Wooster’s Fossil of the Week: A brontothere jaw fragment (Miocene of South Dakota)

April 6th, 2014

Titanotherium proutiiThis fossil has been sitting in a glass case outside my office door for nearly three decades. Only this year — in the desire to find more Fossils of the Week — did I bother to open the cabinet and take it out for a looksie. On the reverse was a 19th century label: “Titanotherium proutii, Badlands, SD”. That started me on a complicated journey through the literature to see just what sort of creature bore these magnificent molars, as well as the history of its discovery.
Titanotherium proutii occlusalIn this occlusal (meaning the biting surface) view you can see in the beautiful flowing lines of the hard (dark) and softer (light) enamel that there are some serious cracks repaired with a dodgy yellowish glue. The specimen is very fragile — that glue has probably been holding it together for well over a century. These are classic plant-eating teeth for both cutting and grinding leaves, roots and small branches.
TitanotheriumThe animal represented here is a titanothere, a large extinct mammal common in what would become the Badlands of South Dakota during the Paleogene. Above is a recreation of a relative of our species: Megacerops (Titanotherium) robustum. (The artist who drew this illustration in 1912 was Robert Bruce Horsfall, 1869-1948.) The titanotheres, now better known as brontotheres, were roughly the size and shape of rhinoceroses, but were actually more closely related to horses. They had elephant-like feet, inwardly-curved skull caps, and impressive horns on the nose.

I usually delight in tracking down taxonomic histories (the technical history of scientific names), but Titanotherium proutii has defeated me. The history of this taxon is convoluted beyond recovery — a sad tale of mistakes, misplaced fossils, specimens given multiple names, and over-zealous “splitting” of taxa. In other words, typical middle 19th century vertebrate paleontology. Mader (1998) says that Titanotherium Leidy 1853 (or 1852?) is a nomen dubium or “doubtful name”. Even the species, which later became Palaeotherium proutii, is nomen dubium. The names are simply worthless to science. I have been unable to figure out what the accepted name for our fossil now is.
Joseph_LeidyJoseph Leidy (1823-1891) named Titanotherium and (maybe) T. proutii (there is dispute as to who named it first). Leidy was a well known American biologist and paleontologist who taught first at the University of Pennsylvania and then Swarthmore College. He described and named the first nearly-complete dinosaur skeleton, Hadrosaurus foulkii. (It was found in the Cretaceous of New Jersey and he named it in 1858.) Leidy was also an early supporter of Charles Darwin and his the new theory of evolution, early enough for this to be an unpopular position. Edward Drinker Cope was one of his students, which forever places him at the beginning of the famous “Bone Wars” between Cope and Othniel Charles Marsh, which raged from 1877 to 1892. That epic conflict actually began in the New Jersey marl pits where Leidy’s hadrosaur was found. Leidy thus leaves us with a mixed legacy of discoveries, innovations and insights mixed with errors and folly. Just the sort of character we would expect on the frontier of a new science in a new country.
Leidy1Leidy’s 1853 (Plate XVI) figure of a jaw fragment of “Titanotherium proutii“.


Leidy, J. 1853. The ancient fauna of Nebraska: or, a description of remains of extinct Mammalia and Chelonia, from the Mauvaises Terres of Nebraska. Smithsonian contributions to knowledge, vol. 6. Washington, Smithsonian Institution.

Mader, B.J. 1998. Brontotheriidae. In: C.M. Janis, K.M. Scott, and L.L. Jacobs (eds.), Evolution of Tertiary Mammals of North America 1: 525-536.

Mihlbachler, M.C., Lucas, S.G. and Emry, R.J. 2004. The holotype specimen of Menodus giganteus, and the “insoluble” problem of Chadronian brontothere taxonomy. Paleogene Mammals. New Mexico Museum of Natural History and Science Bulletin 26: 129-135.

Warren, L. 1998. Joseph Leidy: the last man who knew everything. Yale University Press, New Haven; 303 pages.

Wooster’s Fossil of the Week: Thoroughly encrusted brachiopod from the Upper Ordovician of Indiana

March 30th, 2014

1 Rafinesquina ponderosa (Hall) ventralLast week was an intensely bored Upper Ordovician bryozoan, so it seems only fair to have a thoroughly encrusted Upper Ordovician brachiopod next. The above is, although you would hardly know it, the ventral valve exterior of a common strophomenid Rafinesquina ponderosa from the Whitewater Formation exposed just south of Richmond, Indiana (locality C/W-148). I collected it earlier this month on a trip with Coleman Fitch (’15).
2 Rafinesquina ponderosa (Hall) dorsalThis is the other side of the specimen. We are looking at the dorsal valve exterior. Enough of the brachiopod shows through the encrusters that we can identify it. Note that both valves are in place, so we say this brachiopod is articulated. Usually after death brachiopod valves become disarticulated, so the articulation here may indicate that the organism had been quickly buried. This brachiopod is concavo-convex, meaning that the exterior of the dorsal valve is concave and the exterior of the ventral valve is convex.
3 Protaraea 032314Returning to the ventral valve, this is a close-up of the encruster that takes up its entire exterior surface. It is the colonial heliolitid coral Protaraea richmondensis Foerste, 1909. (Note the species name and that it was collected just outside Richmond, Indiana.) This thin coral is a common encruster in the Upper Ordovician. Usually it is a smaller patch on a shell. This is the most developed I’ve seen the species. The holes, called corallites, held the individual polyps.
4 Bryo on Protaraea 032314The encrusting coral has an encruster on top of it. This is a trepostome bryozoan, which you can identify by the tiny little holes (zooecia) that held the individuals (zooids). The patch of coral it is occupying must have been dead when the bryozoan larva landed and began to bud.
5 Trepostome 032314Now we’re returning to the concave dorsal valve with its very different set of encrusters. This is a close-up of another kind of trepostome bryozoan, this one with protruding bumps called monticules. They may have functioned as “exhalant current chimneys”, meaning that they may have helped channel feeding currents away from the surface after they passed through the tentacular lophophores of the bryozoan zooids. For our purposes, this is a feature that distinguishes this bryozoan species from the one on the ventral valve.
6 Cuffeyella 032314There is a third, very different bryozoan on the dorsal valve. This blobby, ramifying form is a well-developed specimen of Cuffeyella arachnoidea (Hall, 1847). It is again a common encruster in the Upper Ordovician, but not usually so thick.
7 Cuffeyella on hinge 032314If we look closely at the hinge of the brachiopod on the dorsal side, we can see a much smaller C. arachnoidea spreading on the ventral valve.
8 Encrusted edge 032314Finally, this is a side view of the brachiopod with the ventral valve above and the dorsal valve below. We’re looking at the junction of the articulated valves, the commissure. For the entire extent of the commissure, the encrusting coral grows to the edge of the ventral valve and no further. This is a strong indication that the brachiopod was alive when the coral was growing on it. The brachiopod needed to keep that margin clear for its own feeding.

The paleoecological implications here are that the coral was alive at the same time as the brachiopod. This means that the convex exterior surface of the ventral valve was upwards for the living brachiopod. The concave exterior surface of the dorsal valve faced downwards. The coral and bryozoan encrusting the top of the living brachiopod were exposed to the open sea; the bryozoans encrusting the undersurface of the living brachiopod were encrusting a cryptic space. We are thus likely seeing the living relationships between the encrusters and the brachiopod — this encrustation took place during the life of the brachiopod.

Further, this demonstrates that this concavo-convex strophomenid brachiopod was living with the convex side up. This has been a controversy for decades in the rarefied world of brachiopod paleoecology. This tiny bit of evidence, combined with some thorough recent studies (see Dattilo et al., 2009; Plotnick et al., 2013), strengthens the case for a convex-up orientation. Back when I was a student these would be fighting words!


Alexander, R.R. and Scharpf, C.D. 1990. Epizoans on Late Ordovician brachiopods from southeastern Indiana. Historical Biology 4: 179-202.

Dattilo, B.F., Meyer, D.L., Dewing, K. and Gaynor, M.R. 2009. Escape traces associated with Rafinesquina alternata, an Upper Ordovician strophomenid brachiopod from the Cincinnati Arch Region. Palaios 24: 578-590.

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

Mõtus, M.-A. and Zaika, Y. 2012. The oldest heliolitids from the early Katian of the East Baltic region. GFF 134: 225-234.

Ospanova, N.K. 2010. Remarks on the classification system of the Heliolitida. Palaeoworld 19: 268–277.

Plotnick, R.E., Dattilo, B.F., Piquard, D., Bauer, J. and Corrie, J. 2013. The orientation of strophomenid brachiopods on soft substrates. Journal of Paleontology 87: 818-825.

Wooster’s Fossil of the Week: Intensely bored bryozoan from the Upper Ordovician of Kentucky

March 23rd, 2014

Bored Bryo 1 585Yes, yes, I’ve heard ALL the jokes about being bored, and even intensely bored. I learn to deal with it. This week we continue to highlight fossils collected during our productive expedition to the Upper Ordovician (Cincinnatian) of Indiana (with Coleman Fitch ’15) and Kentucky (with William Harrison ’15). Last week was Coleman’s turn; this week it is William’s.

The beautiful fan-like bifoliate (two-sided) trepostome bryozoan above was collected from the lower part of the Grant Lake Formation (“Bellevue Limestone”) at our locality C/W-152 along the Idlewild Bypass (KY-8) in Boone County, Kentucky (N 39.081120°, W 84.792434°). It is in the Maysvillian Stage and so below the Richmondian where Coleman is getting most of his specimens. I’ve labeled it to show: A, additional bryozoans encrusting this bryozoan; B, a very bored section; C, a less bored surface showing the original tiny zooecia, monticules, and a few larger borings.
Bored Bryo 2 585The other side of this bryozoan is more uniform. It has an even distribution of small borings and no encrusters. This likely means that at some point after the death of the bryozoan and subsequent bioerosion this side was placed down in the mud while the exposed opposite side was encrusted.
Encruster Bored Bryo 031314_585A closer view of the upwards-facing side (with the encrusting bryozoan at the top) shows just how intense the boring was prior to encrustation. Some of the borings are close to overlapping. The encrusting bryozoan has its own borings, but far fewer and significantly larger.
Close borings 031314_585In this close view of the downwards-facing side we see lots of the small borings. Some are star-shaped if they punched through the junction of multiple zooecia. Note that these borings are rather evenly spread and seem to have about the same external morphology and and erosion. Likely they were all produced about the same time. It must have been a crowded neighborhood when all those boring creatures were home.

The questions that are provoked by this specimen are: (1) Were there any borings produced while the host bryozoan was still alive? (We may find elements of bioclaustration with some holes); (2) Why are zones B and C in the top image so different in the amount of bioerosion? Could zone C have still been alive at the time and resisted most bioeroders? Maybe zone C was covered by sediment? (But the margin is very irregular); (3) Why are the later encrusting bryozoans (zone A) so much less bioeroded?; (4) How do we classify such tiny pits that are between microborings and macroborings in size? (Trypanites is becoming a very large category) (5) What kind of organism made so many small pits? Were they filter-feeders as we always say, or was something else going on? (Sectioning specimens like this may reveal some internal connections between the pits.)

William has plenty of fun work ahead of him!


Boardman, R.S. and Utgaard, J. 1966. A revision of the Ordovician bryozoan genera Monticulipora, Peronopora, Heterotrypa, and Dekayia. Journal of Paleontology 40: 1082-1108

Bromley, R.G. 1972. On some ichnotaxa in hard substrates, with a redefinition of Trypanites Mägdefrau. Paläontologische Zeitschrift 46: 93–98.

Erickson, J.M. and Waugh, D.A. 2002. Colony morphologies and missed opportunities during the Cincinnatian (Late Ordovician) bryozoan radiation: examples from Heterotrypa frondosa and Monticulipora mammulata. Proceedings of the 12th International Conference of the International Bryozoology Association. Swets and Zeitlinger, Lisse; pp. 101-107..

Kobluk, D.R. and Nemcsok, S. 1982. The macroboring ichnofossil Trypanites in colonies of the Middle Ordovician bryozoan Prasopora: Population behaviour and reaction to environmental influences. Canadian Journal of Earth Sciences 19: 679-688.

Taylor, P.D. and Wilson. M.A. 2003. Palaeoecology and evolution of marine hard substrate communities. Earth-Science Reviews 62 (1-2): 1–103.

Vogel, K. 1993. Bioeroders in fossil reefs. Facies 28: 109-113.

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: Bryozoan bored and bryozoan boring in the Upper Ordovician of Indiana

March 16th, 2014

Bored Bryo on Brach top 585This week and next we will highlight fossils collected during our brief and successful expedition to the Upper Ordovician (Cincinnatian) of Indiana (with Coleman Fitch ’15) and Kentucky (with William Harrison ’15). We found what we needed to pursue some very specific topics.

Above is a trepostome bryozoan collected from the Liberty Formation (we should be calling it the Dillsboro Formation in Indiana; our locality C/W-149) on IN-101 in southeastern Indiana (N 39.48134°, W84.94843°). You can see the regular network of tiny little holes representing the zooecia (zooid-bearing tubes) of the calcitic zoarium (colony) of the bryozoan. The larger, irregular holes (still pretty small!) are borings cut by worm-like organisms into the bryozoan skeleton shortly after the death of the colony.
Bored Bryo on Brach bottom 585Flipping the specimen over we see the most interesting parts. On the left is a remnant of the original calcitic strophomenid brachiopod shell that was encrusted by the trepostome bryozoan. On the right the shell has broken away, exposing the encrusting surface of the trepostome. We are thus looking here at the inside of a brachiopod valve and the underside of the bryozoan that encrusted it.

This is just what we hoped to find for Coleman’s project on interpreting half-borings in brachiopod shell exteriors. This specimen demonstrates two crucial events after encrustation: First, the borings in the bryozoan extended down to the brachiopod shell and turned sideways to mine along the shell/bryozoan junction (note half-borings in the bryozoan base on the right), and second, the bryozoan broke mostly free of the brachiopod shell, with only a bit remaining on the left. Somewhere there is or was a fragment of that brachiopod with an exterior showing half-borings and no bryozoan encrustation. Thus a brachiopod without bryozoan encrusters may have actually been encrusted at some point, but the bryozoans were later detached. We’ve added a bit to the uncertainty of the encrusting fossil record — even calcitic skeletal evidence on this small scale can go missing. We’ve also started on a good story about the behavior of the tiny critters that bored into this shelly complex.
Ctenostome closer 031314_585A bonus in this specimen can be seen in this closer view of that brachiopod shell interior above. That branching network is a complex ctenostome bryozoan boring called Ropalonaria. This is a particularly well developed specimen with thicker, shorter zooids than I’ve seen before. This kind of boring is the subject of a previous Fossil of the Week entry.

Coleman has a great start on his Independent Study project with specimens like these. He has a lot of sectioning and adequate peeling ahead of him!


Brett, C.E., Smrecak, T., Parsons-Hubbard, K. and Walker, S. 2012. Marine sclerobiofacies: Encrusting and endolithic communities on shells through time and space. In: Talent, J.A. (ed.) Earth and Life, International Year of Planet Earth, p. 129-157. Springer.

Pohowsky, R.A. 1978. The boring ctenostomate Bryozoa: taxonomy and paleobiology based on cavities in calcareous substrata. Bulletins of American Paleontology 73(301): 192 p.

Smrecak, T.A. and Brett, C.E. 2008. Discerning patterns in epibiont distribution across a Late Ordovician (Cincinnatian) depth gradient. Geological Society of America Abstracts with Programs 40:18.

Wilson, M.A., Dennison-Budak, C.W. and Bowen, J.C. 2006. Half-borings and missing encrusters on brachiopods in the Upper Ordovician: Implications for the paleoecological analysis of sclerobionts. Geological Society of America Abstracts with Programs 38:514.

Wooster’s Fossil of the Week: A whale ear bone (Neogene)

March 9th, 2014

Whale inner earbonesThis is another fossil that has sat in a display case for decades in Scovel before I really examined it. Unlike last week’s specimen, though, it has no identifying label on its reverse. This is always a serious disappointment for science — no location! I show the fossil above with a front and back view (as much as there is a front or back). We are looking at an auditory bulla (part of the middle ear system) of an ancient whale. The most we can say is that this may be from a type of sperm whale that lived during the Neogene. Likely this specimen was collected on the east coast of the United States, maybe Maryland or Virginia.

Surprisingly, whale ear bones are rather common in the later fossil record. They seem to have been of denser bone than the rest of the whale skeleton, so they were better preserved. The auditory bulla is a bony cover for the delicate middle ear bones and tissues. In humans it is part of our temporal bone. Whales have several adaptations in their ears for hearing underwater. They have no external ear opening. They use instead the lower jawbone to transmit vibrations to the ear complex (something like what many snakes do). They have a pad of fat to enhance these vibrations for the tiny ear bones (tiny relative to the massive size of the whale). You can learn much more about fossil whale ear bones at this excellent blog post from the Virginia Museum of Natural History.

You are asking, though, fine enough, but how can I use a fossil whale ear bone? There’s a video to train you! These bones have “ancient, ancient memory” that is “preserved sonically”. Just be sure to hold it in your non-dominant hand and remember that “this is an art”. Do it correctly and you will have tapped into the wisdom of our ancient whale brothers and sisters. To think that every day I walked blithely by this portal to the Knowledge of the Ages.


Fraser, F.C. and Purves, P.E. 1960. Hearing in cetaceans: evolution of the accessory air sacs and the structure and function of the outer and middle ear in recent cetaceans. Bulletin of the British Museum (Natural History) 7: 1-140.

Ketten, D.R. 1997. Structure and function in whale ears. Bioacoustics 8: 103-135.

Wooster’s Fossil of the Week: An agate-replaced coral from the Oligocene-Miocene of Florida

March 2nd, 2014

DSC_3384_585I long thought of this beautiful specimen as more rock than fossil. It is a scleractinian coral that has had its outer skeleton replaced by the silicate material agate and its interior skeleton completely hollowed out. The result is a geode that happens to also be a fossil.
FLMNH_585Then during last month’s North American Paleontological Convention in Gainesville, Florida, I saw the above specimens on display in the Florida Museum of Natural History. These fossils were so striking that I decided to highlight our single example.
DSC_3388_585This is a view of the top surface of the Wooster specimen. In the upper left is an array of holes with crystals radiating away from them. These are remnants of the original corallites, and there is just enough information there for us to conclude the likely genus is Montastraea. This piece thus becomes an example of Florida’s official state stone. Here’s the official definition: “… a chalcedony pseudomorph after coral, appearing as limestone geodes lined with botryoidal agate or quartz crystals and drusy quartz fingers, indigenous to Florida.” Our specimen came from the Hawthorn Group of rocks near Tampa, Florida.
DSC_3393_585The outside of the fossil shows horizontal banding remaining from the original growth lines in the coral, which is another clue that this is Montastraea. The coral made its skeleton of aragonite around 30 million years ago. After death and burial, silica-rich groundwater began to replace the aragonite on the surface of the coral with what later became banded agate. The interior dissolved away into a hollow cavity.

The common name for this fossil is “agatized coral“, and it is a collector’s item. It is apparently Florida’s only native gemstone. Pretty cool that their state rock and gemstone is a fossil!


Scott, T.M. 1990. The lithostratigraphy of the Hawthorn Group of peninsular Florida. World Phosphate Deposits 3: 325-336.

Wooster’s Fossil of the Week: An interlocking rugose and tabulate coral (Devonian of Michigan)

February 23rd, 2014

Hexagonaria percarinata colony viewThis beautifully polished fossil looks like half of an antique bowling ball. Normally I hate polished fossils because the external details have been erased, but in this case the smooth surface reveals details about the organisms and their relationship. We have here a large colonial rugose coral with a smaller tabulate coral embedded within it. The specimen is from the Devonian of Michigan. It may look familiar because it is a large “Petoskey Stone“, the state stone (not fossil!) of Michigan. The large rugose coral is Hexagonaria percarinata (Sloss, 1939).
Hexagonaria percarinata close view 585In this closer view you can see the multiple star-like corallites of this coral. Each corallite held a tentacular feeding polyp in life. The radiating lines are thin vertical sheets of skeleton called septa. The corallites in this type of coral shared common walls and nestled up against each other as close as possible. In the lower center of the image you can see a very small corallite that represents a newly-budded polyp inserting itself as the colony grew. If rugose corals were like modern corals (and they probably were), the polyps were little sessile benthic carnivores catching small passing organisms with a set of tentacles. They may also have had photosymbionts to provide oxygen and carbohydrates through photosynthesis.
Tabulate coral intergrown with HexagonariaIn the midst of the rugose coral is this irregular patch with another type of coral: a tabulate coral distinguished by numerous horizontal partitions in its corallites (and no septa). It was likely a favositid coral, sometimes called a “honeycomb coral”. It was clearly living in the rugosan skeleton and not pushed into it by later burial. Note, though, the ragged boundary between the two corals. The rugose coral has the worst of it with some corallites deeply eroded. What seems to have happened is that the rugose coral had an irregular opening in its corallum (colonial skeleton) after death and the tabulate grew within the space, eventually filling it. The tabulate likely stuck out far above the rugose perimeter, but the polishing shaved them down to the same level. This is thus not a symbiotic relationship but one that happened after the death of the rugose coral.
Stumm, Erwin C   copyThe rugose coral species, Hexagonaria percarinata, was named in 1939 by Laurence Sloss, a famous sedimentary geologist with an early start in paleontology, but it is best known through the research of Erwin Charles Stumm (1908-1969; pictured above). Stumm was at the end of his life a Professor of Geology and Mineralogy and the Curator of Paleozoic Invertebrates in the Museum of Paleontology at the University of Michigan. Stumm grew up in California and then moved east for his college (George Washington University, ’32) and graduate (PhD from Princeton in 1936) education. He taught geology at Oberlin College up the road for ten years, and then moved to Michigan to start as an Associate Curator and Assistant Professor. I knew his name because in 1967 he was President of the Paleontological Society. He is said to have been a dedicated teacher of undergraduates and effective graduate advisor. It is fitting that his name is connected to such a popular fossil as Hexagonaria percarinata.


Sloss, L. 1939. Devonian rugose corals from the Traverse Beds of Michigan. Journal of Paleontology 13: 52-73.

Stumm, E.C. 1967. Growth stages in the Middle Devonian rugose coral species Hexagonaria anna (Whitfield) from the Traverse Group of Michigan. Contributions from the Museum of Paleontology, The University of Michigan 21(5): 105-108.

Stumm, E.C. 1970. Corals of the Traverse Group of Michigan Part 13, Hexagonaria. Contributions from the Museum of Paleontology, The University of Michigan 23(5): 81-91.

Wooster’s Fossil of the Week: A tubeworm-encrusted parasitic gastropod (Silurian of Indiana)

February 16th, 2014

Platyostoma1_585Last week three Wooster geology students and I visited Ken Karns, an enthusiastic citizen scientist who has developed an extraordinary fossil collection in his home in Lancaster, Ohio. Ken is a man of prodigious energies and skills as he not only is an expert fossil collector and preparator, he also has a world-class curated collection of Ohio beetles! He was introduced to us by our friend Brian Bade, a man with similar enthusiasms and skills. The students were Steph Bosch (’14), Lizzie Reinthal (’14) and Ian Tulungen (’15). Our goals were to meet Ken, see his magnificent collection with Brian and other friends, and then focus on a project for Ian’s future Independent Study work. Success on all counts, and the specimen above is evidence. Ken was very generous in loaning this specimen to us along with several others for Ian’s work.

The above specimen is from the type section of the Waldron Shale Member (Silurian, Wenlockian, Homerian, about 430 million years old) of the Pleasant Mills Formation near St. Paul, south-central Indiana. Ken Karns collected and prepared it. It is a platyceratid snail of the genus Platyostoma Conrad 1842. It is probably of the species P. niagarense Hall 1852, but there is another species in the same unit (P. plebeium Hall 1876). I’m not quite sure of the differences between these species because platyceratids are notoriously variable. It is possible they are synonymous. Unlike most gastropods, platyceratids had calcite shells instead of aragonite, so they are very well preserved. For an excellent taxonomic review of the genus Platyostoma and its founder, Timothy Abbott Conrad, please see Tony Edger’s blog entry. (We’ve talked about Conrad in this blog as well.)
Platyostoma2_585In this different angle on the specimen you can see additional encrusters (sclerobionts) on the surface of the Platyostoma shell. In the lower right is a remnant of a sheet-like bryozoan, but the most prominent sclerobionts are the tubeworms Cornulites proprius Hall 1876. These encrusters interest us very much.
Cornulitids on Platyostoma_585In this closer view it is apparent that several of the cornulitids are aligned with their apertures pointing in the same way. This is a pattern we’ve seen on many of these snails. Platyostoma was a parasitic snail that lived attached to crinoids, which were abundant in the Waldron fauna. They lived high on the calyx of the crinoid firmly fixed to its skeleton. These cornulitids and other encrusters were thus living high off the substrate perched on the snails. They were filter-feeders like the crinoids, so they may have been feeding on some suspended food fraction missed by the crinoid arms, or they were competing for nutrients and added to the parasitic load on the poor crinoids. The cornulitids were further living on a living snail shell, from what we can tell, so they grew with a substrate slowly growing underneath them. This produces all sorts of delicious paleoecological questions to sort out!
Platyostoma long cornulitid_585Check out the size of this specimen of Cornulites proprius attached to another Platyostoma niagarense. Clearly these tubeworms could do very well under these conditions! This is the largest cornulitid I’ve seen.

Ken_Karns_preparatory_labHere is Ken Karns in his fossil preparation laboratory, which he assembled himself. The box with the armholes is for air-abrading specimens to remove matrix.

Display cases KenThis is one section of the display cases Ken has in his basement museum. Most of the specimens shown here are from the Waldron Shale.

Platyostoma collection displayedA closer view of a display of Platyostoma from the Waldron Shale. Note the many encrusters.

Lizzie Brian KenLizzie Reinthal, Brian Bade and Ken talk about fossil preparation with some Waldron material. The cases are full of curated specimens.

Encrusted crinoid rootsThere are so many treasures in Ken’s collections. I am fascinated by this little slab showing the holdfast of a crinoid with sheet-like bryozoans encrusting it. The bryozoans show that the roots were at least partially exposed at some point.

Thank you again to Brian Bade for arranging this trip, and Ken Karns for being such a fantastic host. We are looking forward to many Waldron projects in the future!


Baumiller, T.K. 2003. Evaluating the interaction between platyceratid gastropods and crinoids: a cost–benefit approach. Palaeogeography, Palaeoclimatology, Palaeoecology 201: 199-209.

Baumiller, T.K. and Gahn, F.J. 2002. Fossil record of parasitism on marine invertebrates with special emphasis on the platyceratid-crinoid interaction. Paleontological Society Papers 8: 195-210.

Brett, C.E., Cramer, B.D., McLaughlin, P.I., Kleffner, M.A., Showers, W.J. and Thomka, J.R. 2012. Revised Telychian–Sheinwoodian (Silurian) stratigraphy of the Laurentian mid-continent: building uniform nomenclature along the Cincinnati Arch. Bulletin of Geosciences 87: 733–753.

Feldman, H.R. 1989. Taphonomic processes in the Waldron Shale, Silurian, southern Indiana. Palaios 4: 144-156.

Gahn, F.J. and Baumiller, T.K. 2006. Using platyceratid gastropod behaviour to test functional morphology. Historical Biology 18: 397-404.

Gahn, F.J., Fabian, A. and Baumiller, T.K. 2003. Additional evidence for the drilling behavior of Paleozoic gastropods. Acta Palaeontologica Polonica 48: 156-156.

Hall, J. 1881. Descriptions of the Species of Fossils Found in the Niagara Group at Waldron, Indiana. In: Indiana Department of Geology and Natural Resources, Eleventh Annual Report, p. 217-345. [PDF of the text downloadable here.]

Liddell, W.D. and Brett, C.E. (1982). Skeletal overgrowths among epizoans from the Silurian (Wenlockian) Waldron Shale. Paleobiology 8: 67-78.

Peters, S.E. and Bork, K.B. 1998. Secondary tiering on crinoids from the Waldron Shale (Silurian: Wenlockian) of Indiana. Journal of Paleontology 72: 887-894.

Sutton, M.D., Briggs, D.E.G., Siveter, D.J. and Siveter, D.J. 2006. Fossilized soft tissues in a Silurian platyceratid gastropod. Proceedings of the Royal Society B: Biological Science 273(1590): 1039-1044.

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

Wooster’s Fossils of the Week: Bioclaustration-boring structures in bryozoans from the Upper Ordovician of the Cincinnati region

February 9th, 2014

Chimneys 149aAnother bioerosion mystery from those fascinating Upper Ordovician rocks around Cincinnati. Above you see a flat, bifoliate trepostome bryozoan (probably Peronopora) with pock holes scattered across its surface. At first you may think, after reading so many blog posts here, that these are again the simple cylindrical boring Trypanites, but then you note that they are shallow and have raised rims so that they look like little meteorite craters. These holes thus represent tiny organisms on the bryozoan surface while it was alive. The bryozoan grew around these infesters, producing the reaction tissue of the rims. This is a kind of preservation called bioclaustration (literally, “walled-in life” from the same root in claustrophobia and cloisters). The specimen is from locality C/W-149 (Liberty Formation near Brookville, Franklin County, Indiana; 39º 28.847′ N, 84º 56.941′ W).
Chimneys 153aThis is another trepostome bryozoan with these rimmed pits. It is from locality C/W-153 (Bull Fork Formation near Maysville, Mason County, Kentucky; 38º 35.111′ N, 083º 42.094′ W). The pits are more numerous and have more pronounced reaction rims.
Chimneys 153bA closer view. One of the interesting questions is whether these pits are also borings. Did they cut down into the bryozoan skeleton at the same time it was growing up around them? We should be able to answer that by making a cross-section through the pits to see what their bases look like. The bryozoan walls should be either cut or entire.
Chimneys 153cThis is an older image I made back in the days of film to show the density of the rimmed pits in the same bryozoan as above. If we assume that the pit-maker was a filter-feeding organism, how did it affect the nutrient intake of the host bryozoan? Maybe the infester had a larger feeding apparatus and took a larger size fraction of the suspended food? (This could be a project where we apply aerosol filtration theory.)  Maybe the bryozoan suffered from a cut in its usual supply of food and had a stunted colony as a result? These are questions my students and I plan to pursue this summer and next year.

It is good to get back to the glorious Cincinnatian!


Ernst, A., Taylor, P.D. and Bohatý, J. 2014. A new Middle Devonian cystoporate bryozoan from Germany containing a
new symbiont bioclaustration. Acta Palaeontologica Polonica 59: 173–183.

Kammer, T.W. 1985. Aerosol filtration theory applied to Mississippian deltaic crinoids. Journal of Paleontology 59: 551-560.

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

Rubinstein, D.I. and Koehl, M.A.R. 1977. The mechanisms of filter feeding: some theoretical considerations. American Naturalist 111: 981-994.

Tapanila, L. 2005. Palaeoecology and diversity of endosymbionts in Palaeozoic marine invertebrates: trace fossil evidence. Lethaia 38: 89-99.

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

Wooster’s Fossils of the Week: Mysterious borings in brachiopods from the Upper Ordovician of the Cincinnati region

February 2nd, 2014

Half borings 152a1Above is a well-used brachiopod from the Upper Ordovician of northern Kentucky (C/W-152; Petersburg-Bullittsville Road, Boone County; Bellevue Member of the Grant Lake Formation). It experienced several events on the ancient seafloor during its short time of exposure. Let’s put a few labels on it and discuss:

Half borings 152a2Our main topic will be those strange ditch-like borings (A) cut across into the exterior of this brachiopod shell. This is an example of bioerosion, or the removal of hard substrate (the calcitic shell in this case) by organisms. These structures were likely created by worm-like filter-feeders. The shell also has a nice trepostome bryozoan (B) encrusting it (and partially overlapping the borings) and the heliolitid coral Protaraea richmondensis (C), which is distinguished by tiny star-like corallites. The borings are what we need to make sense of in this tableau. Here’s another set on another brachiopod:

Half borings 152bThis closer view of a brachiopod shell exterior from the same locality shows two of these horizontal borings. The mystery is why we see only half of the boring. These are apparently cylindrical borings of the Trypanites variety, but they should be enclosed on all sides as tubes. Why is half missing? It is as if the roofs have been removed. I think that is just what happened.

Half borings 152cThis encrusted and bored brachiopod, again from the same locality, gives us clues as to what likely happened. Here we see an encrusting bryozoan and those borings together. The borings cut through the bryozoan down into the brachiopod shell. Could it be that encrusting bryozoans provided the other half of the borings?

BoringXsectHere’s a test of that idea. Above is a cross-section through the boundary between an encrusting bryozoan (above) and a brachiopod shell (below). It was made by cutting through the specimen, polishing it, and then making an acetate peel. The bryozoan shows the modular nature of its colonial skeleton, and the brachiopod displays its laminar shell structure. The two round features are sediment-filled borings running perpendicular to the plane of the section. The boring on the left is completely within the brachiopod shell; the one on the right is cut along the interface of the bryozoan and brachioopod. Remove the bryozoan and we would have a half-boring as discussed above.

Half borings 152eIf that postulate is true, it means that the encrusting byozoans must have been removed from the brachiopod shells, taking the other halves of the borings with them. We should thus find bryozoans that “popped” off the shells with the equivalent half-borings on their undersides. You know where this is going. The bryozoan above (same locality) shows its upper surface. Note that there are a scattering of tiny borings punched into it.

Half borings 152fThis is the underside of the bryozoan. We are looking at its flat attachment surface. It was fixed to a shell of some kind (I can’t tell what type) and became detached from it. You see the half-borings along with vertical borings drilled parallel to the attachment surface. It appears that small organisms drilled into the bryozoan zoarium (colonial skeleton) on its upper surface, penetrated down to the boundary with the brachiopod shell, and then turned 90° and excavated along the boundary between brachiopod and bryozoan. This makes sense if they were creating a dwelling tube (Domichnia) that they would want surrounded by shell. Punching straight through the bryozoan and brachiopod would leave them in a tube without a base. What would this look like from the inside of the brachiopod shell?

Half borings 152dThis time we’re looking at the interior of a brachiopod shell (same location) that has been exfoliated (some shell layers have been removed). The horizontal borings are visible running parallel to the shell.

Horizontal in bivalveThis view of an encrusted bivalve shell may help with the concept. In the top half you see an encrusting bryozoan. In the bottom you see bivalve shell exposed where the bryozoan has been broken away. Cutting into that shell are the horizontal borings. Their “roofs” were in the now-missing parts of the bryozoan.

There are two conclusions from this hypothesis: (1) There was a group of borers who drilled to this interface between bryozoan and brachiopod skeleton, detected the difference in skeleton type, and then drilled horizontally to maintain the integrity of their tubes; (2) the bryozoans were cemented to the brachiopods firmly enough that the borers could mine along the interface, but later some bryozoan encrusters were removed, leaving no trace of their attachment save the half-bored brachiopod shell. This latter conclusion is disturbing. A tacit assumption of workers on the sclerobionts (hard-substrate dwellers) of brachiopods and other calcitic skeletons is that the calcitic bryozoans cemented onto them so firmly that they could not be dislodged. We could thus record how many shells are encrusted and not encrusted to derive paleoecological data about exposure time, shell orientations and the like. But if the robust bryozoans could just come off, maybe that data must be treated with more caution? After all, bryozoans that were removed from unbored brachiopods could leave no trace at all of their former residence.

Two students and I presented these ideas at a Geological Society of America meeting eight years ago (Wilson et al., 2006), but we never returned to the questions for a full study. Now a new generation of students and I have started a project on this particular phenomenon of sclerobiology. It will involve collecting more examples and carefully dissecting them to plot out the relationship between the borings and their skeletal substrates. We also want to assess the impact these observations may have on encruster studies. Watch this space a year from now!


Brett, C.E., Smrecak, T., Hubbard, K.P. and Walker, S. 2012. Marine sclerobiofacies: Encrusting and endolithic communities on shells through time and space, p. 129-157. In: Talent, J.A. (ed.), Earth and Life; Springer Netherlands.

Smrecak, T.A. and Brett, C.E. 2008. Discerning patterns in epibiont distribution across a Late Ordovician (Cincinnatian) depth gradient. Geological Society of America Abstracts with Programs 40:18.

Wilson, M.A., Dennison-Budak, C.W. and Bowen, J.C. 2006. Half-borings and missing encrusters on brachiopods in the Upper Ordovician: Implications for the paleoecological analysis of sclerobionts. Geological Society of America Abstracts with Programs 38:514.

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