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.”

Descent with Modification

At this time last year, Fernbank Museum of Natural History was hosting the Darwin exhibit. On loan from the American Museum of Natural History, this exhibit was a major coup for the museum and the Atlanta area, which has enjoyed a growing culture of celebrating science during the past few years. Along with this exhibit, the museum also planned and concurrently displayed an evolution-themed art show, appropriately titled Selections, which I wrote about then here.*

Descent with Modification (2011), mixed media (colored pencils and ink) on paper, 24″ X 36.” Although this artwork might at first look like a tentacled creature infested with crustaceans and living on a sea bottom, its main form actually mimics a typical burrow system made by ten-legged crustaceans (decapods). Yet it’s also an evolutionary hypothesis. Intrigued? If so, please read on. If not, there are plenty of funny cat-themed Web sites that otherwise require your attention. (Artwork and photograph of the artwork by Anthony Martin.)

One unusual feature of this art show was that five of the eight artists were also scientists (full confession: I was one of them). Furthemore, one of the other artists was married to a scientist (fuller confession: that would be my wife Ruth). The show stayed up for more than three months, which was also as long as the Darwin exhibit resided at Fernbank. Thus we like to think it successfully exposed thousands of museum visitors to the concept that scientists, like many other humans, have artistic inspirations and abilities, neatly refuting the stereotype that not all of us are joyless, left-brained automatons and misanthropes.

Last week I was reminded of this anniversary and further connections between science and art during a campus visit last week by marine biologist and crustacean expert Joel Martin (no relation). Dr. Martin was invited to Emory University to give a public lecture with the provocative title God or Darwin? A Marine Biologist’s Take on the Compatibility of Faith and Evolution. His lecture was the first of several on campus this year about the intersections between matters of faith and science, the Nature of Knowledge Seminar Series. This series was organized as a direct response to the university inviting a commencement speaker this past May who held decidedly strong and publicly expressed anti-science views.

Dr. Martin, who is also an ordained elder in his Presbyterian church and has taught Sunday school to teenagers in his church for more than 20 years, gave an informative, organized, congenial, and otherwise well-delivered presentation to an audience of more than 200 students, staff, faculty, and other people from the Atlanta community. In his talk, Martin effectively explored the false “either-or” choice often presented to Americans who are challenged by those who unknowingly misunderstand or deliberately misrepresent evolutionary theory in favor of their beliefs. Much of what he mentioned, he said, is summarized in a book he wrote for teenagers and their parents, titled The Prism and the Rainbow: A Christian Explains Why Evolution is Not a Threat.

I purposefully won’t mention any of the labels that have been applied to the people and organizations who promote this divisiveness between evolutionary theory and faith. After all, words have power, especially when backed up by Internet search engines. Moreover, it is an old and tired subject, of which I grow weary discussing when there is so much more to learn from the natural world. Better to just say that Martin persuasively conveyed his personal wonder for the insights provided by evolutionary theory, how science informs and melds with his faith, and otherwise showed how science and faith are completely compatible with one another. You know, kind of like science and art.

Previous to his arrival, his host in the Department of Biology asked Emory science faculty via e-mail if any of us would like to have a one-on-one meeting with Dr. Martin during his time here. I leaped at the chance, and was lucky enough to secure a half-hour slot in his schedule. When he and I met in my office, we had an enjoyable chat on a wide range of topics, but mostly on our shared enthusiasm for the evolution of burrowing crustaceans, and particularly marine crustaceans.

Ophiomorpha nodosa, a burrow network in a Pleistocene limestone of San Salvador, Bahamas. In this instance, the burrows were probably made by callianassid shrimp, otherwise known as “ghost shrimp,” and are preserved in what was a sandy patch next to a once-thriving reef from 125,000 years ago. (Photograph by Anthony Martin.)

Interestingly, during this conversation we also touched on on how art and science work together, and I was pleasantly surprised to find out that Dr. Martin is a talented artist, too. It turns out he has illustrated many of his articles with exquisite line drawings of his beloved subjects, marine crustaceans. Yes, I realize that some artists like to draw a line (get it?) between being an “artist” and an “illustrator,” with the latter being held in some sort of disdain for merely “copying” what is seen in nature. If you’re one of those, sorry, I don’t have the time or inclination to argue about this with you. (Now go back to putting a red dot on a white canvas and leave us alone.)

Cover art of branchiopod Lepidurus packardi from California, drawn by Joel W. Martin, for An Updated Classification of the Recent Crustacea, also co-authored by Joel W. Martin and George E. Davis: No. 39, Science Series, Natural History Museum of Los Angeles County, Los Angeles, California.

During our discussion in my office, I pointed out a enlarged reproduction of a drawing of mine depicting the burrow complex of an Atlantic mud crab (Panopeus herbstii). He immediately recognized it as a crustacean burrow, for which I was glad, because it is an illustration of just that in my upcoming book, Life Traces of the Georgia Coast.

Burrow complex made by Atlantic mud crab (Panopeus herbstii), originally credited to a snapping shrimp (Alpheus heterochaelis). Scale = 5 cm (2 in). (Illustration by Anthony Martin, based on epoxy resin cast figured by Basan and Frey (1977).

After his campus visit, though, I realized that an even more appropriate artistic work to have shown him was the following one made for the Selections art exhibit last fall, titled Descent with Modification. This title in honor of the phrase used by Charles Darwin to describe the evolutionary process, but also is a play on words connecting to the evolution of burrowing crustaceans.

Descent with Modification again, but this time look at it as an evolutionary chart, where the burrow junctions in the burrow system reflect divergence points (nodes) from common ancestors. For example, from left to right, the ghost shrimp is more closely related to a mud shrimp, and both of these are more closely related to the ghost crab (middle) than they are to the lobster and freshwater crayfish (right). The main vertical burrow shaft represents their common ancestry from a “first decapod,” which may have been as far back as the Ordovician Period, about 450 million years ago.

The image shows five burrowing crustaceans, or to be more specific, ten-legged crustaceans called decapods. Along with these is a structure, which has a burrow entrance surrounded by a conical mound of excavated and pelleted sediment, a vertical shaft connecting to the main burrow network, and branching tunnels that lead to terminal chambers. A burrowing crustacean occupies each chamber, and these are, from left to right: a ghost shrimp (Callichirus major), a mud shrimp (Upogebia pusilla), a ghost crab (Ocypode quadrata), a marine lobster (Homarus gammarus), and a freshwater crayfish (Procambarus clarkii).

Here’s the cool part (or at least I think so): this burrow system also serves as an evolutionary chart – kind of a cladogram – depicting the ancestral relationships of these modern burrowing decapods. For example, lobsters and crayfish are more closely related to one another (share a more recent common ancestor) than lobsters are related to crabs. Mud shrimp are more closely related to crabs than ghost shrimp. Accordingly, the burrow junctions show where these decapod lineages diverged. So the title of the artwork is a double entendre with reference to Darwin’s phrase describing evolution as a process of “descent with modification,” along with burrowing decapods undergoing change through time as they descend in the sediment.

Modern decapod burrows and trace fossils of probable decapod burrows support both the science and the artwork, too. Here are a few examples to whet your ichnological and aesthetic appetites:

Thalassinoides, a trace fossil of horizontally oriented and branching burrow systems made by decapods in Early Cretaceous rocks (about 115 mya) of Victoria, Australia. In this case, these burrows were likely by freshwater decapods, such as crayfish, which had probably diverged from a common ancestor with marine lobsters more than 100 million years before then. Scale = 10 cm (4 in). (Photograph by Anthony Martin.)

Thalassinoides again, but this time in limestones formed originally in marine environments, from the Miocene of Argentina. Note the convergence in forms of the burrows with those of the freshwater crayfish ones in Australia. Think that might be related to some sort of evolutionary heritage? Scale = 15 cm (6 in). (Photograph by Anthony Martin.)

Horizontally oriented burrow junction of a modern ghost shrimp – probably made by a Carolina ghost shrimp (Callichirus major) – exposed along the shoreline of Sapelo Island, Georgia. Note the pelleted exterior, which is also visible on the burrow networks of the fossil ones in the Bahamas, pictured earlier. So if fossilized, this would be classified as the trace fossil Ophiomorpha nodosa. Scale in centimeters. (Photograph by Anthony Martin.)

Two ghost-shrimp burrow entrances on a beach of Sapelo Island, Georgia, with the one on the right showing evidence of its occupant expelling water from its burrow. No scale, but burrow mound on right is about 5 cm (2 in) wide. (Photograph by Anthony Martin.)

Burrow entrance and conical, pelleted mound made by a freshwater crayfish (probably a species of Procambarus) in the interior of Jekyll Island, Georgia. Scale = 1 cm (0.4 in). (Photograph by Anthony Martin.)

So the take-away message of all of these musings and visual depictions is that evolution, faith, science, art, trace fossils, modern burrows, and burrowing decapods can all co-exist and be celebrated, regardless of whether we sing Kumbaya or not. So let’s stop dividing one another, get out there, and learn.

*I’m also proud to say that my post from October 17, 2011, Georgia Life Traces as Art and Science, was nominated for possible inclusion in Open Laboratory 2013. Thank you!

Further Reading

Basan, P.B., and Frey, R.W. 1977. Actual-palaeontology and neoichnology of salt marshes near Sapelo Island, Georgia. In Crimes, T.P., and Harper, J.C. (editors), Trace Fossils 2. Liverpool, Seel House Press: 41-70.

Martin, A.J. In press. Life Traces of the Georgia Coast: Revealing the Unseen Lives of Plants and Animals. Indiana University Press, Bloomington, IN: 680 p.

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.

Martin, J.W. 2010. The Prism and the Rainbow: A Christian Explains Why Evolution is Not a Threat. Johns Hopkins University Press, Baltimore, MD: 192 p.

Martin, J.W., and Davis. G.E. 2001. An Updated Classification of the Recent Crustacea, No. 39, Science Series, Natural History Museum of Los Angeles County, Los Angeles, California: 132 p.


The Traces We Leave Behind: A Tribute to Jordi Maria de Gibert

Paleontologists have an odd relationship with death. We often joke about how our livelihoods depend on what has died before us, or we experience great delight when we find an exquisite fossil, which probably was buried alive for it to be so well preserved. We also blithely talk about “death assemblages” and happily explain this gruesome term to non-paleontologically inclined students, friends, spouses, and partners without much thought about how it sounds to people outside of our field.

For ichnologists, who mostly study the tracks, burrows, and other vestiges of these lives that preceded us, our perspectives become even more skewed. Once-live animals, through their behavior, made trace fossils. Yet we almost never see what made them. Hence we also spend much of our time among the living, watching them make traces that we can use as analogs for those trace fossils left by their ancestors. Sometimes we find ourselves identifying with modern animals, developing empathy for what they experience as they form traces, a sensitivity that can extend to their trace-fossil equivalents. Hence for ichnologists, these parts of the fossil record become just a bit less removed from death, and we end up feeling for our tracemakers, both long gone and extant.

Jordi Maria de Gibert, contemplating and lamenting the loss of dinosaurian tracemakers from mass extinctions. The window display was in Basel, Switzerland, one of many places visited by Jordi in his quest to learn all things ichnological. (Photograph by Anthony Martin, who is also pictured in the reflection, along with ichnologists Luis Buatois and Gabriela Mángano, taken in 2003.)

In this sense, our small and close-knit international community of ichnologists was shocked to learn about the sudden loss of one of our own “tracemakers” this past weekend, Jordi Maria de Gibert. His death was unexpected and its impact accentuated because he and the rest of us had just gathered together only last month for the International Congress of Ichnology (Ichnia) in St. Johns, Newfoundland. We also anticipated seeing him again in his home city of Barcelona in 2016, where he died on Sunday. None of us had prepared ourselves to reflect on his legacy, let alone contemplate the possibility that his cognitive traces would cease any time soon.

The aftermath of the first Ichnia football match (sometimes known as “soccer” to you Yanks) between ichnologists of Team Gondwana and Team Laurasia, which took place on a pitch near Trelew, Argentina. Jordi, in the middle of the back row, is either signaling “Peace,” “Victory,” or, most likely, ordering two beers: one for him, and one for you so he can sit down to argue about trace fossils with you. (Photograph by Anthony Martin, taken in 2004.)

Most of our dismay about Jordi’s departure is because we loved him as a person, but it is also surely connected to our witnessing an ascendancy cut short. For instance, at the end of the meeting in St. Johns, Jordi addressed all of us as the newly elected president of the International Ichnological Association, and he had volunteered to serve as the main organizer for the next Ichnia meeting four years from now. His larger-than-life personality was on full display during his informal and impromptu speech: enthusiastic, cheerful, witty, earnest. In the days before then, he delivered multiple presentations on ongoing research projects, most of which revolved around his continuing interests in crustaceans and their traces, as well as those of marine bioeroders, animals that make a living by boring into rocks. Jordi was a prolific publisher of peer-reviewed papers on these topics, and was well known for his cooperative spirit, coauthoring with many ichnologists and other types of paleontologists on these papers.

Jordi (right, seated), in his preferred life habit, talking about fossils with colleagues (and friends) at an outcrop. And this wasn’t just any outcrop, but was at Mistaken Point, Newfoundland, which has one of the most spectacular Ediacaran fossil assemblages in the world. This had to have been a dream come true for him, as it was for many of us.

Jordi showing off his “Bama booties,” required footwear for the sacred ground of Mistaken Point, as some other ichnologist vainly attempts to “photobomb” him with his own blue-footed bootie. (Photograph by Ruth Schowalter.)

I had known Jordi since 1995, having first met in Bornholm, Denmark at a small ichnological meeting there. He and I were still new to our discipline (we were about the same age) and quite green, but eager to learn from our elders. As is typical with many academic friendships, over the next 17 years we would see each other at various meetings, and by my count we saw trace fossils and toasted one another in six countries (Denmark, U.S., U.K., Switzerland, Poland, Canada). Each time together, I grew more impressed with his intense and tenacious will to seek out more knowledge, digest it, and pass it on to others. He was a fierce intellectual who relished the debating of ideas, and was never satisfied with a conversation if he did not leave it wiser. This, of course, benefited all who were brave enough (and lucky enough) to enter into such discussions with him.

A happy time at the Ichnia 2004 banquet in Trelew, Argentina, with (from left to right) Renata Guimarães Netto, Jordi, and Ludvig Loewemark, where the exchange of ideas and good cheer flowed nearly as fast as the wine.

Jordi was young as far as ichnologists go, and as I argued in my previous post, the best ichnologists are the most experienced ones. So he knew as well as any of us that gaining more experience in the field was essential, and traveled to many places and studied traces of all ages – from the Ediacaran to the present – and traces of all kinds, from plant roots to dinosaur tracks. Accordingly, because of his dedication and broad interests, he had already become one of our best. In this vein, one of the metaphorical jokes ichnologists tell is how our academic success can be measured by how deeply we can burrow: shallow tiers are the least successful, whereas the deepst tiers are the most successful. Jordi was assuredly well on his way to the deepest tier, but we are all saddened about his unexpectedly reaching the historical layer before so many of us.

Los quatros amigos, posing happily toward the end of an ichnology field trip in Switzerland in 2003: from left to right, ichnologists Noelia Carmona, Gabriela Mángano, Luis Buatois, and Jordi, sporting some distinctive headgear and proudly flouting conformity. (Photograph by Anthony Martin.)

I learned about Jordi’s death on Sunday through our mutual ichnologist friend, Renata Guimarães Netto, who had likewise known Jordi for more nearly 20 years. Quickly the word spread through social media, e-mails, and phone calls, our sadness multiplying and magnifying worldwide. Only last month, we had celebrated with him, and now we mourned him, and expressed our sorrow to his family members, and close friends.

To ease some of this pain and enjoy an otherwise beautiful Sunday in Decatur, Georgia, my wife Ruth and I went for a walk. Without thinking, I suggested that we meander in one of the largest, quietest green spaces in Decatur, which turned out to be its cemetery. (Yes, I know. All I can say is that the subconscious is more powerful than we know.) While we strolled, I thought about times spent with Jordi on field trips and in conferences, while also recalling papers he had written and discoveries he had made. As mentioned earlier,  Jordi’s interests were varied, but perhaps his favorite research topic was crustacean burrows, especially the burrows of crabs, shrimp, lobsters, and other 10-legged crustaceans. Too bad we were nowhere near the Georgia coast, I thought, as it would have been a fitting and comforting homage for Ruth and I to take in the many decapod burrows of the Georgia beaches and salt marshes, which Jordi had never seen in person.

That’s when an eerie coincidence happened. During our walk, we spotted a former pond on the cemetery grounds, now drained for dredging. There’s something about a big pit of mud that appeals to every ichnologist, so I excitedly suggested that we go take a look to see what traces were there. We expected to find lots of tracks, such as those of birds, raccoons, squirrels, and coyotes, and maybe a few other urban fauna. Instead, though, the muddy surface was perforated by decapod tracks and burrows.

Need to see some crustacean traces, but you live in the landlocked part of Georgia? Just go to a dried pond and look for tracks like these.

These were the traces of crayfish, decapods that diverged from a common ancestor to modern lobsters more than 250 million years ago to live in freshwater environments as their brethren dispersed throughout the seas. A few years back, I studied Cretaceous crayfish and their burrows in Australia, but had never seen a live crayfish in its burrow here in Georgia, let alone seen so many of their tracks in one place. We even saw some crayfish (probably a species of Procambarus) poking their heads and claws out of their burrows, or walking around on the mudflat. So it turned out we did not need to go to the Georgia coast after all to see traces reminding us of Jordi: they had been right here with us the whole time.

A crayfish at its burrow entrance in the mudpit now in Decatur Cemetery, either defending its territory, or waving goodbye to people who study its kind and their traces. Your choice, but I know which one I’m picking.

In April, Jordi began writing about ichnology and invertebrate zoology for a more public audience through his cleverly titled blog, Infaunal Epiphany. His first entry was titled First Post, Hope Not Last!, in which he expressed a growing aspiration to connect with more than just his academic colleagues:

We scientists produce science. We scientists consume science. Most of us do that. We do our research, we publish it and other scientists read it. We are keeping all the fun for ourselves!!! It is true that there are scientists, journalists and writers who devote an effort to popularize science results. They are the ones building a bridge to society and I think it is fair to do that as many of us are investigating on public money.

Jordi wanted to share the fun of science, and in that respect, field trips with him were always a delight. These are probably what I will miss most about him, a pang that becomes particularly acute when I realize that one of our last conversations was about his some day visiting the Georgia coast to see its modern traces with me and our like-minded friends.

Lastly, in the light of his most recent life departing us, perhaps Jordi’s most poignant post on his nascent blog was Seven Reasons to Reincarnate as a Cephalopod. I won’t spoil it for anyone who hasn’t read this wonderful piece, but will just say that this post alone showcases how Jordi’s fine sense of humor blended readily with his science.

We will never know whether Jordi’s wish came true, let alone which cephalopod he might have become, or whether some element of his considerable spirit is now somehow connected to one of his beloved crustacean tracemakers or bioeroders in the past or present. But we can be assured that he will continue to live with us through his works and our memories of him. When our ichnological community meets again in his home town of Barcelona four years from now, his traces will all around us, continuing to inspire us to learn and live more.

Salud y un abrazo grande, mi amigo Jordi. Vaya con las trazas.

Correction: Someone pointed out to me that the newly elected International Ichnological Association (IIA) president is actually Alfred Uchman, not Jord. Jordi only seemed presidential to me because of his inspiring report given at Ichnia 2012 as outgoing secretary of the IIA and his agreement to host Ichnia 2016. (I am pleased to report that Alfred likewise gave an excellent speech to those gathered.) Apologies for the mistake, and thanks (as always) to anyone who points them out to me.