Flight of the Quahogs

Let’s try a science-education experiment. Give a child a live clam and ask, “Can this animal fly?” and I predict her or his answer – accompanied by much giggling – will be “No!’ But if you ask, “Can you fly?”, the answer may change, especially if this child has already flown on an aircraft. So of course humans can fly, but to do this, they require machines, paragliders, or other technological aids in order to move through the air and – this is important – arrive on the ground safely.

Shattered-Quahogs-Pier-Jekyll-IslandFor clams that try to fly, they end up with more than shattered dreams. How did these clams (Mercenaria mercenaria, also known as quahogs or “hard clams”) end up doing Humpty-Dumpty impressions on a wooden pier? Please read on. (Photograph by Anthony Martin, taken on Jekyll Island, Georgia.)

In a similar way, clams can fly. They just need a little help from other animals that can fly and willingly give them a temporary lift from the earth they and their molluscan relatives have known for all of their evolutionary history. Compared to most of our forays into the air, though, these flights are much more limited. Clam aerial exploits are brief and mostly vertical, with little time for them to appreciate the view from above or otherwise experience unusual sensations. They go up, then they come down, and fast.

Clams do not have landing gear. So they can hit the ground hard, especially if their free fall happened after a lengthy trip up into the air and the ground surface is hard: think of a sandflat at low tide, a paved parking lot, or a wooden boardwalk. A a result, the most common end to clam flights is a shattered shell, which is quickly followed by the demise of the clam as it is consumed by the very same animal that bestowed it with flight, however brief and self-serving.

Impact-Trace-Seagull-Clam-DropTraces of a unidirectional vertically oriented clam flight (otherwise known as “falling”) that did not end well for the clam, but worked perfectly for the flying animal that took it for a ride. Notice the impact trace on the hard sandflat, outlining the ribbed shell of the clam (probably Dinocardium robustum) and bits of shell. Most of the probably-still-alive-but-definitely-dying animal  was dragged off to a nearby spot so that its soft parts could be eaten by the same perpetrator that took it for a ride. (Photograph by Anthony Martin, taken on Sapelo Island, Georgia.)

So just what flying animals do such dastardly deeds, taking hapless clams up for a ride, only to drop them to a certain death? By now the gentle reader has probably figured out birds are responsible for this blatant bivalvicide, and some may have already known that seagulls are the most likely culprits. In some coastal areas and during low tides, some seagulls fly over exposed sandflats and mudflats, searching for the outlines of clams buried below the surface. These avian ichnologists then swoop down, land, pick up the clam with their beaks, take off, and then once high enough, they drop them, serving up instant raw clam on the half (or quarter, or eighth) shell. Typically all that is left is a jigsaw puzzle of clamshell pieces and the seagull perpetrator’s footprints, but with the latter only evident on muddy or sandy surfaces amenable to preserving tracks.

Seagull-Tracks-Eaten-ClamIchnological evidence of who killed the clam, provided by the tracks a laughing gull (Larus altricilla).The other half of the shell was broken by its falling onto the sandflat elsewhere, then the gull carried its clam on the half-shell to a more scenic place for its meal. (Photo by Anthony Martin, taken on Little St. Simons Island, Georgia.)

I found this behavior so compelling that I started my book Life Traces of the Georgia Coast (2013) with a story about a laughing gull (Larus altricilla) and the traces of its unwitnessed predation on an Atlantic cockle (Dinocardium robustum), seagull behavior on the Georgia coast. I was not the first person to note this method of clam-smashing by seagulls, as it has been documented by other scientists in parts of the U.S. and abroad, and has been caught on video. Amazingly, though, despite more than 15 years of visiting the Georgia coast, I had never actually witnessed seagulls dropping clams. instead I had only performed post-mortem forensics, in which I would find broken clamshells on hard sandflats accompanied by seagull tracks, telling tales of murder most fowl.

Video footage of a western gull (Larus occidentalis) picking up a clam, flying up about 10 meters (> 30 feet), and dropping it onto rocks to crack it open. After this doesn’t work the first time – and after shooing away a potential clam-stealing rival – it tries again, and is presumably successful. It’s almost as if this gull is using a scientific methodology, isn’t it? (The videographer is only credited as ‘Trisera’ on the YouTube page, and I don’t know where it was filmed, but suppose it’s on the western coast of the U.S.)

Seagull-Cockle-Predation-DiagramHere’s the first illustration a reader will see in my book, Life Traces of the Georgia Coast (2013, Indiana University Press), which I drew to provide a visual forensic analysis of how an Atlantic cockle met its demise at the hands of – er, I mean, wings and bill of – a laughing gull. Part (a) depicts the gull landing after recognizing the outline of the cockle from the air, stopping, and extracting it from the sandflat. Part (b) shows where the cockle was dropped and broken successfully, accompanied by the gull landing and trampling the area as it enjoyed its clam dinner.

This meant I was more than overdue to get visual confirmation of gulls killing clams, which was finally granted just a few weeks ago during a recent trip to Jekyll Island (Georgia). It was the day after I had given an invited talk at the annual meeting of The Initiative to Protect Jekyll Island environmental group, and while my wife Ruth and I were relaxing before leaving the island, but of course were also observing whatever nature we could.

In that spirit, and while sitting on a deck on the west side of the island and looking at a mudflat (in between swatting sand gnats), we noticed a seagull flying about 10 meters (>30 feet) above a wooden pier. At one point, it paused its ascent, and we saw an object fall from its mouth and down toward the pier. Thunk! We clearly heard the impact of the object correlate with what we saw, and with much excitement realized that we had just witnessed seagull clam-cracking for the first time.

Jekyll-Island-Mudflat-Dead-Clams A mudflat replete with mud snails (probably Ilyanassa obseleta), grazing away and making gorgeous meandering trails on the western side of Jekyll Island (Georgia). But wait, what are those big white chunks on the same surface?

Dead-Clams-Mudflat-Jekyll-IslandWhy, look at that: hard clams (Mercenaria mercenaria) in an unnatural state, i.e., disarticulated, broken, and dead on the surface of the mudflat. These clams normally burrow into and live under the mud, and usually manage to stay intact if they stay below the surface. The pieces of clams here must have bounced off the wooden pier, which is casting a shadow in the lower right-hand side of the picture. (Both preceding photographs by Anthony Martin and taken on Jekyll Island, Georgia.)

What was most surprising to me about this broken-shell assemblage on the pier was how it was represented only by the hard clam, or quahog (Mercenaria mercenaria). These thick-shelled clams are quite common in sparsely vegetated muddy areas of salt marshes, burrowing into the mud and connecting their siphons to the surface so that they can filter out suspended goodies in the water during high tides. During low tides, however, they become vulnerable to avian predation. Despite being “hidden” in the mud, somehow the seagulls spotted them from the air, landed next to them on the mudflat, and pulled them out of the mud. They then used the nearby pier as an anvil, and the clam’s hard, thick shell unwittingly became its own hammer when they hit the pier after falling from a fatal height.

Shattered-Quahogs-Jekyll-Pier-MartinThe horror, the horror: a clam killing “ground,” thoughtfully supplied by humans for seagulls in the form of a long, hard, wooden pier. (Photograph by Ruth Schowalter and Yours Truly for scale, taken on Jekyll Island, Georgia.)

OK, now it’s time to think about broken clams and deep time. If you found such an assemblage of broken shells of the same species of thick-shelled clams in a geologic deposit, how would you interpret it? Would you think of these broken shells as predation traces, let alone ones made by birds? Which also prompts the question, when did seagulls or other shorebirds start using flight and hard surfaces to open clams? Did it evolve before humans, and if so, was it passed on as a learned behavior over generations as a sort of “seagull culture”?

All of these are good questions paleontologists should ask whenever they look at a concentration of broken fossil bivalves that are all of the same species, and overlapping with the known geologic range of shorebirds. In short, these may not be “just shells,” but evidence of birds using gravity-assisted killing as part of their predation portfolio.

On the 9th Day of Ichnology, My Island Gave to Me: 9 Molluscans Hiding

For today’s entry in the holiday-and-ichnology inspired countdown of Georgia-coast traces, we will move from the maritime forest to the shoreline, where a trace made through the behavior of one species of molluscan – the knobbed whelk (Busycon carica) – influenced the behavior (and hence traces) of another molluscan – the dwarf surf clam (Mulinia lateralis).

These traces then attracted shorebirds, which added their tracks and beak probe marks to the molluscan traces. This is a excellent modern example of how the interaction of one species of animal with a sediment can affect the interactions of other species with that same sediment, leading to their creation of composites traces.

Whelks-Dwarf-Surf-Clams-Burrowing-JekyllSee all of the knobbed whelks (Busycon carica) in this photo? I know, you don’t actually see their shells because they buried themselves, but you see their outlines on this sandy beach surface because of the many dwarf surf clams (Mulinia lateralis) that burrowed around them. Also look for all of the bird tracks and probe marks around the whelks. (Photograph by Anthony Martin, taken on Jekyll Island, Georgia.)

I’ve already written about these composite traces and the ecological story they tell, so for those details, go to this link and this link. But the summary version goes like this:

  • Low tide stranded the whelks and clams on the beach.
  • Whelks used their muscular feet to pull themselves into the still-wet sand to avoid desiccation and predation.
  • Clams took advantage of disturbed (liquified) sand around the whelks and buried themselves, also to avoid predation and desiccation.
  • Shorebirds saw whelk-shaped concentrations of clams, chowed down on them.

The story gets more complicated in places, especially when seagulls decided they also wanted to eat the whelks, but that’s most of it. So next time you’re on a beach and you see a triangular-shaped concentration of small clams, take a second look to see whether there’s a live whelk underneath it: traces begetting traces.

Further Reading

Busycon carica: Knobbed Whelk. Smithsonian Marine Station at Fort Pierce.

Mulinia lateralis: Dwarf Surf Clam. Smithsonian Marine Station at Fort Pierce.

Links to Previous Posts in This Theme

On the 12th Day of Ichnology, My Island Gave to Me: 12 Snails Grazing

On the 11th Day of Ichnology, My Island Gave to Me: 11 Plovers Probing

On the 10th Day of Ichnology, My Island Gave to Me: 10 Beetles Boring

The Paleozoic Diet Plan

Given the truth that the Atlantic horseshoe crab (Limulus polyphemus) is more awesome than any mythical animal on the Georgia coast (with the possible exception of Altmaha-ha, or “Altie”), it’s no wonder that other animals try to steal its power by eating it, its eggs, or its offspring. For instance, horseshoe-crab (limulid) eggs and hatchlings provide so much sustenance for some species of shorebirds – such as red knots (Calidris canutus) and ruddy turnstones (Arenaria interpres) – that they have timed their migration routes to coincide with spawning season.

Ravaged-Limulid-SCISomething hunted down, flipped over, and ate this female horseshoe crab while it was still alive. Who did this, what clues did the killer leave, and how would we interpret a similar scenario from the fossil record? Gee, if only we knew some really cool science that involved the study of traces, such as, like, I don’t know, ichnology. (Photograph by Gale Bishop, taken on St. Catherines Island, Georgia, on May 4, 2013.)

Do land-dwelling birds mammals eat adult horseshoe crabs? Yes, and I’ve seen lots of evidence for this on Georgia beaches, but from only three species: feral hogs (Sus crofa) and vultures (Coragyps atratus and Cathartes aura: black vultures and turkey vultures, respectively). In all of these interactions, no horseshoe-crab tracks were next to their bodies, implying they were already dead when consumed; their bodies were probably moved by tides and waves after death, and later deposited on the beach. This supposition is backed up by vulture tracks. I’ve often seen their landing patterns near the horseshoe-crab bodies, which means they probably sniffed the stench of death while flying overhead, and came down to have an al fresco lunch on the beach.

Nonetheless, what I just described is ichnological evidence of scavenging, not predation. So I was shocked last month when Gale Bishop, while he was monitoring for sea-turtle nests on St. Catherines Island (Georgia), witnessed and thoroughly documented an incident in which a raccoon (Procyon lotor) successfully preyed on a live horseshoe crab. Yes, that’s right: that cute little bandit of the maritime forest, going down to a beach, and totally buying into some Paleozoic diet plan, a passing fad that requires eating animals with lineages extending into the Paleozoic Era.

Limulid-Death-Spiral-SCISo what’s the big deal here? Horseshoe crab comes up on beach, gets lost, spirals around while looking for the ocean, and dies in vain, a victim of its own ocean-finding ineptitude: the end. Nope, wrong ending. For one thing, those horseshoe crab tracks are really fresh, meaning their maker was still very much alive, then next thing it knows, its on its back. Seeing that horseshoe crabs are not well equipped to do back-flips or break dance, I wonder how that happened? (Photograph by Gale Bishop, taken on St. Catherines Island, Georgia, and you can see the date and time for yourself.)

Here is part of the field description Gale recorded, which he graciously shared with me (and now you):

“Female Horseshoe Crab at 31.63324; 81.13244 [latitude-longitude] observed Raccoon feeding on upside-down HSC [horseshoe crab] on south margin of McQueen Inlet NO pig tracks. Relatively fresh HSC track. Did this raccoon flip this HSC?”

Raccoon-Tracks-Pee-Limulid-Eaten-SCIWell, well. Looks like we had a little commotion here. Lots of marks made from this horseshoe crab getting pushed against the beach sand, and by something other than itself. And that “something else” left two calling cards: a urination mark (left, middle) and just above that, two tracks. I can tell you the tracks are from a raccoon, and Gale swears the urination mark is not his. (Photograph by Gale Bishop, taken on St. Catherines Island, Georgia, and on May 4, 2013.)

I first saw these photos posted on a Facebook page maintained by Gale Bishop, the St. Catherines Island Sea Turtle Program (you can join it here). This was one of this comments Gale wrote to go with a photo:

GB: “This HSC must have been flipped by the Raccoon; that was NOT observed but the fresh crawlway indicates the HSC was crawling across the beach and then was flipped – only tracks are Rocky’s!”

[Editor’s note: “Rocky” is the nickname Gale gives to all raccoons, usually applied affectionately just before he prevents them from raiding a sea-turtle nest. And by prevent, I mean permanently.]

My reply to this:

AM: “VERY fresh tracks by the HSC, meaning this was predation by the raccoon, not scavenging.”

In our subsequent discussions on Facebook, Gale agreed with this assessment, said this was the first time he had ever seen a raccoon prey on a horseshoe crab, and I told him that it was the same for me. This was a big deal for us. He’s done more “sand time” on St. Catherines Island beaches than anyone I know (every summer for more than 20 years), and in all my wanderings of the Georgia barrier island beaches, I’ve never come across traces showing any such behavior.

(Yes, that’s right, I know you’re all in shock now, and it’s not that this was our first observance of this phenomenon. Instead, it is that we used Facebook for exchanging scientific information, hypotheses, and testing of those hypotheses. In other words it is not just used for political rants, pictures of cats and food, or political rants about photos of cat food. Which are very likely posted by cats.)

Now, here’s where ichnology is a pretty damned cool science. Gale was on the scene and actually saw the raccoon eating the horseshoe crab. He said it then ran away once it spotted him. (“Uh oh, there’s that upright biped with his boom stick who’s been taking out all of my cousins. Later, dudes!”) And even though I trust him completely as a keen observer, excellent scientist, and a very good ichnologist, I didn’t have to take his word for it. His photos of the traces on that Georgia beach laid out all of the evidence for what he saw, and even what happened before he got there and so rudely interrupted “Rocky” from noshing on horseshoe-crab eggs and innards.

Raccoon-Galloping-Limulid-Death-Spiral-Traces-SCIAnother view of the “death spiral” by the horseshoe crab, which we now know was actually a “life spiral” until a raccoon showed up and updated that status. Where’s the evidence of the raccoon? Look in the middle of the photos for whitish marks, grouped in fours, separated by gaps, and each forming a backwards “C” pattern. Those are raccoon tracks, and it was galloping away from the scene of the crime (toward the viewer).

Raccoon-Galloping-Pattern-SCISo you don’t believe me, and need a close-up of that raccoon gallop pattern? Here you go. Both rear feet are left, both front feet are right, and the direction of movement was to the left; when both rear feet exceed the front, that’s a gallop, folks. Notice the straddle (width of the trackway) is a lot narrower than a typical raccoon trackway, which is what happens when it picks up speed. When it’s waddling more like a little bear, its trackway is a lot wider than this. Conclusion: this raccoon was running for its life.

Although this is the only time Gale has documented a raccoon preying on a horseshoe crab – and it is the first time I’ve ever heard of it – we of course now wonder whether this was an exception, or if it is more common that we previously supposed. The horseshoe crab was a gravid female, and was likely on the beach to lay its eggs. Did the raccoon somehow know this, and sought out this limulid so that – like many shorebirds – it could feast on the eggs, too, along with some of the horseshoe crab itself? Or was it opportunistic, in that it was out looking for sea-turtle eggs, saw the horseshoe crab, and thought it’d try something a little different? In other words, had it learned this from experience, or was it a one-time experiment?

All good questions, but when our data set is actually a datum set (n = 1), there’s not much more we can say about this now. But given this new knowledge, set of search patterns, and altered expectations, we’re more likely to see it again. Oh, and now that you know about this, so can you, gentle reader. Let us know if you see any similar story told on the sands of a Georgia beach.

You want one more reason why this was a very cool discovery? It shows how evolutionary lineages and habitats can collide. Horseshoe crabs are marine arthropods descended from a 450-million-year-old lineage, and likely have been coming up on beaches to spawn all through that time. In contrast, raccoons are relative newcomers, coming from a lineage of land-dwelling mammals (Procyonidae) that, at best, only goes back to Oligocene Epoch, about 25 million years ago. When did a horseshoe crab first go onto land and encounter a land-dwelling raccoon ancestor? Trace fossils might tell us someday, especially now that we know what to look for.

So once again, these life traces provided us with a little more novelty, adding another piece to the natural history of the Georgia coast. Moreover, a raccoon preying on a horseshoe crab was another reminder that even experienced people – like Gale, me, and others who have spent much time on the Georgia barrier islands – still have a lot more to learn. Be humble, keep eyes open, and let the traces teach you something new.

(Acknowledgement: Special thanks to Dr. Gale Bishop for again spotting something ichnologically weird on St. Catherines Island, documenting it, and sharing what he has seen during his many forays there.)

Darwin, Worm Grunters, and Menacing Moles

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Further Reading

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

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

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

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

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

Shorebirds Helping Shorebirds, One Whelk at a Time

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, 417:936938.

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