Boobies

The northern gannet (Morus bassanus). Photo by Alan Wilson.

The Taxon of the Week post is a day late this week because, of course, yesterday was a public holiday here in Australia (as far as I can make it out, it seems a bunch of poms were so glad to see land after six months in a leaky boat [trying hard to keep afloat] that they've been celebrating ever since - the people already here may have had a different view of matters, but nobody ever asked them). Not only is it late, but it'll be short, too, because the Taxon of the Week is the Sulidae, which has been covered only recently by Darren Naish (here and here) with all the innuendo that is unavoidable when dealing with a group of birds going by the common name of "boobies" (though he did omit mentioning the close connection between boobies and shags).

The Sulidae are the boobies and gannets, a small but fairly cosmopolitan assortment of seabirds. Among the living sulids, the gannets form the genus Morus, and most boobies belong to the genus Sula. The exception is Abbott's booby (Papasula abbotti) which was originally included in Sula, and is still commonly referred to as such (at least in popular sources), but sits on the gannet rather than the booby side of the divergence between the two main genera (Friesen & Anderson, 1997). The distinctions between the three genera are not huge, and some authors in the past have referred to all living sulids as Sula. In the recent fauna there is a clear geographical division between gannets and boobies - Morus is found in the northern and southern temperate zones while Sula and Papasula are tropical or subtropical - but this does not appear to have always been so in the past. The Pliocene Pisco Formation of Peru has provided species assigned to both Sula and Morus (Stucchi & Urbina, 2004).


A sulid on the Galapagos Islands. This photo comes from here, where it is identified as a masked booby (Sula dactylatra). I don't think it is - the orange beak identifies it as a Nazca booby (Sula granti), a species that was only distinguished from S. dactylatra in 1998 (Pitman & Jehl, 1998). Don Roberson's site has a photo by Bob Pitman of the two species standing side by side to show the difference.

Eight fossil genera have been assigned to Sulidae (as well as fossil species of Sula and Morus*) - the Eocene Masillastega and Eostega, the Oligocene Empheresula, the Miocene Microsula (now a synonym of Morus), Miosula, Enkurosula and Sarmatosula, and the Pliocene Palaeosula and Ramphastosula. The Cretaceous Elopteryx did a stint as a close relative of the sulids, but is now regarded as a dinosaur of the Troodontidae and not even a bird (for most definitions of the word "bird"). Eostega lebedinskyi is known from a single mandible that was recently redescribed by Mlíkovský (2007), who reasserted its sulid nature (past authors have disagreed). Masillastega rectirostris is known from a skull from the famed Messel formation, and differs from living sulids in having a comparatively long beak (Mayr, 2002). It is also distinct in having seemingly inhabited a freshwater environment, while all modern species are exclusively marine. Mayr (2002) only tentatively regarded Masillastega as a sulid, and it may be a stem-member of the family (Mayr, 2005). Mlíkovský (2007) synonymised the two Eocene genera on the basis of a lack of significant differences between them.

*Apart from a single taxon described as a subspecies of the modern species (Papasula abbotti costelloi [ha ha]), there don't appear to be any fossil species assigned to Papasula. Considering the only recent distinction of Papasula from Sula, one wonders whether any fossils of the former are masquerading as the latter.

Empheresula arvernensis is represented by a pelvis from France (the original material also included a sternum, but Lambrecht later indicated that the sternum was not even sulid - Mlíkovský, 2002). The French Oligocene also provided Sula ronzoni, which, if correctly assigned (which seems to be debatable*), would suggest that the living sulid lineages had diverged by that point (Friesen & Anderson, 1997, used a molecular clock to estimate a divergence time for Sula vs. Morus/Papasula of 23 million years ago, which is also consistent with this, but the age calculation methods used by Friesen & Anderson can only be described as [ahem] dated). The European Miocene genera Enkurosula and Sarmatosula are both known from isolated humeri, and are both doubtfully distinct from Morus (Nelson, 2006; Olson, 1984, suggests that Microsula (=Enkurosula) pygmaea may be conspecific with the contemporaneous Microsula avita of Maryland in the United States, which has itself been since reassigned to Morus), as are the Californian genera Miosula and Palaeosula.

*I'm rather confused here. Nelson (2006) notes that Sula ronzoni has four notches on the sternum, and indicates that this would place it on the sulid stem. However, Mlíkovský (2002, 2007) states that the type material of S. ronzoni is an incomplete pelvis, so what is Nelson talking about?



The wierdest of all sulids, though, is the Pliocene Peruvian Pisco Formation's Ramphastosula ramirezi. The genus name means "toucan-booby" and is undeniably appropriate as Ramphastosula, instead of having a dagger-like straight beak like all other sulids, had a deep beak with a distinct arch as shown in the reconstruction above from Stucchi & Urbina (2004). The skull of Ramphastosula is also more robust than in other sulids, seemingly to support the enlarged beak. Ramphastosula was obviously pursuing a different lifestyle to other sulids, as it looks as if it would be ill-suited to catching fish by plunge-diving. Stucchi & Urbina (2004) suggest that its robust skull indicates greater diving ability than other sulids, so perhaps Ramphastosula was more inclined to pursue its prey underwater than its modern relatives. Unfortunately, no post-cranial material is known as yet for this species.

REFERENCES

Friesen, V. L., & D. J. Anderson. 1997. Phylogeny and evolution of the Sulidae (Aves: Pelecaniformes): a test of alternative modes of speciation. Molecular Phylogenetics and Evolution 7 (2): 252-260.

Mayr, G. 2002. A skull of a new pelecaniform bird from the Middle Eocene of Messel, Germany. Acta Palaeontologica Polonica 47 (3): 507-512.

Mayr, G. 2005. The Paleogene fossil record of birds in Europe. Biological Reviews 80: 515-542.

Mlíkovský, J. 2002. Cenozoic Birds of the World. Part 1: Europe. Ninox Press: Praha.

Mlíkovský, J. 2007. Taxonomic identity of Eostega lebedinskyi Lambrecht, 1929 (Aves) from the middle Eocene of Romania. Annalen des Naturhistorischen Museums in Wien 109A: 19-27.

Nelson, J. B. 2006. Pelicans, Cormorants, and Their Relatives: The Pelecaniformes. Oxford University Press.

Olson, S. L. 1984. A brief synopsis of the fossil birds from the Pamunkey River and other Tertiary marine deposits in Virginia. In Stratigraphy and Paleontology of the Outcropping Tertiary Beds in the Pamunkey River Region, Central Virginia Coastal Plain: Guidebook for the Atlantic Coastal Plain Geological Association 1984 field trip (L. W. Ward & K. Krafft, eds.) pp. 217-223. Atlantic Coast Plain Geological Association.

Pitman, R. L., & J. R. Jehl, Jr. 1998. Geographic variation and reassessment of species limits in the "masked" boobies of the eastern Pacific Ocean. Wilson Bulletin 110: 155-170.

Stucchi, M., & M. Urbina. 2004. Ramphastosula (Aves, Sulidae): a new genus from the early Pliocene of the Pisco. Journal of Vertebrate Paleontology 24 (4): 974–978.