The Death Crawl of a Jurassic Crinoid

Think of a crinoid, and you will likely visualize one of these gorgeous echinoderms looking like a colorful, delicate flower on a brightly lit seafloor, aptly justifying its nickname as a “sea lily.” Take your crinoidal fantasy just a bit further, and imagine its fine, feathery arms gently waving in harmony with ocean currents passing through them, its stalk bending with each current, but otherwise staying firmly attached to a sea bottom. If you know a little more about crinoids, though, you might also think of one without a stalk – a “feather star” – swimming above the ocean floor, performing an aquatic dance reminiscent of the Hindu Mother Goddess Durga.

A swimming stalkless crinoid (“feather star”) at the Tanjung Papaya dive site, Mandado Bay, Indonesia, recorded by Pim Van Schendel in October 2015. Notice how its barely touches the sandy bottom, leaving few clues of its behavior in the sediments below.

You might also let your dreams go back to the ancient past, when crinoid “meadows” blanketed shallow-marine environments throughout the world, starting about 450 million years ago in the Ordovician Period and continuing through the Paleozoic Era. By the Carboniferous Period, crinoids were so abundant that their body parts contributed to limestones we now use for buildings. Following the mass extinction at the end of the Paleozoic, crinoids became more rare, but several lineages persisted through a few more mass extinctions, including the living ones that delight us today.

Ordovician-Crinoid-Slab-Cincinnati-MuseumAn Ordovician crinoid “meadow,” buried by a tropical storm about 440 million years ago. Slab is about 2 m (6.6 ft) wide, and more than a hundred exquisitely preserved  are preserved on it. Specimen is at the Cincinnati Museum Center (Cincinnati, Ohio), and was discovered, recovered, and donated by Dan Cooper of The Dry Dredgers fossil club to the museum. (Photograph by Anthony Martin.)

Now, instead of such idyllic reveries, think of a crinoid experiencing a slow, agonizing death. Imagine it imitating the clichéd image of a man crawling through a desert and croaking the word “water” as it pulls itself along a barren and air-filled landscape, searching for a comforting sea. If this is not a jarring enough of a picture for you, don’t worry, it gets worse. The crinoid doesn’t quite make it to the sea, and when it can’t move any further, it tries in vain to attach itself to the land beneath it. Minutes later, it dies. A few hours pass before it is finally submerged (too late) by the next high tide, its body put to rest under a blanket of sediments.

The real twist to this story, though, is how 170 million years later, some upright bipedal primates – at least a few of whom were quite fond of crinoids – spotted this one on a ground surface in the middle of present-day Portugal, still connected to its last trail of life.


The only known trace fossil of a crawling crinoid in the geologic record in a limestone bed of the Chãi des Pias Formation (Middle Jurassic, about 170 mya) near São Bento, Portugal. The red arrow points to the trail, evident here as a shallow, meandering, and slightly darker groove. How do we know a crinoid made it? Because it ends with a crinoid. Its last three decisions were to move to the right, then to the left, and stop forever. (Photograph by Anthony Martin.)

I was lucky enough to see the only known trace fossil of a crawling crinoid and the crinoid that made it during a paleontological field trip last month in Portugal. The trip was connected to the International Ichnological Congress, a once-every-four-year meeting simply known as Ichnia. The field-trip stop with the crinoid and its trail was at a relatively modest outcrop of Middle Jurassic limestone near the Portuguese parish of São Bento (Porto de Mós municipality).

Despite constant rain that day and the small area exposed at the field site, its trace and body fossils grabbed our attention, then held us as willing captives for more than an hour. Among the ichnological treats offered by this Jurassic tidal-flat deposit were crab trackways – including what might be the longest invertebrate trackway in the geologic record – long snail trails, crustacean burrows, fish trails, and much more. For those people who love body fossils, and especially of echinoderms, the rock also held beautifully preserved sea stars and spiny sea urchins. It was marvelous, and for days afterwards, all of the trip participants talked about this place and its bounty.

Given such fossil riches, it is tempting for me to share them all with you here. Nonetheless, I will instead focus on the star: not a sea star, but a relative. Despite the rain, its trail was easy to spot, as it was located inside of a white-yellow strip on the surface of an otherwise dull-gray limestone. The area surrounding the trace fossil had been inadvertently brightened by the researchers studying it. When they made a latex mold of the trace fossil, the latex took some of the weathered surface with it. From my perspective, it looked like a landing strip. Which, in a sense, it was.

Jurassic-Crinoid-Trail-Portugal-1Crinoid coming in for a landing, the hard way. The bright area marks where paleontologists made a latex mold of the crinoid trail, but also. shows how it crawled for more than 2 m (6.6 ft) along the tidal-flat surface. The crinoid is at the far end of the strip. Based on all of the paired shoes with people wearing, one might conclude that the rain was doing a poor job of dampening our enthusiasm. (Photograph by Anthony Martin.)

Jurassic-Crinoid-Trail-Portugal-2A close-up of the first part of the crinoid trail. Once stranded on the tidal flat, it began dragging itself across the originally soft sediments, leaving a groove from its stalk and impressions from its arms (arrow). Scale has centimeters on the left and inches on the right. (Photograph by Anthony Martin.)

Jurassic-Crinoid-Trail-Portugal-3Although this fossilized crinoid trail is underwater here, it wasn’t when the crinoid made it. Exposed on the tidal flat, it tried to get back to the sea by pulling itself forward with its arms, its stalk dragging behind it. Its arms left a wake of disturbance on either side of a thinner central groove from its stalk. (Photograph by Anthony Martin.)

Jurassic-Crinoid-End-of-Trail-PortugalThe end of the trail, which was not a happy one for the crinoid, but ultimately a fulfilling one for paleontologists and ichnologists. (Photograph by Anthony Martin.)

Trace fossils that represent the last moments of an animal’s life – corroborated by a direct association of the animal with its trace – are rare, but known. Such traces, whether modern or fossil, are called mortichnia (mort = “death” and ichnia = “traces”), and this trail with its crinoid maker definitely qualifies as one. Based on this trace fossil and many other geological clues at the outcrop, it was on what was originally a tidal flat, an environment well outside of a crinoid’s comfort zone. It may have been happily filter-feeding offshore, but was uprooted by waves and washed up by a high tide.

Because this is a Jurassic crinoid, you might wonder (as I did) if any modern stalked crinoids can crawl like this, using their arms to drag themselves along a sedimentary surface. The answer is yes, they can. But this is easier for a crinoid to do when underwater, where it is more buoyant. As far as I know, no one has experimented with modern stalked crinoids to see whether they can do this on land, let alone documented any of these animals getting dumped onto a tidal flat and then trying to make it back home.

Footage of a crawling stalked crinoid, albeit one underwater. This one was observed in about 400 m (1,300 ft) deep water off Little Bahama Bank, as reported by Baumiller and Messing (2007). For a detailed analysis of its movement and the traces that would result from this, read their article here.

Although the species of crinoid that made its death crawl is not yet identified, the researchers who studied it concluded it is an isocrinoid. Paleontologists who have studied stalked crinoids figured that the first ones capable of crawling may not have evolved until the Devonian Period, about 350-400 million years ago. Until then, they were sedentary. What changed, giving crinoids good reason to get up and walk away? Probably predation. Predators, which likely included fish but also other echinoderms, must have found crinoids easy and tasty targets, which would have favorably selected for more mobile forms. As Eddie Vedder might say, it’s evolution, baby.

Ordovician-Crinoids-Cincinnati-Museum-1Once Ordovician crinoids settled down, they had no place to go. (Photograph by Anthony Martin, taken at the Cincinnati Museum Center.)

I won’t go through all the details of the report on this crinoid and the other extraordinary trace and body fossils at this site in Portugal. For that, you can read the original research article by Carlos Neto de Carvalho – who was also one of the field-trip leaders – and his colleagues, published earlier this year. All of them deserve to be famous for this extraordinary discovery, and I and my colleagues who were there all felt privileged to have seen it for ourselves, 170 million years after a crinoid went on its final journey.

Many thanks (muito obrigado) to Carlos Neto de Carvalho and Joana Rodrigues for organizing and leading such memorable field trips before and after Ichnia 2016, giving us all an appreciation for the wonderful paleontology and culture of Portugal. For more information, photos, and videos about stalked crinoids, check out Christopher Mah’s excellent post Stalked Crinoid Roundup! and other crinoid-related posts at his appropriately named blog, Echinoblog.)


Baumiller, T.K., and Messing, C. 2007. Stalked crinoid locomotion, and its ecological and evolutionary implications. Palaeontologia Electronica, 10, 2A. (PDF of open-access article here.)

Neto de Carvalho, C., Pereira, B., Klompmaker, A., Baucon, A., Moita, J.A., Pereira, P., Machado, S., Belo, J., Carvalho, J., and Mergulhão. 2016. Running crabs, walking crinoids, grazing gastropods” behavioral diversity and evolutionary implications of the Cabeço da Laderia Lagerstätte (Middle Jurassic, Portugal). Communicações Geológicas 103, Especial 1, 39-54. (PDF of open-acccess article here.)

“Worm Burrows” as a Geological Cliché

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Further Reading

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

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

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

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