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 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.

Wooster’s Fossil of the Week: an encrusted nautiloid (Upper Ordovician of Kentucky)

March 4th, 2012

Two fossils this week in our series. The large segmented cone is a bisected nautiloid cephalopod from the Upper Ordovician of northern Kentucky. The original shell (made of the mineral aragonite) has been dissolved away, leaving the sediment that filled it (making an internal mold). Encrusting the nautiloid mold is a grayish, bumpy layer called Dermatostroma Parks, 1910.

The nautiloid belongs to a subclass of cephalopods still with us today. This Ordovician fossil is in the Family Orthoceratidae McCoy, 1844, which existed from the Early Ordovician (490 million years ago) through the Triassic (230 million years ago). It had a straight, conical shell with walls inside separating chambers (camerae) and a central tube (the siphuncle) connecting them. They were swimming (nektic) predators that could control their buoyancy through a mix of gases and liquids in the camerae mediated by the siphuncle.
Reconstruction of an orthocerid nautiloid by Nobu Tamura.

The fact that the mold is encrusted is interesting in itself. The encrusting organism (Dermatostroma) had to grow over the mold after the aragonitic shell had dissolved and the sediment cemented up. This must have happened on the seafloor, not long after the death and partial burial of the nautiloid. Such rapid dissolution and cementation is characteristic of Calcite Sea conditions, a situation we don’t have in today’s oceans.

Dermatostroma is a genus of stromatoporoid sponge named and described by William Arthur Parks in 1910. It is always very thin and often distinguished by a field of tiny bumps (meaning this species is likely Dermatostroma papillatum). It was a filter-feeding organism, and its fossils are often overlooked.

William Arthur Parks (1868-1936) was a Canadian paleontologist from Hamilton, Ontario. He was a professor at the University of Toronto for most of his career. In 1927, he was elected President of the Paleontological Society. Parks did detailed work on the almost microscopic details of fossil stromatoporoid sponges, and then made a dramatic field change and became an accomplished dinosaur paleontologist. The small ornithopod dinosaur Parksosaurus is named after him.


Palmer, T.J., Hudson, J.D. and Wilson, M.A. 1988. Palaeoecological evidence for early aragonite dissolution in ancient calcite seas. Nature 335 (6193): 809–810.

Parks, W.A. 1910. Ordovician stromatoporoids of America. University of Toronto Studies, Geology Series 7, 52 pp.

Sweet, W.C. 1964. Nautiloidea — Orthocerida, in Treatise on Invertebrate Paleontology. Part K. Mollusca 3, Geological Society of America, and University of Kansas Press, New York, New York and Lawrence, Kansas.

Wooster’s Fossil of the Week: A mysterious sponge (Late Ordovician of Ohio)

February 5th, 2012

I’ve been collecting and studying fossils from the Upper Ordovician of the Cincinnati region for three decades now, but I’ve never seen another specimen like the one pictured above. An amateur collector, Howard Freeland, generously donated this rock to Wooster late last year. He found it in Cincinnatian limestones cropping out in Brown County, Ohio.

At first Howard understandably thought he had found fish bones, which would be extraordinary for this age of rock and place of deposition. He took the slab to the Smithsonian Institution for identification by a vertebrate paleontologist. Not bones, was the answer, but they didn’t know how else to classify these finger-like fossils. When Howard showed them to me I suggested they were fossil sponges, and so here we are. I could be wrong so I hope the web community has some other ideas.

I believe these are sponge pieces because they were originally hollow (now they are filled with sediment), fibrous in structure, and had small holes irregularly preserved on their surfaces. They look in texture like the hexactinellid sponge Brachiospongia, but they do not have their distinctive thick extensions and radiating shape.

Small, irregular holes on fossil surface. They could be sponge incurrent pores. I would expect them to be more regular, though.

My search of the Ordovician sponge literature (what there is of it) has not turned up anything similar. I’ve gone to the usual websites for the Cincinnatian (like Steve Holland’s excellent Cincinnatian fossil catalog and the Dry Dredger’s webpages), but no luck.

Sometime during the existence of this webpage someone will come across these images and post their solution in the comments. I look forward to learning from them!


Carrera, M.G. and Rigby, J.K. 1999. Biogeography of Ordovician sponges. Journal of Paleontology 73: 26-37.

Wooster’s Fossil of the Week: A cornulitid (Late Ordovician of Indiana)

December 18th, 2011

This may look like just another wormtube on a shell — a recurring theme on this blog — but it is special, of course. This is the common Paleozoic genus Cornulites Schlotheim 1820, specifically Cornulites flexuosus (Hall 1847). It was found in the Whitewater Formation (Upper Ordovician) during a College of Wooster field trip to southeastern Indiana.

Above is a larger view of the substrate for this wormtube: the ventral valve exterior of a strophomenid brachiopod. If you look closely at the costae (fine radiating lines) of the brachiopod you can see that it was alive when the cornulitid landed on its shell. As both animals continued to grow, the wormtube bent toward the commissure (opening) of the brachiopod, no doubt to snatch some suspended food from its feeding current. The cornulitid was thus a parasite on the host brachiopod. (See Morris and Rollins, 1971; Vinn and Mutvei, 2005; and Vinn and Wilson, 2010, for much more detail on cornulitid paleoecology.)

Suggested cornulitid internal anatomy (from Olev Vinn).

Cornulitids (Ordovician – Carboniferous) belong to a large group of tube-dwelling organisms that, surprisingly, may be closely related to brachiopods and bryozoans. Cornulitids, along with fellow tube-dwellers the microconchids, tentaculitids and hederelloids, have a foliated shell ultrastructure with various other features indicating they may be part of a larger group called the lophophorates (see Taylor et al., 2010). Much work still needs to be done on their systematics and paleoecology to sort out the evolutionary relationships here, but we have a good start.
The genus Cornulites was described and named by Ernst Friedrich, Baron von Schlotheim (1764-1832), a German palaeontologist and politician born in Almenhausen, Thuringia, Germany. As a noble, he was home-schooled (as we’d say now) and then sent to the Gymnasium (like a high school) in Gotha, Germany. After graduation, he attended Göttingen where he studied political administration and the natural sciences with Johann Friedrich Blumenbach. He enjoyed geology very much and so went off to Freiburg to learn from the famous Abraham Gottlob Werner of Neptunist fame. One of his friends was the scientist-explorer Alexander von Humboldt. After this extraordinary education, Schlotheim entered the civil service in Gotha in 1792, eventually rising all the way up to Lord High Marshal a few years before his death. During his administrative work, though, he continued serious paleontological studies, being one of the first paleontologists to use Linnean binomial nomenclature, making fossils much more useful for stratigraphy and later evolutionary studies. Schlotheim had some very progressive ideas about what we would later call uniformitarianism, and he recognized that geology could tell a history of the Earth quite different from that outlined by theological scholars.

Here’s to the intellectual innovations and courage of Baron von Schlotheim and the little fossil wormtube that reminds us of him!


Morris, W. R., and H. B. Rollins. 1971. The distribution and paleoecological interpretation of Cornulites in the Waynesville Formation (Upper Ordovician) of southern Ohio. The Ohio Journal of Science 71: 159-170.

Schlotheim, E.F. von. 1820. Die Petrefakten-Kunde auf ihrem jetzigen Standpunkte durch die Beshreibung seiner Sammlung versteinerter und fossiler Ueberreste des their-und Planzenreichs der Vorwelt erlaeutert. Gotha, 437 p.

Taylor, P.D., Vinn, O. and Wilson, M.A. 2010. Evolution of biomineralization in ‘lophophorates’. Special Papers in Palaeontology 84: 317-333.

Vinn, O. and Mutvei, H. 2005. Observations on the morphology and affinities of cornulitids from the Ordovician of Anticosti Island and the Silurian of Gotland. Journal of Paleontology 79: 726-737.

Vinn, O. and Wilson, M.A. 2010. Abundant endosymbiotic Cornulites in the Sheinwoodian (Early Silurian) stromatoporoids of Saaremaa, Estonia. Neues Jahrbuch für Geologie und Paläontologie 257: 13-22.

Wooster’s Fossil of the Week: an orthid brachiopod (Upper Ordovician of Indiana)

November 27th, 2011

This beautiful brachiopod is Vinlandostrophia ponderosa (Foerste, 1909), an orthid brachiopod from the Maysvillian (Upper Ordovician) of southern Indiana. Until recently it had been traditionally known as Platystrophia ponderosa until a critical paper by Zuykov and Harper (2007) investigated the “Platystrophia plexus” of species and convincingly made P. ponderosa the type species of Vinlandostrophia.

Brachiopods are filter-feeding, bivalved marine invertebrates who have been with us since the Cambrian Period. They were among the most common animals of the Ordovician. The fossils of the Cincinnatian Series in southern Indiana, southwestern Ohio and northern Kentucky have extraordinary numbers and varieties of fossil brachiopods — so many they roll under your feet in some places.

August F. Foerste (1862-1936) described what he called Platystrophia ponderosa in 1909. He was a pioneering paleontologist who grew up and worked in the Dayton area. Foerste went to Denison University where he was a very successful undergraduate, publishing several geological papers. He returned to Dayton after graduation with a PhD from Harvard, teaching high school for 38 years. When he retired he was offered a teaching position at the University of Chicago, but instead went to work at the Smithsonian Institution until the end of his life.

This is, by the way, the 500th post of the Wooster Geologists blog. It is great fun.


Foerste, A.F. 1909. Preliminary notes on Cincinnatian fossils. Denison University, Scientific Laboratories, Bulletin 14: 208-231.

Zuykov, M.A. and Harper, D.A.T. 2007. Platystrophia (Orthida) and new related Ordovician and Early Silurian brachiopod genera. Estonian Journal of Earth Science 56: 11-34.

Wooster’s Fossil of the Week: A receptaculitid (Middle Ordovician of Missouri)

September 18th, 2011

This week’s fossil is a long-standing paleontological mystery. Above is a receptaculitid from the Kimmswick Limestone (Middle Ordovician) near Ozora, Missouri. I think I found it on a field trip with Frank Koucky in the distant mists of my student days at Wooster, but so many outcrops, so many fossils …

Below is a nineteenth century illustration of a typical receptaculitid fossil. They are sometimes called “sunflower corals” because they look a bit like the swirl of seeds in the center of a sunflower. They were certainly not corals, though, or probably any other kind of animal. Receptaculitids appeared in the Ordovician and went extinct in the Permian, so they were confined to the Paleozoic Era. Receptaculitids were bag-like in form with the outside made of mineralized pillars (meroms) with square or diamond-shaped heads. The fossils are usually flattened disks because they were compressed by burial. You may notice now that the fossil at the top of this post is a mold of the original with the dissolved pillars represented by open holes. (Paleontologists can argue if this is an external or internal mold.)So what were the receptaculitids? When I was a student we called them a kind of sponge, something like a successor of the Cambrian archaeocyathids. In the 1980s a convincing case was made that they were instead a kind of alga of the Dasycladales. Now the most popular answer is that they belong to that fascinating group “Problematica”, meaning we have no idea what they were! (Nitecki et al., 1999). It’s those odd meroms that are the problem — they appear in no other known group, fossil or recent.

I find it deeply comforting that we still have plenty of fossils in the Problematica. We will always have mysteries to puzzle over.
Another Wooster receptaculitid specimen, this time seen from the underside showing side-views of the meroms.
Diagram of a receptaculitid in roughly life position showing its inflated nature and pillar-like meroms. From Dawson (1880, fig. 25): a, Aperture (probably imaginary here). b, Inner wall. c, Outer wall. n, Nucleus, or primary chamber. v, Internal cavity.

Finally, this is what a typical receptaculitid looks like in the field (Ordovician of Estonia). Note that nice sunflower spiral of the merom ends.


Dawson, J.W. 1880. The chain of life in geological time: A sketch of the origin and succession of animals and plants. The Religious Tract Society, 272 pages.

Nitecki, M.H., Mutvei, H. and Nitecki, D.V. 1999. Receptaculitids: A Phylogenetic Debate on a Problematic Fossil Taxon. Kluwer Academic/Plenum, 241 pages.

Fossils in the Wild: Invertebrate Paleontology Field Trip

September 11th, 2011

CAESAR CREEK LAKE, OHIO–The 2011 Invertebrate Paleontology class had a productive field trip on a beautiful Ohio day. Thunderstorms roamed the state, but we saw them only when we were comfortably on the bus.

We worked in the emergency spillway at Caesar Creek Lake in southwestern Ohio, roughly halfway between Cincinnati and Dayton. This site is maintained by the US Army Corps of Engineers as a fossil-collecting preserve. You obtain a free permit at the visitor center, agree to follow the rules, and extraordinary fossils await your picking. (Last time I was here it was very cold.)

The fossils are in the Arnheim, Waynesville, Liberty and Whitewater Formations of the Richmondian Stage in the Cincinnatian Series of the Ordovician System. These are shaly units with shell-rich limestones formed during storms. Brachiopods, bryozoans, crinoids, trilobites, clams, snails, nautiloids, corals — the whole Ordovician menagerie. Perfect for student collections and our later exercises.

Brachiopod-rich storm layer in the Liberty Formation. Note the circular bryozoan attachment.

Bryozoan colony and brachiopod shell interior from the Waynesville Formation.

Our fancy bus. The design insures that the back seats are rather bouncy.

Last of the summer flower field photos! It was such a beautiful day.

Wooster’s Fossil of the Week: A Biserial Graptolite (Middle Ordovician of Tennessee)

August 28th, 2011

This week’s fossils are graptolites (from the Greek for written rocks) I found many years ago in the Lebanon Limestone near the town of Caney Springs south of Nashville, Tennessee. They are of the genus Amplexograptus and probably belong to the species A. perexcavatus (Lapworth, 1876).

Graptolites were colonial organisms consisting of hundreds and sometimes thousands of tiny zooids (individuals) connected together in a flexible proteinaceous skeleton (the rhabdosome). They first appeared in the Late Cambrian (around 510 million years ago) and disappeared forever in the Early Carboniferous (around 350 million years ago). Amplexograptus colonies were probably attached to floats so they could drift through the ancient oceans filtering out organic particles; they would be officially “passively mobile planktonic suspension feeders”. They belong to the Phylum Hemichordata, although there have always been disputes about their actual evolutionary relationships. This matters because graptolites are important index fossils for sorting out the age relationships of Lower and Middle Paleozoic rocks.

Graptolites are usually preserved as thin carbonaceous films on dark shales, making them rather hard to see (as my paleontology students will readily agree). The great 18th Century naturalist Linnaeus even said that they were “pictures resembling fossils rather than true fossils”. Sometimes, though, they are found in lighter-colored rocks like limestones, as above. Goldman et al. (2002) found Amplexograptus in limestones preserved in three dimensions, possibly because the limestones were cemented early around them before they collapsed with decay. They even studied this same species from the Lebanon Limestone. The 3-D preservation allows for a much more detailed analysis of the tiny cups (thecae) which held the individual zooids. It is possible that I could dissolve the limestone shown above and retrieve some delicate three-dimensional graptolites — but I could also just as easily destroy them.

Amplexograptus perexcavatus was originally described in 1876 by the famous geologist Charles Lapworth (1842-1920), who referred it to the genus Diplograptus. Actually, he had two species in his D. perexcavatus group, so it took some taxonomic detective and legal work to fix the current naming system. Lapworth, who I’ve figured below with an inset of his not-very-helpful diagram of the original D. perexcavatus, is well known by paleontologists for his work with graptolites as index fossils. Scientists and historians of science know him as the man who invented the Ordovician Period in 1879 to solve a bitter dispute between Roderick Murchison and Adam Sedgwick who each claimed the same rock interval in Wales for the Silurian and Cambrian periods respectively. Lapworth’s primary biostratigraphic argument for the Ordovician as a separate period was the distribution of graptolites, including our friend Amplexograptus perexcavatus. (Murchison and Sedgwick were long gone by the time their dispute was settled.)

(Charles Lapworth. Image courtesy of The Lapworth Museum of Geology.)


Goldman, D., Campbell, S.M. and Rahl, J.M. 2002. Three-dimensionally preserved specimens of Amplexograptus (Ordovician, Graptolithina) from the North American mid-continent: taxonomic and biostratigraphic significance. Journal of Paleontology 76: 921-927.

Lapworth, C. 1876. The Silurian System in the South of Scotland, p. 1–28. In: Armstrong, J. Young, J. and Robertson, D. (eds.), Catalogue of Western Scottish Fossils. Blackie and Son, Glasgow.

Wooster’s Fossil of the Week: A trilobite hypostome (Upper Ordovician of southern Ohio)

August 21st, 2011

We had a familiar trilobite last week, so this week we’ll look at a poorly-known part of a trilobite: the hypostome. Above is an incomplete forked, conterminant hypostome of the large trilobite Isotelus. (Isotelus, by the way, is the state fossil of Ohio. Do you know your state fossil?)

Hypostome means “under mouth”. On trilobites it is found underneath the cephalon (head) near what we think was the mouth. They are not common in the fossil record. It is obvious from their color and composition that they are part of a trilobite, but most people don’t know about this little plate on the otherwise soft underside (the ventral side) of the animal. The hypostome is important in some new taxonomic schemes for sorting out the trilobites (Fortey, 1990), and they are useful for interpreting a particular trilobite’s feeding habits (Fortey and Owens, 1999).
Trilobite hypostome forms from Wikipedia (via Obsidian Soul). The small green plates are the hypostomes seen against the gray cephalon above. A – Natant: Hypostome not attached to doublure; aligned with front edge of glabella (shown in red broken lines). B – Conterminant: Hypostome attached to rostral plate of doublure. Aligned with front edge of glabella. C – Impendent: Hypostome attached to rostral plate but not aligned with glabella.

The hypostome of Isotelus is attached to the anterior edge of the skeleton (thus “conterminant”) and has two distally-directed prongs (making it “forked”). Hegna (2010) has recently suggested this hypostome with its unusual shape and terraced outer structure may have been used for grinding food rather than serrating it. Turns out our hypostome has a unique form among the common trilobites!


Fortey, R.A. 1990. Ontogeny, hypostome attachment and trilobite classification. Journal of Paleontology 33: 529-576.

Fortey, R.A. and Owens, R.M. 1999. Feeding habits in trilobites. Palaeontology 42: 429–65.

Hegna, T.A. 2010. The function of forks: Isotelus-type hypostomes and trilobite feeding. Lethaia 43: 411-419.

« Prev - Next »