A new specimen of Parapontoporia

Last fall I made a couple posts detailing an excavation (here and here) of a new odontocete skull from a relatively young (middle-late Pliocene) horizon in the Purisima Formation. This specimen was collected over a six hour period, and the excavation was pulled off and completed just before sunset. The specimen is still not completely prepared, but it does include a complete braincase, the posterior half of the rostrum, both petrotympanics (articulated petrosal and tympanics), and part of one of the lower jaws.

The new skull of Parapontoporia sternbergi in oblique dorsolateral view; bottom photo is labeled.

Three species of Parapontoporia have been described: Parapontoporia pacifica, from the late Miocene Almejas Formation in Baja California, Parapontoporia sternbergi, from the Pliocene San Diego Formation, the Mio-Pliocene San Mateo Formation, and possibly from the Pliocene Wilson Grove Formation, and of course, Parapontoporia wilsoni from the Mio-Pliocene Purisima Formation. Most of the Purisima material is referable to P. wilsoni, which is characterized by a very deep "basin" at the base of the rostrum (shown poorly in the photo below) and by a facial region that is longer than it is wide. P. sternbergi has a shallower basin, and has a facial region wider than it is long (and also appears to me to be smaller in general; P. wilsoni crania can be up to 20% larger than those of P. sternbergi). P. pacifica is not known from a complete braincase, and it has a flat base of the rostrum (i.e. no rostral basin). The new specimen exhibits a shallow rostral basin, and has a braincase that is wider than long, and is damn tiny - all suggesting that it is assignable to Parapontoporia sternbergi rather than P. wilsoni. "But the Purisima Fm. species is Parapontoporia wilsoni!", you might say. Just like modern cetaceans, fossil cetaceans likely had a cosmopolitan distribution - and Parapontoporia sternbergi is primarily known from late Pliocene rocks (specimens identified as P. sternbergi from Miocene strata should probably instead be called Parapontoporia sp.). The type locality of P. wilsoni is about 5.3 Ma, slightly older than the San Diego Fm. Likewise, the new specimen from the Purisima is from a horizon about 10 meters below an ash bed dated at 3.35 Ma, and therefore probably late Pliocene also.

Parapontoporia sternbergi, the most completely known species within Parapontoporia, exhibits an extremely elongate rostrum, filled with about 80 teeth per quadrant; that's a total of 320 teeth. I don't know the specifics for other toothy odontocetes (such as Eurhinodelphinids), but this strikes me as being a terrifyingly large number of teeth for a mammal, and I would not be surprised if Parapontoporia was the toothiest of all mammals. Additionally, the skull of parapontoporia is asymmetrical, unlike Pontoporia and more like the now extinct chinese river dolphin Lipotes. Parapontoporia was obviously named because it is closely related to the La Plata River Dolphin, Pontoporia, right? Right? Believe it or not, this new specimen weighs in on the phylogenetic relationships of Parapontoporia.

A reconstruction of a nearly complete skull from the San Diego Formation referred to Parapontoporia sternbergi by Barnes (1985).

The holotype of Parapontoporia wilsoni from the Purisima Formation.

When Larry Barnes published is major study of Parapontoporia in 1985 (the year I was born...), there were no skulls of Parapontoporia with associated earbones. Because of the similarity of Parapontoporia to modern river dolphins like Pontoporia and Lipotes, he looked through museum collections and tried to identify possible petrosals that could be referred to this new taxon. Oddly enough, one set of six petrosals from the San Diego Formation appeared very similar to modern Lipotes, although Barnes felt that the skull was more similar to Pontoporia.

Isolated petrosals from the San Diego Formation referred to Parapontoporia sternbergi by Barnes (1985).

Because of this discrepancy, there has been some disagreement (not in the published literature - in discussion only) about what the real petrosal of Parapontoporia looks like. At a UCMP visit once, Nick Pyenson showed me another petrosal which he felt might be a better match than the ones Barnes referred. This is a fairly serious matter, because these isolated petrosals have been used in various cladistic analyses to fill in the data matrix - and given the key phylogenetic position of Parapontoporia as a sister taxon to Delphinoidea - means that if the petrosals are not in fact from Parapontporia, there could be some serious errors in previously phylogenetic studies.

The petrotympanic complexes of the new Purisima Formation specimen of Parapontoporia sternbergi in dorsal/cerebral aspect.

The petrotympanic complexes in (near) ventral aspect.

This specimen is the first skull of Parapontoporia with well-preserved petrotympanic complexes associated with it. These petrosals are identical to those referred to this taxon by Barnes (1985), and confirm his referral. Interestingly, this also confirms the Lipotes-like morphology of the earbones. Previous cladistic analyses (e.g. Muizon 1988) have resulted in Parapontoporia being the sister taxon to Lipotes rather than Pontoporia, and several authors have suggested such a relationship, despite the hand-drawn cladogram and assertions of Barnes (1985). This new find indicates the Lipotes-like ear morphology does belong to Parapontoporia.

Petrosals of A) Inia, B) Pontoporia, and C) Lipotes.

Another Purisima Fm. skull of Parapontoporia wilsoni with a petrotympanic complex, in a private collection.

I'd be lying if I said that this were the first specimen ever found with petrosals and tympanics; two other specimens are known. One is a specimen of Parapontoporia pacifica from the Capistrano Formation at the San Diego Natural History Museum, which has a crushed tympanic and petrosal. The other specimen is also from the Purisima Formation, shown above, but has been unavailable for study. The petrosal is barely exposed and stuck within the concretion. This specimen remains in a private collection. This no longer bothers me, as the new specimen has both petrotympanics fully freed from the skull, and one of which has already been CT-scanned at UT Austin for a study by Manuel Martinez and Jonathan Geisler (among other authors, which I am dead last for this minor contribution). Look out for it at SVP!

References and further reading:

BARNES, L. G. 1984. Fossil odontocetes (Mammalia: Cetacea) from the Almejas Formation,
Isla Cedros, Mexico. Paleobios 42:1–46.

BARNES, L. G. 1985. Fossil pontoporiid dolphins (Mammalia: Cetacea) from the Pacific coast
of North America. Contributions in Science, Natural History Museum of Los Angeles
County 363:1–34.

FORDYCE, E., AND C. DE MUIZON. 2001. Evolutionary history of cetaceans: a review. Pages
169–233 in J. -M. Mazin and V. de Buffrenil, eds. Secondary adaptation of tetrapods to
life in water. Verlag Dr. Friedrich Pfeil, Munich, Germany.

GEISLER, J. H., AND A. E. SANDERS. 2003. Morphological evidence for the phylogeny of
Cetacea. Journal of Mammalian Evolution 10:23–129.

MUIZON, C. de. 1988. Les relations phylog`en´etiques des Delphinida (Cetacea, Mammalia).
Annales de Paleontologie 74:159–227.

PYENSON, N. D. 2009. Requiem for Lipotes. Marine Mammal Science 25:714-724.

Return to the thanksgiving odontocete locality

Over winter break Chris Pirrone and I returned to the locality where, a month prior, we had excavated a complete odontocete skull. If you recall, when we collected the specimen we had to excavate and pedestal the skull sideways, rather than the normal way (downwards) that most paleontologists are used to. When I had seen the skull at the end of the summer, it was only a few feet above the sand. I knew that over thanksgiving, there would be a very good chance that the sand would all be gone due to the rough weather which tends to erode away the beaches.

Luckily, at thanksgiving there was a three foot-wide 'rim' of the summer beach left, and closer to the ocean, the beach sand was about 8 feet lower. Fortunately, we had just enough time to collect the fossil. When we returned in January, this sliver had been completely eroded away, leaving the excavation pit about 9 feet above the beach and well out of reach. In the above photo, you can see it about left center.

And here I am pointing to it. It would still have been accessible by ladder, but that would have been a real pain, and I'm pretty sure after standing on a ladder for five hours, it's almost a guarantee that you'd fall off.

Comparison between Thanksgiving 2010 and January 2011. The two photos are taken from nearly the same position (note the rock in the foreground) but the more recent photo is taken from further away from the cliff.

Otherwise, after initial preparation, the fossil appears to be a juvenile Parapontoporia, with a partial dentary, and both tympanoperiotics. The tympanoperiotics are now fully prepared, and were articulated, and sitting above the bony nares and the vertex of the skull (i.e. in contact with the dorsal surface). As the skull was upside down, the earbones must have fallen out, and then the skull rolled over on top of them. I'll have pictures soon.

Fossil preparation - odontocete tympanic

About two or three summers ago I collected a beautiful little odontocete tympanic from the Purisima. Problem was, I only found out it was beautiful (past tense) after it sat in about twenty or thirty pieces. Because the part that was exposed looked like some other type of bone (and not an odontocete tympanic) I mis-estimated how sturdy the fossil was, and it exploded as I carved matrix away from it. I have since not repeated the mistake. Anyway, the fossil has sat in pieces in a plastic bag for two years, and I finally got the courage to glue it back together. I say courage because 1) I was somewhat embarassed by this damage, and 2) I was nervous to piece together all these tiny fragments. Below is a photo of what I had to work with.

Tympanic fragments prior to preparation.

I began by finding the pieces of the robust involucrum, which is the thick portion of the cetacean tympanic. There are more or less three major portions of the tympanic: the involucrum (frequently the only preserved part), the posterior process, which attaches to the posterior involucrum, and the paper-thin involucrum (which in odontocetes is usually under 1-1.5mm thick, hence the overall fragility of these elements). Then, I started finding matching pieces, and gluing these to the involucrum.

The tympanic after 30 minutes of preparation; some of the outer lip fragments have been glued in place. Lateral view (top photo) and anterior view (bottom photo) - note the matrix-free tympanic cavity.

After a couple more hours, I was able to finish gluing back most of the outer lip of the tympanic, as well as the posterior process.

Tympanic before (left) and after (right) addition of the sigmoid and posterior process.

Tympanic in dorsal (left) and lateral (right) aspects.

After the fossil was glued together, it became very obvious that this was a tympanic from the "river dolphin" Parapontoporia wilsoni, which has a small posterior process, a sharp anterior apex of the bulla, and most characteristically a laterally inflated outer lip, not seen in any other Purisima odontocete (for which tympanics are known, and out of the given tympanic sample from the Purisima Fm.). As far as crania, jaws, periotics, tympanics, and parts thereof go, Parapontoporia is by far the most common Purisima odontocete (i.e. between collections at UCMP, SCMNH, and LACM go, there are roughly a dozen nearly complete crania known, rostrum not included).

Newly prepared tympanic (right) side by side with another very well preserved Parapontoporia wilsoni tympanic.

All in all, I was extremely pleased; in one afternoon I had turned a pile of fragments (which I had virtually no hope for) into a beautiful little specimen. All but three tiny fragments under 5mm in size were glued on; the other ones probably attached to the margin of the outer lip, which may require fragments lost during collection (or, conversely, pieces pulverized). But let's not split hairs here - this by far was the most damage I've ever done to an odontocete tympanic, to the point where I was embarassed to even think about it; and now, it's one of the nicest I have. The positive side to this inadvertently destructive mode of

collection was that all the matrix was absent from the tympanic cavity, unlike the specimen on the left in the above photo (where the matrix inside was actually phosphatized, but phosphatic 'cementation' had fortuitously not formed an overgrowth around the rest of the bulla - best case scenario!). This is one of about a dozen and a half or so odontocete bullae I've recovered from the Purisima.

New lower jaw of the extinct lipotid Parapontoporia

Before I dive back into fossil odobenids, I'd like to show off something I just finished preparing. I collected this specimen from underwater in July in Santa Cruz County; I arrived at the exposure several days after I initially discovered it and coated it with vinac. Unfortunately, upon my return when I intended to collect it, the tide wasn't low enough; the specimen was exposed on the apron of a cliff, and was about 6" underwater. Unfortunately, I was there at low tide, and in the intervening days, the sand on the beach had locally eroded, allowing waves to go *just* a bit higher on this 5m stretch of beach.

Oblique view of the lower jaw of Parapontoporia wilsoni.

Anyway, after forty minutes of carving out a pedestal, cursing like mad because I thought I was going to destroy the fossil, and being investigated (and probably secretly laughed at) by a sea otter and a sea lion, I decided to undercut the pedestal. The pedestal was about 14" long and 5" wide, and I was worried that it might crack in half during undercutting - any bone exposed in that crack might fall out (and be swept away by the surf), and then I wouldn't be able to connect the bone from the two pieces of the pedestal back together. Needless to say I was shocked (and endlessly pleased) when the pedestal popped off perfectly.

Dorsal aspect of the fused dentaries of Parapontoporia wilsoni.

Lateral aspect of the fused dentaries of Parapontoporia wilsoni.

Parapontoporia is a very conspicuous member of late Neogene marine vertebrate assemblages in California and Baja California, and has also been reported from Japan. In California, it is known from late Miocene (Tortonian - 9 Ma) through late Pliocene (Piacenzian - 2 Ma) strata, including the San Mateo, San Diego, Capistrano, Pismo , Purisima , and Wilson Grove Formations. Three species are known - Parapontoporia pacifica from the late Miocene Almejas Formation of Baja (Barnes, 1984), Parapontoporia wilsoni from the Mio-Pliocene Purisima Formation (Barnes, 1985), and Parapontoporia sternbergi from the San Diego Formation (Barnes, 1985).

The cranium and jaws of Parapontoporia sternbergi, on display at the San Diego Natural History Museum.

Since description, P. pacifica is still only known from one partial cranium, while P. sternbergi is now represented by about a dozen well preserved crania, and several nearly complete lower jaws. While only the partial holotypic cranium of Parapontoporia wilsoni has been described, however, there are now probably around two dozen crania known, in addition to around 50-75 periotics. Only two lower jaws are known, though - one crappy fragment at UCMP, and a neat (but highly abraded) fragment of an articulated rostrum with teeth at CAS. Two more well preserved jaws are known, both from the early Pliocene of the Purisima - one I collected with my girlfriend in 2006, and the specimen I collected this summer. The 2006 specimen has one tooth, but is better preserved than this specimen.
This specimen may not represent P. wilsoni; the P. wilsoni holotype is about a million years older, and it is certainly possible that crania from this stratum represent P. sternbergi due to their younger age; description of material from the San Mateo Formation is needed to investigate this further. In fact, a huge body of fossils of Parapontoporia need to be described.

Closeup of the teeth of the new jaw of Parapontoporia wilsoni.

Parapontoporia was originally named for its similarity to the extant La Plata River dolphin, or Franciscana (Pontoporia blainvillei; Barnes, 1984, 1985). However, subsequent studies have placed it within the Lipotidae, as the sister taxon of the now extinct Yangtze River Dolphin (Lipotes vexillifer; Geisler and Sanders, 2003; Muizon, 1988), which was only described in 1918. Parapontoporia has an extremely long rostrum and mandibular symphysis, and *may* have the most teeth of any mammal (which, if it isn't Parapontoporia, I'm sure it's some kind of eurhinodelphid or other longirostrine odontocete from the Chesapeake Group of the east coast).

Wherever Parapontoporia occurs, it dominates the odontocete assemblage - in the Purisima, up to 38% of isolated periotics are referable to Parapontoporia. The most abundantly known odontocete crania from the Purisima belong to this taxon. Interestingly, despite decades of construction in San Diego, there are now more crania of this taxon known from the Purisima than from the San Diego Formation. Many of these Purisima crania are still in concretions, but nonetheless, they exist, and an excellent opportunity for a study of ontogenetic and stratigraphic variation is possible given this sample (a project Nick Pyenson was bugging me to do, but I simply didn't have the time as an undergrad). In fact, I picked up two partial crania this summer (both in nodules, though; one weighed about 55 pounds).

Nick Pyenson (2009) recently published a pretty neat (albeit depressing) paper in marine mammal science about the consequences of the extinction of Lipotes, given its 'colorful' evolutionary history. But this post is long enough as is, and I could do several more posts just on Parapontoporia; I'll save discussion of that paper for later.

BARNES, L. G. 1984. Fossil odontocetes (Mammalia: Cetacea) from the Almejas Formation, Isla Cedros, Mexico. Paleobios 42:1–46.
BARNES, L. G. 1985. Fossil pontoporiid dolphins (Mammalia: Cetacea) from the Pacific coast of North America. Contributions in Science, Natural History Museum of Los Angeles County 363:1–34.
GEISLER, J. H., AND A. E. SANDERS. 2003. Morphological evidence for the phylogeny of Cetacea. Journal of Mammalian Evolution 10:23–129.
MUIZON, C. de. 1988. Les relations phylog`en´etiques des Delphinida (Cetacea, Mammalia). Annales de Paleontologie 74:159–227.
PYENSON, N.D. 2009. Requiem for Lipotes: an evolutionary perspective on marine mammal extinction. Marine Mammal Science 25:714-724.