Wooster’s Fossil of the Week: An almost planispiral gastropod from the Middle Jurassic of southern Israel

August 11th, 2013


Discohelix tunisiensis apical copyAdd this to the list of fossils that have confused me. This summer, during a Wooster expedition, Lizzie Reinthal and Steph Bosch collected the above specimen from the Matmor Formation (Middle Jurassic, Callovian) of southern Israel. I simply assumed it was an ammonite, especially because we were anxious to find ammonites to further reinforce our biostratigraphic framework (how we tell in which geological time interval our fossils belong). When I later tried to identify it by searching through the Jurassic ammonite literature, though, I could find nothing like it. I then sent a photograph to my friend Zeev Lewy, a prominent ammonite expert recently retired from the Geological Survey of Israel. His answer was a surprise: this fossil is the gastropod Discohelix tunisiensis Cox 1969.
Discohelix tunisiensis adapicalHow could this be a snail when it looks so much like a cool, multi-whorled planispiral ammonite, complete with ribs? Well, it is not planispiral, now that I look at it again. Above you see the other side of the specimen, with its slightly depressed center. Most ammonites don’t show such asymmetry. This actually is a gastropod, and it represents an ancient group (the clade Vetigastropoda — don’t get me started on the complications of gastropod systematics!) with primitive features reminiscent of Paleozoic marine snails (from a group I learned to call “archaeogastropods“). It is not as much that the snail has converged on an ammonite style of shell, it’s that the ammonites developed a similar shell much later for entirely different reasons (swimming, for example). Discohelix was likely an herbivore grazing in patchy coral reefs like we have represented in the Matmor Formation. It has become a useful index fossil for the Jurassic of the Tethyan Realm, although this is the first time I’ve found it in Israel.
Pseudotorinia (Architae-group) retiferaThe above is the marine snail Pseudotorinia (Architae-group) retifera. It used to be called Discohelix retifera, and you can see why. It may not be in the same genus, but you can see that this modern group and Discohelix are closely related. Discohelix itself is now known only from the fossil record.
DunkerDiscohelix was named as a genus in 1847 by Wilhelm Bernhard Rudolph Hadrian Dunker (1809-1885), a German natural scientist with interests in geology, paleontology and marine zoology. (I love that middle name of “Hadrian”.) Like so many 19th Century paleontologists, Dunker started with a practical training in mining engineering and then followed a passion for fossils and modern shells. He had a huge collection of materials that eventually ended up in the Museum für Naturkunde in Berlin. He traded and corresponded with many top scientists of his day, including Charles Darwin. He also published many monographs on modern and fossil molluscan taxa. In 1846, he and Hermann von Meyer established the journal Palaeontographica. This journal survives to this day in two descendants: Palaeontographica A (Paleozoology, Stratigraphy) and Palaeontographica B (Paleobotany).


Cox, L.R. 1969. Gasteropodes Jurassiques du Sud-Est Tunisien [Jurassic gastropods from SE Tunisia]. Annales de Paleontologie, Invertebres 55: 241-268.

Grundel, J. 2005. The genus Discohelix Dunker, 1847 (Gastropoda) and on the content of the Discohelicidae Schroder, 1995. Neues Jahrbuch fur Geologie und Palaontologie-Monatshefte 12: 729-748.

Tëmkin, I., Glaubrecht, M. and Köhler, F. 2009. Wilhelm Dunker, his collection, and pteriid systematics. Malacologia 51: 39-79.

Wendt, J.1968. Discohelix (Archaeogastropoda, Euomphalacea) as an index fossil in the Tethyan Jurassic. Palaeontology 11: 554-575.

Wooster’s Fossil of the Week: An irregular echinoid from the Middle Jurassic of southern Israel

August 4th, 2013

Holectypus depressus adoral 585From the view above, this fossil from the Matmor Formation (Middle Jurassic, Callovian) of southern Israel looks like your standard echinoid (a group that contains sea urchins and sand dollars), but turn it on its side (see below) and you see it is unusual. Echinoids have two large categories: those that are globular in shape (like sea urchins) are called “regular“, and those that are flattened (like sand dollars) are “irregular“. (I know, oddly value-laden terms, these.) This specimen belongs to a group that is rounded on its top portion and flattened on its bottom (oral) surface. This between-ness makes it a fun little specimen.
Holectypus depressus Side 585I could not identify this echinoid, which we collected on our Israel expedition this summer, because I could not find the most important diagnostic features. Fortunately my colleague Andrew Smith, recently retired from the Natural History Museum in London, quickly knew what it was. (This is not surprising — he’s the world’s expert on fossil echinoids. Check out his incredible Echinoid Directory.) Andrew identified this specimen as belonging to the genus Holectypus Desor, 1842, and probably the species Holectypus depressus (Leske, 1778).
Holectypus depressus apical system 585The first feature Andrew noticed was the apical disk on the very top of the test (the term for an echinoid skeleton). In the above image (where the black scale bar = 200 microns) I’ve labelled the four gonopores (where gametes exit, as you might have guessed) and the madreporite (a sieve plate at the opening of the water vascular system). This arrangement is characteristic of the genus.
Holectypus depressus oral 585Most surprising to me was Andrew’s identification of the most obvious defining feature of Holectypus, the periproct (the anal opening). I couldn’t find it, but Andrew knew where to look. In this view of the oral surface, it is the gap labeled “P”. Looks just like a place where the test is broken, right?
Callovian France HolectypusHere is the oral surface of an unbroken Holectypus specimen from the Callovian of France. The large periproct is immediately visible as the whole at the bottom. Now the broken margin of the periproct on our specimen makes sense.

Holectypus belongs to a group of irregular echinoids still around today. They are sometimes characterized as having “conservative” evolution, meaning they have not changed much over long periods. The irregular echinoids appeared earlier in the Jurassic as a modification of their regular ancestors. They became flattened and bilateral, the periproct moved out of the apical disk, their ambulacra (rows of tube feet visible as tiny holes radiating from the apical disk on the top image) pulled up away from the mouth, and their spines were reduced in size and increased in number. These were primarily adaptations for burrowing into the sediment. Holectypus has retained its inflated upper portion, has relatively large spines (some still cling to our specimen), and still is circular in outline. It was a deposit feeder but not specialized for burrowing.

We wouldn’t want to call this a “transitional fossil”, but it is a nice example of the gradient of adaptations present when there is a major outbreak of innovation as during the rise of the irregular echinoids in the Jurassic.


Kroh, A. and Smith, A.B. 2010. The phylogeny and classification of post-Palaeozoic echinoids. Journal of Systematic Palaeontology 8: 147-212.

Rose, E.P.F. and Olver, J.B.S. 1985. Slow evolution in the Holectypidae, a family of primitive irregular echinoids, p. 81-89. In: Keegan, B.F. and O’Connor, B.D.S. (eds.), Proceedings of the Fifth International Echinoderm Conference, Galway, 24-29 September, 1984.

Saucède, T., Mooi, R. and David, B. 2007. Phylogeny and origin of Jurassic irregular echinoids (Echinodermata: Echinoidea). Geological Magazine 144: 333-359.

Wooster’s Fossil of the Week: An infected crinoid from the Middle Jurassic of southern Israel

July 28th, 2013

CrinoidGalls03 copyThis weathered beauty is a stem fragment of the articulate crinoid Apiocrinites negevensis from the Matmor Formation (Middle Jurassic, Callovian) of the Negev, southern Israel. The regular divisions you see making up the stem are the columnals, which look a bit like a stack of poker chips. You can even make out the crenulations on the articulating faces of the columnals, seen as tiny zig-zags. What is unusual about this stem, of course, are the large swellings with multiple holes. These appear to be something like the galls you sometimes see in plant stems formed when a parasite is surrounded by living plant tissue.
CrinoidGalls02 copySenior Independent Studies student Lizzie Reinthal (’14) is working on these odd structures (we have dozens of examples) as part of her investigation of the taphonomy of A. negevensis in the Matmor Formation. We know that the swellings were made by the interaction of some sort of organism with the living crinoid, but we don’t yet know the timing or mechanism. It could be that the holes were drilled first into the stem and the crinoid grew the extra skeletal tissue to essentially push them away, or the swellings could have been the equivalent of galls and some sort of enclosed animal bored its way out of the structure. (And an extra point to those of you who spotted the barnacle boring! Note that it has no swelling around it and thus was likely drilled after the death of the crinoid.)

These infected crinoid stems were first described from the Matmor by Feldman and Brett (1998). They suggested they were from parasitic myzostome worms, which are usually found on crinoid arms and have a long fossil record (see Meyer and Ausich, 1983, and Hess, 2010). They could also be from some sort of embedded organism like that represented by Phosphannulus on Paleozoic crinoid stems (Welch, 1976).

Lizzie will be pursuing the mystery by careful sectioning some of these swellings and seeing if she can relate the crinoid skeletal growth patterns to either a borer or an embedded parasite. Unfortunately that means we must destroy some specimens to better understand the phenomenon, a classic dilemma paleontologists sometimes face.


Feldman, H.R. and Brett, C.E. 1998. Epi- and endobiontic organisms on Late Jurassic crinoid columns from the Negev Desert, Israel: Implications for co-evolution. Lethaia 31: 57–71.

Hess, H. 2010. Myzostome deformation on arms of the Early Jurassic crinoid Balanocrinus gracilis (Charlesworth). Journal of Paleontology 84: 1031-1034.

Meyer, D.L. and Ausich, W.I. 1983. Biotic interactions among recent and fossil crinoids, p. 377–427. In: Tevesz, M.J.S. and McCall, P.L., eds., Biotic interactions in recent and fossil benthic communities. Plenum Press, New York.

Welch, J.R. 1976. Phosphannulus on Paleozoic crinoid stems. Journal of Paleontology 50: 218-225.

Wooster’s Fossils of the Week: Dinosaur footprints of unknown provenance

July 14th, 2013

DinosaurFootprintConcaveEpireliefThese are the only fossils in the Wooster collection I feel some shame about. They are tridactyl theropod dinosaur footprints. They are not spectacular, but they do the job for classes and visits by schoolchildren. I regret that we have them, frankly, because it means at some time in the distant past (well before me) someone chopped them out of a red sandstone, losing in the process all context including location and age. Footprints like these are most valuable, both scientifically and aesthetically, in their original places alongside the rest of the trackway. Now I can only guess that these footprints came from the Jurassic of the southwest, probably Arizona or Utah.

At least we can use them for some trace fossil terminology. (A trace fossil is evidence of organism behavior, such as a trail, burrow, boring or scratch marks.) The top image is what we expect from a fossil footprint: an indentation in the top of the sandy bed now turned to stone. It extends into the rock and is on the surface, so we call it a concave epirelief. (“Epi-” means surface.) The footprint below is also of a three-toed theropod dinosaur, but it extends out of the rock and is on the underside of the bed. We thus call it convex hyporelief. (“Hypo-” means underneath.)

WaterFilledFootprintOne quick way to see the outlines of an indistinct footprint is to fill a concave epirelief with water, as above. This provides much more contrast.
WilsonBoysFootprintThis is how dinosaur footprints should be appreciated: in the field as part of trackways. The kids above are three of my brothers a long, long time ago. I think this is near Tuba City, Arizona, and these are Jurassic theropod footprints like the Wooster specimens. Note that to increase contrast the footprint is filled with … let’s just say “water”.


Miller, W.E., Britt, B.B. and Stadtman, K. 1989. Tridactyl tracks from the Moenave Formation of southwestern Utah (pp. 209-215). Gillette, D.D., and Lockley, M..G, eds., Dinosaur tracks and traces: Cambridge, Cambridge University Press.

Goodbye, Makhtesh Gadol (for now)

July 7th, 2013

GoodbyeMakhteshGadol070713MITZPE RAMON, ISRAEL–Today Team Israel 2013 had its last visit to Makhtesh Gadol, marking the end of Lizzie Reinthal’s and Steph Bosch’s fieldwork. We collected our last specimens from the Matmor Formation, which is exposed only in the center of this magnificent structure. The students above are looking down into the northern part of the makhtesh from the viewpoint at Mount Avnon.

MakhteshGadol070713This is a view from the same spot looking south along the western wall of the makhtesh. You can see a bit of the curvy, narrow road below that we used to enter and exit the makhtesh. This road was built by the British during World War II when they thought there might be oil underneath this breached anticline.

MG_BronzeAgeStructure070713This ring of stones is the remnant of a Bronze Age livestock pen, along with a probable small shelter for the shepherd in the lower left. This features are found throughout Makhtesh Gadol, usually up on the flanks of the walls or the Matmor Hills. I happened to come across this one in today’s walkabout.

LastCollecting070713Finally, here are students collecting at our last site — the southernmost exposure of the echinoderm-rich subunit we’ve found to be so productive. This morning we found … wait for it … two more bryozoans! This is usually not big news in most of the places I’ve worked, but it sure is here. They are again runner-types, but not Stomatopora. Much more to report on these after we get back home with them.

We still have five more working days in Israel. One will be devoted to finishing Oscar’s fieldwork, one to finishing smaller projects in the Makhtesh Ramon area, one to exploration of geological sites Yoav has chosen for us, one for a trip to Jerusalem and the headquarters of the Geological Survey of Israel, and a final day to make sure we’ve done all we came to do.

Bryozoans finally make an appearance in the Jurassic of southern Israel

July 5th, 2013

MatmorCollecting070513MITZPE RAMON, ISRAEL–The concentrated effort of all four Wooster Geologists in Israel finally paid off in fossil bryozoans today. Steph Bosch (center) is studying the bryozoans of the Matmor Formation (Middle Jurassic) in Makhtesh Gadol for her Independent Study project. We have bryozoan specimens collected on earlier expeditions to the Matmor, but on this trip have been stymied in our efforts to add to the meager collection. Bryozoans are rare in these equatorial sediments, even though the diversity of other fossils here is very high. We want to describe this fauna for the first time and place it in its environmental and evolutionary context.

This afternoon we visited a site at the far northeastern end of the makhtesh where we knew lots of crinoids were preserved. (The locality is creatively known as “GPS 004″; N 30.94916°, E 35.01110°.) Our working hypothesis is that bryozoans preferred the same conditions as echinoderms in the Matmor Sea. The strategy worked: we found two runner-type bryozoans on crinoid columns. They are the stomatoporid-type of uniserial runner, giving us at least two different species of bryozoan now from the Matmor. Undoubtedly there are more such specimens on the many crinoid ossicles we collected to wash and examine later in the Wooster lab. They are far too small to photograph here in my hotel room!

Steph, appropriately, found the first specimen. Oscar Mmari (on the left) found the second just as we were leaving. Lizzie Reinthal (on the right) is actually working on crinoid taphonomy, so she was very pleased by all the specimens we collected today.

MatmorCollectingView070513A wider view of our collecting site in the Matmor Formation, with the northeastern wall of Makhtesh Gadol in the background.

It feels good to now have all three Wooster seniors with materials they’ve collected for their Independent Study projects. A nice way to end our first week of work in Israel!

Phosphate success while Jurassic bryozoans remain elusive

July 4th, 2013

OscarOZsection070413MITZPE RAMON, ISRAEL–Oscar Mmari celebrated on his second outcrop today the completion of his stratigraphic columns for his project on the phosphate-bearing portions of the Mishash Formation in the Negev of southern Israel. It is certainly the least pretty section we’ve worked with because it is in the industrialized mining zone near the eastern outlet of Makhtesh Gadol, but we got the work done. This particular place is at N 30.94072°, E 35.03784° (for those of you following along). Look at this place on Google Earth and you’ll see how extensive the phosphate mines are here. Oscar is not yet done with fieldwork because we will come back to his sections for further observations and collecting.

LizzieStephMatmor070413Later in the afternoon we visited outcrops of the Matmor Formation in the northern part of Makhtesh Gadol. Just above Lizzie Reinthal’s right shoulder above (she’s in the center) the outlet from the makhtesh is visible as a break in the surrounding walls. Steph Bosch on the right is bravely hiding her disappointment because after lots of careful peering at the encrusted undersides of beautiful fossil corals, the four of us failed to come up with a single bryozoan. This is curious in itself. The encrusters are well preserved and diverse. (Olev Vinn and I wrote a paper on the various serpulid and sabellid worm tubes; Cezary Krawczynski and I have a paper on the thecideide brachiopods from here.) Every place we think we will see bryozoans we find instead worm tubes and calcisponges. The few Matmor bryozoans collected on previous field trips have been on echinoderm ossicles and holdfasts. Since echinoderms are common where corals are not, and vice versa, it may be that the bryozoans preferred the same conditions the crinoids and echinoids did. Later on this trip we will thus have a full-court press in the crinoid thickets and echinoid-rich beds.

PrettyCorals070413Still, the corals here are gorgeous. Here are two varieties of typical colonial scleractinians we saw today. The one on the right has all sorts of encrusters in the nooks and crannies between its corallites. The corals here are remarkably well preserved considering that their original aragonitic skeletons have been replaced by calcite. These are a bit of consolation for the missing bryozoans!

Fossil collecting under a desert sun

July 2nd, 2013

GroupCollecting070213MITZPE RAMON, ISRAEL–Today the Wooster Geologists in Israel collected fossils from the Matmor Formation in Makhtesh Gadol. It was time to begin assembling data for Lizzie Reinthal’s and Steph Bosch’s Independent Study projects, and to follow up on some projects I have going in this Middle Jurassic set of fossiliferous marls and limestones. We do all our work as a team so that everyone is helping with everyone else’s project at some point. In the image above we have everyone at work picking up fossils at one outcrop near the center of the makhtesh (N 30.93369°, E 34.97588°, to be exact).

LizzieCollecting070213Lizzie is working on crinoid taphonomy, which we’re approaching in a very systematic way. She laid out individual square-meter boundaries on her fossiliferous exposures and collected every bit of crinoid present, from single columnals (the disks that make up a crinoid stem) to partial calices (the crowns or “heads” where the feeding arms were attached). She even seived the sediment to get a sample of small fragments like those from the arms. This way we can assess how the crinoids were disarticulated and tossed about on the seafloor, and whether their stems had some characteristic detachment horizons.

MatmorMakhteshGadol070213_585We are here in the Matmor Hills looking south towards the distant wall of the makhtesh. Two geologists are collecting from a yellowish marl. What do you think that small red dot is in the center?

OscarCollecting070213Why it is Oscar Mmari showing his flair for innovation! He brought the umbrella not for rain (that’s not going to happen here in July, for sure) but just for this. He isn’t much for hats, and we all must admit he had the best shade all day.

StephCollecting070213And here is Steph Bosch carefully collecting tiny little specimens — hoping, hoping some will be encrusted by bryozoans. (None yet, but the best sites are yet to come.)

BittenSpines070213So what did we find? I can show you only a few things because fossil photography in a hotel room is more difficult than you might imagine. (Can you guess what background I’m using?) At one location in the Matmor Formation we collected every exposed echinoid spine in an attempt to statistically determine how many were bitten by some sort of predator. You can see that the spines displayed above have been chomped, probably by some sort of fish. Spines like these were part of an earlier Fossil of the Week post.

Nautiloid070213Yoav found this  nautiloid in subunit 52 of the Matmor Formation. The entire shell, including the septa (walls dividing the chambers), has been replaced with a fine-grained calcite. What are the two holes apparently punched through the shell? They could be just erosion, but I like to think they’re bite marks from some toothy reptile! [Added note: I think I’ve just identified this nautiloid. It has a distinctive groove on the venter (not visible here) which may place it in the genus Cymatonautilus. This genus lived only a short time (latest early to middle Callovian Stage of the Jurassic Period) and is found along the shallow shelf of the Tethys. Right interval, right place.]

Field reconnaissance in the northern Negev of Israel

July 1st, 2013

1FoldedPhosphates070113MITZPE RAMON, ISRAEL–This morning Team Israel 2013 met our friend Yoav Avni, a geologist with the Geological Survey of Israel (GSI), and we traveled north to our field localities. We did a survey of the sites so that we could put together an efficient schedule for our work over the next two weeks. We had a four-wheel drive vehicle from the GSI so we could get to places our little Budget rental car could only have nightmares about.

The first locations were for Oscar Mmari’s Cretaceous phosphorite work. The outcrop pictured at the top of this entry is on the east side of Makhtesh Gadol (N 30.93657°, E 035.03312°). We are looking toward the west at an incredibly asymmetric limb of a syncline. In the upper part of the exposure you can see the rocks dipping almost vertically, yet in the foreground they are nearly horizontal. They make an almost 90° bend. The Mishash Formation phosphatic zone is partly exposed as the white rocks along the side of the wadi. The phosphorites here are very thick and chalky.

2MishhashPhosphates070113A second phosphorite exposure for Oscar is in Wadi Havarim (N 30.84269°, E 34.75509°) not too far north of Mitzpe Ramon. The top of the cherty portion of the Mishash Formation is on the left in the middle; the light-colored units above are phosphorites. In the background is Nahal Zin, a deep valley formed by water draining north into the Dead Sea. The base level of the Dead Sea is so low that the wadis leading to it are rapidly downcut.

3OscarPhosphorite070113Here is Oscar getting is first look at the phosphorites at Wadi Havarim. Later this week we will measure at least one section at each locality and take plenty of samples for thin-sectioning and scanning electron microscopy.

4TraceFossil070113Steph Bosch’s hand gives us a scale for a nice set of trace fossils found in the phosphorite at Wadi Havarim. These look like callianassid shrimp burrows to me. We found some preserved as burrow-fills with apparent fecal pellets forming the outer walls. If true then the trace fossil ichnogenus is Ophiomorpha. This is a good indicator of shallow water.

5PhosphateSign070113We also briefly visited two phosphate mining sites east of Makhtesh Gadol. One has this helpful sign outside describing the geology of these deposits. The phosphorites are shown in yellow. Note that they formed in two synclines, both highly asymmetrical (as shown in our top photo).

6PhosphateMinedValleyWe viewed one phosphate mine where virtually the whole valley has been quarried, producing enormous piles of waste materials. Reclaiming mined terrain like this is especially difficult in this arid climate. Oscar will not only be looking at the geology of these phosphate deposits, but also the economics of mining, which now includes remediation and controls on emissions and water pollution.

7MatmorCollectingFirstDay070113After lunch we drove down into the center of Makhtesh Gadol and plotted out future localities for Steph and Lizzie to do their work in the Matmor Formation. (The above site is at N 30.93837°, E 34.97907°.) I’ve been to these sites many times with students, so it was relatively easy to make our plan for collecting crinoids and encrusting bryozoans tomorrow and next week.

8NabateanCistern070113Finally, no fieldwork in Israel is complete without a touch of archaeology. Yoav took us into a Nabatean cistern and showed us the clever engineering (and strategic plastering) necessary to make this hand-cut cavern into a water trapping and storage facility. This cistern is cut into the Menuha Formation, a chalk unit very familiar to Andrew Retzler (’11). This cistern was originally made sometime between 100 BCE and 100 CE. After the nabateans it was used by the Romans, Byzantines and Arabs. It was last used by Israeli pioneers over 60 years ago.

Tomorrow we return to Makhtesh Gadol and work in the hot sunlight filling collecting bags with tiny bits of crinoids and assorted encrusters. We’ve had a very good start.

Wooster’s Fossil of the Week: A barnacle-bored belemnite from the Jurassic of Spain

June 23rd, 2013

Hibolites_barnacle_borings585Two subjects of previous Fossil of the Week posts are combined together here: a belemnite (the elegant Hibolites hastatus) and barnacle borings (the ichnofossil Rogerella). This specimen is from the Jurassic of Moneva Teruel, Spain.

(Living belemnites reconstructed by Bogdanov on Wikipedia)

(Living belemnites reconstructed by Bogdanov on Wikipedia)

Belemnites are extinct cephalopods, oddly enough. The specimen is the guard or rostrum — a calcitic internal skeleton that gave the squid-like animal rigidity. Because they are made of solid, stable calcite, these guards can be extremely common in the fossil record, especially in the Jurassic and Cretaceous. Some people call these “belemnite battlefields”, probably because the fossils look so much like bullets.

de_BlainvilleHibolites hastatus was named by Henri Marie Ducrotay de Blainville (1777-1850) in 1827. He was a French naturalist with many accomplishments, especially in anatomy and systematics. He spent most of his career in the Faculty of Sciences at Paris, where he was a colleague (and eventual ideological enemy) of the epic Georges Cuvier. In 1830 he took over from the “father of invertebrate zoology” Jean-Baptiste Lamarck as the chair of natural history. Clearly he ran in powerful circles.

The yellow features in this close-up above are holes drilled by acrothoracic barnacles, which were then filled with fine-grained dolomite. The trace fossil thus formed is known as Rogerella. It is found from the Devonian all the way to today. Its presence on these belemnite guards shows that these structures laid for enough time on the seafloor that they could be colonized by barnacles. They are thus an indicator of the taphonomy (or history from death to discovery) of these fossils.


Doyle, P. and MacDonald, D.I.M. 1993. Belemnite battlefields. Lethaia 26: 65-80.

Lambers, P. and Boekschoten, G.J. 1986. On fossil and recent borings produced by acrothoracic cirripeds. Geologie en Mijnbouw 65: 257–268.

Mariotti, N. 2002. Systematics and taphonomy of an Early Kimmeridgian belemnite fauna from the Mediterranean Tethys (Monte Nerone, Central Apennines, Italy). Geobios 35: 213-232.

« Prev - Next »