The Ichnology of Jurassic Park

Posted on April 9, 2013 by TMartin

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.)

Aw, 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.

Partially 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.)

Dinosaur 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.

A 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.”

Uh 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.

Here’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.

Want 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.

Trace Evidence for New Book

Posted on January 7, 2013 by TMartin

This past Friday, I very happily received the first complimentary copy of my new book, Life Traces of the Georgia Coast from Indiana University Press. After years of field observations, photographing, writing, editing, drawing, teaching, and speaking about the plant and animal traces described in this book, it was immensely satisfying to hold a physical copy in my hands, feeling its heft and admiring its textures and smells in a way that e-books will never replace. So for any doubters out there (and I don’t blame you for that), here is a photograph of the book:

A photograph, purportedly documenting the publication of at least one copy of my new book Life Traces of the Georgia Coast. Photo scale (bottom) in centimeters.

Still, given that a photograph of the book only constitutes one line of evidence supporting its existence, I knew that more data were needed. So of course, I turned to ichnology for help. After all, a 692-page hard-cover book should also make an easily definable resting trace. Here is that trace, formed by the book in the same spot shown previously.

Ichnological evidence supporting the existence of my new book, Life Traces of the Georgia Coast. Using the “holy trinity” of ichnology – substrate, anatomy, and behavior – as guides for understanding it better: the substrate is a bedspread; the “anatomy” is the 6 X 9″ outline of the book, with depth of the trace reflecting its thickness (and mass); and the behavior was mine, consisting of placing the book on the bedspread and removing it. E-book versions of the book should make similarly shaped rectangular traces, although these will vary in dimensions according to the reading device hosting the book.

However, I also admit that hard-core skeptics may claim that such photos could have been faked, whether through the manipulative use of image-processing software, or slipping the cover jacket onto a copy of Danielle Steel’s latest oeuvre. As a result, the best and perhaps only way to test such a hypothesis is for you and everyone you know to buy the book (which you can do here, here, or here). Or, better yet, ask your your local bookstore to carry copies of it, which will also help to ensure the continuing existence of those bookstores for future book-purchasing and ichnological experiments, including books of other science-book authors.

Lastly, just to make this experiment statistically significant, I suggest a sample size of at least n = 10,000, which should account for inadvertent mishaps that may prevent deliveries of the book, such as lightning strikes, volcanic eruptions, or meteorite impacts. Only then will you be able to assess, with any degree of certainty, whether the book is real or not.

Thank you in advance for your “citizen science,” and I look forward to discussing these research results with you soon.

Suggested Further Reading

Martin, A.J. 2012. Life Traces of the Georgia Coast: Revealing the Unseen Lives of Plants and Animals. Indiana University Press, Bloomington, Indiana: 692 p.

A Sneak Peek at a Book Jacket (with Traces)

Posted on January 4, 2013 by TMartin

After returning from a two-week vacation in California with my wife Ruth, we noticed a cardboard tube awaiting us at home. Intriguingly, the mystery package, which was only about 60 cm (24 in) long and 8 cm (3 in) wide, had been sent by Indiana University Press, the publisher of my new book, Life Traces of the Georgia Coast. We were a little puzzled by it, considering that it couldn’t possibly contain complimentary copies of the book. (As of this writing, I still have not held a corporeal representation of the book, hence my continuing skepticism that it is really published.) What was in this mystery tube?

Front cover and spine of my new book, Life Traces of the Georgia Coast: Revealing the Unseen Lives of Plants and Animals (Indiana University Press). The book, newly released this month, is not yet in stores, but supposedly on its way to those places and to people who were kind enough to pre-order it. But if you didn’t pre-order it, that’s OK: you can get it right here, right now.

Upon opening it, we were delighted to find that it held ten life-sized prints of the book jacket: front cover, spine, back cover, and front-back inside flaps. The cover art, done by Georgia artist Alan Campbell, looked gorgeous, and had reduced well to the 16 X 25 cm (6 X 9″) format, retaining details of traces and tracemakers, but also conveying a nice aesthetic sense. I was also amused to see the spine had the title (of course) but also said “Martin” and “Indiana.” Although I’ve lived in Georgia for more than 27 years, I was born and raised in Indiana, so it somehow seemed fitting in a circle-of-life sort of way to see this put so simply on the book.

Back cover of Life Traces of the Georgia Coast, highlighting a few of the tracemakers mentioned in the book – sea oats, sandhill crane, sand fiddler crab, and sea star – while also providing a pretty sunset view of primary dunes, beach, and subtidal environments on Sapelo Island. (P.S. I love that it says “Science” and “Nature” at the top, too.)

I had no idea what the back cover might be like until seeing these prints, but I thought it was well designed, bearing a fair representative sample of tracemakers of the Georgia barrier islands: sea oats (Uniola paniculata), a sandhill crane (Grus canadensis), sand fiddler crab (Uca pugilator), and lined sea star (Luidia clathrata), as well as a scenic view of some coastal environments. I had taken all of these photos, so it was exciting to see these arranged in such a pleasing way. My only scientifically based objection is that I would have like to see it include photos of insects, worms, amphibians, reptiles, or mammals (these and much more are covered in the book), as well as a few more tracks, trails, or burrows. Granted, I suppose they only had so much room for that 6 X 9″ space, and thus I understood how they couldn’t use this space to better represent the biodiversity of Georgia-coast tracemakers and their traces. (Oh well: guess you’ll have to read the book to learn about all that.)

Inside front and back flap material for Life Traces of the Georgia Coast, which also includes a summary of the book (written by me) and a rare photo of me (taken by Ruth Schowalter) in my natural habitat, which in this instance was on St. Catherines Island, Georgia.

I had written the summary of the book on the inside flap nearly a year ago, so it was fun to look at it with fresh eyes, almost as if someone else had written it for me. Fortunately, I banished my inner critic while reading it, and just enjoyed the sense that it likely achieved its goal, which was to tell people about the book and provoke their interest in it.

In short, this cover jacket symbolizes a next-to-last step toward the book being real in my mind. Now, like any good scientist, all I need is some independently verifiable evidence in the form of tactile data, such as a physical book in my hands. Stay tuned for that update, which I’ll be sure to share once it happens. In the meantime, many thanks to all of the staff at Indiana University Press – who I’ll mention by name next time – for their essential role in making the book happen and promoting it in this new year.

Information about the Book, from Indiana University Press

Life Traces of the Georgia Coast: Revealing the Unseen Lives of Plants and Animals, Anthony J. Martin

Have you ever wondered what left behind those prints and tracks on the seashore, or what made those marks or dug those holes in the dunes? Life Traces of the Georgia Coast is an up-close look at these traces of life and the animals and plants that made them. It tells about the how the tracemakers lived and how they interacted with their environments. This is a book about ichnology (the study of such traces), a wonderful way to learn about the behavior of organisms, living and long extinct. Life Traces presents an overview of the traces left by modern animals and plants in this biologically rich region; shows how life traces relate to the environments, natural history, and behaviors of their tracemakers; and applies that knowledge toward a better understanding of the fossilized traces that ancient life left in the geologic record. Augmented by numerous illustrations of traces made by both ancient and modern organisms, the book shows how ancient trace fossils directly relate to modern traces and tracemakers, among them, insects, grasses, crabs, shorebirds, alligators, and sea turtles. The result is an aesthetically appealing and scientifically accurate book that will serve as both a source book for scientists and for anyone interested in the natural history of the Georgia coast.

Life of the Past – Science/Paleontology

692 pp., 34 color illus., 137 b&w illus.
cloth 978-0-253-00602-8 $60.00
ebook 978-0-253-00609-7 $51.99

More information at:

http://www.iupress.indiana.edu/catalog/806767 ]http://www.iupress.indiana.edu/catalog/806767

Most Intriguing Traces of the Georgia Coast, 2012

Posted on December 18, 2012 by TMartin

The end of another revolution of the earth around the sun brings with it many “best,” “most,” “worst,” “sexiest,” or other such lists associated with that 365-day cycle. Tragically, though, none of these lists have involved traces or trace fossils. So seeing that the end of 2012 also coincides with the release of my book (Life Traces of the Georgia Coast), I thought that now might be a good time to start the first of what I hope will be an annual series highlighting the most interesting traces I encountered on the Georgia barrier islands during the year.

In 2012, I visited three islands at three separate times: Cumberland Island in February, St. Catherines Island in March, and Jekyll Island in November. As usual, despite having done field work on these islands multiple times, each of these most recent visits in 2012 taught me something new and inspired posts that I shared through this blog.

For the Cumberland Island visit, it was seeing many coquina clams (Donax variabilis) in the beach sands there at low tide, and marveling at their remarkable ability to “listen” to and move with the waves. With St. Catherines Island, it was to start describing and mapping the alligator dens there, using these as models for similar large reptile burrows in the fossil record. Later in the year, I presented the preliminary results of this research at the Society of Vertebrate Paleontology meeting in Raleigh, North Carolina. For the Jekyll trip, which was primarily for a Thanksgiving-break vacation with my wife Ruth, two types of traces grabbed my attention, deer tracks on a beach and freshwater crayfish burrows in a forested wetland. So despite all of the field work I had done previously on the Georgia coast, these three trips in 2012 were still instrumental in teaching me just a little more I didn’t know about these islands, which deserve to be scrutinized with fresh eyes each time I step foot on them and leave my own marks.

For this review, I picked out three photos of traces from each island that I thought were provocatively educational, imparting what I hope are new lessons to everyone, from casual observers of nature to experienced ichnologists.

Cumberland Island

Coyote tracks – Coyotes (Canis latrans) used to be rare tracemakers on the Georgia barrier islands, but apparently have made it onto nearly all of the islands in just the past ten years or so. On Cumberland, despite its high numbers of visitors, people almost never see these wild canines. This means we have to rely on their tracks, scat, and other sign to discern their presence, where they’re going, and what they’re doing. I saw these coyote tracks while walking with my students along a trail between the coastal dunes, and they made for good in-the-field lessons on “What was this animal?” and “What was it doing?” Because Cumberland is designated as a National Seashore and thus is under the jurisdiction of the U.S. National Park Service, I’m interested in watching how they’ll handle the apparent self-introduction of this “new” predator to island ecosystems, which may begin competing with the bobcats (Lynx rufus) there for the same food resources.

Ghost Shrimp Burrows, Pellets and Buried Whelk – Sometimes the traces on the beaches at low tide are subtle in what they tell us, and the traces in this photo qualify as ones that could be easily overlooked. The three little holes in the photo are the tops of ghost shrimp burrows. Scattered about on the beach surface are fecal pellets made by the same animals; ghost shrimp are responsible for most of the mud deposition on the sandy beaches of Georgia. The triangular “trap door” in the middle of the photo is from a knobbed whelk (Busycon carica), which has buried itself directly under the sand surface. The ghost shrimp are perhaps as deep as 1-2 meters (3.3-6.6 ft) below the surface, and are feeding on organics in their subterranean homes. These homes are complex, branching burrow systems, reinforced by pelleted walls. Hence these animals and their traces provide a study in contrasts of adaptations, tiering, and fossilization potential. The whelk trace is ephemeral, and could be wiped out with the next high tide, especially if the waiting whelk emerges and its shallow burrow collapses behind it. On the other hand, only the top parts of the ghost shrimp burrows are susceptible to erosion, meaning their lower parts are much more likely to win in the fossilization sweepstakes.

Feral Horse Grazing and Trampling Traces – Probably the most controversial subject related to any so-called “wild” Georgia barrier islands is the feral horses of Cumberland Island, and what to do about their impacts on island ecosystems there. A year ago, I wrote a post about these tracemakers as invasive species, and discussed this same topic with students before we visited in February. But nothing said “impact” better to these students than this view of a salt marsh, overgrazed and trampled along its edges by horses. This is a example of how the cumulative effects of traces made by a single invasive species can dramatically alter an ecosystem, rendering it a less complete version of its original self.

St. Catherines Island

Suspended Bird Nest – I don’t know what species of bird made this exquisitely woven and suspended little nest, but I imagine it is was a wren, and one related to the long-billed marsh wren (Telmatodytes palustris), which also makes suspended nests in the salt marshes. This nest was next to one of several artificial ponds with islands constructed on St. Catherines with the intent of helping larger birds, such as egrets, herons, and wood storks, so that they can use the islands as rookeries. These ponds are also inhabited by alligators, which had left plenty of tracks, tail dragmarks, and other sign along the banks. With virtually no chance of being preserved in the fossil record, this nest was a humbling reminder of what we still don’t know from ichnology, such as when this specialized type of nest building evolved, or whether this behavior happened first in arboreal non-avian dinosaurs or birds.

Ant Nest in Storm-Washover Deposit – As you can see, the aperture of this ant nest, as well as the small pile of sand outside of it, did not exactly scream out for attention and demand that its picture be taken. But its location was significant, in that it was on a freshly made storm-washover deposit next to the beach. This “starter nest” gives a glimpse of how ants and other terrestrial insects can quickly colonize sediments dumped by marine processes, such as storm waves. These sometimes-thick storm deposits can cause locally elevated areas above what used to be muddy salt marshes. This means insects and other animals that normally would never burrow into or traverse these marshes move into the neighborhood and set up shop, blissfully unaware that the sediments of a recently buried marginal-marine environment are below them. Ant nests also have the potential to reach deep down to those marine sediments, causing a disjunctive mixing of the traces of marine and terrestrial animals that would surely confuse most geologists looking at similar deposits in the geologic record.

Alligator Tracks in a Salt Marsh - These alligator tracks, which are of the left-side front and rear feet, along with a tail dragmark (right) surprised me for several reasons. One was their size: the rear foot (pes) was about 20 cm (8 in) long, one of the largest I’ve seen on any of the islands. (As my Australian friends might say, it was bloody huge, mate.) This trackway also was unusual because it was on a salt pan, a sandy part of a marsh that lacks vegetation because of its high concentration of salt in its sediments. (According to conventional wisdom, alligators prefer fresh-water environments, not salt marshes.) Yet another oddity was the preservation of scale impressions in the footprints, which I normally only see in firm mud. Finally, the trackway was crosscut by trails of grazing snails and burrows of sand-fiddler crabs (Uca pugilator). This helped me to age the tracks – probably less than 24 hours old, and not so fresh that I should have reason to get worried. (Although I did pay closer attention to my surroundings after finding them.) Overall, this also made for a neat assemblage of vertebrate and invertebrate traces, one I would be delighted to find in the fossil record from the Mesozoic Era.

Jekyll Island

Grackle Tracks and Obstacle Avoidance – These tracks from a boat-tailed grackle (Quiscalus major), spotted just after sunrise on a coastal dune of Jekyll Island, are beautifully expressed, but also tell a little story, and one we might not understand unless we put ourselves down on its level. Why did it jog slightly to the right and then meander to the left, before curving off to the right again? I suspect it was because the small strands of bitter panic grass (Panicum amarum), sticking up out of the dune sand, got in its way. Similar to how we might avoid small saplings while walking through an otherwise open area, this grackle chose the path of least resistance, which involved walking around these obstacles, rather than following a straight line. If we didn’t know about this from such modern examples, but we found a fossil bird trackway like this but didn’t look for nearby root traces, how else might we interpret it?

Acorn Worm Burrow, Funnels and Pile – When I came across the top of this acorn-worm burrow, which was probably from the golden acorn worm (Balanoglossus aurantiactus), and on a beach at the north end of Jekyll, I realized I was looking at a two-dimensional expression of a three-dimensional structure. Acorn worms make deep and wide U-shaped vertical burrows, in which they quite sensibly place their mouth at one end and their anus at the other. These burrows usually have a small funnel at the top of one arm of the “U,” which is the “mouth end.” The “anus end” is denoted by a pile of what looks like soft-serve ice cream, which it most assuredly is not, as this is its fecal casting, squirted out of the burrow. What was interesting about this burrow is the nearby presence of a second funnel. This signifies that the worm shifted its mouth end laterally by adding a new burrow shaft to the previous one, superimposing a little “Y” to that part of the U-shaped burrow.

Ghost Crab Dragging Its Claw – As ubiquitous and prolific tracemakers in coastal dunes of the Georgia barrier islands, and as many times as I have studied their traces, I can always depend on ghost crabs (Ocypode quadrata) to leave me signs telling of some nuanced variations in their behavior. In this instance, I saw the finely sculpted, parallel, wavy grooves toward the upper middle of its trackway, made while the crab walked sideways from left to right. A count of the leg impressions in the trackway yielded “eight,” which is the number of its walking legs. This means the fine grooves could only come from some appendage other than its walking legs: namely, one of its claws. Why was it dragging its claw? I like to think that it might have been doing something really cool, like acoustical signaling, but it also might have just been a little tired, having spent too much time outside of its burrow.

So now you know a little more about who left their marks on the Georgia barrier islands in 2012. What will 2013 bring? Let’s find out, with open eyes and minds.

Tracking Wild Turkeys on the Georgia Coast

Posted on November 20, 2012 by TMartin

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.

Different Coastlines, Same Traces, and Time

Posted on October 25, 2012 by TMartin

This past week, I visited North Carolina for varied reasons, but all related to paleontology and geology. First, I gave a well-attended evening lecture about polar dinosaurs, graciously invited and hosted by the Department of Geography and Geology at the University of North Carolina-Wilmington (UNCW). Later in the week, I presented a poster at the Society of Vertebrate Paleontology (SVP) meeting in Raleigh (covered last week here), while also taking in a couple of days of talks, posters, and enjoyably catching up with paleo-friends while meeting neo-friends. Regrettably, I had to leave the meeting early, but with good reason, which was for a field trip to look at fossils in a Pleistocene outcrop near Wilmington with faculty and students from UNCW. Overall, it was a fulfilling week, teeming with paleontological and social variety.

This pithy summary, though, omits lots of details (and if it didn’t, then it wouldn’t be pithy). But one item worth explaining a bit more here was a brief trip to Wrightsville Beach, which is a barrier island was just east of Wilmington. Dr. Doug Gamble, a geography professor in the UNCW Department of Geography and Geology, offered to take me there just before my talk, which I eagerly accepted. Considering all of the field work I had done on the Georgia barrier islands to the south of there, and that I would be teaching a course on barrier islands next semester, going to this beach was an opportunity to learn more about the similarities and differences between Georgia and North Carolina beaches.

Panorama of Wrightsville Beach on the coast of North Carolina, replete with human locomotion traces and dwelling structures. These features make it very different from most beaches in Georgia. But what about other traces? Don’t you just love rhetorical questions? Including this one? (Photograph by Anthony Martin.)

Many North Carolina beaches are famous (or infamous) as examples of what can go wrong with unrestrained development of barrier islands. Many such case studies have been explored through the research, writings, and activism of geologist Dr. Orrin Pilkey of Duke University, as well as other coastal geologists who have looked at the effects of human alterations of these habitats. Wrightsville Beach is such a barrier-island beach, having been heavily modified by human activities during the past 150 years or so. When comparing it in my mind to the Georgia barrier islands, it most resembled Tybee Island, which is also next to a relatively large city (Savannah), easily accessible by a bridge, and developed as a sort of “vacation destination” for people who like beaches, but also want them to have all of the amenities of the places they left behind. Otherwise, it held little resemblance to the mostly uninhabited and undeveloped beaches I prefer to peruse on the Georgia barrier islands.

After driving over the bridge to the island, we walked onto the beach in several places, and I began looking for traces. At first there was little to see, which was a direct result of there being too much to see. Because it was a pleasant day and we were visiting in the afternoon, much of the beach had been heavily trampled by humans, with more than a few of these people aided in their bioturbation by canine companions. Obvious restructuring of the beach included a jetty at the north end that combined a concrete wall and boulders, and pilings of concrete blocks at the south end. Dunes were modest, low-profile, and capped by sparse stands of sea oats (Uniola paniculata), and behind these were hotels, condominiums, and houses, all chock-a-block. It would be too strong to say this beach was alien to me, let alone post-apocalyptic, but it did seem like an altered reality compared to my experiences in Georgia.

A jetty at Wrightsville Beach (North Carolina) composed of concrete and rocks, intended to preserve sand on the beach, which it is doing here, but also results in an imbalanced distribution of sand along it. Note the abundant human and canine tracks on the right, shouting out any other animal traces that might have been in the sand. (Photograph by Anthony Martin.)

Another view of the jetty at Wrightsville Beach, sharply contrasting sand deposition and erosion on either side of it. (Photograph by Anthony Martin.)

A pile of broken concrete being used as rip-rap at the south end of Wrightsville Beach in an attempt to slow erosion there. Or something. (Photograph by Anthony Martin.)

Only with more walking toward the south end of the beach did we see less of an overwhelming human-dog ichnoassemblage and start noticing signs of the native fauna. With this, I became comforted by the familiar. These traces included some I had seen many times on Georgia beaches, including: the soda-straw-like burrows of parchment worms (Onuphis microcephala); the volcano-like sand mounds and chocolate-sprinkle-like feces of callianassid shrimp (either Biffarius biformis and Callichirus major); the soft-serve-ice-cream-like fecal mound of acorn worms (Balanoglossus aurantiactus); and the hole-in-the-ground-like burrows of ghost crabs (Ocypode quadrata). (OK, so I ran out of metaphors.) Seagull tracks abounded as well, lending more of a dinosaurian flavor to the trace assemblage.

Two burrows of parchment worms (Onuphis microcephala) on Wrightsville Beach, exposed by a little bit of erosion, with tiny fecal pellets at their bases. Scale in millimeters. (Photograph by Anthony Martin.)

Burrow aperture and fecal pellets of a ghost shrimp (either Biffarius biformis or Callichirus major) on Wrightsville Beach. Scale in millimeters again. (Photograph by Anthony Martin.)

Fecal casting of an acorn worm, and probably that of a golden acorn worm (Balanoglossus aurantiactus) on Wriightsville Beach. One end of its burrow is underneath this pile, and that would be its anal end, which is sensibly located in a different place from its oral end. And I think you know the scale by now. (Photograph by Anthony Martin.)

Ghost crab (Ocypode quadrata) burrow and tracks, out of the intertidal zone and more into the dunes on Wrightsville Beach. (Photograph by Anthony Martin.)

These traces thus showed us that this North Carolina beach, one majorly changed by humankind during the past 150 years, actually was more biodiverse than one might think at first glance. In my mind, then, it became just a bit more wild through these signs of life hinting at what laid beneath our feet.

At this point, I could depress everyone by listing what traces and biota were not there, but that’s not the point, so I won’t. In a more progressive sense, what traces we saw represented traces of hope, of life hanging on despite environmental change, living almost invisibly beneath our feet. So as human development continues on beaches like these, and sea level rises through the rest of this century, I felt assured of their being survivors of this change, and of their traces outlasting our humanity. The trace fossils of the future are now, and recording our effects on the life that makes these traces. How many will wink out with our species, and how many of their marks will outlast us?

An intergenerational stroll – a grandmother and grandson? - alongside the pier on Wrightsville Beach in North Carolina. Did she have memories of this beach in her childhood? How do these compare to what she sees there now? What memories will this child have of it in the future, especially as the sea continues to rise? If these memories are not recorded, what will be left behind? (Photograph by Anthony Martin.)

Further Reading

Pilkey, O., and Fraser, M.E. 2005. A Celebration of the World’s Barrier Islands. Columbia University Press, New York: 309 p.

Thieler, E.R., Pilkey, O., Cleary, W.J., and Schwab, W.C. 2001. Modern sedimentation on the shoreface and inner continental shelf at Wrightsville Beach, North Carolina, U.S.A. Journal of Sedimentary Research, 71: 958-970.

Tracking Bookzilla

Posted on August 31, 2012 by TMartin

The AJC-Decatur Book Festival, an annual event held on Labor Day weekend in my hometown of Decatur, Georgia, begins tonight with a poetry reading from Natasha Trethewey, the new U.S. Poet Laureate and a colleague of mine at Emory University. The festival is one of the largest independent book fairs in the U.S., featuring readings by hundreds of fiction and non-fiction authors, poets, illustrators, spoken-word artists, and other people interested in all things literary, as well as having lots of books for sale.

But along with the authors and other written-word enthusiasts will be an elusive figure, one you may or may not notice among the 80,000 or so people, but whose visage will preside over activities as an iconic talisman. Known as Bookzilla, he – or she, as its gender is uncertain – is apparently the result of a genetic experiment gone awry, mixing the traits of a near-sighted human, theropod dinosaur, and book (pre-Kindle version).

A rare photo of Bookzilla, making a rare appearance in downtown Decatur, Georgia this past June. Note the physical characteristics denoting a voraciously studious consumer of words, and one that doesn’t care whether a book is fiction, non-fiction, new, or used.

Fortunately, as an ichnologist, I don’t have to rely on sightings to know where Bookzilla might be or what he/she was doing at the festival. Bookzilla, much like other cryptozoological beings who are infrequently observed in nature, leaves many traces from his/her behavior. So when I spotted Bookzilla in downtown Decatur earlier this year, I made sure to observe his/her behavior very carefully and take several photos of him/her to aid in my ichnological predictions of Bookzilla traces. After all, every trace is a function of what I often call the “Holy Trinity” of ichnology: an tracemaker’s anatomy, its behavior, and the substrate preserving its traces.

Another view of Bookzilla, giving an overall view of its form and behavior, thus lending to a better appraisal of its tracemaking abilities. An especially important feature to keep in mind is his/her large, forward-leaning, rectangular head, which should cause deeper impressions in the front halves of his/her tracks. Also note the small child cowering nearby, yet being reassured by her father that Bookzilla, and by default books and the knowledge they contain, are to be embraced, not feared.

Close-up view of the foot morphology of Bookzilla. Note the unusual blend of a plantigrade mode, with metatarsals in contact with the walking surface, combined with a trapezoidal (but symmetrical) outline to his/her foot, as well as three sharp unguals (claws) elevated well above the surface by excessive padding in the foot. Such feet, particularly when moved by a vertically oriented biped such as Bookzilla, will result in distinctive trackways that could never be confused with, say, those of a person wearing a clumsy costume.

Based on this admittedly limited sample of Bookzilla’s anatomy and behavior, I can nonetheless reasonably hypothesize what sorts of tracks and trackway patterns Bookzilla would form during normal activities. In a firm substrate, such as moist sand, Bookzilla’s tracks would only register only a vague trapzoidal outline of his/her feet, and claws would not register. However, in a softer substrate, such as a saturated mud or wet cement, Bookzilla would impart shallow impressions of three claws directly in front of the trapezoidal outlines.

Hypothesized Bookzilla tracks, which could be from either the right or left feet owing to bilateral symmetry of the foot structure. (Left) Track outline formed in relatively firm substrate, such as a moist sand, with no claws showing and with the deepest part of the track in the front half because of its weighty, tome-like head. (Right) Track outline formed in soft substrate with finer grain size and higher water content, such as mud. In this instance, claw marks will register, and the track outline will be accompanied by numerous pressure-release structures from the tracemaker deforming the sediment with application and release of foot pressure caused by forward movement. Footprint length (minus claws) = 30-35 cm (12-14 in).

A typical trackway pattern for Bookzilla should show mostly diagonal walking (right-left-right diagonal to one another), but will be punctuated by many side-by-side tracks, representing long periods of stopping and standing while reading books.

Hypothesized Bookzilla trackway pattern, indicating alternate (diagonal) walking, typical of a biped, but also interrupted by frequent stops to look at books and read. Note claw impressions are only visible in parts of the trackway, depending on local variations in substrate conditions; in this instance, stepping into wet cement.

Considering how Bookzilla is well known as an insatiable reader and is constantly hunting for books, no sitting traces, such as those interpreted for some theropod dinosaurs, are known, nor expected. I also cannot yet speculate about about additional traces made by Bookzilla, such as dwelling structures, toothmarks, or scat. Remarkably, his/her consumption of books does not actually result in digestive products, but instead seems to cause increased levels of cognitive pleasure.

So if you are in the Decatur-Atlanta area this weekend and attending the book festival, maybe you’ll be lucky enough to actually see Bookzilla, but you’re much more likely to spot his tracks. In the meantime, take a look at some books and enjoy all of the literary traces that are offered there.

(The AJC-Decatur Book Festival will take place in downtown Decatur, Georgia from September 1-2. The full schedule is here, and it will feature appearances and talks by well-established and emerging authors and illustrators, the latter including my wife Ruth Schowalter. If you can’t make this year’s festival, it is held on Labor Day weekend every year, so plan to make next year’s. And many thanks to Daren Wang for suggesting this several months ago as a blog post!)

Mistaken Point and the Limits of Actualism

Posted on August 23, 2012 by TMartin

Sometimes we paleontologists, especially those who also study modern organisms and their behaviors, get a little too sure of ourselves, thinking we have a clear vision of life during the pre-human past. So it’s good to have that confidence shaken a little, made uneasy by a glimpse at a much deeper past, one that preceded the bulk of fossils that shape our accepted norms and basic expectations in paleontology.

Welcome to the Ediacaran Period, the span of earth history from 635-542 million years ago, and a time when actualism – the precept that the present is the key to the past – becomes a naïve, idealistic dream, a glib summary of a world that has only existed for a mere 12% of earth history.

What are these? They’re fossils, but otherwise I’m not sure what else to tell you: guess I’ve been spending too much time in the present. But for for those people who have studied them and know better than me, they’re called Charniodiscus, and they’re frond-like fossils with holdfasts (those circular parts connected to their stems) that kept them attached to the seafloor about 565 million years ago. All you have to do to see these fossils is go to Newfoundland, Mistaken Point Ecological Reserve in Newfoundland, Canada, get permission from the Reserve to visit them, have a guide accompany you, and walk 40-45 minutes to the site from a car park. Incidentally, there will be absolutely no cafes or toilets on the way there. You know, just like how it was in the Precambrian. (Photograph by Anthony Martin; scale in centimeters.)

These discomforting realizations started a little less than two weeks ago, inspired by a field trip to the Ediacaran-Cambrian rocks of eastern Newfoundland, Canada. Why was I in cool, temperate Newfoundland, instead of sweating it out on the summertime Georgia coast? The occasion was a pre-meeting trip associated with the International Congress on Ichnology, simply known among ichnologists as Ichnia. This was the third such meeting, a once-every-four-years event (coinciding with years of the summer Olympics). The previous two were in Krakow, Poland (2008) and Trelew, Argentina (2004), and thus far these meetings also include fabulous field trips.

For Ichnia 2012, upon seeing an announcement of a field trip to Mistaken Point and other localities associated with the Precambrian-Cambrian boundary, I eagerly signed up for it. You see, Mistaken Point is world famous for its extraordinary preservation of more than 1,000 body fossils of those weird and wonderful fossils known as the Ediacaran fauna, Ediacaran biota, Vendian fauna, or Vendobionts (take your pick). This was the main reason why my fellow ichnologists on the field trip – 16 of us from 9 countries – were along for the ride, despite the trip’s clear emphasis on body fossils.

A rare photo of ichnologists getting really excited about seeing body fossils, which is totally understandable when we’re talking about the Ediacaran fossils at Mistaken Point, Newfoundland. Eventually, though, they later became unruly and started demanding, “Show me your trace fossils!” Fortunately for the sake of international ichnological relations, the field-trip leaders happily obliged that same day. (Photograph by Ruth Schowalter.)

These rare fossils, which are strange enough to even cause paleontologists to question whether or not they are animals (hence the cautious use of the more inclusive term “biota” instead of “fauna”), are abundantly exposed on broad bedding planes in Mistaken Point Ecological Reserve on the southeastern coast of Newfoundland. Discovered in 1967, these fossils have since proved to be one of the best examples of easily visible body fossils from more than 542 million years ago, and the Newfoundland fossils comprise the only such assemblage that originally lived in deep-marine environments. They evidently died in place when suffocated by a layer of volcanic ash that settled onto the seafloor, hence the fossils reflect a probable sample of their original ecosystem. This ash layer neatly preserved the fossils, and its minerals provided a means to calculate absolute age dates for the assemblage, which is from 565 +/- 3 mya (million years ago).

Bedding-plane exposure at Mistaken Point with many frond-like fossils, broadly referred to as rangeomorphs. (Photograph by Anthony Martin, Canadian-themed scale is in centimeters.)

A close-up of one of the more exquisitely preserved rangeomorphs, which I think is Fractofusus misrai. But you really shouldn’t trust this ichnologist with that identification, so it’d be wise to double-check that with a real expert. (Photograph by Anthony Martin.)

Just a few years ago, though, Mistaken Point became paleontologically famous again, and this time for its trace fossils. Researchers from Memorial University in Newfoundland and Oxford University looked at bedding planes near those holding the the body fossils, and were surprised to find a few trails there. At that time, it was the oldest evidence of animal movement from the fossil record, and although these finds have been disputed and others have tried to stake this claim for trace fossils elsewhere, it is still holding up fairly well.

A surface trail, probably made by a < 1 cm wide animal moving along the seafloor about 565 mya. The animal moved from left to right, which is indicated by the crescentic ridges inside the trail, which open in the direction of movement. (Photograph by Anthony Martin, taken at Mistaken Point, Newfoundland.)

Another surface trail, but this one without the internal structure of the other one, and with levees on either side of the central furrow. (Photograph by Anthony Martin, taken at Mistaken Point, Newfoundland.)What’s this? Don’t have a clue. It looks like a series of overlapping trails, some looping, but would have taken me several hours to unravel. Anyway, it generated some good discussion at the outcrop, and they’re probably trace fossils, which made us ichnologists both happy and perplexed. (Photograph by Anthony Martin, taken at Mistaken Point, Newfoundland; scale in centimeters.)

What made these trace fossils? It’s hard to say, and that’s a humbling statement for me to make. In public talks I’ve given about my upcoming book, and in a presentation I gave the following week at Ichnia on the Memorial University campus, I’ve assured how the actualism of the Georgia barrier islands and its traces can reliably serve as models for interpreting many trace fossils formed in different environments, and trace fossils of various geologic ages from around the world. But in this instance, I didn’t have a inkling of what made the Mistaken Point trace fossils. These trace fossils were also made in deep-marine environments, which are lacking from the Georgia coast, and I haven’t learned much about deep-marine trace fossils from elsewhere.

In short, my ignorance was showing, and these trace fossils were completely out of my realm of experience. The only feeble hypothesis I could conjure on the basis of what I’ve seen in modern sediments of the Georgia barrier islands are small marine gastropod trails. Sorry, that’s all I got.

Oooo, look, it’s snail! Making a trail! Isn’t that neat? And if you squint really hard and have a couple of beers, you might agree that it almost resembles one of the fossil trails from Mistaken Point. Don’t see it yet? Here, have another beer. (Photograph by Anthony Martin, taken at Sapelo Island, Georgia; scale in millimeters. )

But if ignorance loves company, I can feel good in knowing that others have grasped at the same straw of actualism and found it far too short. I could tell a few of my ichnological colleagues were likewise a little challenged by what they saw at Mistaken Point, and I knew that for some of them – like me – they normally deal with trace fossils in much younger rocks. But hey, that’s what geology field trips are supposed to do: challenge us with what’s really there in the rock record, right there in front of us, rather than what we wish were there.

Fortunately, a little more information provided during the meeting after the field trip helped my understanding of the trace fossils we saw at Mistaken Point, and actually connected to modern tracemakers. Alexander Liu, the primary author of the paper that first reported the trace fossils, gave a talk that reviewed the evidence for Precambrian trace fossils, including those from Mistaken Point. In experiments he and his coauthors did with living anemones in a laboratory setting, they were able to reproduce trails similar to the Mistaken Point trace fossil with the internal structure. Thus these researchers were able to use actualism to assist in their interpretation, which also meant that neoichnology was not so useless after all when applied to the Ediacaran. That made me feel a little better.

Let’s take a look at that first surface trail again, but this time with the help of my trustworthy colleague Paleontologist Barbie, who was along for the field trip. The crecentic ridges in the interior of the trail may represent marks where the basal disc of a anemone-like animal pushed against the surface as it moved. Even more interesting, the arrow points to an oval impression, which may be a resting trace that shows the approximate basal diameter of the tracemaker. What was the tracemaker? It’s currently identified as a small anemone, which is based on modern traces. Neoichnology rules! (Photograph by Anthony Martin.)

Ediacaran trace fossils still engender debate, though, and especially with people who don’t necessarily accept that animals made trails during the Ediacaran. For instance, about four years ago, some scuba-diving researchers observed a giant protozoan making a trail on a sediment surface in the Bahamas. Accordingly, they proposed that one-celled organisms – not animals – could have made similar trails during the Ediacaran Period. Interestingly, this shows how actualism can produce conflicting results when applied to Ediacaran fossils. After all, it’s still a big world out there, and we humans haven’t really observed everything in it yet.

So I’ll make one last point about Ediacaran fossils here, then will move on to more recent times. If you think that at the very least we paleontologists should be able to tell the difference between trace fossils and body fossils in Ediacaran rocks, you’re also in for some confusion. In the only research article I have ever attempted on Ediacaran fossils, which were much closer to Georgia – coming from the Carolina Slate Belt of North Carolina – my coauthors and I struggled with exactly that question with some fossils found in that area. In the end, we said they were body fossils, not trace fossils. And as everyone knows, I love trace fossils, and I really wanted these to be trace fossils. But they were not. That’s science for you: denying your deepest desires in the face of reality.

So surely the Cambrian would be easier to interpret, right? I meanl, after 542 mya, animals started burrowing merrily, to and fro, hither and tither, with uninhibited and orgiastic abandon, and, well, you get the idea. But, not really. Another part of the field trip involved looking at what happened with the departure of the relatively unbioturbated alien world of the Ediacaran, pre-542 mya, to the more familiar sediment mixing of the Cambrian and Ordovician Periods, post-542 mya. Yet even these rocks and their trace fossils were still not quite like what we see today.

This will be the subject of my next post, which will again explore the theme of how we should approach strict actualism like any scientifically based idea: with a mixture of astonished wonder, but also with a hard-edged look at what is really there.

As we bid adieu to Mistaken Point and began our walk back to the car park, we could swear we saw lifeforms emerging from the mist-covered rocks, resurrected from the deep time and deep water of the Avalonian Precambrian. Then we realized those were just some of our group behind us. Oh well. Maybe next time. (Photograph by Anthony Martin.)

(Acknowledgements: Much appreciation is extended to the field trip leaders – Liam Herringshaw, Jack Matthews, and Duncan McIlroy – for their organization and execution of a fantastic three-day field trip; to Valerie and Richard of the Mistaken Point Ecological Reserve for guiding us to the site; to my ichnological colleagues for their cheery and knowledge-broadening company; and my wife Ruth for being with me and providing an artist’s perspective about her experiences with us crazy ichnologists, shared here and here.)

Further Reading

Fedonkin, M., Vickers-Rich, P. Grey, K., and Narbonne, G. 2007.The Rise of Animals: Evolution and Diversification of the Animalia. Johns Hopkins Press, Washington: 320 p.

Liu, A.G., McIlroy, D., and Brasier, M.D. 2010. First evidence for locomotion in the Ediacaran biota from the 565Ma Mistaken Point Formation, Newfoundland. Geology, 38: 123-126.

Matz, M.V., Frank. T.M., Marshall, N.J., Widder, E.A., and Johnsen, S. 2008. Giant deep-sea protest produces bilaterian-like traces. Current Biology, 18: 1-6

Tacker, R.C., Martin, A.J., Weaver, P.G., and Lawver, D.R. 2010. Trace vs. body fossil: Oldhamia recta revisited. Precambrian Research, 178: 43-50.

Vickers-Rich, P., and Komarower, P. (editors). 2007 The Rise and Fall of the Ediacaran Biota. Geological Society of London, Special Publication 286: 448 p.

Traces of Toad Toiletry and Naming Trace Fossils

Posted on July 12, 2012 by TMartin

Sometimes I envy those people on the Georgia barrier islands who, through sheer number of hours in the field, come upon animal traces that I’ve never seen there. But this was one of those instances where the find was so extraordinary that I will suppress my jealous urges, celebrate the person who found it, marvel at it, and share its specialness with others.

Gale Bishop, a fellow ichnologist who is currently on St. Catherines Island, found an intriguing sequence of traces during a morning foray on its dunes and beaches there last week. In his second life – his first was as a geology professor at Georgia Southern University – he has transformed into an indefatigable sea-turtle-nesting monitor on St. Catherines and coordinator of a teacher-training program. Part of his daily routine there, among many other duties, includes looking for mother-turtle traces – trackways and nests – during the nesting season, which in Georgia is from May through September.

Along the way, with his eyes well trained for spotting jots and tittles in the sand, Gale often notices oddities that likely would be missed by most people, including me. The following photograph, which he shared on the St. Catherines Island Sea Turtle Program page on Facebook, is from a find he made about 7:15 a.m. on Saturday, July 7. Take a look, and please, if you haven’t already, forget the title of this post as you ponder its clues.

A mystery in the dune sands of St. Catherines Island on the Georgia coast, begging to be interpreted. No, not the shovel: those are never mysterious. Look at the traces to the left and above the shovel. What made these, what was it doing, and who else was in the neighborhood afterwards? Oh, and again, stop staring at the shovel. (Photograph by Gale Bishop.)

Gale called me out specifically when he posted this and several other related photos on Facebook, and asked me to tell a story about it. I gave him my abbreviated take in the comments, kind of like an abstract for the research article:

Looks like southern toad (Bufo terrestris) to me. What’s cool is the changes of behavior: hopping, stopping, pooping, and alternate walking (which people don’t expect toads to do – but they do).

That was my knee-jerk analysis, which took a grand total of about a minute to discern and respond. (After all, this was Facebook, a forum in which prolonged and deep thinking is strongly discouraged.) But I also kept in mind that quick, intuitive interpretations later need introspection and self-skepticism, especially when I’m making them. (See my previous post for an example of how wrong I could be about some Georgia-coast traces.) So rather than fulfill some Malcolm Gladwell-inspired cliché through my intuition, I sat down to study the photo with this series of questions in mind:

Why did I say “Southern toad” as the tracemaker for the sequence of traces that start from the lower left and extend across the photo?
What indicates the behaviors listed and in that order: hopping, stopping, pooping, and alternate walking?
What signified the changes in behavior, and where did these decisions happen?
Why did I assume that most people don’t expect toads to walk (implying that they think they just hop)?

The first leap in logic – how did I know a Southern toad (Bufo (Anaxyrus) terrestris) was the tracemaker – was the easiest to make, as I’ve often seen their tracks in sandy patches of maritime forests and coastal dunes. These hardy amphibians leave a distinctive bounding pattern, with the front-foot impressions together and just preceding the rear-foot ones; the toes of their front feet also point inward. In the best-expressed tracks, you will see four toes on the front feet and five toes on the rear.

Close-up of bounding pattern (from lower left of previous photo), showing front-foot impressions just in front of and more central than the rear feet impressions. Direction of movement is from bottom to top of photo. (Photograph enhanced to bring out details, but originally taken by Gale Bishop.)

The only other possible animal that could make a trackway pattern confusable with a toad in this environment is a southeastern beach mouse (Peromyscus polionotus). Still, mice mostly gallop, in which their rear feet exceed their front feet as they move. Mouse feet are also very different from those of a toad, with toes on both feet all pointing forward (remember, toad toes point inward). So although dune mice live in the same environment as these tracks, these weren’t mouse tracks. The only alternative tracemakers would be spadefoot toads (Scaphiopus holbrookii) or a same-sized species of frog, such as the Southern leopard frog (Rana sphenocephala). But neither of these species is as common in coastal dunes as the Southern toad, so I’ll stick with my identification for now.

Mouse tracks – probably made by the Southeastern beach mouse (Peromyscus polionotus) – on costal dunes of Little St. Simons Island, Georgia. The two trackways on the left are moving away from you, whereas the one on the trackway on the right is heading toward you. All three show a gallop pattern, in which the larger rear feet exceeded the front feet. Scale = 10 cm (4 in). (Photograph by Anthony Martin)

The second conclusion – the types of behaviors and their order – came from first figuring out the direction of travel by the tracemaker, which from the lower left of the photo toward its middle. This shows straight-forward hopping up to the point where it stops.

From there, it gets really interesting. The wide groove extends to the left past the line of travel and had to be made by the posterior-ventral part of the toad’s body (colloquially speaking, its butt). This, along with the disturbed sand on either side of the groove, shows that the toad turned to its right (clockwise) and backed up with shuffling movement. That’s when it deposited its scat, which I’ve also seen in connection with toad tracks (and on St. Catherines, no less). This really helped me to nail down the identity of the tracemaker, almost being able to declare, “Hey, I know that turd!”

Southern toad bounding pattern that abruptly stops, followed by clockwise turning, backing up, and, well, making a deposit. (Photograph by Gale Bishop, taken on St. Catherines Island.)

How about the alternate walking? Turns out that toads don’t just hop, but also walk: right side, left side, right side, and so on. This pattern – also called diagonal walking by trackers – is in the remainder of the photo (with the direction of movement left to right). When toads do this, the details of their front and rear feet are better defined, and you can more clearly see the front foot registers in front of the rear and more toward the center line of the body.

This side-by-side movement is also what imparted a slight sinuosity to the central body dragmark, which was from the lower (ventral) part of its body, or what some people would call “belly.” In my experience, most people are very surprised to find out that toads can walk like this, which I can only attribute to sample bias. In other words, they’ve only seen frogs and toads hop away from them when startled by the approach of large, upright bipeds.

Close-up of alternate walking pattern and body dragmark made by Southern toad. Direction of movement is from upper left to lower right. (Photograph enhanced to bring out its details, but original taken by Gale Bishop on St. Catherines Island.)

But wait, what are those two dark-colored depressions in the center of the alternate-walking trackway? Well, it doesn’t take much imagination to figure those out, especially if you’ve already had a couple of cups of coffee. Yes, these are urination marks, and even more remarkable, there are two of them in the same trackway. So not only did this toad do #2, but also #1 twice.

Southern toad urination mark #1, not too long after doing #2. (Photograph by Gale Bishop.)

Urination mark #2 , which you might say was #2 of #1, but with both #1′s after #2, or, oh, never mind.

Notice that the second mark seems to have had less of a stream to it, which makes sense in a way that I hope doesn’t require any more explanation or demonstration.

So to answer to one of the questions above – what signified the changes in behavior – you have to look for the interruptions in the patterns, much like punctuation marks in a sentence. The commas, semi-colons, colons, dashes are all part of a story too, not just the words.

The summary interpretation of what happened. Let’s just say that this frog (or toad, whatever) didn’t come a courtin’.

Through the series of photos Gale shared in an album on Facebook, he also showed that he was following a protocol all good trackers do: he changed his perspective while observing the traces. Likewise, I teach my students to use this same technique when presented with tracks and other traces, that it’s a good idea to walk around them. While doing this, they see changes in contrast and realize how the direction and angle of light on the traces alters their perceptions of it. At some points, a track or other trace may even “disappear,” then “reappear” with maximum clarity with just a few more steps.

A different perspective of the same traces, viewed from another angle. Do you notice something new you didn’t see in the previous photo and its close-ups? (Photograph by Gale Bishop, taken on St. Catherines Island.)

Now, because I’m also a paleontologist, this interesting series of traces also prompts me to ask: what if you found this sequence of traces in the fossil record? Well, it’d be a fantastic find, worthy of a cover story in Nature. (That is, if the tracks somehow went across the body of a feathered dinosaur.) Right now, I can’t think of any trace fossils like this coming from vertebrates – let alone toads or frogs – so let’s go to invertebrate trace fossils for a few examples of similarly interconnected behaviors preserved in stone.

In 2001, a sequence of trace fossils was reported from Pennsylvanian Period rocks (>300 million years old), in which a clam stopped, fed, and burrowed along a definite path, with all of its behaviors clearly represented and connected. The ichnologists who studied this series of trace fossils – Tony Ekdale and Richard Bromley – reckoned these behaviors all happened in less than 24 hours; hence the title of their paper reflected this conclusion.

Ichnologists have a sometimes-annoying and always-confusing practice of naming distinctive trace fossils, giving them ichnogenus and ichnospecies names. (For a detailed discussion of this naming method, I talked about it in another blog from the dim, dark, distant past of 2011 here.) For instance, Ekdale and Bromley stated in their study that three names could be applied to the distinctive trace fossils made by a single clam, with each a different form made by a different behavior: Protovirgularia (burrowing), Lockeia (stopping), and Lophoctenium (feeding).

Along those lines, another ichnologist (Andy Rindsberg) and I also suggested that an assemblage of trace fossils in Early Silurian rocks (>400 million years old) of Alabama, with many different ichnogenera, were all made by the same species of trilobite. The take-home message of that study, as well as Ekdale and Bromley’s, is that a single species or individual animal can make a large number of traces. This also means that ichnodiversity (variety of traces) almost never equals biodiversity (variety of tracemakers).

So let’s go back to the toad traces, put on our paleontologist hats, and think about a “what if.” What if you found this series of traces disconnected from one another: the hopping trackway pattern, the diagonal walking pattern, the urination marks, the groove, and the turd, all found in disparate pieces of rock? Taken separately, such trace fossils likely would be assigned different names, such as “Bufoichnus parallelis,” “B. alternata,” “Groovyichnus,” “Tinklichnus,” and “Poopichnus.” (Please do not use these names beyond an informal, jovial, and understandably alcohol-fueled setting.)

Color, present-day version of the variety of traces made by a Southern toad (above), and a grayscale imagining of it fossilizing perfectly (below). Key for whimsically named ichnogenera in fossilized version: Bp = “Bufoichnus parallelis,” Ba = “Buofichnus alternata,” G = “Groovyichnus,” P = “Poopichnus,” and T = “Tinklichnus.” Please don’t cite this.

Granted, the environment in which Gale noted these traces – coastal dune sands – are not all that good for preserving what is pictured here, but other environments might be conducive to fossilization. To quote comedian Judy Tenuta, “It could happen!”

So if someone does find a fossil analogue to Gale’s evocative find on St. Catherines Island, I will understand their giving a name to each separate part, even if I won’t like it. The most important matter, though, is not what you call it, but what it is. And in this case, the intriguing story of toiletry habits left in the sand one July morning by a Southern toad is worth much more than any number of names.

Further Reading

Ekdale, A.A., and Bromley, R.G. 2001. A day and a night in the life of a cleft-foot clam: Protovirgularia-Lockeia-Lophoctenium. Lethaia, 34: 119–124.

Halfpenny, J.C., and Bruchac, J. 2002. Scats and Tracks of the Southeast. Globe Pequot Press, Guilford, Connecticut: 149 p.

Jensen, J.B. 2008. Southern toad. In Jensen, J.B., Camp, C.D., Gibbons, W., and Elliott, M.J. (editors), Amphibians and Reptiles of Georgia. University of Georgia Press, Athens, Georgia: 44-46.

Rindsberg, A.K., and Martin, A.J. 2003. Arthrophycus and the problem of compound trace fossils. Palaeogeography, Palaeoclimatology, Palaeoecology, 192: 187-219.

Marine Moles and Mistakes in Science

Posted on June 25, 2012 by TMartin

A first day of field work in the natural sciences can be expected to hold surprises, no matter what type of science is being attempted. Sometimes these are unpleasant ones, such as finding out the fuel gauge in your field vehicle – which you are driving for the first time, and in a remote place – doesn’t work. Other times, you make a fantastic discovery, like a new species of spider, a previously undocumented invasive plant, or a fossil footprint. But sometimes you see something that just makes you scratch your head and say, “What the heck is that?”, or more profane variations on that sentiment.

What is this long, meandering ridge making its way through a beach to the high tide mark on Sapelo Island, Georgia, and what made it? If you’re curious, please read on. But if you already know what it is, then you know a lot more than I did the first time I saw something like this. (Photograph by Anthony Martin.)

The last of those three scenarios happened to me on Sapelo Island, Georgia, in June 2004. My wife Ruth was with me, and we had just arrived on the island the previous afternoon, having stayed overnight at the University of Georgia (Athens) Marine Institute, or UGAMI. We decided that our first full morning in the field would be at Nannygoat Beach on the south end of Sapelo, which is a 5-minute drive or a 20-minute walk from the UGAMI.

We drove a field vehicle there (the gas gauge and everything else worked), parked, and took the boardwalk over the coastal dunes. Our elevated view from the boardwalk afforded a good look at many insect, ghost crab, bird, and mammal tracks made in the early morning. Circular holes punctured the dunes, made by ghost crabs (Ocypode quadrata). Sand aprons composed of still-moist sand were next to these burrow entrances, bearing crisply defined ghost-crab tracks, although early-morning sea breezes had already started to blur these.

At some point after walking onto the beach, though, we saw traces that we had not noticed in previous visits to Sapelo, and ones I have rarely seen there or on other Georgia barrier islands since. These oddities were meters-long, slightly sinuous to meandering ridges, about 15-20 cm (6-8 in) wide, extending in the sandy areas from the dunes through the berm and down to the high-tide mark, where they ended abruptly.

Same meandering ridge shown in the first photo, but viewed from the high-tide mark, showing how it connects with the primary dunes. Note how a few holes are punched in the part near me: more about those soon. (Photograph by Anthony Martin, taken on Sapelo Island, Georgia. P.S.: My wife Ruth is the scale in both photos, fulfilling one of the top 10 signs that I might be a geologist.)

Although a few ridges crossed one another, they rarely branched, and if they did, the branches were quite short, only about 10-15 cm (4-6 in). When we followed them to the dunes, they seemed to originate from some unseen place below the sandy surfaces. We investigated further by cutting through some of the ridges to see what they looked like inside. They turned out to be mostly open tunnels with circular cross sections about 5 cm (2 in) wide, slightly wider than a U.S. dollar coin. They were mostly hollow, and only occasionally did we encounter a plug of sand interrupting tunnel interiors. This supposition was backed up by ridges that had collapsed into underlying voids. A few of the ridges stopped with a rounded end the same diameter as the ridge, or as a larger, raised, elliptically shaped “hill.”

Ridge with quite a bit of meander in it. Check out the short branch toward the top right, where the tracemaker must have changed its mind and backed up, then continued digging toward the viewer. Scale = 15 cm (6 in). (Photograph by Anthony Martin, taken on Sapelo Island, Georgia.)

Two separate ridges intersecting, caused by one crossing the other, resulting in “false branching.” Also notice the partial collapse of sand into underlying hollow tunnels and how one of the ridges ends in a rounded mound. Scale = 15 cm (6 in). (Photograph by Anthony Martin, taken on Sapelo Island, Georgia.)

A short ridge ending in a raised, elliptical “hill,” connected to a partially collapsed tunnel that is not otherwise evident as an elevated surface. Same scale as before. (Photograph by Anthony Martin, taken on Sapelo Island, Georgia.)

Ruth and I agreed that these tunnels were burrows, instead of some random features made by the winds, tides, or waves. But by what? Clearly their makers were impressive burrowers, capable of digging through meters of sand. Their bodies also were probably just a little narrower than the burrow interiors, which helped us to think about body sizes. Then we considered where we were – dunes and beach – and what animals were the most likely ones to burrow in these environments.

A process of elimination – determining what they were not – was a good way to start figuring out their potential makers. For example, no way these burrows were from insects, such as beetle larvae, ant lion larvae, or mole crickets, because they were just too big. Insects also have a tough time handling salinity, so once they got to the surf zone with its saturated, saline sand, they would have had problems, or (more likely) an aversive reaction and turned around immediately instead of plowing ahead.

Insect burrow in coastal dune sand, made by a small beetle. Look at both the form and scale, and you’ll see this is not a match for what we were seeing. Scale in centimeters. (Photograph by Anthony Martin, taken on Cumberland Island, Georgia.)

Small mammals, like beach mice (Peromyscus polionotus), didn’t seem like good candidates either. Beach-mouse burrows are totally different from what we were seeing, and their burrows do not run all of the way down to the intertidal zone. Mice, like insects, also don’t like marine-flavored water; even if they might be able to temporarily tolerate it, they wouldn’t continue to burrow through moist, salty sand.

A beach-mouse burrow, with their tracks coming and going. Either the mice dug this burrow, or they occupied an abandoned ghost-crab burrow. Regardless, this also doesn’t match our mystery traces. Scale in millimeters. (Photograph by Anthony Martin, taken on Little St. Simons Island, Georgia.)

This led to an initial hypothesis that these burrows were from one of the most common larger burrowing animals in the area, and one comfortable in dune, berm, and beach environments with saturated, salty sand. These could only be from ghost crabs, I thought, an explanation supported by undoubted ghost crab burrows that perfectly intersected these tunnels, accompanied by undoubted ghost-crab tracks.

Ghost-crab burrows intersecting tunnels, accompanied by lots of ghost-crab tracks. Wow, that’s really convincing circumstantial evidence, wouldn’t you say? (Photograph by Anthony Martin, taken on Sapelo Island, Georgia.)

End of story, right? Well, no. I and a lot of other scientists have said this before, but it bears repeating: part of how science works is that in its practice we do not prove, we disprove. I somehow knew the “ghost crab burrowing horizontally through meters of sand from the dunes to the beach” hypothesis was a shaky one, and it bothered me that it just didn’t seem right. So I started reading as much as possible about ghost-crab burrowing behaviors. I thought I already knew a lot about this subject, but nonetheless was willing to acknowledge that there might be some holes in my learning (get it – holes?) that needed filling (get it – filling? Oh, never mind).

The gentle reader probably surmised what happened next. That’s right: not a single peer-reviewed reference mentioned ghost crabs digging meters-long shallow tunnels from the dunes to the beach. So either I was wrong, or I had documented a previously unknown and spectacular tracemaking behavior in this very well-studied species. A single cut by Occam’s Razor simply said, “You’re wrong.”

You thought I made long horizontal burrows that go all of the way from the dunes to the surf zone? Wow, you primates are dumber than I thought. (Photograph by Anthony Martin, taken on Sapelo Island, Georgia.)

If not a ghost crab then, what else could make meters-long horizontal burrows of the diameter we had seen? This is when I began to reconsider my original rejection of moles as possible tracemakers.

So what am I: chopped liver? (Photograph from Kenneth Catania, Vanderbilt University, and taken from Wikipedia.org here.)

Here’s what was the most interesting about this mistaken interpretation: it was made because of where we were. In other words, our initial mystification about these traces stemmed from their environmental context. Had we seen these burrows winding down a sandy road in the middle of a maritime forest on Sapelo Island, we would not have hesitated to say the word “mole.” Yet because we saw exactly the same types of burrows in coastal dunes and beaches, we said, “something else.”

A long, meandering mole burrow in the sandy road going through a maritime forest on the north end of Sapelo Island. So if you see a burrow like this in the forest, you instantly say “mole.” But if you see it on the beach, you say, “Um, uh, duh…must be something else!” My tracks (size 8 1/2, mens) and 15 cm (6 in) photo scale for, well, scale. (Photograph by Anthony Martin.)

Another long, meandering ridge ended in a rounded “hill,” a trace that no one would hesitate to call a mole burrow, especially because it’s in the middle of a maritime forest. (Photo by Anthony Martin, taken on Sapelo Island, Georgia.)

A trip back to the literature further confirmed the mole hypothesis while also serving up a big slice of humble pie. I was embarrassed to find that these same burrows were described and interpreted as mole burrows in an article published in 1986. Even more mortifying: my dissertation advisor (Robert “Bob” Frey) was the first author on the article; it had been published while I was doing my dissertation work with him; and I had read the article years ago, but didn’t remember the part about mole traces. It was like these burrows were saying to me, “Go back to school, young man.”

OK, so these are mole burrows. Case closed. Now that we’ve identified them, we can stop thinking about them, and go on to name something else. But that ain’t science either, is it? This one answer – mole burrows – actually inspires a lot of other questions about them, which could lead to heaps more science:

• Which moles made these burrows? The Georgia barrier islands have two documented species of moles, the eastern mole (Scalopus aquaticus) and star-nosed mole (Condylura cristata). Of these two, eastern moles are relatively common on island interiors, whereas star-nosed moles are either rare or locally extinct from some of the islands. But star-nosed moles are also more comfortable next to water bodies and seek underwater prey. So could these traces actually signal the presence of star-nosed moles in dune and beach environments? Frey and his co-author, George Pemberton, originally interpreted these as eastern mole burrows, but they also didn’t eliminate the possibility of star-nosed moles as the tracemakers, either.

• What is the evolutionary history of moles on the Georgia barrier islands? Are these moles descended from populations isolated from mainland ones 10,000 years ago by the post-Pleistocene sea-level rise, or do they represent more modern stock that somehow made its way to the islands? A genetic study would probably resolve this issue, but who the heck is going to compare the genetic relatedness of moles from the Georgia barrier islands to those on the mainland?

• What were they eating? Moles don’t just burrow for the exercise, but for the food. While burrowing, they are also voraciously chowing down on any invertebrate they encounter in the subsurface. But what would they eat in beach sands? As many shorebirds know, Georgia beaches are full of yummy amphipods, which would likely more than substitute for a mole’s typical earthworm- and insect-filled diet in terrestrial environments. Yet as far as I can find in the scientific literature, no one has documented mole stomach contents or scat from coastal environments to test whether these small crustaceans are their main prey or not.

• What happened to these moles once their burrows got to the surf zone? Did they turn around and burrow back, or did they go for a swim in the open ocean? The latter is actually not so far fetched, as moles are excellent swimmers, especially star-nosed moles. But how often would they do this?

• Just how common (or rare) are these burrows in beaches? Just because I just perceive these burrows as rare could be an example of sample bias. Yes, I wrote an entire book about Georgia-coast traces and tracemakers and have done field work on the islands since 1998. But I don’t live on the Georgia barrier islands, nor have I spent more than a week continuously on any of them. Keenly observant naturalists who live on the islands or otherwise spend much time there could better answer this question than me. I suspect they’re actually much more common than I originally supposed, and now look for them to photograph or otherwise document whenever I go back to any of the islands.

• Would such burrows preserve in the geologic record? Probably so, especially if they were made in dunes and filled with a differently colored or textured sand. But I’ll bet that nearly every paleontologist or geologist would make the same mistake I did, and reach for a burrowing marginal-marine crab or some other invertebrate as the tracemaker.

Geologists would be further fooled if fossil mole tunnels were intersected by genuine ghost-crab burrows, which would constitute a great example of a composite trace made by more than one species of animal. But why did the crabs burrow into the mole tunnels? Because it was easier. After all, the moles left hollow spaces and loosened sand over wide areas, practically begging ghost crabs to exploit these disturbed areas.

Anyway, I doubt many geologists would think of a small terrestrial mammal as a tracemaker for such burrows in sedimentary rocks formed in marginal-marine environments, although I’d love to be proved wrong on this. I’m hoping my writing about it here will help to prevent such confusion, and that whoever benefits from it will buy me an adult beverage as thanks.

In summary, this example of making a crab burrow out of a mole tunnel thus serves as a cautionary tale of how where we are when making observations in the field can influence our perceptions. But it also goes to show us how our wonderment with what we observe in natural environments can be renewed and encouraged by daring to be wrong once in a while, and learning from those mistakes.

Further Reading

Frey, R.W., and Pemberton, S.G. 1986. Vertebrate lebensspuren in intertidal and supratidal environments, Holocene barrier island, Georgia. Senckenbergiana Maritima, 18: 97-121.

Gorman, M.L., and Stone, R.D. 1990. The Natural History of Moles. University of Chicago Press, Chicago, Illinois: 138 p.

Harvey, M.J. 1976. Home range, movement, and diel activity of the eastern mole, Scalopus aquaticus. American Midland Naturalist, 95: 436-445.

Henderson, R.F. 1994. Moles. Prevention and Control of Wildlife Damage, Paper 49, University of Nebraska, Lincoln: D51-58. (Entire text here.)

Hickman, G.C. 1983. Influence of the semiaquatic habit in determining burrow structure of the star-nosed mole (Condylura cristata). Canadian Journal of Zoology, 61: 1688-1692.

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Unseen lives of the Georgia barrier islands

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