Rose for the the exceptional hospitality they extended to our merry band of paleontologists, geologists, river sweepers, or what have you.

You can’t see it, but I’m standing in a sauropod dinosaur track, which is a little deeper than the rest of the river bed. You also can’t see the invertebrate burrows that are in the limestone bedrock, which is fine, because I can’t show them to you yet anyway. But be patient: you’ll learn about them some day. (Photograph by Martha Goings, taken in Dinosaur Valley State Park, Texas.)

I can’t yet say much more about what I did during that week, as all participants signed an agreement that National Geographic has exclusive rights to research-related information, photos, and video unless approved by them. But if you’re a little curious about the daily happenings of the expedition (which just ended last week), Ray Gildner maintained a blog that succinctly touched on all of the highlights, Glen Rose Dinosaur Track Expedition 2012.

Still, I can say, with great satisfaction, that I did successfully describe and interpret invertebrate trace fossils that were in the same rocks as the dinosaur tracks. Hopefully my colleagues and I will have figured out how these burrows related to environments inhabited by the dinosaurs that walked through what we now call Texas.

All in all, my lone week in the Lone Star State was a marvelously edifying and educational experience, one I’ll be happy to share with many future generations of students and all those interested in learning about the not-so-distant geologic past of the southeastern U.S.

Group photo from the Glen Rose Dinosaur Track Expedition 2012. Names of all participants can be found here, but in the meantime, just sit back and admire those Dinosaur World t-shirts everyone is wearing. (Photograph by James Whitcraft or Ray Gildner: they can fight over who actually took it. Although, the automatic timer on his camera took the photo, so maybe it should get credit instead.)

Further Reading

Bird, R.T. 1985. Bones for Barnum Brown: Adventures of a Dinosaur Hunter. Texas Christian University Ft. Worth, Texas: 225 p.

Farlow, J.O. 1993. The Dinosaurs of Dinosaur Valley State Park. Texas Parks and Wildlife Department, Austin, Texas: 30 p.

Jasinski, L.E. 2008. Dinosaur Highway: A History of Dinosaur Valley State Park. Texas Christian University, Ft. Worth, Texas: 212 p.

Kuban, G.J. 1989. Elongate Dinosaur Tracks. In Gillette, David D., and Martin G. Lockley (editors), Dinosaur Tracks and Traces, Cambridge University Press, Cambridge, U.K.: 57-72.

Pemberton, S.G., Gingras, M.K., and MacEachern, J.A. 2007. Edward Hitchcock and Roland Bird: Titans of Vertebrate Ichnology in North America. In Miller, William, III (editor), Trace Fossils: Concepts, Problems, Prospects. Elsevier, Amsterdam: 32-51.

“Worm Burrows” as a Geological Cliché

This past week, I was privileged to have participated in a marvelous three-day field trip to the Triassic and Jurassic sedimentary rocks in and around St. George, Utah. The field trip, organized by paleontologist Andrew Milner and many others in association with the Society of Vertebrate Paleontology meeting in Las Vegas, Nevada, provided our enthusiastic group of nearly forty professional and amateur paleontologists with a grand geological tour of southern Utah and northern Arizona, along with the fantastic dinosaur tracksites in that area.

Foremost among these places where dinosaurs left their marks was one of the most incredible tracksites have seen anywhere, which, like Lark Quarry in Queensland, Australia, is enclosed within a building to protect it. This place, called the St. George Dinosaur Discovery Site at Johnson Farm, has one of the few sitting-dinosaur trace fossils known from the fossil record, along with the world’s best collection of dinosaur swimming tracks, rare examples of dinosaur tail-drag marks, hundreds of other dinosaur tracks, and thousands of invertebrate trace fossils. All were enthralling as detailed records of daily life in the Early Jurassic Period, from about 195 million years ago.

You would think on a field trip like this that Georgia – countering Ray Charles’ memorialized sentiment – would not be on my mind. Yet the modern traces made by living animals of the Georgia barrier islands habitually creep into my thoughts whenever I travel into the geological past. In this instance, the trigger for my thoughts of Georgia traces was through hearing other field-trip participants utter the most recurring of geological clichés connected to invertebrate trace fossils: “worm burrows.”

Invertebrate trace fossils (left) directly associated with theropod dinosaur footprints (right) from the Moenave Formation (Lower Jurassic), southern Utah. These trace fossils are probably the burrows of larval insects made in moist muddy sand, rather than burrows made by earthworms in soils. So don’t be calling them “worm burrows,” or else a baby kitten will get mildly scolded. (Photograph by Anthony Martin.)

Several people spontaneously spoke this ichnological banality as soon as they saw small burrows preserved in the rock, many of which were directly associated with the exquisitely preserved dinosaur tracks. This happened often enough (which is to say, twice) that I just had to call attention to this geological faux pas. “Stop saying ‘worm burrows’!” I said with mock outrage. I quickly followed my joking admonishment with a brief explanation of how most of the burrows were much more likely to be from insects, rather than worms. Traits of the burrows – such as scratchmarks and short, branching, angled tunnels – implied insect tracemakers, such as the larvae of beetles or flies.

Insect traces associated with dinosaur tracks should not be all that surprising to anyone. After all, insects originated in terrestrial environments about 400 million years ago, meaning they were more than halfway through their evolutionary history by the time these Jurassic trace fossils were made. I had seen many similar burrows made by insects on the Georgia barrier islands and elsewhere in Georgia, which gave me enough confidence to propose their more probable identity.

Insect burrows – probably made by “mud-loving” beetles – along the shore of a freshwater pond on Sapelo Island, Georgia. Notice the burrows are relatively younger than (cross-cut) two tail dragmarks made by resident alligators (Alligator mississippiensis). Sandal as scale, which is size 8 1/2 (men’s). (Photograph by Anthony Martin.)

Of course, once you draw attention to a word or phrase among friends that is guaranteed to provoke annoyance, you should expect them to bring it up more frequently later as fodder for their amusement. Indeed, this happened for the remainder of the field trip, and I did not disappoint my audience as I responded with histrionic cringing, flinching, and groaning each time we encountered more of these “worm burrows” in Triassic or Jurassic rocks and they were identified as such.

Look, worm burrows! Ha-ha! The beautiful invertebrate trace fossils, former burrows filled with white sand that contrasts from the surrounding hematite-stained sand, are also in the Moenave Formation (Lower Jurassic) of southern Utah. (Photograph by Anthony Martin.)

All frivolity aside, the point I was trying to make to my field-trip tormenters was this: whenever we look at sedimentary rocks formed in continental environments, and we happen to notice invertebrate trace fossils in those same rocks, we should think before speaking. In other words, we do better as paleontologists, geologists, or naturalists in general when we reexamine our neat, preconceived labels before applying them loudly and confidently to observed phenomena, and particularly with invertebrate trace fossils.

For example, even the word “burrow” can be too glib for interpreting certain invertebrate trace fossils. Many invertebrates do not move underneath a sedimentary surface but along it; traces of such movements are either trackways, which are made with legs and leave impressions of these, or trails, which are made by whole-body movement without legs, such as those formed by worms or snails.

In my experience, trackways and trails are often lumped in with burrows, despite possessing impressions made by legs, furrows, and levees. For example, some of the trace fossils we saw on the field trip were certainly trails, yet I heard these called “burrows” by a few people. Granted, this sort of confusion is actually more understandable than the “worm-burrow” mistake, because trails can segue into shallow horizontal burrows and vice-versa, or some “trails” actually can have tiny leg impressions, meaning they actually are trackways. Thus the distinction between these end members can become blurred quite easily if you don’t pay attention to the details of a given invertebrate trace.

Modern land snail (pulmonate gastropod) making a trail on surface of a coastal dune, Cumberland Island, Georgia; scale in centimeters. (Photo by Anthony Martin.)

Fossil trail, possibly made by a snail, on a former sand dune in the Navajo Formation (Lower Jurassic) of southern Utah. Research funding for scale. (Photograph by Anthony Martin.)

Insect burrow, probably made by a beetle larva, in which it changes from a shallow burrow to a trackway on the surface of a coastal dune, Little St. Simons Island, Georgia. Scale in millimeters. (Photograph by Anthony Martin.)

In the sands and muds of the Georgia barrier islands, insect burrows in particular have often caused me to keep quiet about what I think made them, versus what really made them. Many times I have seen a little lump at the end of a horizontal burrow, scooped up the tracemaker hiding underneath, and been surprised by what was there. Most of these tracemakers have turned out to be small adult beetles or beetle larvae of various species, but I can’t ever predict which life stage or species will be there based just on their traces. (At least, not yet.)

Shallow burrow with short branches in a coastal dune, Cumberland Island, Georgia. Gee, I wonder what worm made it?

Surprise! It was a tiny adult beetle, found at the end of the burrow. Didn’t see that coming, did you? Well, maybe you did after all of the pedantic foreshadowing. (Both photographs by Anthony Martin.)

As a result of these insect-inspired search images, embedded in my consciousness from years of looking at Georgia-coast insect traces, I cannot ever again look at trace fossils made in formerly terrestrial environments and simply say, “worm burrows,” at least with a clear scientific conscience or a straight face. Hence whenever I see similar burrows in sedimentary rocks that were formed in lakes, streams, or soils from the Devonian Period to the recent, my default hypothesis is “insect burrows,” rather than “worm burrows.” Is this always right? No, as some terrestrial trace fossils, such as Edaphichnium and Castrichnus, were almost certainly made by earthworms, and nematode worms may have formed others, like Cochlichnus. (Although Cochlichnus has also been linked with insect tracemakers – but that’s a another story for another day.) Nonetheless, saying “insect burrows” is more likely to be correct than the alternatives, and in science, it’s good practice to learn from your mistakes.

So geologists and paleontologists everywhere, I beseech you not to limit yourselves descriptively when you encounter the millions of lovely and varied invertebrate trace fossils in sedimentary rocks formed in terrestrial environments. The truth will set you free (or at least put you on parole), and these seemingly simple trace fossils will become more intriguing as you realize their full complexity and potential mystery. Call them something other than “worm burrows,” then see what happens.

Invertebrate trace fossils (burrows) in sandstone from the Moeanave Formation (Lower Jurassic) in St. George, Utah. Do they look a little different to you now that you’re ready to give them a different name than mere “worm burrows”? (Photograph by Anthony Martin.)

(Acknowledgements: Many thanks to Andrew Milner, Jim Kirkland, Tyler Birthisel, Martin Lockley, Brent Breithaupt, Neffra Matthews, and many others for their organizing a most excellent three-day field trip to the Triassic-Jurassic rocks of southern Utah and northern Arizona. We all learned heaps from this direct experience, and greatly appreciate the huge amount of time and effort put into preparing for the field trip.)

Further Reading

Milner, A.R.C., Harris, J.D., Lockley, M.G., Kirkland, J.I., and Matthews, N.A. 2009. Bird-like anatomy, posture, and behavior revealed by an Early Jurassic theropod dinosaur resting trace. PLoS One, 4(3): doi:10.1371/journal.pone.0004591.

Rindsberg, A.K., and Kopaska-Merkel, D. 2005. Treptichnus and Arenicolites from the Steven C. Minkin Paleozoic footprint site (Langsettian, Alabama, USA). In Buta, R. J., Rindsberg, A. K., and Kopaska-Merkel, D. C., eds., = Pennsylvanian Footprints in the Black Warrior Basin of Alabama, Alabama Paleontological Society Monograph No. 1: 121-141.

Smith, J.J., Hasiotis, S.T., Kraus, M.J., and Woody, D.T. 2008. Relationship of floodplain ichnocoenoses to paleopedology, paleohydrology, and paleoclimate in the Willwood Formation, Wyoming, during the Paleocene–Eocene thermal maximum. Palaios, 23: 683-699.

Verde, M., Ubilla, M., Jiménez, J.J., and Genise, J.F. 2006. A new earthworm trace fossil from paleosols: aestivation chambers from the Late Pleistocene Sopas Formation of Uruguay. Palaeogeography, Palaeoclimatology, Palaeoecology, 243: 339-347.