Traces of the Red Queen

The seagull looked peaceful on that beach, lying on its left side with its eyes closed. Yet it was a permanent quietude, as only its head was there.

This disembodied head stuck out as a white spot with a red edge, perched on top of a pile of dull-brown, dead cordgrass. The torso so recently connected to this head was nowhere to be seen, and I could find no tracks belonging to the gull or any other animal nearby. It looked as if it had been placed there as an object of art, ready for erudite admirers – wine glasses in hand – to comment on its broader themes and nuanced metaphors. To a ichnologist, though, it also spoke of a sudden death, and one likely dealt by a aerial predator.

Seagull-Head-Decapitated-WassawThe place where I saw this gruesome sign was on Wassaw Island, Georgia. Wassaw is the only island on the Georgia coast that was never logged or otherwise developed by European or Americans, hence it retains a more primitive feel compared to most other Georgia islands. You can only get there by boat, and in this instance our boat captain and guide – John Crawford – had taken our field-trip group there to learn about its unique natural history. Because of its intact environments and general lack of human influence on the landscape, I was not surprised to see something new on Wassaw. However, I haven’t seen anything like this since.

Within minutes of arriving on the island, this beheaded seagull presented a little mystery for us. As mentioned before, tracks and the rest of the body were not visible, nor were any droplets of blood around its head, either. Moreover, its dry feathers and the freshness of its fatal wound – a clean severing of its neck vertebrate – also meant it had not washed up on shore. Where did it die, and how did it get there?

After ruling out the land and sea, we looked above the beach, and realized that the attack must have been delivered up there, in the air. We then imagined what could have possessed the bulk, ferocity, and other means to chop through a seagull’s neck while in flight. The list of suspects was a short one, and we quickly narrowed it down to one: a bald eagle.

Our hypothesis was not so far-fetched, as bald eagles don’t just eat fish, but also kill and eat other birds, including gulls. This meant the seagull head we saw that morning was very likely a result of bird-on-bird predation. Extending this a bit further into the evolutionary pasts of these birds, it reflected a time when when their non-avian dinosaur ancestors killed and were killed by similar behaviors, but on the ground.

How did birds evolve flight from non-flighted theropod ancestors? No doubt one of many selection pressures exerted on non-avian dinosaurs was predation. Any means for increasing the likelihood of escape from predators also bestowed a greater probability for passing on genes coding for that “escaping trait” to the next generation of not-quite-flighted dinosaurs.

Of course, flight has evolved for many uses in birds. Nevertheless, making a quick getaway from mortal peril is still one of them. Yet flight has also been used as a means for enhancing predation in the birds that kill other birds, exerting new and different selection pressures on prey. This example of an evolutionary back-and-forth “arms race” between predators and prey is often nicknamed the Red Queen hypothesis, named after Lewis Carroll’s character in Alice in Wonderland. Only now I will change her line (said to a fleeing Alice) about running in place:

Now, here, you see, it takes all the running you can do to keep in the same place.

to a more avian-appropriate one:

Now, here, you see, it takes all the flying you can do to keep in the same place.

Still, In this Georgia-coast example, a more appropriate literary allusion would have been to the Queen of Hearts from Alice in Wonderland, a decapitating character famous for uttering the line, “Off with their heads!” In this sense, the Red Queen and Queen of Hearts meet in the arms race between predators and prey.

Will this “Red Queen of Hearts” scenario happen again during eagle and seagull conflicts? Yes: that is, unless the seagulls’ descendants adapt, which may be followed by the eagles’ descendants adapting to these changes. And on it goes, this evolution of the now blending with the then, a reminder that these days of the dead affect those of the living, as well as those not yet alive.

Tracking Wild Turkeys on the Georgia Coast

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Further Reading

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

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

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

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

Of Sandhill-Crane Footprints and Dinosaurs Down Under

Last week, while in Athens, Georgia, I found myself musing about footprints from the barrier islands of Georgia and the Cretaceous rocks of Australia, despite their separation by half a world and more than 100 million years. These seemingly random thoughts came to me during a visit to the Department of Geology at the University of Georgia to give a lecture in their departmental seminar series.

It was a pleasure speaking at the geology department for many reasons, but perhaps the most gratifying was how it was also a homecoming. I had worked on my Ph.D. there in the late 1980’s, and in 1988-1989 had taught introductory-geology classes in the very same lecture hall where I gave my presentation. Several of my former professors, who were junior faculty then, are still there and now comprise a distinguished senior faculty. So seeing them there now, their smiling faces in the audience along with the latest generation of undergraduate and graduate students, generated all sorts of warm-and-fuzzy feelings.

But enough about the present: let’s go back about 100 million years to the Cretaceous Period, which was the subject of my talk. I had actually asked to speak about the modern Georgia barrier islands and their traces: you know, the main theme of this blog and my upcoming book of the same title (Life Traces of the Georgia Coast, just in case you need reminding). Nonetheless, my host and valued friend, paleontologist Dr. Sally Walker, figured that a summary of my latest research on the Cretaceous trace fossils of Victoria, Australia would bring in a wider audience, especially if I used the magical word “dinosaur” in the title (which I did).

For my talk at the UGA Department of Geology, I could have talked about this place – St. Catherines Island, Georgia – and it’s modern traces. After all, it’s only about a four-hour drive and short boat ride from Athens, Georgia.

But instead I talked about this place – coastal Victoria, Australia – and its trace fossils from more than 100 million years ago. Which wasn’t such a bad thing.

In retrospect, she was right, and I thoroughly enjoyed putting together an informative and (I thought) entertaining presentation that shared highlights of fossil discoveries from that part of Australia during the past five years. For the benefit of the students in the audience, basic geology was woven throughout the talk, as I included facets of sedimentology, stratigraphy, geochemistry, paleobotany, paleoclimatology, plate tectonics, evolution, history of science, field methods, and oh yes, dinosaurs. (If you are interested in hearing more about the science and personal experiences behind these recent findings in Australia, these are related in another blog of mine written previous to this one, The Great Cretaceous Walk.)

So how do the barrier islands of the Georgia coast and their animal traces relate to the Cretaceous of Australia? I mentioned the main reason briefly in my talk, but will elaborate more here: I likely owed one of my most important fossil discoveries in Australia to track-imprinted memories gained from field work on the Georgia coast. The fossil find, which happened in June 2010, was of about two dozen thin-toed theropod dinosaur tracks in Cretaceous rocks along the Victoria coast. These tracks represent the best assemblage of dinosaur tracks found thus far in southern Australia, and the largest collection of polar-dinosaur tracks in the Southern Hemisphere. Moreover, some of these tracks just happened to be about the same size and forms of footprints made by sandhill cranes (Grus canadensis).

Comparison between the footprint of a sandhill crane (Grus canadensis), made in moist sand next to a freshwater pond, St. Catherines Island, Georgia (top), and a footprint made by a theropod dinosaur about 105 million years ago on a river floodplain, Victoria, Australia (bottom). Notice the resemblance?

Sandhill cranes do not normally live on the Georgia barrier islands, and nearly all of them simply fly over or stop briefly during their annual migrations from south of Georgia to the Great Plains, or vice versa. However, at least a few have settled on St. Catherines Island, the same place on the Georgia coast where I recently studied gopher tortoise burrows. According to Jen Hilburn, the island ornithologist, some of these cranes found life so comfortable on the island that they stayed. This turned out to be fortunate for me, as I became familiar with their tracks after repeated visits to St. Catherines. Even though these tall, beautiful, and majestic birds restrict themselves to just one island year-round, St. Catherines is big enough to hold a wide variety of habitats and substrates, so I have seen their tracks in salt marshes, next to fresh-water ponds, and along dusty roads throughout the entire length of the island.

Who are you calling a “dinosaur”? A sandhill crane on St. Catherines Island graciously poses for its portrait, helping this ichnologist get a better idea of what an anatomically similar tracemaker might have looked like more than 100 million years ago.

Sandhill-crane trackway on the sandy substrate of a high salt marsh, St. Catherines Island, Georgia. In this environment, its tracks are accompanied by fiddler-crab burrows and feeding pellets, as well as the tracks and dig marks of raccoons hunting the fiddler crabs. Scale (toward the top of the photo) in centimeters.

So to make a long story short, while walking along the Victoria coast last year, I also carried with me mental picture of these tracks in Georgia. These images, I am sure, contributed to my stopping to look at a rock surface that held faint but nearly identical impressions made by dinosaurian feet on the once-soft sediments of a river floodplain. This is how ichnology is supposed to work, and it did.

A comparison between sandhill-crane tracks on the Georgia barrier islands and those of Cretaceous dinosaurs in Australia is actually not as far-fetched as one might think at first. For one, we now know that birds are dinosaurs, evolutionarily speaking. This formerly vague hypothesis is now a certainty, and is based on an ever-improving fossil record of feathered theropod dinosaurs, as well as studies from modern biology that show genetic and developmental affinities between modern birds and theropods. Even so, this idea is not new, either. For example, evolutionary biologist Thomas Huxley (1825-1895), friend and noted proponent of Charles Darwin, readily connected Archaeopteryx, the Late Jurassic bird (or dinosaur, depending on evolutionary perspective) with theropod dinosaurs.

Preceding Huxley, though, was one of the first scientists to formally apply ichnology to fossilized dinosaur tracks, Edward Hitchcock (1793-1864). Hitchcock interpreted the abundant dinosaur tracks of the Connecticut River Valley – many made by theropods – as those of large, flightless birds that lived before humans. Although he never made the evolutionary connection between dinosaurs and birds, his hypothesis reflected anatomical similarities between their feet.

A close-up look at sandhill crane feet while it takes a step. Notice the left foot has a little toe facing backwards, but off the ground. This is the equivalent of our “big toe,” also known as digit I, and it rarely registers in their tracks unless a crane walks in soft mud or sand. Instead, you will see impressions of the other three toes with clawmarks, and the middle toe normally makes the deepest mark.

Theropod dinosaurs, like many modern birds, mostly made three-toed tracks, a condition also called tridactyl. Although theropod tracks are occasionally confused with similar tracks made by ornithopod dinosaurs, they have the following traits: (1) three prominent, forward-facing digit impressions; (2) a footprint that is longer than wide; (3) angles of less than 90° between the outermost digits; and (4) well-defined clawmarks. One of the many changes that happened to bird feet as they evolved from non-avian theropods was the dropping of and rearward projection of their first digit (equivalent to our big toe). This condition was a great adaptation for grasping branches in trees and otherwise getting around off the ground. Bird tracks from the Cretaceous Period also tend to be wider than long, a function of the angles between the outermost toes becoming greater than 90°, and most of these also show the impression of a backward-pointing toe. Sandhill-crane footprint made in firm sand of a high salt marsh, St. Catherines Island, Georgia. Like many bird tracks, this one is wider than it is long, which is unlike most theropod dinosaur tracks. Still, these are very similar to tracks made by certain types of thin-toed theropod dinosaurs during the Cretaceous Period. Scale in centimeters.

Much later in their evolutionary history, though, some lineages of birds became either flightless or otherwise spent more time on the ground than in the trees, such as wading birds and shorebirds. These circumstances resulted in their first digit becoming reduced or absent, or vestigial. Violá, the tridactyl theropod-dinosaur footprint came back in style, so to speak, and now dinosaur ichnologists regularly study the tracks and behaviors of birds with such feet to better understand how their theropod relatives may have moved during the Mesozoic Era.

Comparison of a track made by a greater rhea (Rhea americana, right), which is a large flightless bird native to Argentina, to that of an equivalent-sized theropod dinosaur track (right). Both tracks have three forward-facing digits ending with sharp clawmarks and are longer than wide. Scale = 15 cm (6 in). The dinosaur track is a replica of an Early Jurassic theropod (from about 200 million years ago) from the western U.S. Photograph of the rhea track is by Anthony Martin, and of the dinosaur-track replica is by Ty Butler of Tylight™. Scale in the photo to the left = 15 cm (6 in).

Thus while writing the research paper on the dinosaur tracks, I kept in mind the comparison between sandhill-crane footprints in Georgia and the Australian dinosaur tracks. I also recalled how paleontologists had previously measured theropod skeletons – feet and rear limbs, specifically – and proposed a relationship between foot length and probable hip height.

Based on these studies, you can take a theropod track, multiply it by 4.0, and you get the approximate hip height of its trackmaker. When I applied this calculation to the Australian tracks, their hip heights ranged from about 25 to 60 centimeters (10-23 inches). The smallest of these dinosaurs I imagined as chicken-sized; perhaps these were juveniles of the larger ones. But what might be living today that would compare to the largest of the trackmakers? Immediately I thought of herons, but then it struck me that sandhill cranes provided a more apt analogy.

So I think you know where this is going. Adult sandhill-crane tracks are about 12 centimeters (4.7 inches) long, so if you apply the same formula for theropod-dinosaur tracks to them, their hip heights should be 48 centimeters (19 inches). Would this relationship also hold up on a modern dinosaur, such as a sandhill crane?

Just to satisfy my curiosity, I wrote to Jen Hilburn (St. Catherines Island) and asked her to do me a little favor: could she measure the hip height of a living, adult sandhill crane for me? Fortunately, Jen carried out my unusual request (she said it was not easy, so I definitely owe her), and she wrote back with an answer: 58 centimeters (22 inches). This wasn’t a perfect fit with regard to the footprint formula, but it certainly worked for the size of the Australian dinosaurs I had in mind as trackmakers. Based on my study of the Australian tracks, they were made by small ornithomimids, which likewise made thin-toed tridactyl tracks.

After thanking Jen, I delighted in explaining how her measurement of a Georgia-island-dwelling sandhill crane related to a dinosaur-track discovery on the other side of the world. Furthermore, in the Emory University press release that accompanied the publication of the dinosaur-track discovery in August 2011, the reporter (Carol Clark) used my analogy of the trackmakers as “…theropods ranging in size from a chicken to a large crane.”

Sandhill crane walking down a sand pile next to a fresh-water pond and maritime forest on St. Catherines Island, Georgia, and leaving lovely tracks for an ichnologist to study and keep in mind while tracking non-avian theropod dinosaurs.

Artist conception of Struthiomimus, a Late Cretaceous non-avian theropod dinosaur from western North America. Although not a perfect fit, the tracks of cranes and other similarly sized birds can be compared to those of ornithomimid dinosaurs to better discern the presence and behaviors of these dinosaurs. Artwork by Nobu Tamura and from Wikipedia Commons.

What other modern traces from the Georgia coast will contribute to our better understanding the fossil record? Time will tell, and I hope some day to again share those thoughts at my former home – the Department of Geology at the University of Georgia – with friends, students, and colleagues, new and old.

Further Reading

Elbroch, M., and Marks, E. 2001. Bird Tracks and Sign: A Guide to North American Species. Stackpole Books, Mechanicsburg, PA: 456 p.

Forsberg, M. 2005. On Ancient Wings: The Sandhill Cranes of North America. Michael Foreberg Photography: 168 p.

Henderson, D.M. 2003. Footprints, trackways, and hip heights of bipedal dinosaurs: testing hip height predictions with computer models. Ichnos, 10: 99–114.

Johnsgard, P.A. 2011. Sandhill and Whooping Cranes: Ancient Voices over America’s Wetlands. University of Nebraska Press, Lincoln, NB: 184 p.

Lockley, M.G. 1991. Tracking Dinosaurs: A New Look at an Ancient World. Cambridge University Press, Cambridge, UK: 264 p.

Martin, A.J., Anthony J., Rich, T.H., Hall, M., Vickers-Rich, P., and Gonzalo Vazquez-Prokopec. 2011. A polar dinosaur-track assemblage from the Eumeralla Formation (Albian), Victoria, Australia. Alcheringa: An Australiasian Journal of Palaeontology, article online August 9, 2011. DOI: 10.1080/03115518.2011.597564