How Did Freshwater Crayfish Get on a Barrier Island?

Two weeks ago, during an all-too-brief visit to Jekyll Island (Georgia) over the Thanksgiving holiday weekend, I decided to check in on some old friends. When I was first introduced to them about four years ago (2008), their presence on Jekyll was a big surprise for me. But thanks to their distinctive traces and a little bit of detective work, I now know they’re on other Georgia barrier islands, too.

Why look, miniature volcanoes in the middle of a maritime forest on Jekyll Island! Or, could they be something else? (In science, that’s what we like to call an “alternative hypothesis.”) Photo scale (left) in centimeters. (Photograph by Anthony Martin.)

These “friends” were conical towers, which look like small lumpy volcanoes (stratovolcanoes, that is, not shield volcanoes), were the traces of freshwater crayfish. A few of the structures, composed of piled balls of sandy mud, also had circular holes in their centers, and they had all seemingly popped out of the forest floor along the edge of a pool of fresh water. All I needed to do to find them was look in the same place where I was first introduced to them, which was by a Jekyll Island resident who knew about their whereabouts.

The towers were 10-25 cm (4-6 in) wide at their bases, 7-10 cm (3-4 in) tall, and each of the rounded, oval balls of sediment was about 1-1.5 cm (0.4-0.6 in) wide. The overall appearance of the towers said “still fresh,” having not been appreciably weathered, and all that I saw in the area looked about the same age. Knowing a little bit about crayfish behavior, I figure they were made just after the last rainfall on Jekyll, maybe a week or so before I spotted them.

Close-up of a crayfish tower, with a circular hole in the center (that’s the burrow). Scale in centimeters. (Photograph by Anthony Martin, taken on Jekyll Island, Georgia.)

Crayfish that dig burrows adjust their depth according to the water table, which they must do to stay alive because they have gills. If the water table drops, they burrow deeper, but if the water table rises, they move their burrows up. For example, where I live here in the metro Atlanta area, crayfish towers often pop up in people’s backyards the day after a hard rain. (This also means that these people need to get flood insurance, because their backyards are on a floodplain. Thus also demonstrating yet another practical reason to know a little basic ichnology.)

Burrowing was (and still is) accomplished by crayfish using their prominent claws (chelipeds) as spades, rolling up the balls of sediment and placing them outside of the burrow entrance, and thus building up towers. But they also smooth out burrow interiors with their bodies through up-and-down and back-and-forth movement, resulting in their burrows having near-perfect circular cross sections. Crayfish burrow systems can be complicated, with vertical shafts connecting the surface with the below-ground parts, which can consist of branching horizontal tunnels and chambers; the last of these can even be occupied by multiple crayfish.

When I first saw these these towers and burrow cross-sections on Jekyll Island in 2008, I immediately knew they were from crayfish. My certainty was because such traces had been described in loving detail by crayfish researchers and ichnologists, linking these directly to their crustacean makers. In fact, just a few months ago, I saw an example of this connection between traces and tracemakers in my home of Decatur, Georgia, where the drying of a human-made pond there caused the crayfish to burrow into the former pond bottom and move about on its sediments in a desperate attempt to stay wet.

A high density of crayfish burrows in a recently drained human-made pond in Decatur, Georgia. Note the similarity of the towers, circular burrow cross-sections, and rounded balls of sediment to those of the Jekyll Island crayfish burrows. Scale with centimeters. (Photograph by Anthony Martin.)

“Are you looking at me?” Crayfish, about 5 cm (2 in) across, and probably a species of Procambarus, copping an attitude while guarding its burrow entrance. (Photograph by Anthony Martin, taken in Decatur, Georgia.)

With about 70 species documented in the state, Georgia is quite rich in crayfish diversity, qualifying it and bordering states in the southeastern U.S as a “biodiversity hotspot” for these animals. Freshwater crayfish are also geographically widespread – occurring in North and South America, Europe, Madagascar, Australia, New Zealand, New Guinea – a direct result of plate tectonics, which spread and isolated populations from one another during their evolutionary history.

In terms of that history, these crustaceans (decapods, more specifically) split from a common ancestor with marine lobsters about 240 million years ago, an age based on molecular clocks, which have been integrated with fossil evidence. I’ve also seen trace fossils that look very much like crayfish burrows in Late Triassic rocks, from about 210 million years ago, which suggests that burrowing began in this lineage early in the Mesozoic Era.

In a 2008 article I co-authored and published with six other scientists – three paleontologists and three zoologists – we described fossil burrows in rocks from the Early Cretaceous Period (about 115-105 million years ago) of Australia, and named what is still the oldest fossil crayfish in the Southern Hemisphere, Palaeoechinastacus australanus. In this article, we pointed out how burrowing was an adaptation that likely helped these crayfish survive polar winters in Australia during the Cretaceous, but also how burrowing abilities in general have given crayfish an upper claw, er, hand in making it past environmental crises in the geologic past.

Here’s the oldest known fossil freshwater crayfish in Australia and the rest of the Southern Hemisphere, Palaeoechinastacus australanus (= “ancient spiny crayfish of Australia”), found in 105-million-year-old rocks (Early Cretaceous) of southern Victoria. Not everything is there, but you can see most of its tail to the left and the right-side legs. Specimen is Museum Victoria, Melbourne, Australia. (Photograph by Anthony Martin.)

And here’s a bedding plane (horizontal) view of trace fossils attributed to freshwater crayfish burrows, preserved in 115-million-year-old rocks (also Early Cretaceous) near Inverloch, Victoria (Australia). The burrows were filled with sand originally, which cemented differently from the surrounding sediment, making them stand out in positive relief as they weather on the outcrop. Scale = 10 cm (4 in). (Photograph by Anthony Martin.)

So how did these crayfish get onto the Georgia barrier islands? Before answering that, I can tell you how they did not get there, which was from people. Because these are burrowing (infaunal) crayfish, and not ones that hang out on lake or stream bottoms (also known as epibenthic), it’s not very likely that humans purposefully introduced them on the islands for aquaculture. Let’s just say that digging up each crayfish burrow, which may or may not contain a crayfish, would require too much work to make crayfish etoufee worth the effort, no matter how good your recipe might be.

Mmmmm, flavorful freshwater decapod concoction [drooling sounds]. But first imagine having to dig up every single crayfish for this dish. Just to prevent this from happening, your recipe should have some qualifying statement, such as, “Make sure to use epibenthic crayfish, not infaunal ones!” (Original image, modified slightly by me, from Wikipedia Commons here.)

Another point to remember about crayfish is that they are freshwater-only animals, incapable of tolerating salt-water immersion, let alone swimming kilometers through marine-flavored waters to reach offshore islands. Yet I’ve seen their traces on Jekyll and two other Georgia barrier islands, and crayfish species have been reported from two additional islands. (For now I won’t say which other islands or identify the probable species of these crayfish until they’ve been properly studied. Sorry.)

What might seem strange to most people, though, is that I still haven’t seen a single living crayfish on any of the Georgia barrier islands. Nonetheless, seeing and documenting their traces is good enough for me to know where they’re living and how they’re behaving. This again demonstrates one of the many advantages of ichnology: you don’t actually have to see an animal to know it’s there, just as long as it leaves lots of identifiable traces.

Oh yeah: almost forgot about the title of this post. What’s my explanation for how the crayfish got to the islands, including Jekyll? I think they lived on the islands before they were islands. In other words, present-day crayfish on the islands descended from ones that originally lived on the mainland part of Georgia, but these were cut off from their original homeland by the last major sea-level rise (well before the current one, that is). This rise started as long as 11,000 years ago, when the last great ice age of the Pleistocene ended, shedding water from continental glaciers and expanding the seas.

So think of a salty moat filling in the low areas between what are now the Georgia barrier islands and the rest of Georgia, with crayfish on either side of it, metaphorically waving goodbye to one another with their claws. In this scenario, the crayfish of the Georgia barrier islands may represent relics left behind and isolated from their ancestral populations. They may have even undergone genetic drift and became new species, or are well on their way to reproductive isolation from their mainland relatives. But that’s just speculation on my part right now. Like I said, these critters need to be studied before anything can be said about them.

All of this neatly illustrates how our knowledge of the geological past ties in with the present, as well as how ichnology can be applied to conservation biology. With regard to the latter, these little muddy crayfish towers exemplify one of the dangers associated with any rapid, careless development of the Georgia barrier islands. What if most people aren’t aware of the unique plants and animals on the islands because at least some of this biodiversity lies below their feet? Without such knowledge, unheeded development may very well wipe out rare or previously unknown species that have been part of the ecological legacy of the Georgia coast for the past 10,000 years.

This is one of many reasons why environmental protection of the islands is still needed, even on semi-developed one like Jekyll. Fortunately, motivated people are working toward such protection on Jekyll, and most other Georgia barrier islands are under some sort of state or federal protection, or privately owned as preserves.

Nice maritime forest you got there. It’d be a shame if something happened to it. (Photograph by Anthony Martin, taken on Jekyll Island.)

What’s also happened on Jekyll Island is increased ecotourism, highlighted by the success of the Georgia Sea Turtle Center. The center, which opened in 2007, has a rehabilitation center for injured turtles, educates the public about sea turtles nesting on the Georgia coast, and helps to monitor turtle nests on Jekyll during the nesting season. And just how is this monitoring done? By looking for tracks of the nesting mothers on the beaches of Jekyll during nesting season, of course. (Say, didn’t I say something previously about using ichnology in conservation biology?)

So can a Jekyll Island Crayfish Center be far behind? Um, no. Still, it’s time to start thinking of species on the Georgia barrier islands and their traces as assets, bragging points that can be used to bolster ecotourism on the coast. Barrier-island biodiversity is an economic resource that will continue to pay off as long as the species survive and their habitats are protected, while simultaneously feeding our sense of wonder at how these species, including burrowing freshwater crayfish, got to the islands in the first place.

Further Reading

Breinholt, J., Ada, M. P.-L., and Crandall, K.A. 2009. The timing of the diversification of the freshwater crayfish. In Martin, J.W., Crandall, K.A., and Felder, D.L. (editors), Decapod Crustacean Phylogenetics, CRC Press, Boca Raton, Florida: 343-355.

Hobbs, H.H., Jr. 1981. The Crayfishes of Georgia. Smithsonian Institute Press, Washington, D.C.: 549 p.

Hobbs, H.H., Jr. 1988. Crayfish distribution, adaptive radiation and evolution. In: Holdich, D.M., Lowery, R.S. (editors), Freshwater Crayfish: Biology, Management and Exploitation. Croom Helm, London: 52-82.

Martin, A.J. 2011. Ichnology in a time of climate change: predicted effects of rising sea level and temperatures on organismal traces of the Georgia coast. Geological Society of America, Abstracts with Programs, 43(2): 86. Link here.

Martin, A.J., Rich, T.H., Poore, G.C.B., Schultz, M.B., Austin, C.M., Kool, L., and Vickers-Rich, P. 2008. Fossil evidence from Australia for oldest known freshwater crayfish in Gondwana. Gondwana Research, 14: 287-296.

P.S. So you’d like to hear more details on the crayfish of the Georgia barrier islands? Well, then you’re going to have to read my book, which starts out Chapter 5 (on terrestrial invertebrate traces) with a section titled The Crayfish of Jekyll Island. Yes, that’s a sales pitch, but you can also request your public library to get it, or borrow a copy from a friend. Which makes this more of a “knowledge pitch.”

Gopher Tortoises, Making Deep and Meaningful Burrows

As I wrote this post, I was flying from Atlanta, Georgia to Minneapolis, Minnesota to attend the annual meeting of the Geological Society of America (GSA), where I’ll be with about 7-8,000 geoscientists from across and outside of the U.S. Why am I not doing something else, such as field work on the Georgia coast? Well, other than to learn the latest of what’s happening in the world of geology, seeing old friends, and meeting new ones, I’m here to share new scientific knowledge coming out of the Georgia coast with my fellow geologists and paleontologists. The subject of the presentation I will give tomorrow – Tuesday, October 11 – is about the wondrous burrows of a humble-looking, slow-moving, and seemingly lethargic reptile that actually is an ichnological force of nature: the gopher tortoise (Gopherus polyphemus).

A gopher tortoise in captivity, but living a safe and happy life at the 4-H Tidelands Nature Center on Jekyll Island, Georgia. Although it may not look like a big deal, it is a very impressive tracemaker, deserving the rapt attention of geologists and paleontologists. (Photograph taken by Anthony Martin.)

So you’re probably wondering why geologists and paleontologists should hear about gopher tortoises from me. It’s a good question, because I’m not a biologist, and these animals are famous for their very important role in ecosystems. Specifically, they are well known as keystone species in the sandy soils of longleaf pine-wiregrass communities of the southeastern U.S. Just like the keystone to a building, once you remove gopher tortoises from their ecosystems, a lot of other species disappear with it. Surprisingly, their ecological worth all revolves around their burrows.

And oh, what marvelous and grandiose burrows they make! The lengthiest of their measured burrows approach 14 meters (45 feet) long and as much as 6 meters (20 feet) vertically below the ground surface. These burrows commonly twist to the right or left on their way down, which probably helps protect its tortoise occupant against predators, while maintaining a constant temperature and humidity in the burrow. With so much digging, of course, a lot of sand has to be excavated, so the locations of their burrows are easily spotted by looking for piles of sand in the middle of a grassy field or in a longleaf-pine forest. For female tortoises, these sand piles also serve as nesting sites, where they bury their eggs to incubate.

Satellite view of gopher-tortoise burrows on St. Catherines Island, Georgia. Nearly all of the white spots you see in the photo – indicated by the yellow arrows – are the sand piles (aprons) outside of their burrows. Look closely, and you can see some of the trails worn down by tortoises traveling between burrows. Yes, these are animal traces you can see from space! (Original image from the U.S. Geological Survey and Google Earth, taken in May 2008.)

Close-up view of a sand apron outside of a gopher-tortoise burrow entrance. The large amount of sand tells you that this must be a very deep burrow. Field notebook is about 15 cm (6 in) long. (Photograph taken by Anthony Martin on St. Catherines Island, Georgia.)

In cross-section, their burrows have flat bottoms and rounded tops, similar to a tortoise body. Burrow widths varies with the length of the tortoise, as it needs to be wide enough for the tortoise to turn around in the burrow. So this means a 30-cm (12 in) wide burrow can accommodate a tortoise of that length or less. The powerful front limbs of tortoises are specially adapted for digging, ending in flat, spade-like feet with stout claws. Burrow walls are compacted by the hard shell of the tortoise as it moves up and down the burrow. These burrows descend steeply, at angles of 20-40°, which means they have to be good climbers to get out of their deep burrows.

Down-tunnel view of a gopher-tortoise burrow, with the light at the end of that tunnel not  from an oncoming train, but reflected morning sunlight on the tunnel wall at one of its turns. (Photograph by Anthony Martin, taken on St. Catherines Island, Georgia.)

Now think about a tunnel that’s about 10 m (33 ft) long and 30 cm (12 in) wide, and how much space that represents underneath the ground, and you’ll see what I mean about the vital role of these burrows ecologically, geologically, and (most importantly) ichnologically. In terms of ecology, about 200-300 species of invertebrate and vertebrate animals cohabit these burrows (whether a gopher tortoise is in it or not), including the longest snake in North America, the eastern indigo snake (Drymarchon couperi), the secretive gopher frog (Rana capito), the Florida mouse (Podomys floridanus), and a bunch of different insects. At least a few of the insects and the Florida mice make their own burrows, thus adding their little homes to the main burrow, like small anterooms to a big mansion.

Idealized conceptual sketch showing a cut-away view through a gopher-tortoise burrow with many additional burrows made by other animal species. Note especially the short horizontal tunnels near the burrow top, which would have been made by hatchling tortoises, and the vertical shafts that connect to these, which would have been made by Florida mice. (Illustration by Anthony Martin.)

So now you can see why this ichnologist (that would be me) became rather enamored with these burrows. For one thing, they have great preservation potential in the fossil record. A  general rule in ichnology for the preservation of burrows is “deeper is better,” in that burrows that go to great depths are less likely to be eroded by surface weathering and erosion, and more likely to be fossilized. Secondly, we know that vertebrate animals in the geologic past also made big burrows, such as synapsids and even small dinosaurs. I’ve done research on the few dinosaur burrows interpreted from the geologic record, and am especially interested in how such large burrows might compare with similar burrows made by modern animals, such as gopher tortoises.

But how to study these burrows without digging them out and leaving the tortoises undisturbed? Fortunately, two colleagues of mine at Georgia Southern University – Sheldon Skaggs and Robert (Kelly) Vance – came up with an elegant solution, which was to use ground-penetrating radar, also known by its acronym of GPR. This method uses a portable unit to transmit microwaves underground (don’t worry, not these aren’t intense enough to cook the tortoises), which reflect off surfaces with different qualities, especially the curved, compacted surfaces of burrow walls. Computers then process and render these reflections into three-dimensional images that more-or-less represent the forms and geometries of the burrows.

Sure enough, we tried out this technique on gopher-tortoise burrows on St. Catherines Island of the Georgia coast in January and July this year. Although we can’t share all of our results just yet, we did successfully make three-dimensional images of the burrows, all without us having to burrow ourselves, or bother the tortoises by becoming homewreckers. Veronica Greco, a wildlife biologist on St. Catherines Island who has studied the behavior and breeding of the tortoises, also helped us to better understand the biology of these reptiles.

Although it looks like Sheldon (center) is mowing the lawn and I’m (right) just supervising, he’s actually pushing a portable ground-penetrating radar (GPR) unit over a field that has some gopher-tortoise burrows in it, while I walk alongside to look at the reflection profiles. Kelly (background) is no doubt monitoring our every move, but is also recording our location. (Photograph by Ruth Schowalter, taken on St. Catherines Island, Georgia.)

My talk at the GSA meeting will be about how we used GPR to study the burrows in a non-invasive way, and how our results might be applied to studying similar burrows in the fossil record. After the meeting is over, we plan to summarize our results in a research article, which we’ll submit to a journal later this year for peer review.

Unfortunately, gopher tortoises are endangered because of huge losses in acreage of longleaf-pine forests in the southeastern U.S. during the past 200 years or so. Knowing this makes our study of their burrows even more meaningful, for if these wonderful tracemakers go extinct in the near future, we will not have the chance to study them and their burrows. In this sense then, only geologists and paleontologists who know about their ichnology through studies like ours will be able to study their burrows, which would be a sad thing indeed. Let’s hope they survive and thrive, and we can continue to learn more about these superb burrowing animals and their traces.

(P.S. Many thanks to the St. Catherines Island Foundation for their support of our research!)

Further Reading

Aresco, M.J., 1999. Habitat structures associated with juvenile gopher tortoise burrows on pine plantations in Alabama. Chelonian Conservation and Biology, 3: 507-509.

Doonan, T.J., and Stout, I.J., 1994. Effects of gopher tortoise (Gopherus polyphemus) body size on burrow structure. American Midland Naturalist, 131: 273-280.

Epperson, D.M., and Heise, C.D., 2003. Nesting and hatchling ecology of gopher tortoises (Gopherus polyphemus) in southern Mississippi. Journal of Herpetology, 37: 315-324.

Guyer, C., and Hermann, S.M. 1997. Patterns of size and longevity for gopher tortoise burrows: implications for the longleaf pine-wiregrass ecosystem. Bulletin of the Ecological Society of America, 78: 254.

Jackson, D.R. and Milstrey, E.R. 1989. The fauna of gopher tortoise burrows. In Diemer, J.E. (editor), Proceedings of the Gopher Tortoise Relocation Symposium, State of Florida, Game and Freshwater Fish Commission, Tallahassee, Florida: 86-98.

Jones, C.A., and Franz, R. 1990. Use of gopher tortoise burrows by Florida mice (Podomys floridanus) in Putnam County, Florida. Florida Field Naturalist, 18: 45-68.

Lips, K.R. 1991. Vertebrates associated with tortoise (Gopherus polyphemus) burrows in four habitats in south central Florida. Journal of Herpetology, 25: 477-481.

Martin, A.J., Skaggs, S.A., 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.

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.

Witz, B.W., and Wilson, D.S., and Palmer, M.D. 1991. Distribution of Gopherus polyphemus and its vertebrate symbionts in three burrow categories. American Midland Naturalist, 126: 152-158.