Tuesday, August 2, 2011

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"


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.


Pavel said...

It could be a good idea to compare the bullae from medial (not only from dorsal) view. More differences could be found.


Robert Boessenecker said...

I have made the comparisons - we only illustrated them in dorsal aspect in our poster because we only had room for a couple of figures. The problem is that many of these taxa have incomplete bullae, with the outer lip broken, so features in medial aspect aren't very obvious to begin with.

Pavel said...

Medial aspect is not very diverse at all, so all the differences are very distinct (overall shape, swallen involucral region, etc). Anyway Herpetocetus species seem to differ at your figure! bramblei is very good.

Groucho said...

Yuur work is really great. I am on tha other side of the planet esrth meking the same investigatios. Here also we have small baleonepteridae an balaenidae. I would talk about this.

I am senior researcher from Univarsity of Valencia.
My main work it's abour paleobiology an paleoecology of marine mammals (cetacean, sirenia and pinnipedia -sensu lato- ranged from miocene pliocene thru lower pleistocene. My degree is biologist, geologist and etc.

I've maked some serious studies about paleoecology of this environments at the mediterranean area.
Now I am constructing two museums centered o that stuff.


Davidow said...

Hi Bob,

With respect to Mizuhoptera, while you tossed M. sendaicus as dubious, you didn't mention that there were two other nominal species of Mizuhoptera, M. fujinensis and M. kanayaensis, which are also based on isolated earbones. You may not have had the opportunity to examine the holotypes of these species, but would it reasonable to assume that M. fujinensis and M. kanayaensis are dubious and referable to Herpetocetinae indet. like M. sendaicus?

By the way, congratulations on your paper describing Herpetocetus morrowi! As it stands, the original description of H. morrowi is the first paper to test the phylogenetic position of Peripolocetus, which has been recently placed at Chaeomysticeti incertae sedis.

Robert Boessenecker said...

Hello Davidow, I haven't read Hatai et al. in a while, but the impression I had was that at least one of the other species of "Mizuhoptera" was clearly a balaenopterid whale (i.e. not even from the same family). All, if I recall correctly, are based on incomplete tympanic bullae. While it is true that some bullae are diagnostic to the genus or species level, it's not true for all species, and some have bullae that are damned near identical - thus limiting the diagnostic power of these fragmentary holotypes. The logical solution is to declare them nomina dubia or mysticetes of uncertain affinities.