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Horseshoe Crabs Are So Much More Awesome Than Mermaids

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Given all of the controversy over a recent cable-TV program, in which its broadcasting channel decided mythical marine animals deserved more air-time than real ones, I thought it was important to highlight one extant animal that never fails to surprise me. This animal’s lineage is more ancient than dinosaurs, reptiles, or even amphibians, with its oldest fossils dating from about 450 million years ago. It is also the largest living marine invertebrate animal you are likely to see on beaches of the eastern U.S. and Gulf Coast. And at this time of year, if you see it crawling around on a beach, it’s because of sex. For the past month or so, this animal has been participating in massive orgies. Pictures of this gamete-laden frenzy somehow made it past prudish censors of Facebook and other social-media sites, titillating prurient invertebrate enthusiasts everywhere and filling them with cockle-warming glee.

Juvenile-Limulid-SapeloBehold, a fine juvenile specimen of the Atlantic horseshoe crab (Limulus polyphemus)! Although it lives in the ocean, it can walk on land for hours, like some sort of reverse Aquaman, but totally cooler than him. And some day, if this one lives long enough, it will use those legs to walk on land again, but in pursuit of sex. Sounds to me like this animal deserves its own planet. (Photograph by Anthony Martin, taken on Sapelo Island, Georgia.)

As you already know from reading the title of this post, I’m talking about horseshoe crabs. More properly known as limulids by real marine biologists and paleontologists, these ultra-cool, über-hip, but totally retro critters are more closely related to spiders than they are to true crabs, but their common name is so, well, common, that scientists just sigh and begrudgingly go along with it for the sake of public communication.

Modern limulids are represented by four species, three of which are in Asia, but the grandest of them all is the Atlantic horseshoe crab, Limulus polyphemus. This species is at its largest here in Georgia, which may be a function of the Georgia Bight, an extensive offshore shelf that affords more food and habitat than other areas. How big? I’ve seen some as long as 70 cm (27 in) – tail included – and 40 cm (16 in) wide, big enough to scare both of our cats at home. They grow to these sizes after hatching as little limulids not much bigger than the period on this sentence, an astonishing increase in mass if they make it to adulthood (which most don’t).

Baby-Limulid-TrailThe circuitous trail of a baby limulid, made on a sandflat at low tide. Its body width can be estimated by the width of the interior of the trail, and its body length was slightly more than that, meaning it was smaller than my fingernail. See that central groove? That’s from its tail, but if you want to impress your friends, call it a telson. (Photograph by Anthony Martin, taken on Sapelo Island, Georgia.)

Horseshoe crabs are so astounding that I could go on endlessly about all sorts of facts about them. Fortunately for you, gentle reader, other folks have written entire books about them and heaps of popular and scientific articles. (For starters, try going here.) So I don’t want to needlessly duplicate what others have done, and done well. Instead, I’ll focus on my main interest in these animals – their traces – and will regale you with tales of the traces they can make with their tails.

Horseshoe crab tails are spiky projections called telsons. Based on lots of the traces I’ve seen on the Georgia coast and a few direct observations, the main function of a telson is to help a horseshoe crab to get back on its feet after being knocked onto its back. That is, whenever a limulid is upside-down, it immediately start using its telson as a sort of sideways pole vault to lever itself into a less vulnerable position.

Without a telson, an upside-down horseshoe crab is stuck; its legs run furiously, but to no avail. However, with a telson, it can put the pointy end into the sand or mud underneath its body, and push itself up from a surface. This gives a limulid a fighting chance to get back to where it once belonged and start walking. This strategy works best if it turns to its right or left side, as limulids are longer than wide. They may be wonders of nature, but they’re not doing back flips or somersaults.

Limulid-Telson-Windshield-Wiper-TraceA large adult horseshoe crab that was right-side-up when trying to get back to the sea, got tired, and tried to use its telson to move itself along. In this instance, it didn’t work, but the traces made by the telson show its range of motion, working like a windshield wiper. (Photograph by Anthony Martin, taken on Sapelo Island, Georgia.)

OK, all of the preceding information I already knew. After all, I have: coauthored an edited book chapter about juvenile limulid traces and their close resemblance to trace fossils made by trilobites; coauthored another article on the history of limulid-trace studies (which go back to the 1930s!) that’s now in review; and devoted a lengthy section of a chapter in my book to limulids as tracemakers. So you could say I’ve been feeling pretty cocky about what I knew about these animals as tracemakers. That is, until one horseshoe crab showed me how much I still need to learn about them and what they can make.

The humility-inspiring traces showed up in a photo on a Facebook page I follow (and so should you), the St. Catherines Island Sea Turtle Conservation Program. The program organizers – Gale Bishop and Robert (Kelly) Vance – regularly add photo albums showing sea turtle traces (trackways, body pits, nests), and otherwise report on other facets of natural history they observe on St. Catherines Island beaches. As a result, I live vicariously through these pictures while marooned in the metro-Atlanta area. But they also like to throw me ichnological stunners once in a while, such as the following photo that Kelly posted last week.

Limulid-Telson-Trace-1Who needs made-up animals on TV when traces like these, made by awesome invertebrates like horseshoe crabs, turn up on a Georgia beach? (Photograph by Robert Kelly Vance, taken on St. Catherines Island, Georgia; scale is about 15 cm (6 in) long.)

Kelly found these traces while patrolling the beaches of St. Catherines Island for other traces, namely those of expectant mother sea turtles. Although these distracted briefly from his mission, I was very happy he stopped to document these, as I had never seen anything like them, despite much looking at traces on Georgia beaches.

The holes in the sand, defining a nearly perfect circle, were made by the telson of an adult horseshoe crab that kept on trying to right itself after landing on its back. Each puncture mark shows where it inserted the telson into the sand and then pushed itself up and to its side. Based on the number of holes, direction of sand flung out of each hole, and little “commas” made by extraction of the telson, it tried to flip itself a minimum of 16 times, and all to the right. These separate actions culminated in a 360° clockwise rotation of its body. Also check out the central depression with smaller drag marks; this is where its head shield was in contact with the sand. To imagine the movement represented by these traces, think of a horseshoe crab doing a slow-motion, step-by-step, break-dance backspin.

Seeing the evidence for such persistence was wow-inducing in itself, but in my ichnologically influenced euphoria, I figured the limulid finally succeeded in righting itself. After all, the trackway just to the left of the trace, indicates where it walked away from the scene of its gravitationally challenged situation.

But then I realized there was no “impact mark.” This large horseshoe crab flipping itself onto the sandy surface should have registered an outline of its body before it started walking. Instead, the place where it started walking showed no such impression, meaning it must have made a soft landing, with only its legs and telson digging into the sand. What happened? Did it use mind over matter and levitate itself through telekinesis? Or was it gently picked up and placed on its feet by a merciful mermaid? (Or merman: let’s make sure we’re being inclusive when talking about made-up stuff.)

It turned out that Kelly was the dues ex machina that entered this limulid’s drama, providing divine intervention just when it was needed. When I expressed my puzzlement to Kelly about how this large arthropod finally turned itself over, he confessed to saving it, in which he lifted it and put it back on its feet, where it promptly walked away in a series of tight spirals. The spiraling is something I’ve seen before in their tracks, a method used to find the downslope direction, which normally leads horseshoe crabs to the low-tide mark and the comfort of a watery environment.

Limulid-Telson-Trace-2Another perspective of the “escape” traces made by the limulid’s telson (background), but this time with its tracks, showing how it started spiraling clockwise in an attempt to make its way back to the sea. Check out those telson drag marks in the trackway, doing a little bit of back-and-forth movement as its owner walked. (Photograph by Robert Kelly Vance, taken on St. Catherines Island, Georgia.)

Limulid-Telson-Trace-3OK everyone, start singing “Born Free!” The spiraling helped this limulid (arrow) to find a downslope direction, which took it in the right direction to the sea. But it’s not all sunshine and lollipops for other limulids, some of which are visible in the background, and look like they’re still stuck. Given the tidal range on the Georgia coast – 2.5-3 m (8.2-9.8 ft) – strong wave energy, and wide beaches, lots of big limulids that come in with the flood tide get knocked onto their backs by waves and left behind. It’s almost as if some sort of natural selection is taking place, and something similar might have happened in the geologic past, affecting the evolution of its lineage. (Photograph by Robert Kelly Vance, taken on St. Catherines Island, Georgia.)

In the last photograph, I was glad to see how the story told by these traces promised a happy ending for this limulid that had so stubbornly tried to put itself back on its feet. Yet when you also notice how many of its compatriots did not make it back into the life-nourishing sea, it also serves as a sobering reminder that storybook endings don’t always happen in nature, and what we wish to be true sometimes isn’t.

In this instance, I don’t know whether this horseshoe crab made it back into the sea to live another day or not. Still, the lesson it left for us in the sand lives on, and I am now slightly more confident that if any limulids were stuck on their backs at any point in their 450-million-year history, made similar traces with their tails, and these marks were preserved as trace fossils, we just might recognize them for what they are. For that alone, I am grateful. Thank you, horseshoe crabs, for being real, making traces, and continuing to share this planet with us today.

(Acknowledgement: Special thanks to Drs. Robert Kelly Vance and Gale Bishop for being my ichno-scouts on St. Catherines Island, and feeding my mind with such tasty treats while I am landlocked.)

Further Reading

Brockmann, H.J. 1990. Mating behavior of horseshoe crabs, Limulus polyphemus. Behaviour, 114: 206-220.

Martin, A.J. Life Traces of the Georgia Coast. Indiana University Press, Bloomington, Indiana, 692 p.

Martin, A.J., and Rindsberg, A.K. 2007. Arthropod tracemakers of Nereites? Neoichnological observations of juvenile limulids and their paleoichnological applications. In Miller, W.M., III (editor), Trace Fossils: Concepts, Problems, Prospects, Elsevier, Amsterdam: 478-491.

Shuster, C.N., Jr., Barlow, P.B., and Brockmann, H.J. (editors). 2003. The American Horseshoe Crab. Harvard University Press, Cambridge, Massachusetts: 427 p.

A Mirror Less Distant in Time

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(This post is the third in a series about my recent field experiences in Newfoundland, Canada in association with the International Congress on Ichnology meeting (Ichnia 2012) in August, 2012. The first dealt with the unusualness of the Ediacaran Period and the second was about the transition from the Ediacaran to the Cambrian Period for burrowing animals.)

The Ordovician Period, a time represented by rocks from 488-443 million years ago, is an old (and I mean, really old) friend of mine. In my master’s thesis, I studied Ordovician fossils from southwestern Ohio, and for my Ph.D. dissertation, I described and interpreted Ordovician trace fossils and strata in Georgia and Tennessee. Thus for the formative years of my academic career, the Ordovician had a strong presence in my life, overshadowing most other geologically inspired opportunities in my adopted home state of Georgia.

Nice outcrop, eh? It’s composed of Lower Ordovician sedimentary rocks, formed more than 450 million years ago, and is on Bell Island, just offshore from St. Johns, Newfoundland (Canada). It’s a place I had never visited before last month, but its trace fossils took me back to Georgia. How? Guess you’ll have to read some more to find out. (Photograph by Anthony Martin.)

This Ordovician-dominated worldview contrasted with a much later focus on the present-day Georgia barrier islands. Between when I first arrived in Georgia, in 1985 through 1998, my only foray to its coast was a three-day field trip as a graduate student to Sapelo Island in 1988. Fortunately, I’ve been a more regular visitor to Sapelo and other Georgia barrier islands throughout the past 14 years or so, and my geologic perspective has accordingly traveled more than 400 million years forward to study modern plant and animal traces.

However, as I’ve embraced the present and the lessons it offers, what also happened over those years was a personal distancing from the Ordovician. This separation was unfortunate for several reasons. One is that Ordovician body and trace fossils are a mere 1.5-2 hour drive from where I live in the metropolitan Atlanta area, just south of Chattanooga, Tennessee. In contrast, the Georgia coast takes a minimum of four hours to reach by car.

Granted, northwest Georgia was part of my dissertation field area, so my leaving behind a place already prospected, poked, prodded, and otherwise inspected thoroughly more than 20 years ago is understandable and forgivable. Yet a day trip there with a colleague last spring (March 2011), along with a recent field trip to view Ordovician rocks in Newfoundland, Canada last month, reminded me of what was in my geological backyard, while also provoking new thoughts about the intersections between the Ordovician and the Georgia coast.

So what happened during those 20+ years of not studying the Ordovician rocks close to me in Georgia? Well, I gained lots more experience, went to many places with rocks and trace fossils of varying ages, and thus – I like to think – became a better ichnologist. So that leads to an imperiously pronounced statement, so please read it, take it in, and revel in its truth: Ichnology is a skill-based science.

People who study the earth sciences have an old saying, often stated during field trips to students: “The best geologist is the one who’s seen the most rocks.” The same sentiment might be applied to ichnologists. To excel as an ichnologist, it’s not your publication record (let alone impact factors of journals publishing your work), the number or size of your grants, accolades of your peers, “big-idea” review papers, erudite tomes, or any number of trappings imposed by academia that matter. What really matters in becoming a better ichnologist is how many traces you’ve seen, measured, sketched, journaled, photographed, pondered, argued over, and folded into your consciousness.

Hey, look – it’s ichnologists, trying to learn more by studying trace fossils in the field! (Photograph by Ruth Schowalter, taken on Bell Island, Newfoundland, Canada.)

Sure, peer review from your colleagues is still an important part of this learning process. Otherwise, as a tracking instructor once told me and other nascent trackers, “When you always track by yourself, you’re always right.” You don’t want to be that ichnologist who gets things wrong, then insists every other ichnologist is wrong, while also imagining that they’re teeming with jealousy over your brilliance. You know, the “they laughed at Galileo, too” fallacy.

Behold my genius! Only I can clearly see these are the tracks of an eight-legged river otter. Oh, so you think they’re from two four-legged otters, with one following the other? Dolt! Don’t you know who I am?

So am I the best ichnologist? Not just no, but hell no. The acknowledged master of ichnology is Dolf Seilacher. And the main reason I enthusiastically bestow Dr. Seilacher with a crown of back-filled and spreiten-laden burrows is because of the extraordinary amount of experience he has as an ichnologist. Granted, he’s also done all of that academic-type stuff that persuades far less-accomplished members of tenure-review committees to nod their heads with utmost seriousness and say, “Well, I suppose we can make an exception in this case.” But he also has seen, measured, sketched, journaled, photographed, pondered, argued over many, many trace fossils during his 87 years on this planet. Dolf knows traces.

Dolf Seilacher, the widely hailed master of ichnology in the world. Even when he’s wrong, he’s really good at it. (Photograph by Anthony Martin, taken in Krakow, Poland.)

So let’s go back to the Ordovician, and how it relates to Dolf and my claim about the importance of experience in ichnology. In 1997, I invited Dolf to visit Emory University as a distinguished speaker in an evolutionary biology lecture series we had then (since gone defunct, like many things at Emory). Because he had never before visited Georgia, he insisted that we also arrange a field trip for him to see some trace fossils here. So with my friend and colleague, Andy Rindsberg, we organized a day trip to an outcrop near Ringgold, Georgia to look at the Ordovician trace fossils there. Andy had done his master’s thesis on the Ordovician and Silurian trace fossils in that area, and as mentioned earlier, I had done my Ph.D. dissertation about the Ordovician rocks, in which I interpreted them mostly through an ichnological lens.

Dolf Seilacher in Georgia (USA) for the first time in November 1997, coffee in one hand and a trilobite burrow in the other. See all of those Ordovician rocks in the background? Even though he’d never been there before, he noticed trace fossils in them in less time than most of us take to read a Huffington Post headline. Gee, you think it had anything to do with his experience? (Photograph by Anthony Martin, taken near Ringgold, Georgia. And just so you know, no paleontologists were “Dolfed” in this photo.)

Andy and I knew the rocks and their trace fossils at this outcrop better than anyone in the world. Yet within five minutes of arriving at the outcrop, Dolf laid his hand on a large slab of Ordovician rock and began talking matter-of-factly about the trilobite burrows in it. Andy and I looked at each other, and said (almost simultaneously), “Trilobite burrows?”

Dolf was right. This rock and many others there were filled with circular, back-filled burrows, which were made by small trilobites that burrowed into mudflats more than 400 million years ago. During a futile attempt to disprove him the following year, Andy and I  found these burrows connected to trackways, and one even ended in a resting trace, perfectly outlining the body of a small trilobite. (Did I mention Dolf was right?)

Burrow (upper right, circular structure) connected to tracks made by little legs from a little trilobite. Trace fossils are on the bottom of a sandstone from the Upper Ordovician Sequatchie Formation of northwest Georgia. Scale in centimeters. (Photograph by Anthony Martin.)

Later on that same day, we looked more carefully at some other fossil burrows at the outcrop. These broad, banana-shaped trace fossils were ones that Andy and I had noted in our respective studies, called Trichophycus. Dolf continued his trilobite–tracemaker theme, insisting that these were also trilobite burrows. This idea was supported by scratchmarks on the burrow walls, which linked the burrows to the small legs of whichever arthropod lived in the burrows. Again, trilobites made sense as the tracemakers, and we haven’t yet found a reason why this would be wrong.

Trusted field assistant Paleontologist Barbie, pointing to a cluster of Trichophycus (interpreted as trilobite burrows) in the Sequatchie Formation of northwest Georgia. She is pointing to some scratchmarks on the burrow walls, which are preserved in natural casts of the burrows. (Photograph by Anthony Martin.)

Almost 13 years later, in March 2011, Andy and I went back to this same Ringgold outcrop to re-study the trace fossils there, done in preparation for a presentation he gave the next month at a regional Geological Society of America meeting (abstract here). He and I were surprised at how much the outcrop had changed since we last visited. Vegetation, particularly of the thorny variety, covered the ground and impeded our progress. Nonetheless, we found many excellent examples of trilobite burrows (Trichophycus), a beautiful trilobite resting trace (Rusophycus), and, for the first time for either of us, a sea-star resting trace.

Resting trace of a trilobite (Rusophycus), with a small part of its trackway leading to the trace, in the Upper Ordovician Sequatchie Formation of northwest Georgia. These trace fossils are preserved as natural casts on the bottom of a sandstone, so you’re seeing the underside of where the trilobite hunkered down more than 400 million years ago. (Photograph by Anthony Martin.)

Resting trace of a sea star (Asteriacites) in the Upper Ordovician Sequatchie Formation of northwest Georgia. This trace fossil, like that of the trilobite resting trace, is also preserved as natural casts on the bottom of a sandstone, so you’re looking underneath where the sea star moved into the mud. (Photograph by Anthony Martin.)

Our discovery of the latter two trace fossils – the trilobite and sea-star resting traces – took me from the Ordovician to the Georgia coast and back again. Throughout the late 1980s, I recall my Ph.D. advisor, Robert (“Bob”) Frey placing many of his articles in my graduate-student mailbox, all of which dealt with the traces of the modern Georgia coast. That’s odd, I thought. What did the traces of the modern Georgia coast have to do with these 440-million-year-old rocks?

In my limited worldview at the time, I did not see that the Georgia barrier islands and their traces composed a mirror, however removed by time, for looking into that Ordovician past. But eventually, given enough articles read, field work done, and trace fossils examined at these Ordovician outcrops, I slowly realized these 440-million-year-old rocks had been formed in estuaries, similar to those along the Georgia coast. When I first published an article about these rocks and their trace fossils in 1993 (link here), these strata represented the oldest known estuary deposits in the world, and some of the trace fossils could be readily compared to those on the Georgia coast. The beauty of this realization was that Frey, a master ichnologist in his own right and a contemporary of Seilacher, had allowed me to discover it for myself: he just provided the clues.

Remember that small, circular trilobite burrow with tracks connecting to it? Now compare it to the same sort of traces made by a modern beach mole crab (Albunea paretii), which left its burrow on the right, walked to the left, and is here rapidly burying itself in the sand. Scale in centimeters. (Photograph by Anthony Martin, taken on Sapelo Island, Georgia.)

Resting trace and attached trackway of a juvenile horseshoe crab (or limulid, specifically Limulus polyphemus). So think about a similarly sized trilobite making this, and what the bottom of the trace would like like, then compare it to the Ordovician trilobite resting trace fossil shown earlier. Scale in centimeters. (Photograph by Anthony Martin, taken on Sapelo Island, Georgia.)

Resting trace of a lined sea star (Luidia clathrata), with the original tracemaker just below its trace. This sea star was stuck above the high tide mark, burrowed into the underlying moist sand, but then tried to move to a better place once its spot started to dry out. Now compare this resting trace to the Ordovician trace fossil shown before. No scale, but sea star is about 8-10 cm wide. (Photograph by Anthony Martin, taken on Sapelo Island, Georgia.)

The following year and only a month ago (August 2012), Andy and I had another Ordovician learning opportunity presented to us, but this time in Newfoundland, Canada. A day trip to see Ordovician rocks and trace fossils on Bell Island, just a 30-minute ferry ride from St. Johns, Newfoundland, was a welcome break from the butt-numbing sessions of the previous two days of the Ichnia 2012 conference at Memorial University.

In our first few minutes at the outcrop and its numerous boulders – spoil piles from an iron-ore mine – we realized that one of the dislodged slabs in front of me was loaded with specimens of Trichophycus. It was a pleasant surprise to get reacquainted with this trace fossil, and in a place far away both geographically and experientially from Georgia.

Multiple specimens of Trichophycus in Lower Ordovician rocks of Newfoundland, Canada, preserved as natural casts of the burrows. See all of those scratchmarks on the burrow walls? These were also made by trilobites, but probably different ones from those in Georgia. Scale in centimeters (and that ain’t no real maple leaf.) (Photograph by Anthony Martin.)

Multiple specimens of Trichophycus in the Upper Ordovician Sequatchie Formation of Georgia, USA, also preserved as natural casts of the burrows and showing some scratchmarks on the walls. Do they look familiar to you, too? If so, welcome to the Ordovician. (Photograph by Anthony Martin.)

Here’s that trilobite resting trace (Rusophycus) from Georgia that I showed earlier. Now take a gander at the one below…

Why, that seems to be a trilobite resting trace (Rusophycus), too, but in Lower Ordovician rocks of Newfoundland. Surprise, surprise, surprise! Scale in centimeters. (Photograph by Anthony Martin.)

Suddenly, much of Andy’s and my previous experience with the Ordovician rocks of Georgia came back to us. We were, paradoxically, home, only in this instance, “home” was a time, not a place. Ichnological colleagues who had no idea Andy and I had worked with Ordovician trace fossils stared at us quizzically (and skeptically) as we excitedly discussed the burrows. But once we informed them that we had seen these trace fossils before, our experience was recognized, egos were set aside, and learning was enhanced. Funny how that works sometimes.

So with our trip to Newfoundland, we went from the alien world of the Ediacaran Period, with its trace fossils unlike anything I had seen before, to the more familiar and accommodating Ordovician Period rocks and their trace fossils. What I learned from this trip, combined with many others to Ordovician rocks elsewhere, as well as the modern sediments of the Georgia coast, was that the mirror was not so foggy after all, and that more field experiences can only further clarify these connections between life traces from the present and the not-so-distant past.

Further Reading

Buatois, L.A., Gingras, M.K., MacEachern, J., Mángano, M.G., Zonneveld, J.-P, Pemberton, S.G., Netto, R.G., and Martin, A.J. 2005. Colonization of brackish-water systems through time: Evidence from the trace-fossil record. Palaios, 20: 321-347.

Eldredge, N., 1970. Observations on burrowing behavior in Limulus polyphemus (Chelicerata, Merostomata), with implications on the functional anatomy of trilobites. American Museum Novitates, 2436: 17 p.

Fillion, D. and Pickerill, R.K. 1990. Ichnology of the Lower Ordovician Bell Island and Wabana Groups of eastern Newfoundland. Palaeontographica Canadiana, 7: 1-119.

Martin, A.J. 1993. Semiquantitative and statistical analysis of bioturbate textures, sequatchie formation (upper ordovician), Georgia and Tennessee, USA. Ichnos, 2: 117-136.

Seilacher, A. 2007. Trace Fossil Analysis. Springer, Berlin: 240 p.