Shark-bitten dolphin skull

In 2008 I spent the day before Christmas Eve shivered on a cold, wind-blasted California beach prospecting for vertebrate fossils in the Purisima Formation. I was home on winter break, and although it is far more cold where I go to graduate school in Montana (as I write this I'm looking out at the results of our first winter snow), nothing is worse than being wet and miserably cold out on the foggy, windy coast of the golden state (except perhaps being wet and miserable on the Oregon coast, which I've done).

The thrill (or promise) of discovery is more than enough to keep me fueled in the field during the winter. Indeed, when the birds start singing and the snow melts in the spring, most paleontologists start to get field fever - the field season for most vertebrate paleontologists is during the summer months. Anyone who's ever tried to do coastal fieldwork during the summer, on the other hand, is in for a rude awakening. No erosion takes place during the summer, and many of the outcrops are totally buried. The exposures that are above the beach sand level (which is higher during the summer) are typically covered with dust, sand, and grime, which obscures fossils. The storms in the winter months clean this nasty coating off, and transport beach sand into offshore bars, often exposing strata below the beach (I see new fossil localities every winter this way). Winter is my field season.

Historically, I've had really good luck the day before Christmas Eve. It's my last day before Christmas to make it out in the field. The prior year, I found a humongous Carcharocles megalodon tooth (the only specimen known from the Purisima Formation), and discovered a partially articulated fur seal skeleton.

The Christmas Eve dolphin.

At 4pm, the tide was beginning to come back in, and with little over an hour of daylight, it was looking like I was going to come home empty-handed. I went to one last cove before I turned around to head back to the beach. I walked for a few minutes and spotted something in a boulder I had not seen on my way out: a pair of flat bones joined along an articulation that looked suspiciously (even from 20 feet away) like the palate of a dolphin skull. Upon closer examination, yes indeed! It was a dolphin skull in a mollusk shell bed; the width and flatness of the palate suggested it was not Parapontoporia, the most common odontocete in the Purisima Formation. I set about chopping into the boulder; fortunately, most of it was relatively soft. However, an extremely hard calcium-carbonate cemented concretion the size of a basketball had formed over the dorsal surface of the braincase and rostrum, and this slowed digging down. By dusk, the concretion didn't budge. After another half hour, it finally popped out of the boulder, and I lugged the 45 pound block back to my car. Exhausted, I drove home, drank a couple of hard-earned beers with dinner, and passed out.

View of the facial region of the skull.

When it came time to go back to Montana, I decided I would rather take the fossil as a carry-on than risk checking it and picking up a broken fossil that I had paid 25 bucks for thanks to baggage fees. After arriving in Bozeman (with a very sore back and neck from lugging 65 pounds

of luggage through the Denver airport), I almost immediately began preparation (starting, of course, with acetic acid baths for several weeks to soften the concretionary matrix). It took about two months to prepare, and as you can see from the above photos, it is damn beautiful. I initally identified it as something like Haborophocoena - it bears numerous similarities. However, after showing them photos of the specimen at SVP 2009 in Bristol, UK, Olivier Lambert and Giovanni Bianucci both think this represents a basal delphinid rather than a basal phocoenid. I'm inclined to agree with them, although part of my original ID was based on the presence of premaxillary eminences, which this specimen has (a phocoenid character). However, the ascending process of the right premaxilla is in contact with the nasals while the left is not (a delphinid character). Whatever it is, it will require preparation of the ventral aspect, and more careful analysis of the morphology than what I've been able to do thus far. Whatever it is, it appears to represent a new genus and species, and will make a beautiful holotype specimen in the future. During preparation, one curious thing I noticed was a notch in one of the premaxillary eminences (the large pads/bumps in front of the bony nares). I initially dismissed it as a pathology.

The left premaxillary eminence showing linear gouges (red lines) and missing bone.

Upon closer examination (which admittedly did not occur until yesterday, almost two years after collection) it became apparent that the abnormal area had two distinct, paralell linear gouges, and a short, less distinct third one in the middle (this one is still partly filled with matrix). Around these gouges is an area of exposed cancellous bone, where the bone has been removed.

Additional gouges present near the base of the rostrum.

I also found four more gouges present: two long ones, and two short ones; all but one are parallel. In fact, aside from the one gouge seen above trending towards the upper left corner of the photo, all the gouges are parallel. This is a textbook set of shark-inflicted bite marks. There are a lot of papers on this in the literature, documenting shark bites on dolphins, baleen whales, pinnipeds, sea turtles, other shark teeth, mosasaurs, plesiosaurs, dinosaur bones, sea stars, and probably other marine critters as well.

In fact, the first record of these types of trace fossils were actually first documented in the modern environment: on predated and scavenged sea-otter carcasses from Monterey Bay, and reported by Ames and Morejohn (1980). The reported linear gouges, subparallel wavy small gouges, and a specimen including a shark tooth embedded in a sea otter skull. The morphology of the traces along with the tooth identified the culprit as the Great White Shark, Carcharodon carcharias. Two years later, these exact types of traces were identified by Tom Demere and Richard Cerutti (1982) on a baleen whale dentary (of my favorite whale, Herpetocetus!), and identified as "Carcharodon sulcidens" (a taxon now just considered to be fossil Carcharodon carcharias).

It's not clear what type of shark fed on my poor little dolphin, or if it was a case of predation or scavening; from what I've read, the majority of carcasses that exhibit bites have bite marks on the posterior portion of the body, which is just about as far as you can get from the face. This makes total sense, given how a shark would have to bite into a fleeing dolphin during pursuit. Furthermore, it's interesting to note that this bite would have had to go clean through the dolphin's melon (if it had not already decomposed). Anyway, I interpret these traces as drag marks from the apices of the shark's teeth; I suppose later on I can figure out the relative motion of the shark's mouth during the bite (most likely lateral shake feeding). It'll make for a nice short paper some day...

Ames, J. A., and Morejohn, G.V., 1980, Evidence of white shark, Carcharodon carcharius, attacks on sea otters, Enhydra lutris: California Fish and Game, v. 66, p. 196-209.

Deméré, T.A., and Cerutti, R.A., 1982, A Pliocene shark attack on a cetotheriid whale: Journal of Paleontology, v. 56, p. 1480-1482

Benthic feeding in basal mysticetes, part 1: paleopathology of a Miocene "cetothere"

The last couple years have been relatively good for cetacean paleontology; we saw the description of the protocetid Maiacetus (which I still have to cover...) about a year ago, the rediscription of "Balaenoptera" gastaldii, now recognized to be a fossil gray whale, the fantastic analysis done by T.A. Demere et al. regarding the presence of baleen in toothed mysticetes, Larry Barnes' Albireo monograph, the odontocete cranial lexicon by Jim Mead and Ewan Fordyce, the taphonomic study of the Sharktooth Hill Bonebed by Nick Pyenson and colleagues, Frank Whitmore and L. Barnes' Herpetocetus monograph, Steeman's (2009) paper on Uranocetus, Brian Beatty and Alton Dooley's paper on paleopathology in the Carmel Church Diorocetus, and most recently Erich Fitzgerald's paper on Mammalodon, among many others which I've probably failed to remember.

For the purposes of the next couple posts, I'll be focusing on basal mysticete feeding, and will be discussing Beatty and Dooley (2009), Fitzgerald (2010), and Demere et al. (2008).

The pathologic left dentary of Diorocetus from the Carmel Church Quarry in Virginia, From Beatty and Dooley (2009). The pathologic fracture is directly below the 'c' in 10 cm.

For those of you who pay close attention to Alton Dooley's blog "Updates From the Vertebrate Paleontology Lab", Dooley does quite a bit of fieldwork at the Carmel Church Quarry, an exposure of the middle Miocene Calvert Formation. The Calvert Formation in Maryland and Virginia is famous among amateur paleontologists and fossil collectors for the stunning abundance of easily collected fossil shark teeth. The Calvert Formation is also famous for its incredible cetacean fossil assemblage - primarily chronicled by Remington Kellogg, the father of marine mammal paleontology (although there have been a number of papers recently on Calvert Fm. and Chesapeake Group cetaceans). One of the recent discoveries at Carmel Church is a beautiful skeleton of the archaic mysticete Diorocetus. This skeleton includes a complete skull (which initially was fragmented to hell, but Dooley and the VMNH have managed to put all of the pieces back together, and it looks pretty damn nice), dentaries, anterior vertebral column, and nearly complete set of ribs.

Fracture and pathology in the left dentary, from Beatty and Dooley (2009).

The left dentary was found to have an odd fracture in it; the two parts didn't match up very well, and it did not appear to be a post-depositional fracture, like most of the fractured material at Carmel Church (it is not clear if the jumbling and fracturing of bones is biostratinomic or diagenetic - e.g. peri- or post- burial). Additionally, a callus of bone was identified around this fracture, and can be seen well in x-rays (above). Additional pathologies were noted in the anterior tips of the premaxillae, and the left squamosal, which was significantly less dense and more porous than the right (potentially due to decreased stress during post-injury feeding?). Most interesting is the fact that although the callus formed, the fracture never healed, suggesting repetitive use that kept the bone from healing, i.e., the anterior and posterior portions of the dentary remained as separate elements until death, allowing some motion at the fracture site.

What could cause this sort of a fracture? The authors indicate the most common cause of these sorts of injuries in extant mysticetes are collisions with ships and boats - which obviously did not exist in the middle Miocene. Other possibilities include predation, agonistic (violent) intraspecific behavior, and a collision or impact with seafloor topography. If this is a case of predation, then our friend survived, given the amount of healing (i.e. callus formation). Agonistic behavior among mysticetes is poorly documented, and are largely restricted to injuries on the order of cuts and scrapes. Otherwise, the authors conclude, lies the chance that this injury was caused by an impact with the seafloor, or submarine outcrop (i.e. much of the California coastline I'm used to has rocky points and sea stacks and a topographically complex seafloor with many submarine exposures of rock). Apparently, injuries of this sort are the most commonly observed trauma on dead gray whales, which are benthic suction feeders.

Before I continue, I'll add a quick note about mysticete feeding. Among modern mysticetes, there are three observed modes of feeding: Lunge/engulfment feeding, suction feeding, and skim/ram feeding. Lunge or engulfment feeding is mostly utilized by balaenopterid whales (e.g. Blue, Fin, Sei, Minke, and Humpback whales), and is characterized by the whale opening its mouth and engulfing clusters/'schools' of nektonic organisms (i.e. krill, fish, etc.). The mouth is closed, and water is actively 'pushed' out of the baleen plates (I can't remember if both the tongue and throat are used for this action, or one or the other). Skim feeding is employed by balaenids (and the sole existing neobalaenid, Caperea, the pygmy right whale), and consists of the whale slightly opening its mouth while swimming forward; nekton prey-rich water enters the oral cavity, and during forward movement, water flows passively out of the oral cavity through the baleen, which traps the poor critters inside the mouth; this feeding is often at or near the surface. Benthic feeding, on the other hand, is only observed in the gray whale (Eschrictius robustus), which will filters through muddy substrate; fine sediment is entrained in suspension, and can escape with water through the baleen, trapping benthic organisms (i.e. amphipods) in the oral cavity. The major problem is that the most basal known fossil mysticetes retained teeth, and did not yet have baleen. More on that later, though.

Cross sections of mysticete ribs; A-B is the new Diorocetus specimen; C - undescribed basal edentulous mysticete, Oligocene, Oregon; D-E-toothed mysticete Aetiocetus cotylaveus; F & I - balaenopterids (latter is Eobalaenoptera); G-H - cetotheriid (sensu stricto) Metopocetus; K - Diorocetus hiatus; L- Balaena ricei. All of these have osteoclerotic ribs, with the exception of Eobalaenoptera and Balaena ricei, which are part of the mysticete crown group.

Another interesting feature Beatty and Dooley (2009) noted was the osteosclerotic condition of the ribs in Diorocetus. In contrast, extant cetaceans have postcranial bones that are osteoporotic (yes, like menopausal women). For the purposes of this discussion, there are two types of bone: cancellous, and cortical (spongy and dense, respectively; there are many other types, which I won't go into here; read papers by de Ricqles and Horner for more info on paleohistology). Cortical bone (or the cortex) is the strong, outer portion, while cancellous bone is the very spongy middle part. Osteosclerosis refers to increasing bone density by adding cortical bone toward the center of the bone, making the cancellous inner portion thinner. Pachyostotic bone is where cortex is increased outward, giving the bone an 'inflated' look - sirenians have pachyostotic (and osteosclerotic) bones. Osteoporosis simply refers to bone that is very porous, and generally lense dense - this is simply a condition; in cetaceans it is 'normal', but in adult women it is a bad condition which can lead to fractures. Osteosclerosis, on the other hand, can act as a sort of 'bio-ballast' adaptation for maintaining (or simply attaining) neutral bouyancy - most terrestrial vertebrates are positively bouyant, especially in seawater. Champsosaurs, I just learned in Jack Horner's class, have retained super-dense embryonic bone into the adult stage as a ballast adaptation.

Beatty and Dooley (2009) observe that Diorocetus hiatus is one of the last mysticetes to retain osteosclerotic bone, and that it may be related to bouyancy problems associated with benthic feeding. Indeed, osteosclerotic bone is a plesiomorphic feature among not only mysticetes, but is also characteristic of pelagic archaeocetes as well (I am not referring to the clade Pelagiceti, by the way). So, it is certainly possible that this is an adaptation for benthic feeding. However, it is also possible that this is a case of phylogenetic inertia, similar to the retention of an enlarged mandibular foramen in mysticetes.

The nature of the likely cause of the mandibular injury may also suggest benthic feeding as well (unless this was a freak accident; i.e. a lunge feeding whale impacting the seafloor). While the repetitive feeding behavior that kept the fracture from healing may have been caused by benthic feeding, *if* Diorocetus had been a lunge feeder, the incredible stresses experienced by mysticete dentaries during this action would certainly keep the fracture from healing. In any event, taken as a whole, the benthic feeding idea is very intriguing, and raises some very interesting questions regarding the primitive mode of feeding by baleen-bearing mysticetes. I understand that the authors have received criticism for some of the more speculative ideas in the paper, you'll find none from me; this study brings up some very interesting ideas, and I'll be covering more on the topic of benthic feeding on my next post, regarding the enigmatic toothed mysticete Mammalodon.

Also see:
Alton Dooley's blog post about the article, and Brian Beatty's post as well.


Beatty, B.L. and A.C. Dooley. 2009. Injuries in a mysticete skeleton from the Miocene of Virginia, with a discussion of bouyancy and the primitive feeding mode in the Chaeomysticeti. Jeffersoniana 20:1-28.