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Traces of Toad Toiletry and Naming Trace Fossils

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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

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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.

Darwin, Worm Grunters, and Menacing Moles

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In my most recent previous post, I teased readers with the promise of revealing how Charles Darwin used a piano as a scientific tool for studying the behavior of earthworms. Regardless of whether or not you already looked up the answer through The Google, by reading Darwin’s last book (The Formation of Vegetable Mould through the Action of Worms with Observations on Their Habits), or other means, I will now gladly make connections between the seemingly disparate subjects of Darwin’s musically inclined experimentation, earthworm behavior, and fishermen of the southeastern U.S. catching earthworms as bait.

What makes this earthworm (Diplocardia) run away as fast as its little chetae, mucus, and peristalic movement can carry it through the soil? Let’s just say it’s not picking up good vibrations. Photograph by Bruce A. Snyder, from here, from www.discoverlife.org.

In writing about earthworms and their traces in my upcoming book, I devoted several pages to Mr. Darwin’s fascination with earthworms. In this exploration, I tell how Darwin was on to something when he tried applying sound – which included those made by playing musical instruments – to earthworms he had gathered from the English countryside. These musical performances were not an instance of Darwin trying to entertain these worms, boost their self esteem, or otherwise help them get in touch with their emotions. Rather, he was simply testing whether worms reacted to sound. What happened? Well, instead of me describing his results, I’ll let Darwin’s words inform you directly:

Worms do not possess any sense of hearing. They took not the least notice of the shrill notes from a metal whistle, which was repeatedly sounded near them; nor did they of the deepest and loudest tones of a bassoon. They were indifferent to shouts, if care was taken that the breath did not strike them. When placed on a table close to the keys of a piano, which was played as loudly as possible, they remained perfectly quiet.

– Charles Darwin, The Formation of Vegetable Mould through the Action of Worms with Observations on Their Habits (1881), p. 27.

Hence it was with deep appreciation last month when I gazed at the piano in the drawing room of Down House, the former Darwin family home, and thought about these experiments. Smiling, I imagined Darwin carefully watching a container of worms while he or someone else in his family forcefully banged on the keys of this piano. Of course, you also can’t help but wonder what was played “as loudly as possible.” Were these single, random notes, chords, or actual musical compositions? If the last of these, what pieces were played? Ideally, I like to think Mr. Darwin or one of his family members played a sea shanty learned during his days on The Beagle (or perhaps even songs learned from pirates), rather than just pounded random notes up or down a scale.

As conclusive as Darwin’s paragraph might seem about the lack of earthworm reactions to sound, he, like any good storyteller, then injected a dramatic twist when reporting his results. He followed up the preceding paragraph with one describing how earthworms, although deaf, are extremely sensitive to vibrations transmitted through solid media. Here he revealed exactly which notes were played (C on the bass clef, G in the treble clef, C in the treble clef) while two worms were in pots placed on top of the piano.

The vibrations transmitted through solid media – not air – caused the worms to withdraw from the soil surface, presumably hiding from the source of the vibrations. As an extension of this experiment, Darwin also used a fork to agitate the soil underneath other worms, which then provoked them to move up to the surface. Darwin correctly surmised that this stirring activity, like sound, also sent vibrations through the soil, which likewise produced aversive reactions in the earthworms.

These responses made sense in an evolutionary way, and show how Mr. Darwin was applying his principle of natural selection to the predator-prey relationships that had evolved between earthworms and moles. The behaviors he observed would have favored the survival of earthworms that associated vibrations with their most feared predators, and reacting accordingly, which is to say, fleeing in terror. And just what were their aversion-inducing predators? They were not robins or other species of birds – early, punctual, or otherwise timed – but the earthworm version of graboids: burrowing moles.

Eastern mole (Scalopus aquaticus) emerging from its burrow, seeking earthworms and other fresh food. Photograph by Kenneth Catania, from Fairfax County Schools.

Graboid emerging from its burrow, seeking humans and other prey. Note the eerie resemblance of its behavior to that of an eastern mole, albeit orders of magnitude larger and accompanied by a keen interest in large, surface-dwelling, bipedal prey. Photo from Wikipedia, but originally taken from the greatest ichnologically inspired horror film of all time, Tremors.

So you didn’t know about graboids, those burrowing predators of the underworld? Fortunately, this educational video provides all of the details you need to know. But if you’re interested in studying their neoichnology, be careful, and stay on the pavement.

As yet another example of ‘backyard science,” Darwin observed many traces of the European mole (Talpa europaea) in the fields just outside Down House, most of which were their mounds, or “molehills.” Indeed, last month as I admired one of Darwin’s original wormstones in the pasture behind Down House, I also noticed a good number of molehills on the grounds. Rather stupidly, I neglected to take a photo of one of these. (I mean, how cool would it have been to share images of the traces of moles that descended from those whose traces Darwin noticed?) Nonetheless, some of my photos of the grassy area near the wormstone show 20-30 cm wide bare patches in this otherwise meticulously maintained lawn. These spots, I suspect, are traces of the Down House groundskeepers, who probably level molehills as quickly as they appear, an ichnological version of “whack a mole.”

The pasture just behind Down House (Charles Darwin’s former home), with a “wormstone” in the lower right, and a few bare patches of ground just to the left. Could the latter mark recent sites of mole tunnels and molehills leveled by Down House groundskeepers, or are these just places where grass did not grow, and hence the products of an ichnologist’s overactive imagination? Anyway, I did see molehills out there, but don’t blame y’all for being a bunch of skeptical scientists and wanting more evidence than my just saying so.

OK, now how does all of this wonderfully elucidated Victorian-era science relate to the ecosystems and biota of the southeastern United States? Enter the “worm grunters.” Worm grunters are people who, independently of Darwin, figured out the same adaptive responses of earthworms to underground vibrations. Through their own experiments, worm grunters, who were interested in efficiently gathering many worms in a short time for putting on fishhooks (or making money selling earthworms to people who put them on hooks), rubbed steel slabs across the top of wooden posts stuck in the ground. Much later, researchers interested in finding out how this technique worked calculated frequencies of the seismic vibrations that caused earthworms to flee upward away from perceived predators.

The southeastern U.S., including the Georgia barrier islands, not only has its own species of earthworms (Diplocardia mississippiensis), but also has its own species of moles: the eastern mole (Scalopus aquaticus) and the less common star-nosed mole (Condylura cristata). Both types of moles no doubt strike fear in the multiple hearts of earthworms, and natural selection being how it is, the fastest burrowing moles (who are most likely to catch worms) also cause considerable vibrations from their digging. This accordingly means the earthworms that detect and escape these vibrations live long enough to reproduce and pass on whatever genes that aided in such perceptions.

In getting caught by this mole, this earthworm may have just won the worm equivalent of a Darwin Award, depending on whether it had reproduced or not. (Which it probably did, considering earthworm hermaphroditism means they are at least twice as likely to get lucky.) Photo from University of Illinois Extension; Home, Yard, and Garden Pests Newsletter, here.

Thus a visit to Down House in southern England and consideration of Darwin’s contributions to ichnology and behavioral ecology are not so far removed conceptually from the practical knowledge gained by some people in parts of the southeastern U.S. Moreover, many of these same people are of English, Irish, or Scottish descent, and effectively applied the same knowledge surmised by Darwin about worms and moles, which is kind of neat in a heritage sort of way.

Would all of these findings count as applied science, despite its historical lack of Ph.D.-bearing investigators, grant funding, publications, and press conferences announcing the results? Yup. After all, science is about its methods.

So next week, we’ll take a closer look at the traces moles make on the Georgia barrier islands. Do these moles just go after earthworms in the forests and meadows of those islands? Nope. After all, science is not just about its methods, but also surprises.

Further Reading

Darwin, C. 1881. The Formation of Vegetable Mould through the Action of Worms, with Observations on their Habits. John Murray, London, U.K.: 326 p.

Edwards, C.A., and Bohlen, P.J. 1996. Biology and Ecology of Earthworms (3rd Edition). Springer, Berlin: 426 p.

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

Hendrix, P.F. 1995. Earthworm Ecology and Biogeography in North America. CRC Press, Boca Raton, Florida: 244 p.

Mitra, O., Callaham, M.A., Jr., and Yack, J.E. 2009. Grunting for worms: seismic vibrations cause Diplocardia earthworms to emerge from the soil. Biology Letters, 2009: 16-19.

Of Darwin, Earthworms, and Backyard Science

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On the other hand, I sometimes think that general & popular Treatises are almost as important for the progress of science as original work.

– Charles Darwin, in a letter to Thomas Huxley, written in his home (Down House) on January 4, 1865

A combined blessing and burden that comes with travel, especially to new places, is the memory we carry of other places. The blessing part comes from the opportunity to connect previously disparate bodies of knowledge and experiences. This is always exciting for anyone who likes that sort of thing, while also satisfying purported promoters of “interdisciplinarity” (which was probably not a word until academia invented it, then pretended to reward those who practice it). On the other hand, the burden is that these thoughts of previous places can act as a veil, obscuring or overlaying our perception of novel sensations. In extreme cases, these remembrances can smother original ideas, especially if the places of our past are idealized and held as some worldly standard to which all other things must be compared.

What does this roundish stone, lying in the ground of the English countryside south of London, have to do with life traces of the Georgia coast? Good question, and if you’d like the start of an answer, please read on.

This Janus-like duality of travel occurred to me after my wife (Ruth) and I left Georgia for a few weeks of vacation in the United Kingdom, yet once there, I thought about my original home of Indiana and the barrier islands of Georgia. Ruth had never been to the U.K., and I hadn’t visited since attending an ichnology conference and field trip in Yorkshire, held in 1999. Fortunately, Ruth has a friend on the northeastern side of London who generously offered us a place to stay before we headed elsewhere. This refuge gave us a few days to learn what London had to offer us while we otherwise adjusted to cultural and temporal differences.

Among the myriad of educational opportunities in the London area is one that had been on my mind for quite a while, thanks to my writing about the Georgia coast. This was an intended visit to Down House, the former home of Charles Darwin and his family. Down House is located in a rural setting of the greater London area – Downe Village in the former parish of Kent – well southeast of Big Ben and all of the other typical touristy trappings of downtown London. Still, it can be visited via public transportation, which became doable for us Yanks once we figured out the needed connections in the intricate rail and bus system weaving throughout the London area.

From where we were staying, it took us nearly two hours to reach Down House. It was a mildly aggravating sojourn by train and bus, but made much better once we realized that driving there in London traffic with a hired car would have been far worse for both us and other people sharing the road (or sidewalk, as it may be). After our bus dropped us off in Downe Village, we saw a small sign pointing the way to Down House, and walked for  15 minutes on a quiet, country road before reaching our goal, a stroll only occasionally interrupted by brief terror induced when cars approached from the direction opposite of our expectations.

 When you step off the bus in Downe Village, this is one of the few clues that you’re near Darwin’s home, a place where scientific thought and human history changed in a big way.

A signpost in Downe Village provides a clue that Darwin has something to do with this area, although some horse named “Invicta” gets equal billing, and “St Mary the Virgin” gets bigger typeface. Still, it was nice to see Darwin’s visage there, too.

Blink and you’ll miss it: after walking about 10 minutes down the road, here’s the sign pointing the way to Down House. Personally, I thought it could use a neon fringe, or at least some DayGlo™ colors, but subdued is probably the way Darwin would have liked it.

We were also a little surprised at the subdued signage pointing us in the right direction to our goal, and I mused briefly about the homes of people who had far less impact on the advancement of human knowledge and world perspectives whose homes are accorded far more attention and adulation. (Yes, I’m looking at you, Graceland.)

The front of Down House, the home of Charles Darwin and his family from 1842 and after his death in 1882.

Down House is both modest and grand, not palatial at all, but impressive inside. Rooms on the second floor (or first floor, if you live in the U.K.) hold displays with a neatly presented synopsis of Darwin’s life and scientific findings, starting with his little boat journey in 1831-1836 through his grand synthesis of evolutionary principles. The ground floor of the house is more or less restored to the time when the Darwin family lived there, with particular attention paid to Mr. Darwin’s study, which was his main writing and experimentation room, or what modern-day scientists might call his “research space.” This is where On the Origin of Species and most other books of his were born. Infused with a purely fan-boy sort of joy, I was thrilled to be in the same place where many of his revolutionary ideas about evolution became expressed through words.

However, one item in the family living room (drawing room) intrigued me in a special way. It was a piano. This object was certainly used for the enjoyment of Darwin family members and guests, with the degree of delight of course depending on the proficiencies and musical choices of whoever played it. But then I was reminded – by the disembodied voice of Sir David Attenborough, no less – that this was not just a musical instrument, but also a scientific tool. (Disappointingly, Sir Attenborough volunteered this information in a recorded audio tour provided with admission to Down House, not through clairvoyance in a Sir Arthur Conan Doyle sense.) On this piano in the room and in the nearby Down House backyard are the places where Darwin conducted some of the earliest quantitative experiments in the behavioral ecology and neoichnology of terrestrial infauna. Or, in plain English, Darwin used this piano and a few other tools to measure and test the behavior of earthworms as tracemakers in soil.

The rear of Down House, with the two windows to the left looking into the drawing room, where the Darwin family piano is located. Unfortunately, photographs are not allowed in the interior of Down House, hence the external, voyeuristic perspective.

Darwin enthusiasts know well that the last book Darwin wrote was about a personal passion of his, the biology and behavior of earthworms. This book, titled The Formation of Vegetable Mould through the Action of Worms with Observations on Their Habits (1881), encapsulates many observations and conclusions he made from his long-term study of the oligochaete annelids that lived abundantly in the backyard and gardens of Downe House. As some biographers have noted, Darwin became quite a homebody after his years of voyaging on The Beagle, and he stayed close to Down House for much of his life after moving there in 1842. Nonetheless, this geographically restricted lifestyle did not mean he stopped inquiring about the natural world around him. On the contrary, he carried out intensive studies in and just outside of Down House, some of which dealt with earthworms, a subject that interested him for more than half of his life.

Darwin’s wonderment at worms was jump-started by something he noticed nearly thirty years after he innocuously tried to improve the soil in the pasture behind Down House. Told that he could get rid of mossy areas by laying down cinders and chalk, he obediently did so, and checked those same areas 29 years afterwards. It turned out the anomalous sediments had been buried about 18 cm (7 in) below the surface.

Darwin soon suspected this surface was newly made, formed by generations of earthworms bringing up soil over the preceding three decades. Through the technical support of his son Horace, an engineer, Darwin began to measure just how much earth an earthworm could worm. He already knew that earthworms burrowed through, consumed, and defecated sediment, which resulted in thoroughly mixed and chemically altered soils. So using his geologically inspired sense of time and rates of processes, he also rightly imagined that the daily activities of earthworms, multiplied by millions of worms and enough years, changed the very ground underneath his feet in a way so that it, well, evolved.

Ever the good scientist, though, Darwin tested this basic idea through experimentation. His assessment was accomplished through a precise measuring device invented by his son and flat, circular rocks, nicknamed wormstones, which were set out in the backyard of Down House. Based on my visual and tactile examination of the one wormstone that still lies outside of Down House, it looked like a quartz sandstone. However, out of respect for it and its ichnological and historical heritage, I did no other tests of its composition.

One of Darwin’s original “wormstones” (foreground center) placed in a pastoral setting behind Down House. Paleontologist Barbie (just behind the wormstone), who has accompanied me for much field work on the Georgia coast, helpfully provides scale.

Close-up view of wormstone, showing three metal slots set into a central ring and two rods, which provided the datum for measuring change in the wormstone’s depth over time. £10 note (with Darwin’s portrait on the right) for scale.

The experiment was elegantly simple. Using a device invented by Horace in 1870 (illustrated below, and photo here), the surface of the wormstone was measured relative to the height of the surrounding soil surface. This change in relative horizon was discerned by fitting the device on three metal slots that had been added to the edge of a central hole in the wormstone. Metal rods inserted through this same hole were connected to underlying bedrock, ensuring that these would stay stationary as worms churned the surrounding soil. Thus these rods acted as a horizontal datum through which any changes in the ground surface could be compared.

Illustration of Horace Darwin’s “wormstone measuring instrument,” with “K” pointing to where the instrument was placed to contact with the metal rods; the change with each measurement over time between this and “A” (a metal ring) would then show how much the stone had sunk downward. My source of this figure is from an online PDF by the Bromley Partnerships, Discover Darwin: An Education Resource for Key Stage Two, but its primary source is not cited there, and I could not otherwise find an attribution.

Darwin figured that the burrowing activity of earthworms underneath the stone, as well as sediment deposition at the surface as fecal castings, would result in the stone “sinking” over time, becoming buried from below. He was right. Using the wormstone and Horace’s measuring device, he calculated the approximate rate of sinking (2.2 mm/year). This was also a measure of soil deposition, which he attributed to earthworms depositing the sediment through fecal castings. Extrapolating these results further, he estimated that 7.5 to 18 tons (6.8-16.4 tonnes) of soil were moved by worms in a typical acre (0.4 hectares) of land.

Something very important to remember in Darwin’s approach to this study was that he was not just a biologist, but also an excellent geologist, taught early in his career – and later befriended – by one of the founders of modern geology, Charles Lyell. Consequently, he had a long-term view of how small, incremental changes every year added up to big changes over time. Or, to put it in Darwin’s own words (The Formation of Vegetable Mould, p. 6), when he responded to a critic claiming that earthworms were too small and weak to have any large-scale effect on their surroundings:

Here we have an instance of that inability to sum up the effects of a continually recurrent cause, which has often retarded the progress of science, as formerly in the case of geology, and more recently in that of the principle of evolution.

Darwin wasn’t just a quantitative ichnologist, but he also described and illustrated some of the traces made by earthworms, such as their burrows, aestivation chmabers, fecal pellets, and turrets made by their fecal casts. (Much later, in 2007, South American paleontologists described fossil examples of fecal pellets and aestivation chambers from Pleistocene rocks of Uruguay.) Darwin even noted the orientations and species of leaves earthworms pulled into burrows to plug these (p. 64-82), then he independently tested these results with pine needles and triangles of paper (p. 82-90)!

Illustrations of turrets made by fecal pellets of earthworms, in The Formation of Vegetable Mould through the Action of Worms with Observations on Their Habits (1881): from left to right, Figure 2 (p. 107), Figure 3 (p. 124), and Figure 4 (p. 127).

In short, Darwin, through combining his vast knowledge of biology with geological principles, had all the right stuff to make for a formidable ichnologist. Even better, he was keenly interested in the ichnological processes happening just outside his house, and didn’t feel the need to take a long boat trip to watch these processes in some far-off, exotic land. Unknowingly, he was also providing an example of how to do “backyard science” long before this term became associated with cost-effective means for introducing children to nature observation.

All of this marvelous research done by Darwin, culminating in his writing a book at Down House that ended up being one of his most popular, leads me to a bit of a mini-rant, followed by my connecting this science to my homes of Indiana and Georgia, and ending with a message of hope, if I may.

Darwin’s earthworm research epitomized the sort of long-term, DIY experimentation that seemingly only Darwin could have done, and in his day. In contrast, to show how far science has changed since his time, the current profit-oriented business model afflicting modern research universities might have demanded Darwin write a multi-million dollar (or pound) grant to conduct this study. (I suppose the piano would have been the most expensive item on the equipment list, and the wormstones the least.)

Moreover, in this hypothetical scenario, Darwin only could have written such a grant after “pre-confirming” most of his results by publishing a series of research papers. And not just by publishing these papers, but also by making sure they were in prestigious journals, most of which would require expensive subscriptions to read, ensuring that only a small handful of people would read about his work. (A book written for a popular audience? Please.) Had Darwin been a young man, the completion of a 30-year-long study also would have depended on whether he was granted tenure early on. This likely would have been decided by people with little or no expertise in geological processes, earthworms, and bioturbation, but who could certainly count grant revenue and compare journal impact factors.

Fortunately, though, Darwin was independently wealthy, well established as a senior scientist, and never had to worry about tenure or other such trivial matters. Instead, he could just focus on studying his much beloved worms, then think of how to share his vast knowledge of them with a broader audience. Darwin never used the word “ichnology” in his writings, let alone “neoichnology,” and he wrote a book on this topic for natural-history enthusiasts, rather than through a series of research papers published in inaccessible journals. Nonetheless, in his own way, he surely advanced the popularization of ichnology through his slow, deliberate, careful, and imaginative methods, which he combined with a desire to communicate these results to all who were interested.

How does all of this link with Indiana and Georgia? Well, Darwin’s “backyard science” reminded me of how I, like many naturalists of a certain generation, grew up learning about nature through what was in my own backyard. Today I have no doubt that my fascination with the behavior and ecology of insects, plants, and yes, earthworms in my Indiana backyard all contributed to a subsequent desire to do science outside as an adult. To satisfy this urge, I later picked geology as my main subject of study, but also took advantage of my biological leanings by concentrating on ichnology in graduate school. My living in Georgia since 1985 and other serendipitous events then eventually led to my writing a book about traces of the Georgia barrier islands (being published through Indiana University Press). In one chapter of this book, when I introduce earthworms as tracemakers, I made sure to write at least a few pages about Mr. Darwin and his experiments with earthworms. So although Darwin never traveled to Indiana or the Georgia coast, I carried my boyhood and adult experiences of both places in my mind to his former home.

Now here’s the hopeful message (not to be confused with a “hopeful monster“). Lots of field-oriented scientists spend much of their time outside for their research, and many require only modest amounts of money for their studies. So what they have begun to do is side-step the reigning corporate mentality influencing so-called “big science” at universities, while also making active attempts to better connect their research with more people than their academic peers. Through organized efforts like The SciFund Challenge and other crowd-sourcing methods, scientists are seeking small personal donations from the public, allowing them to better focus on their research, rather than spending much time, energy, and angst in writing massive research grants that have little chance of being funded. Thus much like earthworm castings, these  donations add up over time and provide rich, fertile ground for conducting basic science. (OK, maybe not the best metaphor, but you get the point.)

Another facet of this research is the stated commitment of scientists to report their research progress through blogs, then publish their peer-reviewed results in open access journals, which provide articles free for anyone with an Internet connection and curiosity in a scientific subject. All of this means that small investigations with big implications – like Darwin’s study on earthworms – are more likely to happen, and are better assured of reaching a public eager to learn about these sciences, while giving the opportunity for people to witness the direct benefits of their investments.

So how does the Darwin family piano relate to his study of earthworms? Do the southeastern U.S., earthworms, and Darwin’s study of their behavior somehow intersect? In answer to the first question, it’s interesting, and in answer to the second, yes. But an explanation of both will have to wait until next time.

In the meantime, if you go out for a walk later today, pay attention to the ground beneath you, and think of how it reflects an ichnological landscape, a result of collective traces made by those “lowly” earthworms, and how Charles Darwin clearly explained this fact in 1881. For me, it was an honor to stand in the same area where Darwin made his measurements, used his humble instruments, and applied his fine mind; this despite my later realization that I was standing on a new ground surface relative to where Darwin stood. After all, 130 years has passed since his death, meaning the ground had been recycled by descendants of the same earthworms he watched with his appreciative and discerning eyes. All of which makes for a different kind of descent with modification, one that instead reflects an ichnological perspective well articulated and appreciated by Darwin.

Darwin’s “sandwalk,” a walking route behind Down House he often took to help with his thinking, and a visible trace today of Darwin’s legacy as one of the first popularizers of ichnology.

Further Reading

Darwin, C. 1881. The Formation of Vegetable Mould through the Action of Worms with Observations on Their Habits. John Murray, London: 326 p. (A scan of the original book, converted to a PDF document, is here.]

Pemberton, S. George and Robert W. Frey. 1990. Darwin on worms: the advent of experimental neoichnology. Ichnos, 1: 65-71. (Text for article here.)

Quammen, D. 2006. The Reluctant Mr. Darwin: An Intimate Portrait of Charles Darwin and the Making of His Theory of Evolution. W.W. Norton, New York: 304 p.

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

 

 

Life Traces as Cover Art

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I’ve been a long-time admirer of the artistic appeal of tracks, trails, burrows, nests, and other traces of animal behavior. My fondness for the beauty of traces also no doubt contributes to my science: after all, the longer I look at a trace, the more I learn about it. This sentiment accords with a long-time principle of paleontology, botany, and other facets of natural history, which is, “If you draw it, you know it,” with the added benefit of expressing your appreciation of natural objects to others through visual depictions.

Here it is: the cover for my upcoming book, Life Traces of the Georgia Coast: Revealing the Unseen Lives of Plants and Animals! The book is scheduled to be published by Indiana University Press in the fall of 2012, so be watching out for it then. But in the meantime, look at the beautiful cover art. Who created it, what inspired it, and what science lies behind its aesthetically pleasing composition? Please read on to find out.

My thinking about traces as objects of art is not very original, though, and in fact has been preceded by most of humanity. For example, animal tracks and other traces were common subjects of rock art extending back to the Pleistocene Epoch. Whether made as pictographs or petroglyphs, these traces of traces are in Australia, southern Africa, Australia, and Europe, with some tens of thousands of years old. Based on this tantalizing evidence, one could reasonably propose that the representation of animal traces through art composes an intrinsic part of our heritage as a species. Yes, I know, that’s a tough hypothesis to pursue any further. So I’ll leave it to my paleoanthropologist colleagues to work out (or not).

Petroglyphs that likely represent bird tracks, etched in Triassic sandstone by Native Americans hundreds of years ago (sorry, I’m a paleontologist, not an archaeologist). The pair of marks on the right is similar to the tracks made by a perching bird with three forward pointing toes and one rearward-pointing toe – such as an eagle – whereas those to the right may be like those of a roadrunner, which has an X-shaped foot. Petroglyphs are in northeastern Arizona, near Petrified Forest National Park.

Much more recently, trace fossils similarly inspired renowned ichnologist Dolf Seilacher, who also saw these vestiges of past behavior as lovely objects that fill us with wonder. As a result, in the mid-1990s, he conceived of a traveling exhibit and book showcasing tableaus of trace fossils and other sedimentary structures, titled Fossil Art. For this show – embraced by natural-history venues but mostly rejected by art museums – Seilacher prepared it by: (1) making latex molds of sedimentary rock surfaces; (2) pouring epoxy resin into the molds to make casts mimicking the original bedding planes; and (3) using indirect lighting to enhance details; and (4) assigning creative titles to each piece as if they were works of art.

So these artificial slabs are not human-made art in the traditional sense, but nonetheless invoke marvel, project splendor, and otherwise make us think, engaging the same senses and thought processes that accompany an appreciation of art. Moreover, the slim book Seilacher authored for the exhibit contains explanatory text about each of the objects, illuminated further by his marvelous illustrations and visual interpretations. I remember first seeing a version of this exhibit in Holzmaden, Germany in 1995, near Seilacher’s home in Tubingen, and most lately enjoyed strolling through it with other many ichnologists – and Seilacher himself – in Krakow, Poland in 2008.

World-renowned ichnologist and (oh yeah) Crafoord Prize winner, Dolf Seilacher, lecturing about the planning and execution of Fossil Art as an exhibit while it was showing at the Geological Museum of Jagiellonian University in Krakow, Poland in September 2008. Photograph by Anthony Martin.

A close-up of Wrong Sided Hands, one of the pieces displayed in Fossil Art, cast from a latex mold of a sample from Lower Triassic Buntsandstein of Germany. The piece is so-called because the false appearance of a “thumb” on the outside of the tracks originally led to the mistaken idea that the animal awkwardly crossed its own path with each step. This turned out to be wrong. Also, check out the mudcracks! Photograph by Anthony Martin.

Another close-up of a piece from Fossil Art, titled Shrimp Burrow Jungle (helpfully translated into Polish here). This one is based on burrow systems made by crustaceans during the Late Triassic in Italy, which became densely populated over time and hence contributed to overlapping systems. Photograph by Anthony Martin.

Hence during my writing of a book about the modern traces of the Georgia barrier islands, I was well aware of how some of these traces could likewise lend to artistic expression. Some of this mindfulness was applied to a collaborative artwork done with my wife, Ruth Schowalter, in which we took an illustration of mine from the book and used it as the inspiration for a large watercolor painting depicting traces that would form with rising sea level along the Georgia coast (discussed in detail here).

Nonetheless, it was especially important to think about traces as art when considering a potential cover for the book. Book authors know all too well that a well-designed, attractive cover is essential for grabbing the attention of a potential reader, so I had that practical consideration in mind. But I also wanted a cover that pleased me personally, sharing my love of beautiful traces with others, especially those varied and wondrous tracks, burrows, and trails I had seen and studied on the Georgia barrier islands during the past 15 years.

In such an endeavor, I also faced the added pressure of precedence set by my publisher, Indiana University Press. My book is part of a series by IU Press, called Life of the Past, which is widely admired not only for its comprehensive coverage of paleontological topics, but also for its fine cover art, showcasing works done by a veritable “who’s who” of “paleoartists,” So I knew the cover art for my book needed to both conform to this legacy of artistic excellence, but also stand out from other books in the series because of its unique themes. After all, this would be first book in Life of the Past focusing specifically on ichnology. Moreover, the book is more concerned on modern tracemakers and their environments, rather than plants and animals of pre-human worlds. This was done with the intention of demonstrating how our knowledge of modern traces helps us to better understand life from the geologic past, an intrinsic principle of geology called uniformitarianism.

Ideally, as an ichnological purist, I would have had a cover devoid of any animals, and just shown environments of the Georgia of the Georgia coast with their traces. Indeed, I did just that in some of my illustrations in the book, in which I purposefully omitted animals and left only their traces. This “ichno-centric” mindset actually serves a pedagogical purpose, in that it would echo the truism that many sedimentary rocks are devoid of body fossils, yet are teeming with trace fossils.

Figure 1.3 from Life Traces of the Georgia Coast, conveying a sense of the variety and abundance of traces on a typical Georgia barrier island, from maritime forest (left) to shallow intertidal (right). I purposefully drew this illustration using a more cartoonish technique to introduce broad search images of traces for people who may not ordinarily think about these. But also notice what’s missing from the figure: the animal tracemakers. Instead, only immobile plants are depicted. Would this make for good cover art? No and no, especially if you’ve seen the typical covers done for Indiana University Press books. Illustration by Anthony Martin.

Realistically, though, I also knew that modern traces, particularly those made in places as easy to visit as parts of the Georgia coast, would be more eye-catching if accompanied by some of their charismatic tracemakers in a beautiful, natural setting. After all, the Georgia coast has lengthy sandy beaches, dunes, maritime forests, and salt marshes, inhabited by a wide variety of animals, such as sea turtles, shorebirds, alligators, horseshoe crabs, ghost crabs, and many others.

I also knew that a paleoartist would not be as well suited to the task of creating a cover as someone who works more with modern environments. A better pick would be someone who was familiar with the landscapes, plants, and animals of the Georgia barrier islands, but also a fine artist. I briefly toyed with the idea of doing it myself, but already felt like far too much of the book had been “DIY,” and was not confident enough in my skills to put together a compelling cover in enough time before the book came together. An artfully done photograph was another possibility, so I sent several prospective examples to the editors for their appraisal, but these were all shot down for not having enough aesthetic elements for an attention-getting cover (i.e., traces + landscapes + sky + water = very difficult to get into a single photo).

Fortunately, through social connections that still happen despite the Internet and its incentives for becoming increasingly introverted, I met Alan Campbell through mutual friends in December 2008 at a dinner party on the Georgia coast. Fortuitously enough, our meeting was also just before Ruth and I did three weeks of field work on the barrier islands for the book. It was only fitting, then, that our first meeting was spent dining with both of us facing a Georgia salt marsh, filled with fiddler crab burrows and other such traces. Alan is a Georgia artist with much life experience along its coast, he has often portrayed its environments through gorgeous watercolors, and he has worked with scientists in the field.

Consequently, I kept Alan in mind as a potential cover artist for the next few years, and after I had finished the text and all figures for the book, I contacted him last year about my idea, while simultaneously suggesting him to the editors at IU Press. After much back-and-forth negotiations, with me in the middle, both parties finally came to an agreement, and Alan had a contract to do the artwork for the cover by December 2011.

To help Alan in researching his task, I sent him all of my illustrations and photos used in the book so that he would have an extensive library of trace images on hand for reference. He also had this blog as a source, in which I regularly write about Georgia-coast traces, explanations that are always accompanied by photographs and an occasional illustration. We also exchanged many e-mails and talked on the phone whenever needed. I told Alan my preferred cover would feature a coastal scene, but one filled with traces. He voiced a concern that the painting might become too “busy,” and the details might be lost in reduction of the image to the size

Alan’s contract specified that he would have preliminary study sketches would be done by February 1, and the final cover art was to be finished by March 30. He was only a little late with the study sketches (delayed by a minor operation), and I was delighted to see the following sketch in mid-February.

Study sketch by Alan Campbell for the cover of Life Traces of the Georgia Coast. Reprinted with his permission, and anyone else who want to use it, you have to ask him, too. By the way, every time you use original artwork without permission, a little kitten dies.

After a little bit of feedback from both me and graphic designers at IU Press, Alan went back to the drawing board (so to speak), and came up with the following watercolor painting.

Life Traces of the Georgia Coast, 2012, watercolor on paper, 14” X 18” by Alan Campbell. Again, if you want to use it, you have to ask him first and get permission. Remember those kittens? They’re alive now, but there’s no guarantee they’re going to stay that way.

I gave this artwork a big thumbs up, as did the people at IU Press. So once approved and the scan was sent to IU Press, it was up to the graphic designers there to pick out the typeface, color of the type for the main title, subtitle, author name, and placement of type without covering up the main composition of the painting. I had no say in this, and that’s a good thing, because they really knew what they were doing. It is a very nicely designed cover, and the only thing that would please me more is if they had produced a holographic image of it. (Maybe next year.)

The final cover art for Life Traces of the Georgia Coast revisited. Does it look a little different, now that you know more about how it came about?

I won’t spoil the fun for potential readers, scientists, and art appreciators by explaining in detail all of the ichnological, ecological, and geological elements incorporated into the cover. After all, I’d like to sell a few copies of the book, while also letting readers make their own personal discoveries. But hopefully all of you now have a better appreciation for how traces made by animals, our recognition and admiration for these, and artistic expression of them can all combine to contribute to a book that can be accurately judged by its cover.

Further Reading

Leigh, J., Kilgo, J., and Campbell, A. 2004. Ossabaw: Evocations of an Island. University of Georgia Press, Athens, Georgia.

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

Morwood, M.J. 2002. Visions from the Past: The Archaeology of Australian Aboriginal Art. Allen & Unwin, Sydney, Australia.

Seilacher, A. 2008. Fossil Art: An Exhibition of the Geologisches Institut. Tubingen University, Tubingen, Germany.

Tomaselli, K.G. 2001. Rock art, the art of tracking, and cybertracking: Demystifying the “Bushmen” in the information age. Visual Anthropology, 14: 77-82.

 

The Ichnology of Peeps

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Once a year, around Easter time, an attentive beachcomber might notice the unusual traces of a migratory animal on the sands of the Georgia barrier islands. Based on a few clues, its traces point toward five identically sized and conjoined tracemakers, indicating some sort of obligatory group behavior.

Eyewitnesses swear these tracemakers – nicknamed “peeps” – possess a few superficial avian qualities, yet they lack many of the anatomical traits we normally associate with birds, such as, well, wings and legs. Indeed, they apparently have flat ventral surfaces, which with their forward movement along beach sands cause trails, rather than trackways.

Peep trail, observed on berm of Nannygoat Beach, Sapelo Island, Georgia. Oddly enough, this trail shows both a sudden start and end, almost as if the peeps were placed and removed from the surface, respectively.

As a result, peep trails – which are sometimes sinuous, but always harmonious – consist of five parallel grooves, each spaced equally and separated by six ridges, four on the interior of the trail and one on each side. Lateral movements along the length of a peep trail can vary the height of these ridges, depending on whether the peeps are banking to the right or left as they turn.

Although flying ability in peeps has only been inferred on the basis of their possible avian affinity, peep traces show only very brief periods of airborne activity. These traces indicate a somewhat clumsy strategy when approaching ground surfaces, culminating in abrupt vertical descents best described to laypeople as “crashing.” Ideally, all five peeps leave impressions of their cranial anatomy, which includes rudimentary beaks and foreshortened premaxillas. I have no idea if this facial configuration reflects acquired characteristics – caused by frequent crashes – or are more attributable to their original genotype.

Peep landing trace, in which impressions of the anterior anatomy are preserved. Note the short beak marks and rounded dorsal portion of the torso, but with a thin shelf close to the ventral surface. Sand ridges around the impressions suggest the tracemaker bounced after landing.

Peep resting traces are sometimes subtle, owing to their light weight, which according to some sources is about 85 grams (3.0 ounces) in total, or 17 grams per peep. In such instances where their resting traces are recognized, though, peep ventral anatomy is more clearly discernible. Interestingly, the anterior portion of their bodies is rounded and broad, but tapers into a blunt, narrow posterior with a possible upturned tail, the latter suggested by a thin groove bisecting the dorsal part of this posterior mark.

But perhaps the puzzling aspect of these traces is their lack of feather impressions. This evidence shows that peeps, despite their inferred avian affinity, must have become secondarily featherless, despite a long history of descent from non-avian dinosaurs.

Peep resting trace, barely noticeable owing to the light weight of its tracemakers, yet still apparent through its typical overall five-part form.

As is typical with resting traces, these are often connected directly to traces of other behaviors, such as locomotion or burrowing. Indeed, peep resting traces sometimes segue into or out of shallow burrows, which again have five impressions on their bases. Burrowing is presumably an adaptive strategy to avoid predation, implying delectable qualities.

A peep resting trace that is also a burrow, and connecting to an exit mark (right) in which the peep tails left impressions with movement up and out of the excavation.

Peeps are rarely sighted outside of small, cellophane-wrapped boxes in urban shopping centers. Nevertheless, one spring I was lucky enough to see a gaggle of them (five, of course), exuberantly unbound. on a beach of Sapelo Island, Georgia. Thus I was able to observe them making trails, landing traces, resting traces, and actively burrow just above the intertidal zone, which may very well be their natural habitat.

Five peeps making a trail as conjoined unit on a Sapelo Island beach, a behavior predicted by their traces. Who says ichnology isn’t a real science?

Peep landing marks from a short aerial excursion, with the peep presence a short distance away also supporting the interpretation of their bouncing forward after landing.

Peeps exiting a shallow burrow that was also a resting trace, a blend of behaviors often implied by traces.

Peeps initiating a deeper burrowing strategy, perhaps as a form of predation avoidance. Note how the trail becomes shortened, straight, and produces a large pile of sand in front of the direction of movement.

Never-before-seen evidence of how these legless peeps burrow! They use a combination of minute lateral undulations and forward movement directed downward at a shallow angle. As a result, the trail entering the burrow becomes covered by sand ridges produced by the subsequent behavior.

Success! These peeps have managed to bury themselves, leaving only a small portion of their heads exposed, with all five watching warily for predators,

Peeps have been the subject of intensive research, but much of this work, however valuable, has been laboratory based and highly experimental. Thus the data I’ve presented here on their traces should greatly expand our understanding of their behavior in the context of natural settings. Further insights on the biology of peeps are currently murky, but their traces hold promise of fitting them into a taxonomic category more precise than “looks like little chicks.”

Although trace fossils of peep trails, landing traces, resting traces, and burrows have not yet been discovered, I propose these should have the following ichnogenus and ichnospecies names: Peepichnus quinquecalles (= “Peep trace of five trails”). However, I anticipate some of my ichnological colleagues will want to split the ichnotaxonomy of peep traces on the basis of whether they were moving horizontally versus vertically (the peeps, not my colleagues) and other such nuances. Personally, I think they just need to relax, stop coming up with so many silly, unpronounceable names, and just enjoy the sweetness of these little tracemakers of the Georgia coast.

 

Into the Dragon’s Lair: Alligator Burrows as Traces

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American alligators (Alligator mississippiensis) tend to provoke strong feelings in people, but the one I encounter the most often is awe, followed closely by fear. Both emotions are certainly justifiable, considering how alligators are not only the largest reptiles living on the Georgia barrier islands, but also are the top predators in both freshwater and salt-water ecosystems in and around those islands. I’ve even encountered them often enough in maritime forests of the islands to regard them as imposing predators in those ecosystems, too.

Time for a relaxing stroll through the maritime forest to revel in its majestic live oaks, languid Spanish moss, and ever-so-green saw palmettos. Say, does that log over there look a little odd to you? (Photo by Anthony Martin, taken on St. Catherines Island.)

But what many people may not know about these Georgia alligators is that they burrow. I’m still a little murky on exactly how they burrow, but they do, and the tunnels of alligators, large and small, are woven throughout the interiors of many Georgia barrier islands. Earlier this week, I was on one of those islands – St. Catherines – having started a survey of alligator burrow locations, sizes, and ecological settings.

Entrance to an alligator burrow in a former freshwater marsh, now dry, yet the burrow is filled with water. How did water get into the burrow, and how does such traces help alligators to survive and thrive? Please read on. (Photograph by Anthony Martin and taken on St. Catherines Island, Georgia.)

In this project, I’m working cooperatively (as opposed to antagonistically) with a colleague of mine at Emory University, Michael Page, as well as Sheldon Skaggs and Robert (Kelly) Vance of Georgia Southern University. As loyal readers may recall, Sheldon and Kelly worked with me on a study of gopher tortoise burrows, also done on St. Catherines Island, in which we combined field descriptions of the burrows with imaging provided by ground-penetrating radar (also known by its acronym, GPR). Hence this project represents “Phase 2” in our study of large reptile burrows there, which we expect will result in at least two peer-reviewed papers and several presentations at professional meetings later this year.

Why is a paleontologist (that would be me) looking at alligator burrows? Well, I’m very interested in how these modern burrows might help us to recognize and properly interpret similar fossil burrows. Considering that alligators and tortoises have lineages that stretch back into the Mesozoic Era, it’s exciting to think that through observations we make of their descendants, we could be witnessing evolutionary echoes of those legacies today.

Indeed, for many people, alligators evoke thoughts of those most famous of Mesozoic denizens – dinosaurs – an allusion that is not so farfetched, and not just because alligators are huge, scaly, and carnivorous. Alligators are also crocodilians, and crocodilians and dinosaurs (including birds) are archosaurs, having shared a common ancestor early in the Mesozoic. However, alligators are an evolutionarily distinct group of crocodilians that likely split from other crocodilians in the Late Jurassic or Early Cretaceous Period, an interpretation based on both fossils and calculated rates of molecular change in their lineages.

Archosaur relatives, reunited on the Georgia coast: great egrets (Ardea alba), which are modern dinosaurs, nesting above American alligators (Alligator mississippiensis), which only remind us of dinosaurs, but shared a common ancestor with them in the Mesozoic Era. (Photograph by Anthony Martin, taken on St. Catherines Island, Georgia.)

Along these lines, I was a coauthor on a paper that documented the only known burrowing dinosaur – Oryctodromeus cubicularis – from mid-Cretaceous rocks in Montana. In this discovery, we had bones of an adult and two half-grown juveniles in a burrow-like structure that matched the size of the adult. I also interpreted similar structures in Cretaceous rocks of Victoria, Australia as the oldest known dinosaur burrows. Sadly, these structures contained no bones, which of course make their interpretation as trace fossils more contentious. Nonetheless, I otherwise pointed out why such burrows would have been likely for small dinosaurs, especially in Australia, which was near the South Pole during the Cretaceous. At least a few of these reasons I gave in the published paper about these structures were inspired by what was known about alligator burrows.

Natural sandstone cast of the burrow of the small ornithopod dinosaur, Oryctodromeus cubicularis, found in Late Cretaceous rocks of western Montana; scale = 15 cm (6 in). (Photograph by Anthony Martin, taken in Montana, USA.)

Enigmatic structure in Early Cretaceous rocks of Victoria, Australia, interpreted as a small dinosaur burrow. It was nearly identical in size (about 2 meters long) and form (gently dipping and spiraling tunnel) to the Montana dinosaur burrow. (Photograph by Anthony Martin, taken in Victoria, Australia.)

What are the purposes of modern alligator burrows? Here are four to think about:

• Dens for Raising Young Alligators – Many of these burrows, like the burrow interpreted for the dinosaur Oryctodromeus, serve as dens for raising young. In such instances, these burrows are occupied by big momma ‘gators, who use them for keeping their newly hatched (and potentially vulnerable) offspring safe from other predators.

Two days ago, Michael and I experienced this behavioral trait in a memorable way while we documented burrow locations. As we walked along the edge of an old canal cutting through the forest, baby alligators, alarmed by our presence, began emitting high-pitched grunts. This then provoked a large alligator – their presumed mother – to enter the water. Her reaction effectively discouraged us from approaching the babies; indeed, we promptly increased our distance from them. (Our mommas didn’t raise no dumb kids.) So although we were hampered in finding out the exact location of this mother’s den, it was likely very close to where we first heard the grunting babies. I have also seen mother alligators on St. Catherines Island usher their little ones through a submerged den entrance, quickly followed by the mother turning around in the burrow and standing guard at the front door.

Oh, what an adorable little baby alligator! What’s that? You say your mother is a little over-protective? Oh. I see. I think I’ll be leaving now… (Photograph by Anthony Martin, taken on St. Catherines Island.)

• Temperature Regulation – Sometimes large male alligators live by themselves in these burrows, like some sort of saurian bachelor pad. For male alligators on their own, these structures are important for maintaining equitable temperatures for these animals. Alligators, like other poikilothermic (“cold-blooded”) vertebrates, depend on their surrounding environments for controlling their body temperatures. Even south Georgia undergoes freezing conditions during the winter, and of course summers there can get brutally hot. Burrows neatly solve both problems, as these “indoor” environments, like caves, provide comfortable year-round living in a space that is neither too cold nor too hot, but just right. The burrowing ability of alligators thus makes them better adapted to colder climates than other crocodilians, such as the American crocodile (Crocodylus acutus), which does not make dwelling burrows and is restricted in the U.S. to the southern part of Florida.

• Protection against Fires – Burrows protect their occupants against a common environmental hazard in the southeastern U.S., fire. This is an advantage of alligator burrows that I did not appreciate until only a few days ago while in the field on St. Catherines. Yesterday, the island manager (and long-time resident) of St. Catherines, Royce Hayes, took us to a spot where last July a fire raged through a mixed maritime forest-freshwater wetland that also has numerous alligator burrows. The day after the fire ended, he saw two pairs of alligator tracks in the ash, meaning that these animals survived the fire by seeking shelter, and further reported that at least one of these trackways led from a burrow. The idea that these burrows can keep alligators safe from fires makes sense, similar to how gopher tortoises can live long lives in fire-dominated long-leaf pine ecosystems.

An area in the southern part of St. Catherines Island, scorched by a fire last July, that is also a freshwater wetland inhabited by alligators with burrows. The burrow entrances are all under water right now, which would work out fine for their alligator occupants if another fire went through there tomorrow. (Photograph by Anthony Martin, taken on St. Catherines Island.)

• Protection against Droughts – Burrows also probably help alligators keep their skins moist during droughts. Because these burrows often intersect the local water table, alligators might continue to use them as homes even when the accompany surface-water body has dried up. We saw several examples of such burrows during the past few days, some of which were occupied by alligators, even though their adjacent water bodies were nearly dry.

For example, yesterday Michael and I, while scouting a few low-lying areas for either occupied or abandoned dens, saw a small alligator – only about a meter (3.3 ft) long – in a dry ditch cutting through the middle of a pine forest. Curious about where alligator’s burrow might be, we approached it to see where it would go. It ran into a partially buried drainage pipe under a sandy road, a handy temporary refuge from potentially threatening bipeds. Seeing no other opening on that side of the road, we then checked the other side of the road, and were pleasantly surprised to find a burrow entrance with standing water in it. This small alligator had made the best of its perilously dry conditions by digging down to water below the ground surface.

Alligator burrow (right) on the edge of a former water body. Notice how water is pooling in the front of the burrow, showing how it intersects the local water table. The entrance also had fresh alligator tracks and tail dragmarks at this entrance, showing that it was still occupied despite the lack of water outside of it. (Photograph by Anthony Martin, taken on Cumberland Island, Georgia.)

Alligator burrows (left foreground and middle background) in a maritime forest, also not associated with a wetland but marking the former location of one. Although the one to the left was unoccupied when we looked at it, it had standing water just below its entrance. This meant an alligator could have hung out in this burrow for a while after the wetland dried up, and it may have just recently departed. Also, once these burrows are high and dry, bones strewn about in front of them also add a delicious sense of dread. Here, Michael Page points at a deer pelvis, minus the rest of the deer. (Photograph by Anthony Martin, taken on St. Catherines Island, Georgia.)

What is especially interesting about the American alligator is how the only other species of modern alligator, A. sinensis in China, is also a fabulous burrower, digging long tunnels there too, which they use for similar purposes. This behavioral trait in two closely related but now geographically distant species implies a shared evolutionary heritage, in which burrowing provided an adaptive advantage for their ancestors.

Thus like many research problems in science, we won’t really know much more about alligator burrows until we gather information about them, test some of the questions and other ideas that emerge from our study, and otherwise do more in-depth (pun intended) research. Nonetheless, our all-too-short trip to St. Catherines Island this week gave us a good start in our ambitions to apply a comprehensive approach to studying alligator burrows. Through a combination of ground-penetrating radar, geographic information systems, geology, and old-fashioned (but time-tested) field observations, we are confident that by the end of our study, we will have a better understanding of how burrows have helped alligators adapt to their environments since the Mesozoic.

Juvenile alligators just outside two over-sized burrows, made and used by previous generations of older and much larger alligators. How might such burrows get preserved in the fossil record? How might we know whether these burrows were reused by younger members of the same species? Or, would we even recognize these as fossil burrows in the first place? All good questions, and all hopefully answerable by studying modern alligator burrows on the Georgia barrier islands. (Photograph by Anthony Martin, taken on Sapelo Island, Georgia.)

Further Reading

Erickson, G.M., et al. 2012. Insights into the ecology and evolutionary success of crocodilians revealed through bite-force and tooth-pressure experimentation. PLoS One, 7(3): doi:10.1371/journal.pone.0031781.

Martin, A.J. 2009. Dinosaur burrows in the Otway Group (Albian) of Victoria, Australia and their relation to Cretaceous polar environments. Cretaceous Research, 30: 1223-1237.

Martin, A.J., Skaggs, S., Vance, R.K., and Greco, V. 2011. Ground-penetrating radar investigation of gopher-tortoise burrows: refining the characterization of modern vertebrate burrows and associated commensal traces. Geological Society of America Abstracts with Programs, 43(5): 381.

St. John, J.A., et al., 2012. Sequencing three crocodilian genomes to illuminate the evolution of archosaurs and amniotes. Genome Biology, 13: 415.

Varricchio, D.J., Martin, A. J., and Katsura, Y. 2007. First trace and body fossil evidence of a burrowing, denning dinosaur. Proceedings of the Royal Society of London B, 274: 1361-1368.

Waters, D.G. 2008. Crocodlians. 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: 271-274.

Acknowledgements: Much appreciation is extended to the St. Catherines Island Foundation, which supported our use of their facilities and vehicles on St. Catherines this week, and Royce Hayes, who enthusiastically shared his extensive knowledge of alligator burrows. I also would like to thank my present colleagues and future co-authors – Michael Page, Sheldon Skaggs, and Kelly Vance – for their valued contributions to this ongoing research: we make a great team. Lastly, I’m grateful to my wife Ruth Schowalter for her assistance both in the field and at home. She’s stared down many an alligator burrow with me on multiple islands of the Georgia coast, which says something about her spousal support for this ongoing research.

Coquina Clams, Listening to and Riding the Waves

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A little more than a week ago, I co-led a class field trip to Cumberland Island, Georgia and the nearby Okefenokee Swamp for a course titled Ecosystems of the Southeastern U.S. Although I had been to both places more than a few times, none of the students – and a few of my colleagues – had never visited either, potentially casting these already special places in a more exciting light for them.

Nonetheless, as is typical with any field trip to a Georgia barrier island, I also noticed new phenomena while on Cumberland, once again demonstrating how field trips with students ideally also cause the instructors to be filled with wide-eyed wonder. Even better, a few seemingly lowly small bivalves – coquina clams (Donax variabilis) – provided the intellectual highlight for me while we were on Cumberland Island. This is saying something for an island bearing charismatic livestock as a touristic draw.

Resting traces (or are they escape traces?) of coquina clams (Donax variabilis) in the upper intertidal zone of a beach on Cumberland Island, Georgia. These clams are buried just underneath each bump of sand, but some others are much deeper and safer. How do I know that? You’ll find out. (Photograph by Anthony Martin.)

We first noticed the clams as a death assemblage just above the uppermost part of the surf zone on the beach. Their shells, some evident as single valves and others as pairs still hinged together, had been deposited by waves following a high tide, then moved slightly by the wind. These finely ribbed and polished shells readily showed why the specific name of coquina clams (D. variabilis) is applied to them, as they display a gorgeous variety of colors: yellow, orange, beige, blue, pink, and other schemes that surely would inspire interior decorators seeking paisley themes.

Coquina clams with both valves intact or apart, some partially covered by windblown sand, and variably colored. (Photo by Anthony Martin, taken on Cumberland Island, Georgia.)

Just a little bit lower on the beach and in the freshly scoured intertidal zone, we then noticed many small bumps of sand. Underneath these bumps were living clams that buried themselves, which helps to avoid drying out between tides or predation by ravenous shorebirds. With regard to the latter, these bivalves still would have been easy targets for shorebirds intent on acquiring some fresh clam snacks: think of a person ducking under a blanket to avoid being eaten by a lion and how well that might work as a tactic. (Please, just think about it and don’t actually test this idea.)

However, instead of simply writing off all coquina clams as inept burrowers who deserve to die at the beaks of their avian overlords – a similar fate experienced by dwarf surf clams (Mulinia lateralis) – we should look well below the surface, and I mean the sand surface. Look again at the photo first shown above. See all of those tiny, paired holes in between the bumps? Those are the traces of siphons from more deeply buried coquina clams, which are much more likely to escape from bivalve-munching birds while also keeping moist until the next high tide.

Here’s the same photo as above, but zoomed in so you can see the details. Did you notice all of the little holes in between shallowly buried clams? If so, bravo. If not, oh well. (Despite the cropping, turning, and otherwise shuffling electrons, this photograph is still by Anthony Martin, and was still taken on Cumberland Island.)

Coquina clams are actually accomplished burrowers, a necessary adaptation for nearly any small animal living in the high-energy surf of a Georgia beach. In the event of a wave breaking on a beach and washing away the top layer of sand, thus exposing a coquina clam, it will open its valves only enough to stick out its foot, which it then vibrates rapidly. This movement loosens the wet sand underneath, and the clam’s smooth, streamlined shell does the rest of the job, allowing it to glide into its self-made local pit of quicksand and vanish from the surface.

Once under the sand, this clam remains in a vertical position and projects its siphons upwards, making paired holes visible at the sand surface. The resulting burrow is Y-shaped, with the clam body making the lower part of the “Y” and the two siphons leaving thinner traces above. On the Cumberland Island beach the given day we observed their traces, I suspect the more deeply buried clams had the benefit of wetter sand early on as the tide dropped, then the ones hiding under mere caps of sand did the best they could with less wet sand later.

Vertical sections of the Y-shaped burrows made by a coquina clam, dwarf surf clam, or similar small, burrowing bivalves on the Georgia coast, in which the clam body is removed and sediment filled in the empty spaces from above. Both views, taken at right angles from one another, assume the clam is not moving up or down in the sand, which actually isn’t very realistic. (Illustration by Anthony Martin.)

How do we apply this knowledge to the fossil record? Paleontologists have found similar small, Y-shaped burrows in fine-grained sedimentary rocks, trace fossils named Polykladichnus. Some of these burrows are interpreted as the works of suspension-feeding bivalves, although polychaete worms or small arthropods are possible tracemakers, too. Where the lower parts of bivalves rested in sediment and left impressions of their lower halves, which were later filled by overlying sediments, are trace fossils called Lockeia. As a result, paleontologists can reliably identify a former presence of bivalves in rocks that might not have any of their shells. (By the way, please remember that trace fossil names are not the names of the bivalves that made the traces, but of the trace fossil itself. Yeah, I know, that’s confusing. But if you need more of an explanation, I have one for you here.)

So what else is significant about coquina clams? Well, for one, their shells were abundant enough to have formed the framework for a loosely cemented limestone common in Florida, coquina limestone. This is a rock encountered by nearly anyone who has enjoyed (or suffered through) an introductory geology lab exercise on sedimentary rock identification. Students of mine have often compared these rock samples to popcorn balls or similar sugar-cemented treats, although I’ve noticed that no one has been tempted to eat them, no matter how long I kept them in lab that day.

Coquina limestone from Florida. Notice that some of the pieces are from other species of bivalves with rougher, corrugated shells, so these rocks are not made entirely of coquina clams. (Photograph by Mark Wilson of Wooster College, and taken from Wikipedia Commons here.)

But I saved the best tidbits of information for last, and something I’ll bet most people don’t know about these clams that makes them, like, totally cool. These little bivalves respond to sound and migrate seasonally. Yes, that’s right: these clams have their own form of “listening” and they can move en masse once the seasons change on the Georgia coast. Here’s how they do both:

Stop, Look, Listen – Of course, clams lack ears (although rumors still persist that they have legs). Thus they do not hear in the sense we do, but instead respond to low-frequency vibrations caused by waves striking the shore. Once these vibrations are detected, they react by: popping out of the sand; jumping up into the water; body surfing on the wave; and quickly reburying themselves in the sand once dumped by that same wave. Like some aficionados of heavy metal or punk rock, louder is better, as higher-decibel waves cause more clams to jump up out of the sand and into the water, an aquatic version of a molluscan mosh pit.

Shelled Migration – Coquina clams, like caribou, wildebeest, and arctic terns, migrate. Unlike the vast distances covered by those animals, though, coquina clams simply move up and down the slope of a beach with the changes of the seasons. Using their wave-surfing and burrowing abilities, they move from the lower intertidal zone – which is where they live during the spring, summer, and fall – to the upper part of the beach, which becomes their winter homes.

So I hope that all of this pondering over a few shells, bumps, and holes on a Cumberland Island beach has helped lend an appreciation for the small wonders on any given Georgia barrier island. Who knows what little discovery the next field trip will bring, or whether some facsimile of what is seen might also be preserved in the fossil record? As the old saying goes, time will tell, whether that time is in the present or the geologic past.

Further Reading

Ellers, O. 1995a. Discrimination among wave-generated sounds by a swash-riding clam. Biological Bulletin, 189: 128-137.

Ellers, O. 1995b. Behavioral control of swash-riding in the clam Donax variabilis. Biological Bulletin, 189: 120-127.

Pemberton, S.G., and Jones, B. 1988. Ichnology of the Pleistocene Ironshore Formation, Grand Cayman Island, British West Indies. Journal of Paleontology, 62: 495-505.

Ruppert, E.E., and Fox, R.S. 1988. Seashore Animals of the Southeast. University of South Carolina Press, Columbia, South Carolina: 429 p.

Turner, H.J., Jr., and Belding, D.L. 1957. The tidal migrations of Donax variabilis Say. Limnology and Oceanography, 2: 120-124.

Going Hog Wild on the Georgia Barrier Islands

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(The following is the third part of a series about traces of invasive species of mammals on the Georgia barrier islands and the ecological effects of these traces. Here is an introduction to the topic, the first entry about the feral horses of Cumberland Island, and the second entry about the feral cattle of Sapleo Island.)

Anytime I hear someone refer to a Georgia barrier island as “pristine,” I wince a little bit, smile, and say, “Well, bless your heart.” The truth is, nearly every island on the Georgia coast, no matter how beautiful, is not in a pristine state, having been considerably altered by humans over the past 4,500 years, whether these were Native Americans, Europeans, or Americans. These varying degrees of change are sometimes subtle but nonetheless there, denoted by the loss of original habitats and native species or the addition of non-native species.

Still, one Georgia barrier island comes close to fulfilling this idealistic label: Wassaw Island, which during its 1,000-year geologic history somehow escaped commercial logging, agriculture, animal husbandry, and year-round settlements. Partially because of this legacy, Wassaw is designated as a National Wildlife Refuge, and is reserved especially for ground-nesting birds. One of the ways this island works well as a refuge for these birds is – as of this writing – its “hog free” status, a condition that can be tested with each visit by looking for the obvious traces of this invasive species.

The interior of Wassaw Island, with maritime forest surrounding a freshwater wetland created by alligators, the rightful owners of the island. On Wassaw, there are no tracks or signs of feral hogs, qualifying it as a “pristine” island. (Photograph by Anthony Martin.]

Contrast this with Cumberland Island National Seashore, where hogs run wild and freely. The huge pits here are in an intertidal zone of a beach on the northwest corner of the island. Naturalist Carol Ruckdeschel (background) for scale. (Photograph by Anthony Martin.)

Feral hogs (Sus scrofa) have a special place in the rogue’s gallery of invasive mammals on the Georgia barrier islands, and most people agree they are the worst of the lot. Hogs are on every large undeveloped island – Cumberland, Sapelo, St. Catherines, and Ossabaw – and they wreak ecological havoc wherever they roam. The widespread damage they cause is largely related to their voracious and omnivorous diet, in which they seek out and eat nearly any foodstuff, whether fungal, plant, or animal, live or dead. Their fine sense of smell is their greatest asset in this respect: every time I have tracked feral hogs, their tracks show head-down-nose-to-the-ground movement as the norm, punctuated by digging that uses a combination of their snouts and front hooves to tear up the ground in their quest for food. In other words, they generally act like, well, you know what.

Most importantly from the standpoint of native animals that try to live more than one generation beyond a single hog meal, feral hogs eat eggs. Hence ground-nesting birds and turtles are among their victims, and hogs are quite keen on eating sea turtle eggs. Mothers of all three species of sea turtles that nest on the Georgia coast – loggerhead (Caretta caretta), green (Chelonia mydas), and leatherback (Dermochelys coriacea) – dig subsurface nests filled with 100-150 eggs full of protein and other nutrients, making tempting targets for any free-ranging feral hogs. Similarly, hogs also threaten another salt-water turtle, the diamondback terrapin (Malaclemys terrapin); this turtle lays its eggs in shallow nests near the edges of salt marshes, which hogs manage to find. Conservation efforts to save diamondback terrapins from human predation have mostly succeeded (it used to be a tasty ingredient in soups), but hogs can’t read and don’t discriminate when it comes to eating eggs. Here is where feral hogs are particularly dangerous as an invasive species: unlike feral horses or cattle, which “merely” degrade parts of their ecosystems: feral hogs can contribute directly to the extinction of native species. As I often tell my students, if you want to cause a species to go extinct, stop it from reproducing.

Sea-turtle nest on Sapelo Island, marked by a stake and protected by plastic fencing to prevent feral hog and raccoon depredation of its eggs. An individual raccoon would only eat about 1/3 of the eggs in a sea-turtle nest, whereas pigs would just keep on eating. (Photograph by Anthony Martin.)

As an ichnologist, though, what astounds me the most about these hogs is the extremely wide ecological range of their traces. I have seen their tracks – often made by groups traveling together – in the deepest interiors of maritime forests, in freshwater wetlands, and crossing back-dune meadows, high salt marshes, coastal dunes, and beaches. If their traces became trace fossils, paleontologists would refer to them as a facies-crossing species, in which facies (think “face”) are the identifiable traits of a sedimentary environment preserved in the geologic record. Based on their tracks and sign, they are ubiquitous in terrestrial and marginal-marine environments. Oh, and did I mention they are also good swimmers? Swimming across a tidal channel at low tide is an easy feat for them, enabling hogs to spread from island to island, without the assistance of humans.

Run away, run away! Feral hogs in a St. Catherines Island salt marsh, consisting of two juveniles and an adult, do not stick around to see whether humans are going to shoot them; they just assume so. This sighting, along with their widespread tracks and other traces, show how feral hogs can occupy and affect nearly every environment on a Georgia barrier island. (Photograph by Anthony Martin.)

So to better understand why feral hogs are such successful invaders of the Georgia islands, it’s helpful to think about their evolutionary history. As expected, this history is complicated, just like that of any domesticated species in which selective breeding narrowed the genetic diversity we see today. About 15 subspecies of Sus scrofa have been identified, making its recent family tree look rather bushy. Based on genetic studies, divergence between wild species of Sus scrofa (so-called “wild boars”) and various subspecies may have happened as long ago as 500,000 years ago in Eurasia, although humans did not capture and start breeding them until about 9,000 years ago.

Depiction of a European wild boar from 1658, in The History of Four-Footed Beasts and Serpents by Edward Topsell. Original image from a woodcut, digital image in Wikipedia Commons here.

The closest extant relatives to these hogs native to North America are peccaries, which live in the southwestern U.S., Central America, and South America. However, peccaries are recent migrants to North America, and only one Pleistocene species (Mylohyus nasutus) is known from the fossil record of the eastern U.S. This means that the post-Pleistocene ecosystems of the eastern U.S., and especially those of the Georgia barrier islands, have never encountered anything like these animals. Also, unlike the feral horses of Cumberland Island and the feral cattle of Sapelo Island, the feral hogs of the Georgia barrier islands were likely introduced early in European colonization of the coast, and may have started with the Spanish in the 16th century.

Unfortunately, part of the selective breeding of Eurasian hogs was for early sexual maturity and large litter sizes. Female feral hogs can reach breeding age at 5 months, and litters typically have 4-8 piglets, but can be greater than 12; females also can produce three litters in just more than a year. Do the math, and that adds up to a lot of pigs in a short amount of time. Furthermore, on Georgia barrier islands with few year-round human residents, the only predation pressures young piglets face daily include raptors (no, not that kind of raptor) or alligators. This means young hogs reach sexual maturity soon enough to rapidly overrun a barrier island.

Feral hog trackway in a sandy intertidal zone of Cumberland Island, showing a typical gallop pattern (four tracks together –> space –> four tracks together), symbolizing how they are running roughshod over this and other islands. (Photograph by Anthony Martin.)

Yet as we have learned in North America, and particularly on the Georgia barrier islands, feral hogs rapidly revert to their Pleistocene roots. Similar to the feral cattle of Sapelo Island, these hogs are rarely seen by people, especially on islands where humans regularly hunt them. Every time I have spotted them on Cumberland, Sapelo, St. Catherines, or Ossabaw, they instantly turn around, briefly flash their potential pork loins and ham hocks, and flee. As anyone who has raised hogs can tell you, pigs are smart and learn quickly. Hence I imagine that after only one or two shootings of their siblings or parents, they readily associate upright bipeds with imminent death, especially if these bipeds are carrying “boomsticks.” (Speaking of which, I know of at least one sea turtle researcher who does his part to decrease feral hog populations – while also feeding the local vultures – through his able use of such a baby-sea-turtle-protection device.)

Hence much of what we learn about these free-ranging pigs and their behaviors in the context of the Georgia barrier islands is from their traces. Among the most commonly encountered feral hog traces are:

• Tracks

• Rooting pits

• Wallows

• Feces

Feral hog tracks are potentially confused with deer tracks, as they both consist of paired hoofprints and overlap in their size ranges, which are about 2.5-6 cm (1-2.5 in) long. Nonetheless, feral hog tracks are less “pointed,” have nearly equal widths and lengths, rounded ends, and the two hoofs often splay. Two dew claws – vestigial toes – frequently register behind the hoofs, especially when hogs step into soft sand or mud or are running. Trackways normally show indirect register of the rear foot onto the front footprint in a diagonal walking pattern, but can also display a whole range from slow walk to full gallop patterns. With repeated use of pathways, trackways become trails, although I’m not sure if hogs are merely using and expanding previously existing whitetail deer trails, if they are blazing their own, or a combination of the two. (I suspect the last of these is the most likely.)

Feral hog tracks, showing nearly equal lengths and widths, rounded ends, and splaying of hooves, all three of which help to distinguish these from whitetail deer tracks. Scale in centimeters. (Photo by Anthony Martin, taken on Sapelo Island.)

Feral hog trackway on upper part of a sandy beach (moving parallel to shore), showing slow diagonal walking pattern, verified by hoof dragmarks between sets of tracks. Scale = 10 cm (4 in). (Photo by Anthony Martin, taken on St. Catherines Island.)

Rooting pits are broad but shallow depressions – as much as 5 m (16 ft) wide and 30 cm (1 ft) deep – that are the direct result of feral hogs digging for food. In most instances, I suspect they are going for fungi and plant roots, but they probably also eat insect larvae, lizards, small mammals, and any other animals that live in burrows. These pits are typically in maritime forests and back-dune meadows, but I have seen them in salt marshes and dunes, and, most surprisingly, in the intertidal areas of beaches. What are they seeking and eating in beach sands? I think anything dead and buried that might be giving off an odor. I have even seen their tracks associated with broken carapaces of horseshoe crabs (Limulus polyphemus), a menu item that never would have occurred to me if I had not seen these traces.

Rooting pit in back-dune meadow on St. Catherines Island. Former student, who answers to the parent-given appellation of “Andrew,” for scale. (Photograph by Anthony Martin.)

Evidence of feral hog feeding on a horseshoe crab (Limulus polyphemus). All I can say is, it must have been really hungry. (Photo by Anthony Martin, taken on St. Catherines Island.)

Wallows are similar in size and appearance to rooting pits, but have a different purpose, which is to provide hogs with relief from both the Georgia summer heat and biting insects that invariably go with this heat. These structures are often near freshwater wetlands in island interiors, but I’ve seen them next to salt marshes, too. If these wallows intersect the local water table, they also make for attractive little ponds for mosquitoes to breed, meaning these hog traces indirectly contribute to the potential spread of mosquito-borne diseases.

Wallow in maritime forest, Sapelo Island, with a standing pool of water indicating the local water table at the time. (Photo by Anthony Martin.)

Hog feces may look initially like deer pellets, but tend to aggregate in clusters. Most of the ones I have seen are filled with vegetation, but the extremely varied diets of feral hogs means you should expect nearly anything to show up in their scat.

Feral hog feces on Sapelo Island, which is more clumped than that of whitetail deer. Scale in centimeters. (Photo by Anthony Martin, taken on Sapelo Island.)

Which of these traces would make it into the fossil record? I would certainly bet on at least some of their tracks getting preserved, based on the sheer ubiquity of these traces in nearly every sedimentary environment of a Georgia barrier island. Other likely traces would be their pits and wallows, although their broad size and shallow depths would make them difficult to recognize unless directly associated with tracks. Feces would be the least likely to make it into the fossil record as coprolites, unless these contained a fair amount of bone or other mineralized stuff, which could happen with hogs.

What to do about these hogs, and how to decrease the impacts of their traces? Well, as most people know, pigs are wonderful, magical animals that were domesticated specifically for their versatile animal protein. So one solution is more active and year-round hunting of hogs, and using them to supplement breakfasts, lunches, and dinners of local residents on the Georgia coast, a neat blend of reducing a harmful feral species while encouraging a chic “locavore” label on such food.

However, the sheer numbers of hogs on some of the islands would likely require a more systematic slaughter to make a dent in their numbers, an approach that would probably deter any ecotourism unrelated to hog hunting. (Let’s just say that firearms and bird watching are an uneasy mix.) The introduction of native predators is another possible solution. For example, Cumberland Island has a population of bobcats (Lynx rufus) that was introduced primarily to control the whitetail deer population, but these cats probably also take a toll on the feral hogs (although how much is unknown). I have even heard suggestions of reintroducing red wolves (Canis rufus) to a few of the islands. These pack-hunting predators were native to the southeastern U.S. before their extirpation by fearful European settlers, and probably would reduce feral hog populations, but just how much of an impact they would have is hard to predict.

In summary, the feral horses, cattle, and hogs of the Georgia barrier islands have significant effects on the ecology and geology of the Georgia barrier islands, and will continue to do so until creative solutions are proposed and implemented to reduce and otherwise manage their numbers. In the meantime, though, these invasive species present opportunities for us to study their traces, learn more about their unseen behaviors, and compare these behaviors with their evolutionary histories. More science is always good, and in this respect, the Georgia barrier islands are the gifts that keep on giving.

Traces of feral mammals on Sapleo Island: feral hog tracks strolling past a piece of feral cattle scat in a sandy road next to a maritime forest. What is the fate of these invasive species on the Georgia barrier islands, and how will these environments continue to change because of their presence? (Photo by Anthony Martin, taken on Sapelo Island.)

Further Reading

Ditchkoff, S.S., and West, B.C. 2007. Ecology and management of feral hogs. Human-Wildlife Conflicts, 1: 149-151.

Giuffra, E., Kijas, J.M.H., Amarger, V., Carlborg, Ö., Jeon, J.-T., and Andersson, L. 2000. The origin of the domestic pig: independent domestication and subsequent introgression. Genetics, 154: 1785-1791.

Mayor, J.J., Jr., and Brisbin, I.L. 2008. Wild Pigs in the United States: Their History, Comparative Morphology, and Current Status. University of Georgia Press, Athens, Georgia: 336 p.

Taylor, R.B., Hellgren, E.C., Gabor, T.M., and Ilse, L.M. 1998. Reproduction of feral pigs in southern Texas. Journal of Mammalogy, 79: 1325-1331.

Wood, G.W., and Roark, D.N. 1980. Food habits of feral hogs in coastal South Carolina. The Journal of Wildlife Management, 44: 506-511.

Tracking the Wild Cattle of Sapelo Island

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(The following is part of a series about traces of key invasive species of mammals on the Georgia barrier islands and the ecological effects of these traces. Here is an introduction to the topic from last month, and the first entry was about the feral horses of Cumberland Island.)

If I were pressed to name my favorite Georgia barrier island, it would be a tough choice, but it would be Sapelo. Many reasons support this preference, both practical and emotional, which I will relate before getting to the topic featured in the title.

Trails made by feral cattle traveling far into a salt marsh on Sapelo Island, Georgia. But I thought cows only stayed in grassy fields and chewed their cuds? Please read on. (Photograph by Anthony Martin.)

As I mentioned in a previous entry, Sapelo is an excellent place to take university students for teaching basic coastal ecology, geology, ichnology, and taphonomy. Many ecologists consider it as the birthplace of modern ecology, which happened in the 1950s and ‘60s, and it hosted studies that established many basic principles of neoichnology (the study of modern traces) in the 1970s and ‘80s. For the latter, one of the key figures was Robert (Bob) Frey, who was my Ph.D. advisor when I attended the University of Georgia. Sapelo’s human history is also fascinating, dating back to more than 4,000 years ago – evidenced by a prominent Native-American shell ring – and continues through today with Hog Hammock, the only Gullah (“saltwater Geechee”) community left on the Georgia coast.

I have been to Sapelo dozens of times, with or without students, and each time there, I continue to be surprised and delighted by some new observation that reveals itself to those with open eyes and minds. Thus it has everything a field-oriented scientist could want, especially one who likes to learn something different with each visit.

All of these facts and feelings, though, may also lend to an impression that Sapelo is an idyllic and ecologically “pure” place, a true slice of what a Georgia barrier island should aspire to be. Alas, it is not, and like other Georgia barrier islands, Sapelo has been ecologically altered because of exotic plants and animals introduced there during colonial and post-colonial times. Among these species, the most noteworthy on Sapelo is Bos taurus, the only population of wild cattle on any Georgia barrier island and one of the few in the continental U.S.

Unlike the feral horses on Cumberland Island, nearly everyone agrees on the origin of the wild cattle on Sapelo: they are most likely descended from domestic cattle released on the island by millionaire R.J. Reynolds, Jr. (of carcinogenic fame). Although the details are sketchy as to exactly when and why he did this, Reynolds, who owned most of Sapelo from 1933 until his death in 1964, let loose his dairy cows and bulls in the first half of the 20th century. Many generations of these cattle have bred in the wild since, and still roam the island in sufficient numbers to warrant some attention from wildlife biologists, ecologists, and others interested in learning about their behavior and impacts on the local ecosystems.

In my experience, though, the words “wild” and “cattle” are rarely used in everyday conversations about these animals that, through our domestication of them, provide us with milk, cheese, and meat. Ask someone to describe a cow, for instance, and most people will be unflattering: “slow,” “docile,” and “stupid” are among the most common adjectives applied, which is sometimes followed by a giggling reference to the Midwestern U.S. tradition of cow-tipping.

Thinking of tipping this cow? Be my guest, and be sure to forward the resulting video to Animal Planet for others’ lurid entertainment. The “cow” is actually a feral bull, and it was standing its ground at the edge of a field on Sapelo Island, fully aware that we spindly little bipeds were staring at it, and seemingly daring us to get closer. The poor quality of this photo is because I had my camera on maximum digital zoom: my momma didn’t raise no dumb kid. (Photograph by Anthony Martin.)

Yet these cattle are descended from wild species, aurochs (Bos primigenius) that survived the end-Pleistocene mass extinctions. You know, the same extinctions event that wiped out mammoths, mastodons, giant ground sloths, wooly rhinoceroses, saber-toothed cats, dire wolves, and other formidable megafauna of the Pleistocene. Hence aurochs must have had adaptive advantages over their Pleistocene cohorts. This was perhaps was related to their preferred ecosystems of wetland forests and swamps: remember that point with reference to Sapelo. Following the mass extinction, though, people in Eurasia, Africa, and India domesticated aurochs about 8,000 years ago. Through selective breeding, people came up with the present-day varieties we see of Bos taurus, which is considered a subspecies of B. primigenius.

Painting titled The Aurochs, by Heinrich Harder (1858-1935), probably made in 1920. Image is in the public domain and I found it on this Web site, authored by Peter Maas. Contrast how the artist depicted an auroch fighting off a pack of wolves with current expectations of how domestic cattle should behave in the face of pack-hunting predators, and you’ll get a better sense of the actual behaviors shown by wild cattle on Sapelo Island.

I am reminded of this evolutionary heritage whenever I go to Sapelo, because the cattle there are cryptic creatures of the maritime forest. Yes, that’s right: cryptic and living in the forest. A casual day-trip visitor to Sapelo will almost never see one, let alone any of several small herds that roam the island. Whenever an individual bull or herd is encountered in more open, grassy areas, they seemingly revert to Pleistocene behavior and slip into the woods, quickly concealing themselves from the prying eyes of humans. In short, they are not slow, docile, or stupid, and would never allow a person to get close enough to make an short-lived and ill-fated attempt to tip any of them.

This is about all you’ll see of a recent presence of the feral cattle on Sapelo Island: tracks, and if you are lucky enough to sight one, it will leave a lot more tracks and sign for you to study than that all-too-brief glimpse. Scale is in centimeters, and look closely where the slightly smaller the rear-foot track (manus) registered directly on top of the fron-tfoot (pes) track. (Photograph by Anthony Martin.)

Hence any meaningful study of these cattle and their ecological effects on Sapelo requires the use of – you guessed it – ichnology. Consequently, I have tracked these cattle, sometimes with my students and sometimes by myself, during many visits there. Although these tracking forays have generated many anecdotal yarns of yore about these “wild cows of mystery” worth retelling, I will reluctantly restrict myself here to summarizing their traces and the effects of these traces on the landscapes of Sapelo.

Traces of feral cattle on Sapelo consist largely of their tracks, trails or otherwise trampled areas, feces, and chew marks. In my experience, the vast majority of their traces are on the northern half of the island, although herds or individual bulls will occasionally leave their marks in the southern half when they graze on grassy areas there.

Tracks made by these feral cattle are unmistakable when compared to those of any other hoofed animal on Sapelo – such as white-tailed deer or feral hogs – which is a function of their greater size. Tracks are shaped like robust, upside-down Valentine’s hearts, with two bilaterally symmetrical hoof impressions rounded in the front and back. Tracks are normally about 9-14 cm (3.5-5.5 in) long, although I have seen newborn calf tracks as small as 5-6 cm (2-2.3 in) long; track widths are slightly less (by about 20%) than lengths. These cattle, like deer, spend much of their time walking slowly, so their rear-foot (pes) impressions often overlap behind their front-foot (manus) impressions, but can also overprint in direct register. Trackways typically show a diagonal-walking pattern, although these can be punctuated by frequent “T-stops,” in which tracks form a “T” pattern, with the top of the “T” made by the front feet whenever a trackmaker stopped.

Near-perfect direct register of smaller rear foot into front-foot tracks made by adult feral cow, recorded in exquisite detail in fine-grained sand. Scale in centimeters. (Photograph by Anthony Martin, taken on Sapelo Island.)

Because these cattle, for the most part, obey herding instincts, they habitually follow one another along the same narrow pathways through maritime forests and salt marshes, resulting in well-worn trails that wind between live oaks in forest interiors or cut straight across marshes. Nonetheless, the cattle also like to use the open freeways provided by the sandy roads that criss-cross much of the northern part of the island, which makes tracking them much easier, especially after a hard rain has “cleaned the slate.” When using a road, the cattle break single file and walk parallel or just behind one another, indicated by their overlapping and side-by-side trackways. On forest trails, they often drag their hooves across the tops of logs downed along trails, chipping and otherwise breaking down the wood.

Feral cattle tracks showing different sizes – and hence age structures – of the cattle, with some trackways overlapping (following one another) and some parallel, taking up the entire width of a sandy road on the north end of Sapelo Island. (Photographs by Anthony Martin, composite of three stitched together in Photoshop™.)

Log on feral-cattle trail, showing chipped wood on surface where hooves dragged across the top, possibly over generations of trail use. White-tailed deer do a similar behavior on their trails, but do not cause such obvious traces. (Photograph by Anthony Martin, taken on Sapelo Island.)

OK, here’s a reminder of something I just said and showed in a photo earlier: these cattle also form trails that wind deeply into the salt marshes. Why? Turns out that instead of restricting themselves to a terrestrial-only diet, they are eating smooth cordgrass (Spartina alterniflora), which grows abundantly in the marshes. This feeding results in their leaving many other traces, such as near-ground-level cropping of Spartina with clean tears, accompanied by considerable trampling of grazed areas. Although I was surprised to discover this for myself several years ago, people who raised cattle on the island in the 19th and early 20th centuries, perhaps through necessity, knew about this alternative foodstuff and fed it to cattle as a substitute for hay. Sure enough, historical references verify the use of smooth cordgrass as part of their diet (of the cattle, not the people, that is).

Evidence that feral cattle of Sapelo walk into salt marshes as a herd and eat the smooth cordgrass (Spartina alterniflora) there, based on trampling and overgrazing. Michael Bauman, who was an Emory undergraduate student at the time, for scale. (Photographs by Anthony Martin.)

Close-up of traces left on smooth cordgrass from feral cattle grazing, which are at various heights according to the level of their grazing activity. (Photograph by Anthony Martin, taken on Sapelo Island.)

Of course, among the most obvious traces these cattle leave in their wake are the end products of digestion (pun intended), feces. These “cow patties” vary in size depending on both the size of the tracemaker and liquid content of the scat. The bigger the tracemaker and the greater the water content to the plants, the wider the patties, which can exceed dinner-plate size. Similar to the situation on Cumberland Island with its feral horses and their feces, the native dung beetles must not be able to keep up with such a bounty, as I see many unrecycled, dried patties throughout the island, and have nearly stepped on freshly dropped pies that showed no signs of having been discovered by caring dung-beetle mothers.

Looks like cow scat. Smells like cow scat. Feels like cow scat. Tastes like cow scat. Good thing we didn’t step in it! But notice that the tracemaker did, leaving a bonus trace (track) on top of its impressive pile. (Photograph taken by Anthony Martin, taken on Sapelo Island.)

Given that the northern part of the island has extensive salt marshes flanking the maritime forest, and places with fresh-water sloughs containing more wetland plants, it makes sense that the cattle would stay mostly in that half of the island. The absence of humans on the north end of the island – other than occasional deer hunters, Department of Natural Resources personnel, or crazy ichnologists – is also a plus, as these cattle avoid people whenever possible.

But how does any of this relate to geology and paleontology? Well, because these feral cattle interact so much with Sapelo salt marshes, I actually included these animals as marginal-marine tracemakers in my upcoming book (Life Traces of the Georgia Coast, just in case you needed reminding). This places these bovines in the same category as feral horses – which negatively affect coastal dunes and salt marshes – and feral hogs, which even go into the intertidal zones of beaches for their foraging.

The biggest difference between the cattle and these other two hoofed species, though, is their impact on the marshes. In all of my years of noting cattle tracks and other sign on Sapelo, I have never seen evidence of their going to the beach, or even to the coastal dunes. Instead, they stay in the forests and wetlands, whether the latter are fresh-water or salt-water ones. This possibly reflects how the cattle, within just a few generations, switched back to auroch behaviors of the Pleistocene, preferring to live in wooded wetlands instead of in the terrestrial grasslands we modern humans keep forcing them to graze.

Thus any paleontologists looking into the fossil record of aurochs or their ancestral species – whether of body fossils or trace fossils – might use these present-day clues when prospecting for fossils. This serves as a great example of why I urge paleontologists to pay attention to invasion ecology and conservation biology, in which “ecologically impure” invasive species give us valuable insights on their evolutionary histories.

What else can we learn about these feral cattle and their ecological and geological impacts on Sapelo, especially through studies of their traces? For one, knowing the actual number of cattle on the island would be useful, as their quantity surely relates to how well the island ecosystems can handle present and future populations. But probably more important than this would be better defining their behaviors in the context of these non-native ecosystems. How to do this with a species that stays hidden so well, one that has apparently reverted to a Pleistocene way of life? Fortunately, behaviors can be defined through the ichnological methods I have outlined here. These methods can then easily augment others normally used by conservation biologists, such as trail cameras and direct observation.

Once this is done, we will know much more about these wild cattle than before, and will no longer have to rely on whispered legends of the mysterious bovines of Sapelo Island. Regardless, there is certainly still room for such stories, perhaps even artwork, operas, plays, movies, and music. Cattle have played such an integral role in the development of humanity, there is every reason to suppose that, as long as they continue to live on Sapelo, they and their traces will continue to intrigue us.

Further Reading

Ajmone-Marsan, P., Fernando Garcia, J., and Lenstra, J.A. 2010. On the origin of cattle: how aurochs became cattle and colonized the world. Evolutionary Anthropology, 19: 148-157.

Bailey, C., and Bledsoe, C. 2000. God, Dr. Buzzard, and the Bolito Man: A Saltwater Geechee Talks about Life. Doubleday, New York: 334 p.

McFeeley, W.S. 1995. Sapelo’s People: A Long Walk into Freedom. W.W. Norton, New York: 200 p.

Sullivan, B. 2000. Sapelo Island (GA): Images of America. Arcadia Publishing,  Mt. Pleasant, South Carolina: 128 p.

Teal, M., and Teal, J.M. 1964. Portrait of an Island. Atheneum, New York: 167 p. [reprinted by University of Georgia Press, Athens, in 1997: 184 p.]