Spectacular shrimp burrows from the Miocene of Sicily

June 5th, 2013

Siciliancountryside060513NOTO, SICILY, ITALY–The first stop on our International Bryozoology Association field trip today was a newly-opened quarry near Cugni di Rio in the dry southeastern countryside of Sicily, a view of which is above. New quarries are always interesting to geologists — a new view of the Earth’s bones.

OphiomorphaAshcontact060513This is a portion of the quarry wall with the inevitable volcaniclastic unit of ash and marine sediments shown as the greenish layered unit above and below limestones dated as Tortonian (Upper Miocene). On top of the ash you can see what look like tubes sticking out of the brownish layer of sediment.

OphiomorphaSlab1_060513When that brownish layer is exposed as the underside of the bedding plane, it looks like this. These branching features are infilled tunnels made by marine shrimp. The walls of the tubes are ornamented by pellets placed there by the shrimp in their frenetic activity. Combining these pellets with the branching style we can place this trace fossil in the venerable ichnogenus Ophiomorpha.

OphiomorphaSlab2_060513This is a closer view showing the branched galleries and maybe a bit of the pitted surface showing where the pellets were attached. These tunnels are completely filled, so we refer to this preservation as full relief.

I know, I know, I should be recording the bryozoans from this stop, but they were far from photogenic!

Wooster’s Fossil of the Week: A twisty trace fossil (Lower Carboniferous of northern Kentucky)

January 27th, 2013

My Invertebrate Paleontology students know this as Specimen #8 in the trace fossil exercises section: “the big swirly thing”. It is a representative of the ichnogenus Zoophycos Massalongo, 1855. This trace is well known to paleontologists and sedimentologists alike — it is found throughout the rock record from the Lower Cambrian to modern marine deposits. It has a variable form but is generally a set of closely-overlapping burrow systems that produce a horizontal to oblique set of spiraling lobes. It was produced by some worm-shaped organism plunging into the sediments in a repetitive way, gradually making larger and larger downward-directed swirls.

Zoophycos is a useful indicator of ancient depositional conditions. It give its name, in fact, to an ichnofacies — a set of fossils and sediments characterize of a particular environment. In the Paleozoic it is found in shallow water and slope environments; from the Mesozoic on it is known almost entirely from deep-sea sediments. Our specimen is from the Borden Formation and was found amidst turbidite deposits, so it is probably from an ancient slope system.

There has been much debate about the behavior and objectives of the organisms who made Zoophycos. The traditional view is that it was formed by an animal mining the sediment for food particles, a life mode called deposit-feeding. Some workers, though, have suggested it could have been a food cache, a sewage system, and even an agricultural garden of sorts to raise fungi for food. I think in the end the simplest explanatory model is deposit-feeding, although with such a long time range, a variety of behaviors likely produced this trace.

Zoophycos was named in 1855 by the Italian paleobotanist Abramo Bartolommeo Massalongo (1824-1860). Massalongo was a member of the faculty of medicine at the University of Padua, chairing their botany department. (Medicine had broad scope in those days!) Why was he studying this trace fossil? Like most of the early scientists who noticed trace fossils, he thought it was some kind of fossil plant.
Zoophycos villae (Massalongo, 1855, plate 2)


Bromley, R.G. 1991. Zoophycos: strip mine, refuse dump, cache or sewage farm? Lethaia 24: 460-462.

Ekdale, A.A. and Lewis, D.W. 1991. The New Zealand Zoophycos revisited: morphology, ethology and paleoecology. Ichnos 1: 183-194.

Löwemark, L. 2011. Ethological analysis of the trace fossil Zoophycos: Hints from the Arctic Ocean. Lethaia 45: 290–298.

Massalongo, A. 1855. Zoophycos, novum genus Plantarum fossilum, Typis Antonellianis, Veronae, p. 45-52.

Olivero, D. 2003. Early Jurassic to Late Cretaceous evolution of Zoophycos in the French Subalpine Basin (southeastern France). Palaeogeography, Palaeoclimatology, Palaeoecology 192: 59-78.

Osgood, R.G. and Szmuc, E.J. 1972. The trace fossil Zoophycos as an indicator of water depth. Bulletin of American Paleontology 62 (271): 5-22.

Sappenfield, A., Droser, M., Kennedy, M. and Mckenzie, R. 2012. The oldest Zoophycos and implications for Early Cambrian deposit feeding. Geological Magazine 149: 1118-1123.

Wooster’s Fossil of the Week: A bivalve boring from the Upper Ordovician of southern Ohio

December 16th, 2012

This week’s fossil is from close to home. In fact, it sit in my office. The above is a trace fossil named Petroxestes pera. It was produced on a carbonate hardground by a mytilacean bivalve known as Modiolopsis (shown below). Apparently the clam rocked back and forth on this substrate to make a small trench to hold it in place for its filter-feeding. This particular specimen of Petroxestes was found in the Liberty Formation (Upper Ordovician) of Caesar Creek State Park in southern Ohio. This is a place many Wooster paleontology students know well from field trips.
The original Petroxestes was at first known only from the Cincinnatian Group, but now it is known from many other places and time intervals, even including the Cretaceous and Miocene. It is a good lesson about trace fossils. They are defined by their morphology, not what organisms made them. It turns out that this slot-shaped trace can be made by other animals besides Modiolopsis, which went extinct in the Permian.


Jagt, J.W.M., Neumann, C. and Donovan, S.K. 2009. Petroxestes altera, a new bioerosional trace fossil from the upper Maastrichtian (Cretaceous) of northeast Belgium. Bulletin de l’Institut royal des Sciences naturelles de Belgique, Sciences de la Terre 79: 137-145.

Pickerill, R.K., Donovan, S.K. and Portell, R.W. 2001. The bioerosional ichnofossil Petroxestes pera Wilson and Palmer from the Middle Miocene of Carriacou, Lesser Antilles. Caribbean Journal of Science 37: 130-131.

Pojeta Jr., J. and Palmer, T.J. 1976. The origin of rock boring in mytilacean pelecypods. Alcheringa 1: 167-179.

Tapanila, L. and Copper, P. 2002. Endolithic trace fossils in Ordovician-Silurian corals and stromatoporoids, Anticosti Island, eastern Canada. Acta Geologica Hispanica 37: 15–20.

Wilson, M.A. and Palmer, T.J. 1988. Nomenclature of a bivalve boring from the Upper Ordovician of the midwestern United States. Journal of Paleontology 62: 306-308.

Wilson, M.A. and Palmer, T.J. 2006. Patterns and processes in the Ordovician Bioerosion Revolution. Ichnos 13: 109–112.

Wooster’s Fossil of the Week: Birch wood with beetle borings (Oligocene of Oregon)

November 4th, 2012

We may be at the Geological Society of America annual meeting today, but that doesn’t stop Fossil of the Week! This week’s fossil is a beautifully-detailed piece of petrified birch wood (Betula) with tree rings and insect borings throughout. It was found in the Little Butte Formation (Oligocene) of Linn County, Oregon. This rock unit consists of thick tuffs and volcanic breccias representing volcanic mudflows and nuée ardente deposits that buried diverse hardwood forests. This formation is known for its spectacular silicified fossil wood.
The beetle borings, shown in closer view above, are very similar to those bored in birch trees today. There is little work done on the ichnotaxonomy of these trace fossils, so I can’t yet give them a name, but at least we can see typical beetle activity in the twists and turns. The holes are apparently filled with a cemented mix of insect feces and wood fragments called frass, just like we find in modern birch wod today.


Beaulieu, J.D., Hughes, P.W., and Mathiot, R.K. 1974. Environmental geology of western Linn County, Oregon. Oregon Department of Geology and Mineral Industries Bulletin, no. 84, 117 p.

Rozefelds, A.C. and De Baar, M. 1991. Silicified Kalotermitidae (Isoptera) frass in conifer wood from a mid-Tertiary rainforest in central Queensland, Australia. Lethaia 24: 439-442.

Wooster’s Fossils of the Week: Bivalve Borings (Upper Miocene of Spain)

October 28th, 2012

This beautiful object has a complex history. In the center is a gray limestone cobble that eroded from an underwater ridge and rolled free on a shallow coral reef in an area now near Abanilla, southeastern Spain. It was encrusted by a scleractinian coral, which grew thickly all around the cobble because it was turned continually by wave and current action. Larvae of the bivalve Lithophaga landed on the surface of the coral and quickly began to bore downwards, creating the trace fossil Gastrochaenolites torpedo Kelly and Bromley, 1984. They bored in some cases all the way into the cobble nucleus. The whole set was then buried in transgressive sediments of the Los Banós Formation during the Late Miocene. In the summer of 1989, my student Genga Thavi (“Devi”) Nadaraju (’90) found it as part of her Keck Geology Consortium fieldwork for her Independent Study project. It now resides proudly in the trace fossil collection at Wooster.

Closer view of the gray limestone cobble in the center. Note the remnants of Lithophaga shells still in the borings.

The bivalve boring Gastrochaenolites was named in 1842 by a French geologist with a magnificent name: Alexandre Félix Gustave Achille Leymerie (1801-1878). He was a prolific author with a long career spent primarily studying Cretaceous rocks and fossils in France and northern Spain.


Kelly, S.R.A. and Bromley, R.G. 1984. Ichnological nomenclature of clavate borings. Palaeontology 27: 793-807.

Leymerie, M.A. 1842. Suite de mémoire sur le terrain Crétacé du département de l’Aube. Mémoire des Société Géologique de France 5: 1-34.

Mankiewicz, C. 1995. Response of reef growth to sea-level changes (late Miocene, Fortuna Basin, southeastern Spain). Palaios 10: 322-336.

Mankiewicz, C. 1996. The middle to upper Miocene carbonate complex of Níjar, Almería Province, southeastern Spain, in Franseen, E.K., Esteban, M., Ward, W.C., and Rouchy, J.-M., eds., Models for carbonate stratigraphy from Miocene reef complexes of the Mediterranean regions: Tulsa, SEPM (Society for Sedimentary Geology), p. 141-157.

Nadaraju, G.T. 1990. Borings associated with a Miocene coral reef complex, Fortuna basin, southeastern Spain. Third Keck Research Symposium in Geology (Smith College), p. 165-168.

Taylor, P.D. and Wilson, M.A. 2003. Palaeoecology and evolution of marine hard substrate communities. Earth-Science Reviews 62: 1-103.

Wooster’s Fossil of the Week: a cameloid footprint (Miocene of California)

August 19th, 2012

This fossil is from near my hometown of Barstow, California. It was collected many years ago loose in talus from the Barstow Formation (Barstovian, Miocene). I note this carefully because today collecting such specimens from the Fossil Beds of the Rainbow Basin Natural Area is illegal, as it should be. This is one of the most fossiliferous Miocene deposits in the world, and it has been heavily vandalized over the years.
The Barstow Formation (in a wonderful syncline) at Rainbow Basin, Mojave Desert, California.

This two-toed footprint is Lamaichnum alfi Sarjeant and Reynolds, 1999. It is preserved as a convex hyporelief, which is essentially a filling of the actual footprint. It was made by a camel-like animal (there are many choices) that walked through stiff volcanic mud along a stream during the Miocene. The impression of this foot was quickly filled with later sediment, probably from an overbank flood.

When I was a kid we found dozens of these footprints in long trackways throughout the Barstow Formation at the Fossil Beds. Those fossils are all gone now, most lost to collectors with rock saws and sledge hammers. Fortunately many have been lovingly preserved in the Raymond M. Alf Museum in Claremont, California. You will note that the ichnospecies of our fossil was named for the charismatic Raymond Alf, a legend in the study of vertebrate trace fossils and a spectacular teacher.


Sarjeant, W.A.S. and Reynolds, R.E. 1999. Camelid and horse footprints
from the Miocene of California and Nevada. San Bernardino Museum
Association Quarterly 46: 3-20.

Wooster’s Fossils of the Week: dinosaur gastroliths (Jurassic of Utah, USA)

June 10th, 2012

These rounded stones are labeled in our collections as gastroliths (literally “stomach stones”) from Starr Springs near Hanksville, Wayne County, Utah. I’m featuring them this week in honor of our Utah Project team working right now in the baking Black Rock Desert near Fillmore, Utah.

From their reported location, these stones are likely out of the Summerville Formation (Middle-Upper Jurassic) and, in another plausible supposition, probably from some sort of dinosaur. Sometimes we just have to trust the labels on our specimens, at least for educational purposes!

My friend Tony Martin recently wrote an excellent blog post on gastroliths, so I won’t repeat his insights here. The general wisdom is that these stones were consumed by herbivorous dinosaurs to aid in their digestion. They would have lodged them in the equivalent of a gizzard and used them to grind their food, much like modern birds. (And yes, dinosaurs were birds themselves.) Gastroliths usually have a resistant lithology to be useful as grinders. The gastroliths above are chert, one of the hardest rock types.

Identifying gastroliths correctly is a bit of a challenge if you don’t find them inside a dinosaur skeleton. The most common indicators are that they are very smooth, are in a location where they were unlikely to have been transported inorganically, and are of a lithology unlike the surrounding rock (“exotics” as geologists like to call them). Still, even with all these criteria met, we must be a tad suspicious if we didn’t find them associated with dinosaur bones. I would never, for example, buy a gastrolith in a rock shop. Without context, it could be just a stream-worn stone. I’m trusting the label on ours that we have the real deal!


Stokes, W.L. 1987. Dinosaur gastroliths revisited. Journal of Paleontology 61: 1242-1246.

Wings, O. 2007. A review of gastrolith function with implications for fossil vertebrates and a revised classification. Acta Palaeontologica Polonica 52: 1-16.

Wooster’s Fossil of the Week: a trilobite burrow (Upper Ordovician of Ohio)

May 27th, 2012

This is one of my favorite trace fossils. Rusophycus pudicum Hall, 1852, is its formal name. It was made by a trilobite digging down into the seafloor sediment back during the Ordovician Period in what is now southern Ohio. It may have been hiding from a passing predator (maybe a eurypterid!), just taking a “rest” (what I learned in college), or maybe looking for worms to eat. (There is another example on this blog from the Cambrian of western Canada.)

Rusophycus is always the first trace fossil I introduce in the Invertebrate Paleontology course because it is simple in form and complex in interpretation. It shows that a relatively straightforward process (digging down with its two rows of legs) can have had several motivations. Rusophycus even shows that more than one kind of organism can make the same type of trace. Rusophycus is also found in the Triassic, long after trilobites went extinct. (These were likely made by horseshoe crabs.) It is also good for explaining the preservation of trace fossils. The specimen above is “convex hyporelief”, meaning it is on the bottom of the sedimentary bed and convex (sticking out rather than in). This is thus sediment that filled the open trilobite excavation.

Trilobites making Rusophycus (from http://www.geodz.com/deu/d/Trilobita).

James Hall (1811–1898) named Rusophycus pudicum in 1852. The image of him above is from shortly before his death (photograph credit: The American Monthly Review of Reviews, v. 18, 1898, by Albert Shaw). He was a legendary geologist, and the most prominent paleontologist of his time. He became the first state paleontologist of New York in 1841, and in 1893 he was appointed the New York state geologist. His most impressive legacy is the large number of fossil taxa he named and described, most in his Palaeontology of New York series.

James Hall is in my academic heritage. His advisor was Amos Eaton (1776-1842), a self-educated geologist (he learned it by reading in prison!). One of James Hall’s students was Charles Schuchert (1856-1942), a prominent invertebrate paleontologist. Schuchert had a student named Carl Owen Dunbar (1891-1979) — Schuchert and Dunbar were coauthors of a famous geology textbook. Dunbar had a student at Yale named William B.N. Berry (1931-2011), my doctoral advisor. Thus I feel an intellectual link to old man Hall above.


Baldwin, C.T. 1977. Rusophycus morgati: an asaphid produced trace fossil from the Cambro-Ordovician of Brittany and Northwest Spain. Palaeontology 51: 411–425.

Donovan, S.K. 2010. Cruziana and Rusophycus: trace fossils produced by trilobites … in some cases? Lethaia 43: 283–284.

Hall, J., Simpson, G.B. and Clarke, J.M. 1852. Palaeontology of New York: Organic remains of the Lower Middle Division of the New-York System. C. Van Benthuysen, New York, 792 pages.

Wooster’s Fossils of the Week: Intricate networks of tiny holes (clionaid sponge borings)

May 13th, 2012

The most effective agents of marine bioerosion today are among the simplest of animals: clionaid sponges. The traces they make in carbonate substrates are spherical chambers connected by short tunnels, as shown above in a modern example excavated in an oyster shell. The ichnogenus thus created is known as Entobia Bronn, 1838. I’ve become quite familiar with Entobia throughout its range from the Jurassic through the Recent (with an interesting early appearance in the Devonian; see Tapanila, 2006).
The holes in this Cretaceous oyster are the sponge boring Entobia; the cyclostome bryozoan is Voigtopora. This specimen is from the Coon Creek Beds of the Ripley Formation (Upper Cretaceous) near Blue Springs, Mississippi. (This specimen was collected during a 2010 Wooster/Natural History Museum expedition to the Cretaceous and Paleogene of the Deep South.)
This is a modern clam shell showing Entobia and several other hard substrate dwelling organisms (sclerobionts).
Entobia was named and first described by Heinrich Georg Bronn (1800-1862), a German geologist and paleontologist. He had a doctoral degree from the University of Heidelberg, where he then taught as a professor of natural history until his death. He was a visionary scientist who had some interesting pre-Darwinian ideas about life’s history.


Bromley, R.G. 1970. Borings as trace fossils and Entobia cretacea Portlock, as an example. Geological Journal, Special Issue 3: 49–90.

Bronn, H.G. 1834-1838. Letkaea Geognostica (2 vols., Stuttgart).

Tapanila, L. 2006. Devonian Entobia borings from Nevada, with a revision of Topsentopsis. Journal of Paleontology 80: 760–767.

Taylor, P.D. and Wilson, M.A. 2003. Palaeoecology and evolution of marine hard substrate communities. Earth-Science Reviews 62: 1-103.

Wilson, M.A. 2007. Macroborings and the evolution of bioerosion, p. 356-367. In: Miller, W. III (ed.), Trace Fossils: Concepts, Problems, Prospects. Elsevier, Amsterdam, 611 pages.

Wooster’s Fossil of the Week: the classic bioclaustration (Upper Ordovician of Ohio)

April 29th, 2012

We’re looking at two fossils above. One is the bryozoan Peronopora, the major skeletal structure. The second is the odd series of scalloped holes in its surface. These are a trace fossil called Catellocaula vallata Palmer and Wilson 1988. They at first appear to be borings cut into the bryozoan colony. Instead they are holes formed by the intergrowth of a soft-bodied parasite with the living bryozoan colony. This type of trace fossil is called a bioclaustration. We gave it the Latin name for “little chain of walled pits”.

My good friend Tim Palmer and I found this specimen and many others in 1987 as we explored the Upper Ordovician Kope Formation in the Cincinnati region. We were collecting bioeroded substrates like hardgrounds and shells, and these features were clearly different from the usual borings. They do not actually cut the bryozoan skeleton, for one thing. For another it is apparent that the bryozoan growth was deflected around whatever sat in those spaces. Tim and I called this kind of interaction “bioclaustration”, meaning “biologically walled -up”.
Catellocaula vallata on the Upper Ordovician bryozoan Amplexopora. Note that the scalloped holes have more lobes than those seen in the lead image. This may mean it was a different species of infesting soft-bodied organism.

The infesting parasite on the bryozoan colony was itself colonial, consisting of small clusters connected by extended stolons. The bryozoan grew around the parasite, roofing over the stolons and making walls on the margins of the clusters. We think the parasite was a soft-bodied ascidian tunicate like the modern Botryllus. If true, it is the earliest fossil tunicate known.

This closer view of C. vallata shows the scalloped margins of the pits and the horizontal connections between them.

Another specimen of C. vallata. This view shows the flat floors of the bioclaustration features.

Acetate peels cut longitudinally through the bryozoan and bioclaustrations. On the left you can see that the bryozoan zooecia (long tubes) were deflected sideways as they grew. On the right is a tunnel connecting two pits, with bryozoan zooids forming the roof. (From Palmer and Wilson, 1988.)


Bromley, R.G., Beuck, L. and Taddei Ruggiero, E. 2008. Endolithic sponge versus terebratulid brachiopod, Pleistocene, Italy: accidental symbiosis, bioclaustration and deformity. Current Developments in Bioerosion, Erlangen Earth Conference Series, 2008, III, 361-368.

Palmer, T.J. and Wilson, M.A. 1988. Parasitism of Ordovician bryozoans and the origin of pseudoborings. Palaeontology 31: 939-949.

Tapanila, L. 2006. Macroborings and bioclaustrations in a Late Devonian reef above the Alamo Impact Breccia, Nevada, USA. Ichnos 13: 129-134.

Taylor, P.D. and Voigt, E. 2006. Symbiont bioclaustrations in Cretaceous cyclostome bryozoans. Courier Forschungsinstitut Senckenberg 257: 131-136.

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