Recent fieldwork in the Purisima Formation, Part 2: a possible new species of Herpetocetus?

Back in late May, Dick Hilton (Sierra College) and I did a three day field trip collecting fossil vertebrates from a locality in the Purisima Formation I recently got a permit for. On the second day, excavated a large block of sediment with what I assumed at the time were sirenian bones. I had not seen the bones except in cross section, and because they were somewhat dense, I thought they might be from a sea cow. We quickly carved out a large block, and due to the cohesive nature of the sediment, we were able to wrap it in tinfoil and duct tape. It was one of the first finds of the day, and I thought there could definitely be something neat inside. Because we were only a half mile from the cars, I left my pack with Dick and hoofed it back to the car with the thirty pound block, and also to grab some gatorade I had left in my car. When I returned, Dick was taking a siesta, and after some lunch, we headed further down the beach. Only a few hundred feet down I spotted a distinctly potato-shaped thing sticking out next to a piece of bone: it rather looked to me like a tympanic bulla, and I climbed up a bit to check it out. It was in fact a tympanic, and when the rest of the piece came out, I was able to see that it was in fact a nearly complete squamosal, complete with the bulla and posterior process of the petrosal. Dick and I thought the trip had been a success just because of this specimen, especially because it was from a section of cliffs where neither of us expected to find anything.

Dick Hilton digging up a huge baleen whale tympanic.

I couldn't have been more wrong. Unfortunately, I didn't know that I was until after the SATLW (Aquatic Tetrapods) conference. I did have a day or two before the conference to prepare the squamosal, and it did indeed have a plug-shaped posterior process of the petrosal, indicating it belonged to the Herpetocetinae, which includes Herpetocetus, Nannocetus, (probably) Piscobalaena, and Cephalotropis (according to Steeman, 2007). After the conference, I opened up the duct taped jacket and began preparation. After a couple hours the exposed pieces were still not making sense, and then I found a couple of bones that looked like they were adjacent to one another. When I removed them, there was a tiny neck of bone connecting them – and after a little more preparation, I realized it was a Herpetocetus petrosal and posterior process. Damnit, another goddamn Herpetocetus.

The petrosal and posterior process of the new specimen, with the facial nerve canal labeled. Upper left is ventral, lower left is dorsal, and right is medial view.

The skull with (partially incorrectly) articulated petrosal of the new skull in dorsal (top) and ventral (bottom) views.

Once I had enough of the block prepared, I realized I had quite a bit of the ventral portion of a small braincase preserved. It includes both exoccipitals, one occipital condyle, the basioccipital, the right squamosal, and the complete petrosal. After preparation, the petrosal is most similar to petrosals of Herpetocetus. This may be a bit technical, but herpetocetine baleen whales have several peculiar features that define them as a group. The posterior process of the petrosal – which is typically an elongate strap of bone that connects with the skull posteriorly – is very short and plug-shaped in these animals. Additionally, the posterior process (which is rarely found attached in isolated fossil mysticete petrosals) is flat and contributes to the lateral side of the skull, instead of being 'hidden' in a trench between the squamosal and exoccipital bones. Secondly, some herpetocetines have a flattened anterior process that is blade shaped; this structure is typically conical and robust or knoblike in most other mysticetes. Clearly, this specimen exhibits both of these features. Additionally, Herpetocetus spp. exhibit a large triangular flange on the side of the bone, which overhangs the squamosal – also present in this specimen. Additionally, herpetocetines all have extremely small earbones relative to most mysticetes. Unfortunately, the neck of the posterior process appears to have been deformed slightly, and when the main portion is articulated correctly, the posterior process sits in its trough a little wonky, and when the posterior process is articulated correctly, the main portion doesn't articulate well.

The posterior process, squamosal, and tympanic of Herpetocetus bramblei.

The two alternate articulations of the petrosal showing correct articulation of the posterior process (left) and correct articulation of the body of the petrosal (right).

The temporal region of the skull of Herpetocetus bramblei with the petrosal outlined in red.

However – it shows several features that differentiate it from all species of Herpetocetus as well as other herpetocetines like Nannocetus and Piscobalaena. Firstly, the anterior process is medially oriented – it is usually anteriorly facing instead. Second, the posterior process is very transversely narrow and elongate – it is typically more nearly circular in other species. Lastly, the most bizarre feature is that it has a very long anterior fissure of the facial nerve canal which is contorted into an S-shape – something I have not seen in any mysticete, fossil or modern.

Various mysticete petrosals in ventral view, showing two fossil rorquals (Plesiobalaenoptera and Balaenoptera sursiplana), a modern balaenid (Eubalaena japonica), the new specimen, and two other Herpetocetus specimens.

This is pretty exciting, and I am looking forward to preparing the other specimen, which includes part of a squamosal and a tympanic, and most likely a petrosal. It should not be too difficult to get these specimens written up and described.

Further Reading

Geisler, J. H. & Luo, Z.-X. 1996. The petrosal and inner ear of Herpetocetus sp. (Mammalia: Cetacea) and their implications for the phylogeny and hearing of archaic mysticetes. Journal of Vertebrate Paleontology, 70, 1045–1066.

Steeman, M.E. 2007. Cladistic analysis and a revised classification of fossil and recent mysticetes. Zoological Journal of the Linnean Society 150:875–894.

Steeman, M.E. 2010. The extinct baleen whale fauna from the Miocene-Pliocene of Belgium and the diagnostic cetacean ear bones. Journal of Systematic Palaeontology 8:63-80.

Whitmore, F.C., and L.G. Barnes. 2008. The Herpetocetinae, a new subfamily of extinct baleen whales (Mammalia, Cetacea, Cetotheriidae). In C.E. Ray, D.J. Bohaska, I.A. Koretsky, L.W. Ward, and L.G. Barnes (eds.). Geology and Paleontology of the Lee Creek Mine, North Carolina, IV. Virginia Museum of Natural History Special Publication 14:141–180.

More problems with Herpetocetus

Back in June at the Aquatic Tetrapods conference I coauthored a poster with Joe El Adli (San Diego Natural History Museum) and Jonathan Geisler (New York College of Osteopathic Medicine) on some of the taxonomic problems of Herpetocetus. Herpetocetus, as I've mentioned before, is an enigmatic small bodied mysticete whale which many bizarre and derived features, while retaining some primitive features as well. Fossils of Herpetocetus are fairly common in Northern California, particularly in the Purisima Formation - or maybe I just have a knack for finding them. Thus far, there is only one described species of Herpetocetus from California: Herpetocetus bramblei, named by Whitmore and Barnes (2008) from a very partial skull (basically just a squamosal with part of the exoccipital, parietal, and pterygoid) with a petrosal from the Purisima Formation. In summer 2007, I excavated a nearly complete skull of this same species from near the type locality, and last summer, I excavated a second specimen which lacked the braincase but included a complete rostrum. Since this topotypic material was collected, additional specimens from other localities in the Purisima Formation indicate that two additional undescribed species are present - one of the new species was discovered very recently, and I'll have more on that soon.

*Holotype, for the non-specialist, is the specimen which a new species is based off of. It should be representative of the new species in terms of its anatomy, and should be relatively complete enough to be comparable to other taxa. A type locality is where the holotype specimen originated.

There are several other described species of Herpetocetus from other corners of the globe - all from the Northern Hemisphere. The genus was first described from the Pliocene of Belgium (Herpetocetus scaldiensis) based on a partial dentary. A partial skull from the Pliocene Yorktown Formation was described as Herpetocetus transatlanticus, also by Whitmore and Barnes (2008). In the 1960's, an isolated tympanic bulla from Japan was named as the type specimen of Mitzuhoptera sendaicus, and a fossil mysticete skeleton with a skull, earbones, and dentary shared both the dentary morphology of Herpetocetus scaldiensis as well as the tympanic morphology of Mitzuhoptera sendaicus, and Oishi and Hasegawa (1995) transferred M. sendaicus to Herpetocetus, resulting in the new combination, Herpetocetus sendaicus. Each of these records is from either side of the Pacific (east and west) and the Atlantic (east and west).

How diagnostic are bullae and dentaries? I've already addressed problems with the jaw morphology of herpetocetines (here and here), and mysticetes in general. If you recall, there are two problems concerning the dentary of Herpetocetus spp. in particular: 1) The dentary of the possible sister taxon Nannocetus is not yet known, and dentaries substantially older than Herpetocetus (and possibly belonging to Nannocetus) are nearly identical to Herpetocetus (see below image), indicating that this general morphology is possibly characteristic of a larger group of whales. 2) Some species of Herpetocetus have dentaries that are very difficult to tell apart and lack autapomorphic characters (unique derived features), and thus are not suitable as holotypes. This logically results in the implication that Herpetocetus scaldiensis, which is based on a jaw, is the type species of Herpetocetus, and thus the species and genus may be taxonomically invalid or nomina dubia (means dubious name in latin).

The first figure of our poster, showing comparative drawings of various fossil herpetocetines.

Earbones have long been used for taxonomic purposes, and in many cases have been designated as holotypes. Sir Richard Owen designated many isolated bullae from the Plio-Pleistocene Red Crag of eastern England as holotypes (all of which have been sunk; e.g. Balaena definata). It is unclear how diagnostic earbones are for baleen whales: petrosals (otherwise known as periotics - the inner ear bone) have all sorts holes and knobs and crests and are rather easy to tell apart from genus to genus. A recent paper published by Eric Ekdale, Annalisa Berta, and Tom Demere (2011) indicate that earbones of extant mysticetes are diagnostic to the species and are easily told apart. Additionally, Steeman (2010) reexamined a large suite of earbones previously described by taxonomic mad man P.J. Van Beneden, who is largely responsible for constipating the entire field of mysticete systematics for over 100 years. Steeman (2010) found that many of these earbones - specifically petrosals - may be diagnostic tools, and generally reached a similar conclusion like Ekdale et al. (2011). But what about bullae?

The second figure from our poster, showing variation in tympanic bulla morphology from various herpetocetines. Note the overall similarity between Herpetocetus spp.

Bullae of three species of Herpetocetus have been described: H. scaldiensis, H. transatlanticus, and H. sendaicus. In our poster, we figured all known bullae (described or undescribed), including both the holotype of Mizuhoptera sendaicus and the referred specimen of Herpetocetus sendaicus, and a new bulla of Herpetocetus bramblei. Additionally figured are bullae of Nannocetus and Piscobalaena, also herpetocetines. We concluded, as we hope that you will when looking at this figure, that the bullae of different Herpetocetus species do not vary significantly from species to species. They are, on the other hand, diagnostic at the family level: they are clearly distinct from all other bullae of (described) cetotheriids. However, a bulla that is only distinct at the genus level is inadequate to be used as a holotype. This suggests that Mizuhoptera sendaicus, unsurprisingly, is probably a nomen dubium. It also indicates something interesting is going on with the skulls of mysticetes, or at least cetotheriids: tympanics are slightly less informative than the petrosals. It might be possible someday to quantify how phylogenetically useful different anatomical regions are, aside from just counting up the number of characters used per anatomic region in a cladistic analysis. Who knows, maybe someone has already thought of that and developed a method.

Further Reading:

New published article (Part 1): herpetocetine jaws, and an example of finding a "simple" research project

New published article (Part 2): taxonomic problems with Herpetocetus and "cetotheres"

References:

El Adli, J., Boessenecker, R.W., and J. H. Geisler. 2011. Taxonomic problems of and relationships among species of the fossil baleen whale genus Herpetocetus. Sixth Triennial Conference on Secondary Adaptation of Tetrapods to Life in Water Program with Abstracts: 23.

Ekdale, E.G., A. Berta, and T.A. Demere. 2011. The comparative osteology of the petrotympanic complex (ear region) of extant baleen whales (Cetacea: Mysticeti). PLOS One 6:1-42.

Oishi, M., and Y. Hasegawa 1995. Diversity of Pliocene mysticetes from eastern Japan. The Island Arc 3:436–552.

Steeman, M.E. 2010. The extinct baleen whale fauna from the Miocene-Pliocene of Belgium and the diagnostic cetacean ear bones. Journal of Systematic Palaeontology 8:1:63-80.

Whitmore, F.C., and L.G. Barnes. 2008. The Herpetocetinae, a new subfamily of extinct baleen whales (Mammalia, Cetacea, Cetotheriidae). In C.E. Ray, D.J. Bohaska, I.A. Koretsky, L.W. Ward, and L.G. Barnes (eds.). Geology and Paleontology of the Lee Creek Mine, North Carolina, IV. Virginia Museum of Natural History Special Publication 14:141–180.

Sixth Triennial conference on the Secondary Adaptation of Tetrapods to Life in the Water

Two weeks ago today, I delivered a presentation on the first day of the 2011 Aquatic Tetrapods conference in beautiful San Diego, California. I had eight hours on my drive down to San Diego the day before to worry about how screwed I was: the last time I gave that presentation, it was only 20 slides longer, and took 50 minutes to deliver. I practiced it once Monday morning in front of Joe El Adli (our gracious host), and Yale students Rachel Racicot and Daniel Field - I barely fit it into 20 minutes. It all worked out fine, and in the actual presentation, I finished the last conclusion slide right as it counted down to zero. This was a fresh break from SVP tradition, which dicates that cetacean research is presented on the afternoon of the last day, giving you all week to worry about the presentation. This time, I was able to relax during the entire meeting.

My title slide for my taphonomy presentation.

It would be very difficult to summarize all the research presented, but I might be able to summarize a few of the highlights that stick out in my mind. Julia Fahlke (University of Michigan) gave a fascinating talk Monday Morning about basilosaurid and protocetid cetaceans with asymmetrical crania, and implications for the evolution of hearing underwater. I'll admit, when I read the abstract I was skeptical, but her presentation was pretty compelling - I'll wait to say more until it gets published, though. Brian Beatty presented some details of his research on meningeal ossification in cetacea, which appears to not be homologous to that in many other mammals. Larry Barnes showed us a new Paleoparadoxia skeleton the LACM has been working on from the Monterey Formation; it's virtually complete, with a very large, gnarly looking skull. Olivier Lambert gave a talk coauthored by Giovanni Bianucci on a new large assemblage of bizarre ziphiid fossils dredged from the seafloor off the coast of Spain; boy, there are some real freaks. Our Australian colleague Erich Fitzgerald presented a new juvenile aetiocetid skull from the Oligocene of Washington State which he's been slowly preparing with acid; needless to say, it's a beautiful specimen. Manuel Martinez, a Peruvian who is Christian de Muizon's Ph.D. student, presented on an incredible new toothed mysticete from the Oligocene of Peru - I won't give any details, but lets just say this will be a very, very important specimen.

Frank Fish gives a mini presentation on locomotor adaptations of various marine mammals, using an assortment of articulated limbs (a walrus forelimb is seen in the foreground).

On wednesday, we went over to the San Diego Natural History Museum for an osteology workshop on aquatic tetrapods. Due to the research focus of Tom Demere (Paleo curator) and Annalisa Berta, the majority of material out on display was from modern and fossil marine mammals. There was a great assortment of wonderful stuff out, and it was amazing to be there with so many other marine mammal (and otherwise) researchers there.

Several cetacean researchers are in this photo: Toshiyuki Kimura (foreground), Mette Steeman (behind "Tosh"), and Giovanni Bianucci (background, left) and Joe El Adli (background, right).

At the workshop, I caught Brian Beatty red-handed demonstrating his very technical method to determine the relative height of the bony tentorium (here on a skull of the Amazon river dolphin, Inia geoffrensis).

For a while, Daryl Domning gave a short presentation on the locomotion and forelimb of sirenians (using a Manatee forelimb skeleton).

Here, some of the brightest minds in cetacean paleontology scrutinize one of the weirdest fossil mysticetes: a new species of Herpetocetus under study by Joe El Adli. From left to right: Mette Steeman, Joe El Adli (standing), Felix Marx (leaning over skull), Meredith Rivin (background), Giovanni Bianucci (in glasses), and Olivier Lambert (in red).

Manuel Martinez, a Peruvian researcher studying with Christian de Muizon in Paris, did not waste a chance to photobomb. Here he is photographed with a cast of the skull of the bizarre extinct edentulous* walrus Valenictus chulavistensis.

Daryl Domning again gives a short presentation, this time on sirenian crania. Here he is showing a cast of the skull of the world's largest sirenian, the extinct Pliocene species Hydrodamalis cuestae (the ancestor of the "modern" Steller's Sea Cow).

My (soon to be) coauthor Morgan Churchill photographs an articulated hindlimb of the modern walrus.

Rachel Racicot (and Daniel Field) hung out with me for a bit in the type room at the San Diego Natural History museum while I photographed some fossil pinniped material. Skulls of gigantic Hydrodamalis cuestae sit on the table behind Rachel.

New published article (Part 2): taxonomic problems with Herpetocetus and "cetotheres"

The "cetotheres" have long been a troubled group of fossil baleen whales. Typically, they have throughout there long and confusing taxonomic history been treated as a wastebasket group to include all extinct mysticetes that lack the synapomorphic (i.e. distinguishing) features of the extant groups of baleen whales (gray whales, Eschrichtiidae; rorquals, Balaenopteridae; and right whales, Balaenidae). For a very long time, this group included strange early-diverging mysticetes such as Parietobalaena, Pelocetus, Diorocetus, Aglaocetus (all Miocene mysticetes from the Chesapeake Group of Maryland and Virginia), Cophocetus from Oregon, Cetotherium from the Miocene of the Ukraine, and my favorite mysticete, the problematic Herpetocetus (among others). Many authors during the latter half of the twentieth century doubted that this was a natural grouping, and I suspect that the proliferation of this notion in the literature has more to do with taxonomic laziness on the behalf of mysticete systematists than anything else.

While the taxonomic problems associated with "cetotheres" are a topic for a different post altogether, a brief summary is warranted for the backdrop of the implications in my recently published article. With the advent of cladistics, some studies found that 'cetotheres' are a paraphyletic group of stem-mysticetes (i.e. that they are an unnatural group characterized by primitive rather than derived features). For several years it seemed that the term 'cetothere' should be shit-canned for all eternity, until Bouetel and de Muizon (2006) published a large study on a small Herpetocetus-like 'cetothere' from the Pliocene and latest Miocene of Peru, called Piscobalaena (after the Pisco Formation). They found that some 'cetotheres' form a natural monophyletic group (i.e. a group that is defined on derived features that includes all the descendants of a common ancestor). Because this clade included Cetotherium rathkei, they called this clade the Cetotheriidae sensu stricto, and other 'cetotheres' the cetotheres sensu lato. This same relationship has been supported by several other phylogenetic analyses.

The lectotype jaw of Herpetocetus scaldiensis. From Bouetel and de Muizon, 2006.

Within the true cetotheres, Herpetocetus is the most derived member, and also the youngest surviving member. As previously mentioned, it was based on a lower jaw from the Pliocene of Belgium. When it was described in 1872, a type specimen was never selected, and the lower jaw was selected as a 'lectotype' over thirty years later. The jaw of this animal is pretty distinctive,
Subsequently, many authors have used the distinctive jaw morphology to refer isolated jaws to the genus Herpetocetus. A nearly complete mysticete skeleton from Japan, including a skull, was identified as Herpetocetus due to its jaw morphology. Subsequently, fossils of Herpetocetus have also been reported from the Pliocene and latest Miocene (6-2 Ma) of California, and the early Pliocene (3-5 Ma) of the east coast (North Carolina), basically indicating a 6-2 million year record only in the Northern Hemisphere.

So you can see, when I first thought long and hard about these early Late Miocene (10-12 Ma) Herpetocetus lookalikes, why I was somewhat confused. Herpetocetus also has distinctive earbones and skulls (based on specimens associated with jaws), and there aren't any earbones or skulls with the typical "Herpetocetus morphology" that occur any older than 6 Ma (there is one undescribed skull from the 6.8 Ma Santa Cruz Mudstone I've identified as Herpetocetus aff. bramblei). What is known from the 10-12 Ma Santa Margarita Sandstone, in addition to the jaws in question, is Nannocetus eremus.

The holotype braincase of Nannocetus and a quick and dirty reconstruction based on the rostrum of Herpetocetus sendaicus.

Nannocetus is a really tiny (greatest width across the skull is about 10 inches) weird mysticete, originally described in 1929 by Remington Kellogg. A second specimen from the Santa Margarita Sandstone was described by Whitmore and Barnes (2008), and is the only other known 'true cetothere' from the Santa Margarita. However, Nannocetus is not yet known by a jaw; could Nannocetus be the rightful owner of the two dentaries I described?

If so, then the supposedly distinctive anatomy of the lower jaw of Herpetocetus is not distinctive, and raises important questions about referring isolated dentaries based on their morphology. Additionally, this problem raises an even more important issue: what, then, of fossil baleen whales described solely based on isolated lower jaws? Most of them are probably invalid, because lower jaws *might* only be diagnostic at the supraspecific level (i.e. at the level of a genus or subfamily - whatever the hell those are). "But Bobby, the type species of Herpetocetus is based only on a lower jaw!" Aw, crap. That's right. We've now come full circle: Herpetocetus may or may not be a valid name in the first place, if jaw morphology is insufficient for taxonomic purposes.

Before us mysticete taxonomists go off ready to sink Herpetocetus as a nomen dubium, there are a few important things I pointed out in the article which should be remembered: 1) The fossil dentaries DO show a couple of features distinct from Herpetocetus, including a mandibular foramen with a a lanceolate opening, unlike Herpetocetus. 2) Although highly likely, it is possible that these dentaries are not Nannocetus. However the age discrepancy does mean something in and of itself. 3) Distinctive skull fragments showing some synapomorphies of Herpetocetus were also in the "type series" described in 1872, so it is unfair to say that it was based only on a lower jaw.

With those exceptions in mind, I hope my new article has established some caveats for mysticete workers. Additionally, this work has identified the possibility that mysticete jaws are perhaps diagnostic to the generic level; this still means they are unsuitable as holotypes, but that they are by no means useless - the jaws of mysticetes tell us quite a bit about the animal's feeding and its relationships (although they are not as fine-tuned as, say, parts of the skull). Hopefully future fossil mysticete holotypes will be designated only on material that is really diagnostic, and hopefully will include comparable elements like earbones, braincases, and (also hopefully) the posterior end of the lower jaw.

New published article (Part 1): herpetocetine jaws, and an example of finding a "simple" research project

The new issue of PaleoBios, the paleontology journal published by UCMP at UC Berkeley, includes my new article on herpetocetine jaws from the Santa Margarita Sandstone in central California. During the fall of 2008, I was in my first semester of graduate school, and was taking a difficult, time-intensive course on advanced stratigraphy. At the time, I already had two articles I had been working on: one on my undergraduate research concerning a new fossil vertebrate assemblage from the Purisima Formation, and another on Plio-Pleistocene fur seals from the Wildcat Group in northern California (both are currently in press). However, neither was in any shape to be published anytime soon - one was only half written, and the one that was nearly finished needed a lot of work (i.e. quality control). I needed a new manuscript to work on, to give me something to do that semester aside from stratigraphy, which had started to eat away at my brain. In other words, I needed to start something fresh from scratch, with a clear beginning, a clear end, and a clear message.

The right dentary of UCMP 85431. Scale bar =10cm. From Boessenecker (2011).

A month earlier, I had given a presentation (coauthored with Jonathan Geisler) on a new skull of Herpetocetus bramblei I had collected from the Purisima Formation in 2007. While ruminating on possible projects, I suddenly remembered a partial lower jaw from the Santa Margarita Sandstone that looks a lot like the lower jaw of Herpetocetus*. The problem is, the Santa Margarita Sandstone is early Late Miocene in age (10-12 Ma), while the oldest known bona fide specimens of Herpetocetus are latest Miocene to earliest Pliocene in age (~5 Ma). In fact, the oldest known described specimen is the fragmentary type specimen of H. bramblei, which is right about 5.33 Ma. So, this specimen (UCMP 85429) is MUCH older than any known specimen of a "true" Herpetocetus.

*All species of Herpetocetus have relatively similar lower jaws, and currently the lower jaw has only been described for the type species, Herpetocetus scaldiensis (the type specimen of which is a lower jaw - more on this later).

The left dentary of UCMP 85429; scale bar = 10cm. From Boessenecker (2011).

The lower jaw of baleen whales has long been assumed to be a fairly diagnostic element, at least in certain groups. Herpetocetus scaldiensis certainly has a very distinctive mandible. Other fossil mysticetes certainly have distinctive mandibles as well. Many fossil mysticetes (H. scaldiensis, Balaenoptera davidsonii, Archaeschrictius ruggieroi, etc.) have been described just off of their lower jaws. Are dentaries really that diagnostic? Perhaps. Demere (1986) used mandibular features to reevaluate "Eschrichtius davidsonii" from the San Diego Formation, which was a chunk of a 30% complete mandible, missing the anterior and posterior ends. Being able to refer a new dentary to this taxon, he demonstrated that the "davidsonii" morphotype was actually a rorqual, and assignable to the genus Balaenoptera, which led to its recombination as Balaenoptera davidsonii. Clearly, mandibular morphology is important, and can be used to assess the taxonomy of certain groups, and 'fix' the taxonomy of certain problem taxa. Are these specimens the oldest known records of Herpetocetus?

Reconstructed lower jaw of Herpetocetinae genus and species indet., based on UCMP 85429 and 85431. From Boessenecker (2011).

Reflecting upon the known fossil record of "true cetotheres" - relatives of Cetotherium and Herpetocetus (an entire other topic worthy of its own post) - I was able to formulate some interesting questions I could ask (and attempt to answer), which would make a nice core of an article. With these objectives in mind, I started taking copious notes on the anatomy of the fossil specimens, and eventually typed these notes up into an anatomical description for the article. By the end of the semester, I had a manuscript that was about three-quarters finished. During spring 2009, I wrapped up the discussion, and constructed some figures. I had a few people look at it and make some comments. Morgan Churchill looked at it late in spring, and had some of the most constructive and useful edits. That summer, I TA'd the Geology Field Course for MSU, and during some of the weeks, I took my manuscript out into the field with me, along with a red pen. In between helping students, I sat out in the hot sun, and critically examined every sentence in the manuscript. I even drafted one of the figures (on vellum, with nice pens) around the campfire one night after a few beers. By the end of the second to last project at field camp, I had all my edits finished, and I sent it off to PaleoBios for review.



To be continued...

Boessenecker, R.W. 2011. Herpetocetine (Cetacea:Mysticeti) dentaries from the Upper Miocene Santa Margarita Sandstone of Central California. PaleoBios 30:1:1-12.

Deméré, T.A. 1986. The fossil whale, Balaenoptera davidsonii (Cope 1872), with a review of other Neogene species of Balaenoptera (Cetacea: Mysticeti). Marine Mammal Science 2:277–298.

Trip to New York

New York City and Central Park from the 4th Floor of the
American Museum of Natural History.

A few weeks ago I took a trip to New York in order to be officially hired as a writer for a website on whale evolution that Jonathan Geisler (New York College of Osteopathic Medicine) is collaborating on with John Gatesy (UC Riverside) for an NSF grant. This trip was convenient because Jonathan and I are collaborating on a couple of research projects together, including the description of a fragmentary pilot-whale like skull from the Purisima Formation, as well as working on a large body of Herpetocetus material from Northern California. Once the website is up, I'll go into quite a bit more detail about it.

In 2008 I presented a talk (my first oral presentation) at SVP on a new skull of Herpetocetus bramblei I had found the previous year. The skull is pretty nice, and has about half the rostrum and a complete braincase, with earbones (tympanic, petrosal, stapes, incus, malleus). Because of its completeness and preservation, it will be a daunting task just to describe it. Fortunately (or unfortunately, depending upon how you look at it!) I have since collected another skull with a complete rostrum, and another (slightly less complete) skull from higher up in the Purisima Formation which may represent a younger species. So, including a fragmentary braincase at UCMP, there are four Herpetocetus crania, about a dozen petrosals, several tympanics, a half dozen dentaries, and some postcrania that we need to describe. Anyway, we didn't really work on this project at all.

One day during the trip we went in to the AMNH to look at some modern odontocete crania for comparison with the fossil pilot whale. A year and a half before, when I presented my poster on it at SVP (the 2009 Bristol meeting), I had identified it as Globicephala sp. at the time; fossil delphinid expert Giovanni Bianucci mentioned to me that he was not quite convinced that it belonged in that taxon. At the time I just identified it as best I could, without really having access to crania of other globicephalines like Pseudorca, Peponocephala, and Feresa. During our AMNH trip, we came to a similar conclusion as Giovanni - it's got too many differences to be in Globicephala, as it shares some other similarities with Pseudorca and Feresa.

Jonathan Geisler puzzles over fossil and modern globicephaline skulls.

However, it still definitely falls within the clade Globicephalinae, and that's what's important any way. I'll go into more detail about the ramifications of this fossil later on once we've at least submitted our paper - hopefully sometime early summer I'll have this one (manuscript #6) off.

Here I am (with appropriate shirt for the visit) with a beluga (Delphinapterus) skull.

Perhaps in a later post I'll post some of the pictures I took on a 15 minute dash through the fourth floor exhibits (which was all the time I had before we wanted to leave).

Herpetocetus skull #3, part 3 - Excavation, Day 2

The next day I returned to the locality, and with the help of my buddy Chris Pirrone, we finished undercutting the pedestal; due to the extreme cohesiveness of the sediment, we were able to undercut the pedestal by 3 inches on all sides - which is generally a rare opportunity; with many other fossiliferous units I've worked in, the side of the pedestal would collapse or calve off into the trench.
Within two hours of arriving, we were ready to begin the jacketing process. Above, you can see the top jacket.

Here's a view from the top of the cliff of the plaster jacket and the hole. You can definitely see that with the size of the hole, I was prepared for something sticking further into the cliff.
Soon after, we flipped the jacket over and I began removing excess rock from the base of the pedestal. After removing several inches, I used a jigsaw to cut off the plaster 'lip' that was left from the base of the pedestal.
Chris Pirrone, with the jacket.
And finally, myself with said jacket. I have more pictures of the completed jacket, but they would give away the location, which I'm not terribly keen on doing. All in all, this excavation totaled about 8 hours. By comparison, the well preserved Herpetocetus bramblei skull I collected nearby in 2007 took about 12 hours of excavation, and four hours of jacket transport. Thanks to the much more convenient location of this fossil, jacket removal and transport took a total of two and a half minutes; the tide was way out, so we were able to walk (comfortably) with the jacket to a conveniently placed set of stairs, and then up. The battle Chris and I waged in July 2007 to get the other Herpetocetus skull off the beach is quite a story, and one that I don't think I've detailed here previously.

Anyway, as I probably already mentioned, this specimen includes the first complete rostrum of
a Herpetocetus ever collected, and will feature prominently in my future studies of this enigmatic fossil mysticete.

Herpetocetus skull #3, part 2 - Excavation Day 1

Unfortunately, when I discovered this fossil in its mutilated state, it was two days before I planned on driving back to MSU at the end of the summer, and my last scheduled day of fieldwork for summer 2009, so it was absolutely out of the question to excavate the fossil then. So, I had to return sometime to excavate it. I didn't have the option of doing so over Thanksgiving, Winter, or Spring Break, so I waited a whole year. As it turned out, not much erosion had occurred locally over since initial discovery; just to make sure, I'd visit the fossil every couple of weeks.

I finally had the time to excavate it in mid August; I had some ambitious plans for my last week of summer in California: Saturday and Sunday up at Lake Tahoe, monday and tuesday excavating the fossil, wednesday at the Santa Cruz Museum of Natural History, Thursday at California Academy of Sciences, Friday at University of California Museum of Paleontology, packing up my car on Saturday, and Sunday thru Tuesday on the road.I found out a couple days before hand I would be able to enable to enlist the help of my longtime friend and avocational paleontologist, Chris Pirrone, a local attorney in the San Jose area. Chris helped me excavate the first Herpetocetus skull (which, if you attended my SVP talk in 2008, was the focal point of that research), and has helped me with a few other excavations in the Santa Cruz area.On Monday I was all by myself removing overburden. Surprisingly, after only about four hours of chopping at the cliff face with a railroad pick, all the overburden was gone. So, I started to carefully remove rock an inch at a time by tapping my clam knife with a rock hammer. Eventually, I found the premaxillae, and was able to determine the angle the rostrum went into the cliff (and, more importantly, the fact that the rostrum did in fact exist). If the rostrum paralleled the cliff face, then it would make for an easier excavation, as I would not need to dig as large a hole. Unfortunately, the rostrum dove in there pretty steeply. Here's what it looked like after the first day of the excavation: admittedly there's not much there - and that's exactly the way I prefer it for an excavation. The more fossil bone you expose during the excavation, the higher the chances of damaging that bone. So it's important to figure out during an excavation what the fossil is, which way it's oriented, and roughly where each feature should be; surprises during excavations are usually made by damaging bone.

I've outlined in red the anterior portion of the rostrum (premaxillae are the skinny medial elements, while the maxillae are the wide lateral elements). I exposed some bits of the premaxillae here and there, but just enough to know 1) the angle of the skull going into the cliff and 2) the tip of the rostrum. Oddly enough, the midline of the rostrum didn't match up with the midline of the skull exposed in the cliff. As it turns out, there was a fracture running obliquely through the rostrum - you can see it, near where the red lines end. It's actually offset by about five or six inches.

To be continued...

Herpetocetus skull #3, part 1 - tragedy and mutilation

Hey folks,
Sorry for being absent over the last month or so; I've spent an inordinate amount of time up at Lake Tahoe with my family (i.e. ~5 trips totaling over 1 month) and the rest of the time I spent collecting thesis data and fossils in Santa Cruz, and working on manuscripts. With regards to the latter, I received preliminary acceptance of two different manuscripts in the space of one week; you'll hear more about these papers later. In mid-July, I wrapped up a manuscript I've been working on for 5 years; I decided I was finished with it, and submitted the damn thing. Within the same day, I re-submitted a manuscript for Palaios that was returned for major revisions. And about a week later, I received preliminary acceptance of a paper for JVP on Plio-Pleistocene fur seals (which had only been in review for a month and a half, and another month for the editorial decision, which must be some sort of world record for JVP).

One of my many projects this summer was to collect and excavate a small mysticete skull I had discovered on my last day of fieldwork last summer (2009). Here's what I saw in the field:
Difficult to make out what it is? Well, I had seen this before, but it was covered up with algae, and I just thought it was a concretion. I want you to look at this picture again, and look for all the broken, mutilated bone in the "trench". Done? Now, I've labeled relevant anatomical structures on the same photo, below.
1= Left zygomatic process of squamosal; 2= foramen magnum; 3= right occipital condyle; 4= supraoccipital shield; 5= left supraorbital process of frontal. Yes, that was a complete mysticete skull, keyword: was. I cannot begin to explain how pissed off I was when I saw this. Even worse, then I realized that I recognized that morphology from somewhere:Yes, it was a complete skull of Herpetocetus bramblei. Here's my beautiful specimen from nearby (which was actually collected properly).

So, I don't know who committed this crime, and I don't want to know who it was. I know who it wasn't - I know a half dozen very experienced, extremely competent amateur collectors in the Santa Cruz area who would have never done something this awful to a fossil. I'm not going to use this as an opportunity to scold, though it would be completely warranted. Instead, if you're reading this - let this serve as a plea:

Please don't do this. There are people who can help. I would love to teach you correct excavation methods. This fossil is now mutilated, and the braincase is completely destroyed, and those features and anatomical data are now lost forever. You can still get help. Please, if you find something like this in the future, just ask someone - I'm perfectly happy to help collect it without destroying the fossil or parts thereof.

Additionally, other collectors who do not know me, or if you don't even do collecting in California, it doesn't matter where you are or the color of your fossil: please avoid this at all costs. There are plenty of people willing to help you. Just please, please back away from the edge. I mean fossil.

Partial mysticete jaw (more fossil preparation)

About two years ago I came across a partial lower jaw of a small baleen whale in a cliff. The outwardly bowed part of the jaw was facing the outside of the cliff, and the middle had eroded away, leaving the anterior and posterior ends. I immediately collected the posterior end, which was (and currently still is) in a small nodule, and awaits further preparation. However, what is preserved indicates it belongs to Herpetocetus - my favorite cetacean. The posterior mandible of Herpetocetus is very distinctive; the three main features of this part of the jaw (the coronoid process, mandibular condyle, and angular process) are all anteroposteriorly elongate relative to other mysticetes, with the angular process projecting posteriorly as a flat flange. Anyway, as the specimen isn't prepared yet, I'm not going to show you pictures of the interesting part.

After I collected the posterior part, I tried to excavate the anterior portion; I successfully removed two segments which broke along natural cracks, and a third portion (which I figured at the time was the anterior tip) which was stuck in a nodule, and was rather stubborn. I decided to leave that part in the cliff, and return later under more favorable conditions. The next few visits it remained, and I checked up on it; I figured noone else would disturb it (or even spot it). Well, I got lazy, and over thanksgiving, I returned with the intention of collecting it, and some fresh pick marks were around it, and whoever it was had chipped away a little bit of bone, much to my chagrin. Anyway, I excavated the rest of it immediately; fortuitously the other person had started a trench around the concretion, so it took a mere 5 minutes to carve around it and pop it out - and it did make a 'pop' sound and land with a resounding thud on the beach sand, to which I said to myself "if I had known it would have been that friggin easy, I woulda done it two years ago!".The anterior dentary before and after chiseling.

The problem was, I had no idea whether or not the two pieces would even connect, given the damage done by the other collector, and two years of erosion. And I had to fly with the fossil in my duffel bag all the way back here to Montana to find out. I "gingerly" chipped away some pieces of the concretion with a rock hammer and chisel, which split the rock from the bone perfectly.

The anterior 1/3 of the Herpetocetus mandible.

Then, when I got back to Bozeman, I took the fossil to campus and nervously matched up the two sides - a lot of bone was definitely missing, but (thank god) the two pieces matched up along the ventral side of the bone, preserving the natural length (which is kind of moot anyway, given that I'm missing the middle - kind of, because I measured the missing distance when I was in the field). Some major acid preparation is in order, as well as some serios airscribing and microblasting on all three pieces. These specimens are very similar to mandibles of Piscobalaena, a cetotheriid from the Pliocene of Peru, and the probable sister taxon to Herpetocetus (Bouetel and Muizon, 2006). To be totally honest, the piece of the mandible I've shown you looks pretty damn similar across much of Chaomysticeti (baleen bearing mysticetes), with the possible exception of balaenids.

The mandible of Piscobalaena nana from the Early Pliocene of Peru (from Bouetel and Muizon, 2006)

This is one of five partial Herpetocetus dentaries I've collected from the Purisima Formation (none are currently in museum collections at UCMP or SCMNH), and is the second most complete specimen; one is a complete, ~4' long, very large and robust dentary, and the others are all posterior dentaries (i.e. posterior 1/3). Two of these are extremely small (i.e. one is a fragment where the shaft of the dentary was only 2.5cm high), and likely represent neonates or extremely young individuals. These specimens, along with a nearly complete skull, a partial skull, half a dozen petrosals, and several tympanics will be the subject of a study by Jonathan Geisler and myself. In addition, there are two more possible (one definite) Herpetocetus crania in-situ, which will (hopefully) be excavated over winter break.

For more information on cetotheriids, see Alton Dooley's recent post on cetotheriids at Updates from the Vertebrate Paleontology Lab.

Bouetel, V., and C. de Muizon. 2006. The anatomy and relationships of Piscobalaena nana (Cetacea, Mysticeti) a Cetotheriidae s.s. from the early Pliocene of Peru. Geodiversitas 28:319-395.

Summer Adventures Part 5: new Herpetocetus cranium

Hey gang, I've finally returned from SVP and London, and finally beaten (nearly) this bad cold I've had (several of my office mates keep insinuating that I got swine flu; some other MSU students did - I was not so unlucky). So, in keeping with the attitude of my most recent posts, I will continue on with my summer adventures.

Earlier this summer I made a quiz type post, and never bothered following up on it (except in the comments). Well... for those of you too lazy to figure out what it was, now I'll show you.
I've shown the dorsal surface of the unprepared fossil adjacent to a photo of the well preserved cranium of Herpetocetus bramblei that I presented on at SVP in 200, with certain features lettered. A=zygomatic process of the squamosal; B= vertex; C = occipital condyles; D = exoccipital 'crest'. Herpetocetus, while exhibiting some decidedly 'primitive' features (primitive isn't politically correct, but given the nature of the cetacean cranium, its really convenient, and since this isn't peer reviewed, I don't care about 'PC'), also has a radically telescoped cranium. The radical telescoping isn't very obvious from the posterior cranium, though, and instead the primitive features are more salient. These include the extremely narrow and triangular supraoccipital shield, and the very anteriorly oriented zygomatic processes. Not terribly obvious from this specimen (because they are worn off), but very obvious on the H. bramblei cranium I have are vertically oriented, very well developed lambdoidal crests; these are typically laterally oriented in mysticetes, overhanging the temporal fossa. Now I'm just yapping; maybe I should really focus my chi into writing up that description...

In any event, the new fossil as exposed shows several diagnostic features of Herpetocetus (outlined above[ish]), and some features (such as the narrow supraoccipital) that are unique as far as Pliocene (and latest Miocene) mysticetes go. This last photo shows the slab (very, ver, very) slowly dissolving away in an acetic acid bath. I let it go while I was gone at SVP... and it didn't look any different when I got back. Acetic acid doesn't really degrade Purisima Formation nodules; it pretty much just softens them up to make airchisel prepping easier.

Stay tuned, folks - I still have tales to tell of fieldwork with Dick Hilton in Oregon, shark teeth lodged in sea otter skulls, and some other less interesting stuff I can't seem to remember at the moment.

Oh, and I got to be the 6,000th visitor to this blog - I rule.