Tag Archives: Georgia barrier islands

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.

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.

 

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.

Tracking the Wild Horses of Cumberland Island

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(The following post is one of a series about traces of important invasive species of mammals on the Georgia barrier islands and the ecological effects of these traces. An introduction to this topic from last week is here.)

Perhaps the most charismatic yet problematic of non-native animals on any of the Georgia barrier islands are the wild horses (Equus caballus) of Cumberland Island. These horses are the source of much controversy, which becomes even more apparent whenever anyone tries to apply some actual science to them. So I will talk about them here from my perspective as a paleontologist and geologist in the hope that this will add another dimension to what is often presented as a two-sided and emotional argument.

Ah, the wild horses of Cumberland Island, Georgia, roaming free since the time of the Spanish in a pristine, unspoiled landscape, grazing contently on the sea oats and strolling through the coastal dunes, in perfect harmony with nature. How much of the preceding sentence is wrong? Almost all of it. If you want to find out why, please read on. But if your mind is already made up about the feral horses of Cumberland and you don’t want to hear anything bad said about them, then you might like this site. (Photograph by Anthony Martin.)

Cumberland Island, much of which is part of the U.S. National Park system as a National Seashore, is the only Georgia barrier island with a population of feral horses. Nevertheless, despite their uniqueness and fame – the latter figuring as key attractions in advertisements about Cumberland and inspiring dreamy book titles – their origins remain murky. One of the recurring romanticized claims is that these horses descended from livestock brought there by Spanish expeditions in the 16th century. This idea is reassuring to the people who repeat it for two reasons:

(1) It establishes horses as living in the landscape for a long time (especially by American standards), meaning that their presence there now is considered “natural.”

(2) It lends itself to the comforting thought that the horses connect to a European cultural heritage, putting an Old World imprint on a New World place.

However, once said enough times, such just-so stories become faith-based and any evidence contradicting them is not tolerated. Thus even when genetic studies of the Cumberland horses show they are not appreciably different from populations of horses on other islands of the eastern U.S. (arguing against a purely Spanish origin), any questioning of the stated premise – in my experience – provokes angry responses from its defenders.

I suspect this virulent reaction is a direct result of challenging both the “naturalness” and “cultural heritage” of the horses on Cumberland. In reality, though, these are opposing values. After all, an admission that these feral horses came from European stock at any point during the past 500 years supports how they clearly do not belong on Cumberland Island, or anywhere else in the Western Hemisphere if we’re talking about the last 10,000 years or so. In other words, the point is moot whether the current horse population originated in the 16th, 17th, 18th, 19th, or 20th century, or is a mixture of older and newer stock. If only horses could talk, then we would know for sure. (A detailed history of the horses on Cumberland Island is provided here for anyone interested in learning more about this.)

Arguments of heritage aside, these horses are newcomers in a geological and ecological sense. The fossil record of the modern Georgia barrier islands backs this up, as some of the islands (including Cumberland) have sediments more than 40,000 years old, but none have body or trace fossils of horses, or anything like a horse. Although three species of horses were living on the mainland part of North America during the Pleistocene Epoch until their respective extinctions more than 10,000 years ago, none were known to have inhabited any of the barrier islands, Pleistocene or recent. The closest ancient analogue to horses on any of the Georgia barrier islands would have been bison (Bison bison), but their bones are rare. This scarcity leads paleontologists to wonder whether the islands ever had self-sustaining populations of large herbivores.

So with all of that human history and pre-history in mind, the traces made by the feral horses of Cumberland and their ecological effects are exceptional to it and every other Georgia barrier island, and hence worth our attention. Just to keep this simple, I will cover three primary types of traces made by these horses. What these traces all have in common (other than being made by a horse, of course) is the decidedly negative impacts these have on the native plants and animals of Cumberland, including keystone species in the oft-labeled “pristine” ecosystems of the island.

• Tracks and trails – These traces are the abundant and easily spotted on Cumberland, even to someone with little or no training in ichnology. Horses are unguligrade, which means they are walking on their toenails (unguals), and the ungual (more popularly called a hoof) is on a single digit. Hooves make circular to slightly oval compression shapes, but if preserved in the right substrate – like a firm mud or fine sand – they will show a “Pac-Man”-like form. Front-foot (manus) tracks are slightly larger than rear-foot (pes) tracks; manus impressions are 11-14 cm (4.3-5.5 in) long and 10-13 cm (4-5.1 in) wide, whereas pes impressions are 11-13 cm (4.3-5.1 in) long and 9-12 cm (3.5-4.7 in) wide, with variations in size depending on ages of the horses making the tracks.

Trackway of feral horse moving through the coastal dunes of Cumberland Island. Note the diagonal walking pattern and how front- and rear-foot impressions merge to make oblong compound traces.

An important point to keep in mind when tracking horses or any other hoofed animals is that their feet readily cut through sediments and vegetation, leaving much more sharply defined and deeper impressions than padded feet of an equivalent-sized animal. Because Georgia-coast sands contain whitish quartz and darker heavy minerals, these contrasting sand colors help to outline horse tracks on surfaces and in cross-section as deep and sharply defined structures that cut across the bedding.

When asked to think about horses in motion, it might be tempting to imagine them galloping, especially along a beach at sunset. Nonetheless, a horse would tire quickly if it galloped all day, especially for no valid reason. Instead, its normal gait is a slow walk, which causes the rear foot to register partially on top of the front-foot impression, but slightly behind; with a slightly faster walk, the rear foot will exceed the front-foot impression. The overall trackway pattern then is what many trackers call “diagonal-walking,” as the right-left-right alternation of steps can be linked with imaginary diagonal lines. Trackway width, also known as straddle, is about 20-40 cm (8-16 in) if a horse was just walking normally, but narrows noticeably once it starts picking up speed.

Feral-horse tracks on Cumberland Island, a close-up of the same trackway shown in the previous photo. This one was likely doing a slow walk, with indirect register of the rear foot just behind and onto the front-foot impression. The scale (my shoe) is a size 8½ mens. (Photograph by Anthony Martin.)

Given enough back-and-forth movement along preferred paths, repeating and overlapping trackways result in trails, which can be picked out as linear bare patches of exposed sand or mud cutting through vegetation. Because horses are much larger than the native white-tailed deer (Odocoileus virginianus) on Cumberland, their trails are considerably wider.

Feral-horse trail along the edge of a low salt marsh where they have trampled and overgrazed the smooth cordgrass in that marsh (Spartina alterniflora). (Photograph by Anthony Martin, taken on Cumberland Island.)

• Chew marks – Horses are grazers and low-level browsers, and they eat a wide variety of vegetation on Cumberland. The most important plant species they eat through grazing are smooth cordgrass (Spartina alterniflora), sea oats (Uniola paniculata), and live oak (Quercus virginiana).  All three of these plants are keystone species in their respective ecosystems: smooth cordgrass predominates in the low salt marshes, sea oats are the mainstay plants of coastal dunes, and live oaks are the largest and most long-lived trees in the maritime forests. Their effects of horses consuming  smooth cordgrass and sea oats is straightforward, as these plants hold in sediments in place keep them from eroding, but how do horses affect live oaks? They eat the seedlings, which means that older oaks are being replaced by younger ones at a slower rate.

Grazing traces consist of clean cuts of vegetation within a vertical swath and over a broad area. Horses, unlike white-tailed deer, have teeth on both their upper and lower jaws, thus they shear plants on the branches, stems, or leaves. In contrast, deer leave more ragged marks, as they only have teeth on their lower jaws and hence have to pull on vegetation to break it off. Horses also can make a browse line, which is an abrupt horizontal line of decreased vegetation at a certain consistent height that more-or-less correlates with the average head height of the horses.

• Dung – During any given stroll on Cumberland, you cannot avoid seeing, smelling, and stepping in horse feces. This abundance of fecal material means that the feces are not being recycled quickly enough into the ecosystems, which implies that native populations of dung beetles are overwhelmed by such abundance. I have seen a few traces of dung beetles in fresh piles of feces, but no matter how hard I have looked, I have yet to witness great thundering herds of beetles rolling balls of dung across the Cumberland Island landscape.

An impressive collection of horse dung, which was probably dropped by a single horse. Note the small holes in the middle, which were likely made by dung beetles that tunneled into this rich supply of food for their offspring.

Close-up of those probable dung-beetle burrows, some with short trails attached. The white quartz sand sprinkled on top shows how it was pulled up by beetles from underneath the dung pile and onto the top surface, thus giving a minimum depth of the burrows. (Both photographs by Anthony Martin, taken on Cumberland Island.)

One of the more interesting ecological consequences of horse dung I have seen on Cumberland is how it influences the behavior of smaller animals as pellets or piles form a microtopography. For example, on some of the dunes near Lake Whitney on Cumberland – the largest body of fresh water on any of the Georgia barrier islands – I was surprised to see that small lizards – probably skinks – were moving around the dung piles or burrowing under them.

Horse droppings as a part of the landscape for small lizards. Here their tracks, accompanied by tail dragmarks, wind around partially buried feces in a sand dune. (Photograph by Anthony Martin, taken on Cumberland Island.)

Small lizard burrow entrance immediately below a horse pellet, showing its use as a sort of roof. This could probably inspire some clever statement on shingles and, well, you know, but I’ll refrain for now. (Photograph by Anthony Martin, taken on Cumberland Island.)

All three categories of traces – tracks, chew marks, and dung – can be found together in ecosystems wherever horses are trampling, grazing, and defecating, respectively.

So now let’s put on our paleontologist or geologist hats (not to be confused with archaeologist hats) and ask ourselves about the likelihood of such traces making it into the fossil record, and how we would recognize them if they did. Their likelihood of preservation, in order, would be tracks, feces, and chew marks. Tracks would be evident as large compression shapes in horizontal bedding planes or deep disruptions of bedding planes in vertical section. Feces, or their fossil versions called coprolites, might get preserved, although herbivore feces, filled with vegetative material, is less likely to make it into the fossil record compared to carnivore feces, which may have lots of bone material in it. The last of these – chew marks – would be nearly impossible to tell from normal tearing and other degradation of plant material before it became fossilized. Good luck on that.

But could the ecological damage caused by an invasive species, in which the introduction of a species serve as a sort of trace fossil in itself? In the case of horses or ecologically similar animals, subtle changes to the landscape over time might take place. This experiment actually has been done on Assateauge Island (North Carolina), which also has a feral horse population. In areas where horses were excluded by fences, the dunes were on average 0.6 meters (2 ft) feet higher than those of overgrazed and trampled dunes. Geologists conducted another study done on Shackleford Banks (North Carolina) in which they examined areas where fences had separated non-horse from horse-occupied parts of the island. These geologists similarly found that horses caused dunes to be less than 1.5 m (5 ft) high, whereas dunes without horses were as much as 3.5 m (11.5 ft) high. This meant that storms more easily penetrated the barriers provided by coastal dunes, more commonly resulting in storm-washover fans.

This change in the coastal geology of back-dune areas also means that ground-nesting shorebirds will become less common, as their nests and nestlings will be drowned or buried more frequently. Horses also are known to step on shorebird eggs and nests, or can scare away parents from nests, which increases the likelihood of egg or nest predators taking out the next generation of shorebirds.

If any horses made it to the Georgia barrier islands during the Pleistocene and established breeding populations, a geologic sequence following their arrival would look like this, from bottom to top: high dunes suffused with root traces (before horses); lower dunes corresponding with fewer root traces and deep disruptions of bedding (horse tracks); increased numbers of storm-washover fans; and a high salt-marsh. In short, a geologist would see an overall progression from a dune-dominated shoreline to a high salt marsh. Similarly, a paleontologist might see a decrease in root trace fossils and shorebird nests, eggshells, and tracks, possibly culminating in local extinctions of each.

This is your Georgia coast.

This is your Georgia coast with horses. Any questions?

Top panorama is of high-amplitude coastal dunes and well-vegetated back-dune meadows on Sapelo Island, whereas the bottom panorama is of low-amplitude dunes with no appreciable back-dune meadows on Cumberland Island. (Both panoramas based on photos taken by Anthony Martin.)

Based on what we know then, should the feral horses of Cumberland Island be removed? Yes. Will they be removed? Probably not. However, regardless of happens, I will keep teaching about the horses of Cumberland Island and their traces, both as an educator and a concerned citizen. Perhaps with enough awareness, circumstances will change for the better so that Cumberland Island can not only remain a beautiful place, but also will become more like what it was before the arrival of horses there.

(Next week in this series about invasive mammal species of the Georgia barrier islands and their traces, I’ll cover a less inflammatory but still intriguing topic: the feral cattle of Sapleo Island.)

Further Reading

Buynevich, I.V., Darrow, J.S., Grimes, T.A.Z., Seminack, C.T., and Griffis, N. 2011. Ungulate tracks in coastal sands: recognition and sedimentological significance. Journal of Coastal Research, Special Issue 64: 334-338.

De Stoppalaire, G.H., Gillespie, T.W., Brock, J.C., and Tobin, G.A. 2004. Use of remote sensing techniques to determine the effects of grazing on vegetation cover and dune elevation at Assateague Island National Seashore: impact of horses. Environmental Management, 34: 642-649.

Dilsaver, L.M. 2004. Cumberland Island National Seashore: A History of Conservation Conflict. University of Virginia Press, Charlottesville, Virginia: 304 p.

Elbroch, M. 2003. Mammal Tracks and Sign: A Guide to North American Species. Stackpole Books, Mechanicsburg, Pennsylvania: 779 p.

Goodloe, R.B., Warren, R.J., Osborn, D.A., and Hall, C. 2000. Population characteristics of feral horses on Cumberland Island and their management implications. The Journal of Wildlife Management, 64: 114-121.

Sabine, J.B., Schweitzer, S.H., and Meyers, J.M. 2006. Nest Fate and Productivity of American Oystercatchers, Cumberland Island National Seashore, Georgia. Waterbirds, 29: 308-314.

Turner, M.G. 1987. Effects of grazing by feral horses, clipping, trampling, and burning on a Georgia salt marsh. Estuaries and Coasts, 10: 54-60.

Turner, M.G. 1988. Simulation and management implications of feral horse grazing on Cumberland Island, Georgia. Journal of Range Management, 41: 441-447.

 

 

 

Alien Invaders of the Georgia Coast

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(This is the first in a series of posts about invasive species on the Georgia barrier islands, their traces, the ecological impacts of these traces, and why people should be aware of both their traces and impacts.)

Paleontologists like me face a challenge whenever we study modern environments while trying to learn how parts of these environments might translate into the geologic record. Sure, we always have to take into account taphonomy (fossil preservation), through which we acknowledge that nearly none of the living and dead bodies we see in a given environment will become fossilized; relatively few of their tracks, trails, burrows, or other traces are likely to become trace fossils, either.

Because of this pessimistic (but realistic) outlook, paleontologists often rub a big eraser onto whatever we draw from a modern ecosystem, telling ourselves what will not be there millions of years from now. We then retroactively apply this concept – a part of actualism or, more polysyllabically, uniformitarianism – to what happened thousands or millions of years ago. When paleontologists do this, they assume that today’s processes are a small window through which we can peer, giving insights into processes of the pre-human past.

Feral horse (Equus caballus) tracks crossing coastal dunes on Cumberland Island, Georgia. During their evolutionary history, horses originated in North America and populations migrated to Asia, but populations in North America went extinct during the Pleistocene Epoch about 10,000 years ago. Using the perspective of geologic time, then, could someone argue that horses are actually “native,” and these feral populations are restoring a key part of a pre-human Pleistocene landscape? (Photograph by Anthony Martin.)

However, a huge complication in our quest for actualism is this reality: nearly every ecosystem we can visit on this planet is a hybrid of native and alien species, the latter introduced – intentionally or not – by us. Thus when we watch modern species behaving in the context of their environments, we always need to always ask ourselves how non-native species have cracked the window through which we squint, through the past darkly.

This theme is considered in Charles C. Mann’s most recent book, 1493: Uncovering the New World Columbus Created, in which he argues how nearly all terrestrial ecosystems occupied by people were permanently altered by the rapid introduction of exotic species worldwide following Columbus’s landfall in the Western Hemisphere. Going even further back, though, the introduction of wild dogs (dingoes) into mainland Australia by humans about 5,000 years ago irrevocably changed the environments of an entire continent. Examples like these show that European colonization and its aftermath in human history during the last 500 years was not the sole factor in the spread of non-native species, and hints at how species invasions have been an integral part of humanity and its movement throughout the world.

Something tells me we’re not in Georgia any more. A male-female pair of dingoes (Canis lupus dingo) pose for a picture in Kakadu National Park, Northern Territory, Australia. Although now considered “native,” dingoes are an example of an invasive species that had a huge impact once brought over by people from southeast Asia about 5,000 years ago. For one, its arrival is linked to the extinction of native carnivorous mammals in the mainland Australia, such as thylacines (Thylacinus cynocephalus) and Tasmanian devils (Sarcophilus harrisii). (Photograph by Anthony Martin.)

Well-meaning (but deluded) designations of “pristine,” “untouched,”and “unspoilt” aside, the Georgia barrier islands are no exception to alien invaders. Moreover, like many barrier-islands systems worldwide, they differ greatly from island to island in: which species of invaders are there; numbers of individuals of each species; and the degree of how these organisms impact island ecosystems and even their geological processes.

Feral cat tracks in back-dune meadows of Jekyll Island, Georgia. Jekyll is one of the few Georgia barrier islands with a significant human presence year-round, hence these cats are descended from domestic cats that were either purposefully or accidentally let loose by residents. What impact do these cats have on native species of animals and ecosystems, and are these effects comparable to those of other invasive species on other islands? Scale = 15 cm (6 in). (Photograph by Anthony Martin.)

This is one of the reasons why I devoted several pages of my upcoming book, Life Traces of the Georgia Coast, to the traces of invasive species – tracks, trails, burrows, and so on – despite their failing an “ecological purity test” for anyone who might prefer to focus on native species and their traces. With regard to invasive species, the genie is out of the bottle, so we might as well study what is there, rather than apply yet another metaphorical eraser to species that are drastically shaping modern ecosystems and affecting the behavior of native species, thus likewise altering their traces.

A large pit of disturbed sand in a back-dune meadow caused by feral hogs (Sus crofa) on St. Catherines Island, Georgia. Because feral hogs are wide-ranging omnivores with voracious appetites, they cause considerable alterations to island habitats, from maritime forests to intertidal beaches. How do these traces affect the behavior and ecology of other species, especially native ones, in such a broad range of environments on the Georgia barrier islands? Can their traces actually alter the geological character of the islands? (Photograph by Anthony Martin.)

What are some of these invasive species? What makes for an “invasive species” versus a mere “exotic species”? How do the traces of invasive species affect native species on the Georgia barrier islands, and the ecology and geology of the islands themselves? And how do paleontologists and geologists figure into the study of invasive species?

These are all questions that I hope to explore in upcoming weeks here, and for the sake of simplicity, I will showcase an invasive species of mammal and its traces each week. Some of the photos shown here serve as a visual teaser of the invasive species and their traces that will be covered: feral horses (Equus caballus), cattle (Bos taurus), hogs (Sus crofa), and cats (Felis domestica). Yes, I know, there are many others, but these four are among the most ecologically significant species, they consist of animals that nearly everyone knows, and – best of all – they make easily identifiable traces. So these fours species will provide a starting point in our learning how the Georgia barrier islands can be used as case studies in the traces and ecological effects of traces made by invasive species.

Trail made by feral cattle (Bos taurus) cutting through a salt marsh and extending to the horizon, providing a clue of how this forest-dwelling animal can travel deeply into and affect marginal-marine environments. How might such traces show up in the geologic record, and was there a species that might have made similar traces on the islands in the recent past? (Photograph by Anthony Martin.)

Shorebirds Helping Shorebirds, One Whelk at a Time

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How might the traces of animal behavior influence and lead to changes in the behavior of other animals, or even help other animals? The sands and the muds of the Georgia barrier islands answer this, offering lessons in how seemingly inert tracks, trails, burrows, and other traces can sway decisions, impinging on individual lives and entire ecosystems, and encourage seemingly unlikely partnerships in those ecosystems. Along those lines, we will learn about how the traces made by laughing gulls (Larus altricilla) and knobbed whelks (Busycon carica) aided sanderlings (Calidris alba) in their search for food in the sandy beaches of Jekyll Island.

A roughly triangular depression in a beach sand on Jekyll Island, Georgia, blurred by hundreds of tracks and beak-probe marks of many small shorebirds, all of which were sanderlings (Calidris alba). What is the depression, how was it made, and how did it attract the attention of the sanderlings? Scale = size 8 ½ (men’s), which is about 15 cm (6 in) wide. (Photograph by Anthony Martin.)

Last week, we learned how knobbed whelks (Busycon carica), merely through their making trails and burrows in the sandy beaches of Jekyll Island, unwittingly led to the deaths of dwarf surf clams (Mulinia lateralis), the latter eaten by voracious sanderlings. Just to summarize, the dwarf surf clams preferentially burrowed around areas where whelks had disturbed the beach sand because the burrowing was easier. Yet instead of avoiding sanderling predation, the clustering of these clams around the whelks made it easier for these shorebirds to eat more of them in one sitting. Even better, this scenario, which was pieced together through tracks, burrows, and trails, was later verified by: catching whelks in the act of burying themselves; seeing clams burrow into the wakes of whelk trails; and watching sanderlings stop to mine these whelk-created motherlodes of molluscan goodness.

Before and after photos, showing how the burrowing of a knobbed whelk caused dwarf surf clams to burrow in the same small area (top), which in turn provided a feast for sanderlings (bottom); the latter is evident from the numerous tracks, peak-probe marks, and clam-shaped holes marking where these hapless bivalves formerly resided. (Both photographs by Anthony Martin, taken on Jekyll Island, Georgia.)

Was this the only trace-enhanced form of predation taking place on that beach? By no means, and it wasn’t even the only one involving whelks and their traces, as well as sanderlings getting a good meal from someone else’s traces. This is where a new character – the laughing gull (Larus altricilla) – and a cast of thousands represented by the small crustaceans – mostly amphipods – enter the picture. How these all come together through the life habits and traces these animals leave behind is yet another example of how the Georgia coast offers lessons in how the products of behavior are just as important as the behavior itself.

Considering that knobbed whelks are among the largest marine gastropods in the eastern U.S., it only makes sense that some larger animal would want to eat one whenever it washes up onto a beach. For example, seagulls, which don’t need much encouragement to eat anything, have knobbed whelks on their lengthy menus.

So when a gull flying over a beach sees a whelk doing a poor job of playing “hide-and-seek” during low tide, it will land, walk up to the whelk, and pull it out of its resting spot. From there, the gull will either consume the whelk on the spot, fly away with it to eat elsewhere (“take-out”), or reject it, leaving it high and dry next to its resting trace. An additional trace caused by gull predation might be formed when gulls carry the whelk through the air, drop them onto hard surfaces – such as a firmly packed beach sand – which effectively cracks open their shells and reveals their yummy interiors.

Paired gull tracks in front of a knobbed whelk resting trace, with the whelk tracemaker at the bottom of the photo. Based on size and form, these tracks were made by laughing gulls (Larus altricilla). The one on the left is likely the one that plucked the whelk from its resting trace, as its feet were perfectly positioned to pick up the narrow end of the whelk with its beak. The second gull might have seen what the first was doing and arrived on the scene soon afterwards, hoping to steal this potential meal for itself. For some reason, though, neither one ate it; instead, they discarded their object of desire there on the sandflat. For those of you who wondered if I then just walked away after taking the photo, I assure you that I threw the whelk back into water. At the same time, though, I acknowledged that the same sort of predation and rejection might happen again to that whelk with the next tidal cycle. Other shorebird tracks in the photo are from willets and sanderlings. (Photograph by Anthony Martin, taken on Jekyll Island.)

Sure enough, on the same Jekyll Island beach where we saw the whelk-surf clam-sanderling interactions mentioned last week, and on the same day, my wife Ruth Schowalter and I noticed impressions where whelks had incompletely buried themselves at low tide, only to be pried out by laughing gulls. Although we did not actually witness gulls doing performing, we knew it had happened because their paired tracks were in front of triangular depressions, followed by more tracks with an occasional discarded (but still live) whelk bearing the same dimensions as the impression.

My wife Ruth aptly demonstrates how to document seagull and whelk traces (foreground) while on bicycle, no easy feat for anyone, but a cinch for her.  Labels are: GT = gull tracks; WRT = whelk resting trace; KW = knobbed whelk; SU = spousal unit; and LCEFV = low-carbon-emission field vehicle. (Photograph by Anthony Martin, taken on Jekyll Island, Georgia.)

With this search image of a whelk resting trace in mind, we then figured out what had happened in a few places when we saw much more vaguely defined triangular impressions. These were also whelk resting traces, but they were nearly obliterated by sanderling tracks and beak marks; there was no sign of gulls having been there, nor any whelk bodies. Hence these must have been instances of where the gulls flew away with their successfully acquired whelks to drop them and eat them somewhere else. But why did the sanderlings follow the gulls with the shorebird equivalent of having a big party in a small place?

Yeah, I did it: so what? A laughing gull, looking utterly guiltless, stands casually on a Jekyll Island beach, unaware of how its going after knobbed whelks also might be helping its little sanderling cousins find amphipods. (Photograph by Anthony Martin.)

Although many people may not know this, when they walk hand-in-hand along a sandy Georgia beach, a shorebird smorgasbord lies under their feet in the form of small bivalves and crustaceans. The latter are mostly amphipods (“sand fleas”), which through sheer number of individuals can compose nearly 95% of the animals living in Georgia beach sands. Amphipods normally spend their time burrowing through beach sands and eating algae between sand grains or on their surfaces.

Close-up view of the amphipod Acanthohaustorius millsi, one of about six species of amphipods and billions of individuals living in the beach sands of the Georgia barrier islands, all of which are practically begging small shorebirds to eat them. Photo from here, borrowed from NOAA (National Oceanic and Atmospheric Administration – a very good use of U.S. taxpayer money, thank you very much) and linked to a site about Gray’s Reef National Marine Sanctuary, which is about 30 km (18 mi) east of Sapelo Island, Georgia.

Because amphipods are exceedingly abundant and just below the beach surface, they represent a rich source of protein for small shorebirds. But if you really want to make it easier for these shorebirds to get at this food, just kick your feet as you walk down the beach. This will expose these crustaceans to see the light of day, and the shorebirds will snap them up as these little arthropods desperately try to burrow back into the sand. This, I think, is also what happened with the gulls pulling whelks off the beach surface. Through the seemingly simple, one-on-one predator-prey act of a gull picking up a whelk, it exposed enough amphipods to attract sanderlings, which then set off a predator-prey interaction between the sanderlings and amphipods, all centered on the resting trace of the whelk.

Two whelks near one another resulted in two resting traces, and now both are missing, which likely means they were taken by laughing gulls. Notice how all of the sanderling trampling and beak marks have erased any evidence of the gulls having been there. (Photograph by Anthony Martin, taken on Jekyll Island.)

So as a paleontologist, I always ask myself, how would this look if I found something similar in the fossil record, and how would I interpret it? What I might see would be a dense accumulation of small, overlapping three-toed tracks – with only a few clearly defined – and an otherwise irregular surface riddled by shallow holes. The triangular depression marking the former position by a large snail, obscured by hundreds of tracks and beak marks, might stay unnoticed, or if seen, could be disregarded as an errant scour mark. The large gull tracks would be gone, overprinted by the many tracks and beak marks of the smaller birds.

Take a look again at the scene shown in the first photograph, and imagine it fossilized. Could you piece together the entire story of what happened, even with what you now know from the modern examples? I’m sure that I couldn’t. Scale bar = 15 cm (6 in). (Photograph by Anthony Martin.)

Hence the role of the instigator for this chain of events, the gull or its paleontological doppelganger, as well as its large prey item, would remain both unknown and unknowable. It’s a humbling thought, and exemplary of how geologist or paleontologist should stop to wonder how much they are missing when they recreate ancient worlds from what evidence is there.

Cast (reproduction) of a dense accumulation of small shorebird-like tracks from Late Triassic-Early Jurassic rocks (about 210 million years old) of Patagonia, Argentina. These tracks are probably not from birds, but from small bird-like dinosaurs, and they were formed along a lake shoreline, rather than a seashore. Nonetheless, the tracemaker behaviors may have been similar to those of modern shorebirds. Why were these animals there, and what were they eating? Can we ever know for sure about what other animals preceded them on this small patch of land, what these predecessors eating, and how their traces might have influenced the behavior of the trackmakers? (Photograph by Anthony Martin; cast on display at Museo de Paleontológica, Trelew, Argentina.)

Another parting lesson that came out of these bits of ichnological musings is that all of the observations and ideas in this week’s and last week’s posts blossomed from one morning’s bicycle ride on a Georgia-coast beach. Even more noteworthy, these interpretations of natural history were made on an island that some scientists might write off as “too developed” to study, its biota and their ecological relationships somehow sullied or tainted by a constantly abundant and nearby human presence. So whenever you are on a Georgia barrier island, just take a look at the life traces around you, whether you are the only person on that island or one of thousands, and prepare to be awed.

Further Reading

Croker, R.A. 1968. Distribution and abundance of some intertidal sand beach amphipods accompanying the passage of two hurricanes. Chesapeake Science, 9: 157-162.

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

Grant, J. 1981. A bioenergetic model of shorebird predation on infaunal amphipods. Oikos, 37: 53-62.

Melchor, R. N., S. de Valais, and J. F. Genise. 2002. The oldest bird-like fossil footprints. Nature, 417:936–938.

Wilson, J. 2011. Common Birds of Coastal Georgia. University of Georgia Press, Athens, Georgia: 219 p.

“Worm Burrows” as a Geological Cliché

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Further Reading

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

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

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

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

Ghost Crabs and Their Ghostly Traces

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The ghost crabs of the Georgia barrier islands – all belonging to the species Ocypode quadrata – are among my favorite tracemakers anywhere, any time. My ichnological admiration for them stems from the great variety of behaviors they record in the beach and dune sands of the islands, telling many fascinating tales of what they were doing while no one was watching. Thus I thought it only appropriate that a blog entry posted close to Halloween deserved a story about an animal that not only has the word “ghost” in its common name, but one that also leaves mystifying marks of its unseen behavior.

On the dawn of June 22, 2004 on Sapelo Island (Georgia), my wife Ruth and I were presented with one of the most intriguing of ghost-crab mysteries related to their vestiges. We were scanning the freshly scoured surfaces of Nannygoat Beach on the south end of the island; high tide only a few hours before had cleansed the beach of the previous day’s traces. The low-angle rays of early-morning sunlight were optimal for contrasting any newly made animal signs on the beach, which is why we were there then. We went to the beach with our minds open to anything novel; indeed, my experience with the Georgia barrier islands is that no matter how many times you visit them, they always hold previously unsolved puzzles.

Sure enough, within about 15 minutes of stepping foot on the beach, Ruth paused and asked one of the most simple – yet important – of scientific questions: “What is this?” She pointed to a depression on the sandy surface, and what I saw was astonishing. It was a trace perfectly outlining the lower (ventral) half of a ghost crab, preserving in detail: impressions of all eight walking legs (pereiopods), including their pointed ends (dactyli); its smaller claw (inferior cheliped) and larger claw (superior cheliped); and its main, rectangular body.

A perfect outline of the bottom side of a ghost crab (Ocypode quadrata), found just after dawn and high tide on Nannygoat Beach, Sapelo Island, Georgia. Why would a ghost crab make such a trace? (Scale in centimeters, and photograph taken by Anthony Martin.)

Even more strangely, only one set of tracks connected with this body imprint, leading away from it, and none moved toward it. This was not an impression made by the dead body of a crab. Instead, the tracks showed that the crab was very much alive when it made its resting trace and immediately afterwards. But what happened just before then? It looked as if the crab floated through the air, dropped vertically, made a perfect 10-point landing, sat there for a while, and walked away.

Another exquisitely defined ghost-crab body impression, and with tracks leading away from it, showing this is not a crab “death mask,” but one made by a live crab. (Scale in centimeters, and photograph taken by Anthony Martin.)

The same ghost-crab impression as above, but this time with the crab anatomy labeled and direction of movement after it stopped and sat down on the sand. What happened to the tracks that must have led to its resting spot? And what’s with that word “hydration”? Let’s just say this is what you call “foreshadowing” in the story. (Scale in centimeters, and photograph taken by Anthony Martin.)

Knowing that ghost crabs can do a lot of things, but not aerial acrobatics, we wondered how this could have happened. Well, single observations can be the start of good science, but for this inquiry to progress any further, we had to see if this seemingly unusual observation could be repeated. So we walked further south along the beach to test whether this was an isolated incident, or if we could find any other ghost-crab outlines with single trackways attached. With such a search image in mind, we quickly found about a dozen more such marks made by crabs of various sizes, but showing an identical behavior. Even better, all were located just below the high-tide mark of the previous night.

Yet another beautiful ghost-crab resting trace, surrounded by a scoured beach surface. Lot of these traces and all just below the high-tide mark meant something was happening that could be answered by the awesome power of science. (Scale in centimeters, and photograph taken by Anthony Martin.)

Time to think. These crabs must have walked to their resting places, but why didn’t they leave any tracks? We soon realized that the tracks were certainly made, but not preserved. So like all other surface traces on the beach, they must have been made erased during high tide. Yes, that was it! The crabs walked to the surf zone just after the high tide, sat down, waited long enough for the tide to drop a little bit, and walked away.

Mystery solved? Well, not quite. This was an incomplete explanation, one with a big, unanswered question. Why did the ghost crab walk to – and sit down in – the surf? (With a prompt like that, feel free to create your own intertidal-crab equivalent of “chicken-crossing-road” punch lines.) Ghost crabs normally spend much of their time in deep, J- or Y-shaped burrows close to or in the dunes, and above the high-tide mark. They are most active at night, when they come out of their burrows to scavenge delectable dead things dumped on the beach by waves and tides, or to prey on smaller invertebrates, like dwarf surf clams (Mulinia lateralis). They also leave their burrows to seek mates, which might involve one crab enticing another to check out its burrow.

A seemingly indignant and defiant ghost crab outside of its burrow during the day, either looking for new territory, food, mates, or all three. They’re greedy that way. In this instance, though, it was mostly running away from me and my camera. (Photograph taken by Anthony Martin.)

None of the crabs that made these traces were scavenging, preying, or mating, yet something in the surf was life-sustaining enough for them to risk becoming meals for early-morning predatory shorebirds. I searched my memory for what I had read previously about ghost crabs and their biological needs, and finally realized what could have driven them to the surf in the middle of the night: they were thirsty.

You see, ghost crabs are living examples of so-called “transitional animals” that evolution-deniers insist do not exist, having an interesting mixture of adaptations to different environments. These crabs are descended from fully marine crabs, so they still have gills that allow them to filter oxygen from marine water. Yet they also have little lungs and can breathe air, enabling them to stay out of the water for hours. Having both gills and lungs makes them semi-terrestrial, living in a world between the land and ocean, and dependent on both realms. They live close to the sea for their food, reproduction (females lay their fertilized eggs in sea water), and water, but their main livelihood is gained from the beach and dunes.

In this respect, ghost-crab burrows in the upper parts of beaches and lower parts of dunes provide protection against predators, but also keep the crabs hydrated. One of the functions of a ghost-crab burrow – which can be more than one meter (3.3 feet) deep, is to intersect the water table below. That way, when a crab needs water for proper respiration, it crawls down the burrow to that saturated area and replenishes it bodily fluids. But they can’t stay down there as the tide rises, so they move higher up the burrow to where there’s some air. Unlike blue crabs (Callinectes sapidus), which have completely developed gills and hence fully marine, if you keep a ghost crab in sea water too long, it drowns.

The previous night was a higher tide than normal, which probably flooded many of the ghost-crab burrows and causing these crabs to abandon their homes. This meant the crabs spent most of the night outside of their burrows, resulting in dehydration, but having to wait out the high tide. As soon as the tide turned and began to drop, the crabs ran to the surf zone, settled down into the wet sand, and soaked up water through small openings where the legs connect to the main body. Spiky “hairs” (setae) on their legs help with this water up-take, drawing up moisture from the sand through capillary action.

My legs? Sorry, I meant to shave. Guess you’ll have to deal with it. Hey, wait a minute: does that pose look like it could make anything you’ve already seen, like, oh, I don’t know, a resting trace? Keep reading. (Photograph by Anthony Martin.)

Ghost crabs are amazingly efficient at pulling water out of sand. So their hunkering down onto a saturated sandy surface with waves breaking on top of them must have been like the ghost-crab equivalent of drinking from a funnel, quenching their thirst in a most satisfying way. Meanwhile, waves washed away their tracks leading to these resting spots. They stayed a while, long enough for the tide to drop and expose the sandy beach surface. Only then did they get up and walk away, fully rehydrated, refreshed, and ready to go back to their burrows or dig new ones.

This was a detailed explanation, but one based entirely on traces and what little I knew about ghost crabs from the scientific literature. How else to test it and see whether it was right or not?

If you just said, “By directly observing this interpreted behavior in a ghost crab,” you would be right. A little more than a month later, on July 30, 2004, I actually got to witness this behavior, and on Nannygoat Beach. Back without Ruth this time, and by myself, I was looking for more traces following a high tide, when I saw a small, wraith-like movement out of the corner of my eye. It was a beautiful adult ghost crab, flat-out running in full daylight and heading straight from the dunes to the surf zone. I stood back and watched it reach the surf, where it promptly sat down and became still.

Here’s a ghost crab that doesn’t mind getting a soggy bottom. This one sprinted from the dunes to the surf, stopped abruptly, and sat a spell. (Photograph by Anthony Martin.)

I took photos while walking toward this crab, expecting it to bolt at any moment. Instead, I was instead surprised to see it remain where it sat, even as its eye stalks rotated to look warily at me. Amazed, I grasped that this one must have been thirsty enough to risk being eaten or stomped. The photo you see shows just how close I got to it, and I was thrilled to see it in exactly the same position depicted by the traces Ruth and I had seen the month before.

Although scientists aren’t always right, if you practice good science, you sometimes hit the nail on the head. Or the crab on the sand. Or, well, never mind. Anyway, this ghost crab is making a trace just like the ones documented the month before and in the same place, and it is a direct result of the same behavior interpreted from just the traces and some knowledge of their physiology. It’s almost as if science has predictive power. Who’d have thought? (Photograph by Anthony Martin.)

With the “resting trace = rehydration” hypothesis now supported by both traces and direct observation, I wrote the results into a formal, peer-reviewed paper. Unexpectedly, such traces had never been documented for ghost crabs, and especially from the perspective of a paleontologist. In the paper, published in 2006, I pointed out that this behavior would explain similar-looking trace fossils in the geologic record, or the preservation of crab bodies frozen in the same position by death, perhaps reaching the surf too late and being buried by wave-borne sands. The geological significance of such trace fossils would be their value in pointing exactly to where the surf may have washed across an ancient shore, millions of years ago. Geologists really like this kind of precision, and become grateful to ichnologists who give them such tools they can easily use in the field.

A fossil crab from the Miocene Epoch (about 15 million years old), preserved in a sandstone bed cropping out on a beach near Comodora Rivadavia, Argentina. This crab and others like it in the sandstone were all preserved the same way: nearly entire, implying they were buried quickly, and parallel to the original sandy surface on which they settled. Could these have died after dehydration near the surf, and then been buried? How long ago did some crabs evolve to become semi-terrestrial? I don’t know, but now we have a hypothesis that can be applied to fossils like these and tested. (Coin is about 2.5 cm (1 in) wide; Photograph by Anthony Martin.)

Since then, I have seen these resting traces on the beaches of every Georgia barrier island, in the Bahamas, and other places where ghost crabs dwell. Although trace fossils echoing this behavior in ghost crabs or their ancestors have not yet been found, I predict that with the right images now in mind, geologists and paleontologists will recognize them some day.

So with this ichnological lesson from ghost-crab traces, I hope they have become just a bit less “ghostly” and much more alive in your imaginations.

Further Reading

Duncan, G.A. 1986. Burrows of Ocypode quadrata (Fabricus) as related to slopes of substrate surfaces. Journal of Paleontology, 60: 384-389.

Martin, A.J. 2006. Resting traces of Ocypode quadrata associated with hydration and respiration: Sapelo Island, Georgia, USA. Ichnos, 13: 57-67.

Wolcott, T. G. 1978. Ecological role of ghost crabs, Ocypode quadrata (Fabricius) on an ocean beach: Scavengers or predators? Journal of Experimental Marine Biology and Ecology, 31: 67-82.

Wolcott, T. G. 1984. Uptake of interstitial water from soil: mechanisms and ecological significance in the ghost crab Ocypode quadrata and two gecarcinid land crabs. Physiological Zoology, 57: 161-184.

Georgia Life Traces as Art and Science

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This past Friday evening (October 14), Fernbank Museum of Natural History in Atlanta, Georgia hosted the official opening of Selections, a visual-art show themed on evolution, especially as it relates to Charles Darwin. Many other art shows or other creative ventures have revolved around evolutionary themes, especially in 2009, which marked the 150th anniversary of On the Origin of Species and the 200th of Darwin’s birth. But two aspects of this display make it distinctive: (1) it was planned more than two years in advance to accompany the traveling exhibit Darwin, on loan at Fernbank from the American Museum of Natural History; and (2) five of the eight participating artists, all local to the Atlanta area, are also scientists.

Other than once again disproving the notion that artists and scientists live in divergent intellectual realms, once lamented by C.P. Snow in 1969 (for a few other examples of how this false dichotomy is becoming less and less defensible, look here, here, here, here, and here), I am pleased to share that my wife Ruth Schowalter and I are two of the artists in this show. Seven drawings and paintings of ours are on display, with three of those collaborative works, in which we freely mixed scientific concepts with our respective artistic expressions.

Here I will focus on just one of those works, a collaborative piece titled Abstractions of a Rising Sea (2011). My reason for taking a closer look at this one exclusively is because of its having been visually inspired by plant and animal traces of the Georgia barrier islands. Also, in keeping with a Darwinian theme, it depicts how changing environments – in this case, rising sea level – can likewise impact the survival of species, thus affecting the types of traces that are formed and preserved in a given place.

Abstractions of a Rising Sea (2011), by Ruth Schowalter and Anthony Martin: watercolor on paper, 66 X 101 cm (26” X 40”), on display at Fernbank Museum of Natural History until January 1, 2012. But this isn’t just abstract art: it’s also a scientific hypothesis. How so? Please read on. (Photograph taken by Anthony Martin.)

Although this painting may look abstract to most viewers, given its strange, funky shapes and patterns expressed with a colorful palette, its basic elements actually embody an evidence-based prediction. The artwork design, shown below, originated as a conceptual drawing I made for my upcoming book, Life Traces of the Georgia Coast; in fact, it will be the last illustration in the book. The drawing, which I later scanned and modified slightly with Adobe Photoshop™, portrays a vertical sequence of traces made by plants and animals on a typical Georgia shoreline, but considerably altered as sea level went up along that shoreline. In short, it reflects my prognosis of how a coastal dune will become inundated by the sea over the next few decades, with traces of marine animals succeeding those of terrestrial plants and animals.

The original illustration that inspired the artwork, which I drew to portray the sequence of traces that would be made in a given place on the Georgia coast as sea level goes up in the next few hundred years. (Illustration by Anthony Martin.)

So if you’ll bear with me for a few minutes, here’s a more detailed explanation. The traces at the bottom of the illustration represent those of a coastal dune, with plant-root traces, insect burrows, and sea-turtle nests. Just above, those traces are replaced by the burrows of ghost crabs, which are semi-terrestrial animals, but dependent on the sea. A typical Y-shaped burrow of a ghost crab (Ocypode quadrata), viewed in longitudinal section in the eroded face of a coastal dune on Sapelo Island, Georgia. This formerly open burrow was filled from above by sand of a slightly different composition, making it easier to spot. But also note that it cuts across the layering (bedding) of the dune, showing that the crab burrow is relatively younger than the dune deposit. (Photograph by Anthony Martin.)

Next are burrows made by marine invertebrates that live in the intertidal and shallow subtidal areas of a beach, such as polychaete worms, sea cucumbers, and acorn worms.

A variety of abandoned polychaete worm burrows, all washed out of their original places by a vigorous waves and tides and found along a beach on Sapelo Island, Georgia. Although each burrow is distinctive, what they share are behavioral adaptations to living in sandy environments dominated by the surf, shown by their reinforced walls. All four species of worms also orient their burrows vertically, which helps prevent too-frequent exhumation. (Photograph by Anthony Martin.)

Accompanying these is a snail shell (lower third, center) with a drillhole, a cannibalism trace made when a moon snail preyed on its own kind.

Drillhole in the shell of a common moon snail (Neverita duplicata) caused by another moon snail, a trace of both predation and cannibalism: Sapelo Island, Georgia (Photograph by Anthony Martin.)

A broken clam shell to the right of the snail is a likewise a predation trace, but attributable to a seagull. (The bird flew up with the clam in its beak, dropped it onto a hard-packed beach sand at low tide, and dined on its freshly killed contents.)

Broken shell of the giant Atlantic cockle (Dinocardium robustum), caused by a sea gull that picked it up, flew with it, and dropped it onto a sandflat at low tide on Sapelo Island, Georgia. Scale in centimeters. (Photograph by Anthony Martin.)

The upper half of the figure is then dominated by traces of marine invertebrates that live fully submerged offshore, such as ghost shrimp and other crustaceans, other polychaete worms, sea urchins, and brittle stars.

Labeled version of the illustration, depicting an overall progression from onshore traces (bottom) to offshore traces (above). If this sequence of sand and mud were to fossilize, this is how paleontologists and geologists would interpret it. (Illustration by Anthony Martin.)

The preceding artistic-scientific deconstruction should also help a viewer to better understand how geologists think when they look at a vertical sequence of sedimentary rock. For example, geologists follow several basic principles when trying to figure out the relative timing of different events in the geologic past.

One of these is called superposition, in which the effects of the oldest (first occurring) event in a given sequence of sedimentary rock are at the bottom, and the effects of subsequent events are recorded in progressively younger rocks toward the top.

The second principle is cross-cutting relationships, in that whatever is cutting across a previously existing structure must be younger than it. Think about how an animal burrow may cut across burrows made by previous generations of animals, and how you could unravel the sequence of “burrowing events” by simply observing which intersects which burrow.

A third principle is Walther’s Law, named after German geologist Johannes Walther (1960-1937) which states (more-or-less) that laterally adjacent environments succeed one another vertically. In other words, where a maritime forest and coastal dune are next to one another today on the Georgia coast, a drop in sea level means that coastal dunes might by succeeded vertically by the forest. Conversely, sea level going up implies that sediments of offshore environments, which are currently next to the beach and dunes, will some day overlie those of the dune.

Hence the illustration shows all three principles at play with a rising sea. For example, ghost-crab burrows cut across a sea-turtle nest from above, vertical burrows of a polychaete worm in turn dissect ghost-crab burrows below them, and a ghost-shrimp burrow from above interrupts one limb of a U-shaped acorn-worm burrow. Even better, a trained ichnologist can look at this sequence of traces and discern the environmental change that happened over the time represented by the sediments.

You can test this supposition by showing the illustration to other ichnologists, and I predict they will say, “Looks like sea level went up.” As a result, seemingly abstract patterns can become meaningful as we apply these images within the context of time passing, a concept we think Darwin – as a geologist and biologist – would have appreciated.

When I first showed this illustration to Ruth, she was quite taken by its forms and compositions, and she imagined what it would look like made much larger and in color. So we got to work on it, purposefully choosing a large piece of watercolor paper, onto which I drew the ichnological design. She then composed the color scheme, using a combination of water-color pencils and brushes, and I painted in a few details here and there, but most of the hard work was hers.

Ruth and my artistic styles are quite different – she’s a visionary artist, whereas I’m a more of a surrealist – but we both agree that meaningful art should provoke thought. So we very much like how this artwork also addresses and combines two contentious issues in American society: evolutionary theory and global-climate change. In Georgia, as in many other places in the U.S., scientists and science-educators still encounter resistance to the teaching of evolution, despite its extensive testing during the past 150 years and its consequent acceptance by virtually all scientists worldwide. Likewise, in recent years, so-called “global-warming deniers” have put much effort into rebuffing, ignoring, or otherwise downplaying the effects of human-caused climate change – despite near-universal scientific consensus – resulting in the twisting of scientists’ words or outright censorship.

For the plants, animals, and people who live on the Georgia coast, politically charged arguments become pointless as the shoreline moves up and over the land. As global climate continues to change and sea level goes up along the Georgia coast, how will life respond to these changes, especially if the sea rises faster than most organisms can adapt? This is a question we could have put to Charles Darwin, and one we attempt to pose through this synthesis of art and science.

(Acknowledgements to my wife and art-science collaborator, Ruth Schowalter, for her invaluable input on this post: thank you! Selections, featuring the artwork discussed here as well as others by us and six other artists, will be showing at Fernbank Museum of Natural History in Atlanta, Georgia until January 1, 2012. Admission to the museum includes viewing of the artwork, permanent exhibits, and the Darwin exhibit.)

Further Reading

Pilkey, O.H., and Fraser, M.E., 2005. A Celebration of the World’s Barrier Islands. Columbia University Press, New York: 400 p.

Purcell, W.S., and Gould, S.J., 2000. Crossing Over: Where Art and Science Meet. Three Rivers Press, New York: 159 p.

Trusler, P., Vickers-Rich, P., and Rich, T.H., 2010. The Artist and the Scientists: Bringing Prehistory to Life. Cambridge University Press, Cambridge, U.K.: 320 p.

Why Study Traces in Georgia? A Celebration of the Familiar

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For those of us who live in Georgia, we either forget or don’t know about the ecological and geological specialness of this part of the U.S. For example, my undergraduate students here in Atlanta often talk dreamily about their desire to visit the Amazon River basin, Costa Rica, Kenya, Australia, or other places far removed from Georgia, beguiled as they are by the exotic “other” qualities of those places with their biota and landscapes. On the other hand, almost none of these students have been to the Okefenokee Swamp, the Blue Ridge Mountains, the Cumberland Plateau, the long-leaf pine forests of Ichauway, or the Georgia barrier islands, unless my colleagues or I have taken them there on field trips. Yet these places, especially those with freshwater ecosystems, collectively hold a biodiversity nearly matching that of the Amazon River basin, an evolutionary consequence of the long geologic history of the Appalachian Mountains.

To be fair, I have likewise found myself succumbing to such place-based deflection and lack of appreciation for what is more-or-less in my backyard. In 2001, I realized that I had been to Brazil (three times) more often than Fernbank Forest (two times), even though Fernbank was only a five-minute bicycle ride from home in Decatur, Georgia. This imbalance was soon corrected, though, and many visits later, I learned to appreciate how this old-growth southern Appalachian forest in the middle of metropolitan Atlanta is a gem of biodiversity, every native species of plant and animal a facet testifying to their long evolutionary histories. Still, I wonder why we often ignore what is nearby, even if it is extraordinary?

Related to this quandary is one of the most common questions I encountered from friends, family, and colleagues while writing my book – Life Traces of the Georgia Coast – which was, “Why are you, a paleontologist and geologist, writing about the traces of modern plants and animals in Georgia?” This is a legitimate inquiry, but my answer surprises most people. I tell them that my main reason for staying here in Georgia to study the tracks, trails, burrows, nests, and other traces of its barrier islands is because these traces and their islands are world-class and world-famous. This high quality is directly linked to the biodiversity of the Georgia barrier islands, but also their unique geological histories compared to other barrier-island systems. Furthermore, these islands have inspired more than a few major scientific discoveries related to modern ecology and geology, some of which, made nearly 50 years ago, are still applicable to diagnosing the fossil record and the earth’s geologic history. In short, the Georgia barrier islands and their traces also reflect a legacy recognized by scientists far outside the confines of Georgia.

How so? I’ll explain in upcoming posts, and hope to demonstrate how the marvelous ecosystems of the Georgia coast and its geological processes are the proverbial gift that keeps on giving, continually helping us to better understanding the earth’s geologic past. Now that’s special!

Burrows at dawn: a partial view of the thousands of ghost-shrimp burrows dotting a Georgia beach at low tide, their entrances looking like tiny volcanoes. What makes these burrows so important, scientifically speaking, and why are they something that would cause scientists from outside of Georgia to travel and see in person? Photo by Anthony Martin and taken on Sapelo Island, Georgia.