Protocetidae

Protocetids Temporal range: Early Eocene–Middle Eocene PreЄ Є O S D C P T J K Pg N
Maiacetus
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Synapsida
Class:	Mammalia
Order:	Artiodactyla
Infraorder:	Cetacea
Family:	†Protocetidae Stromer 1908
Subfamilies and genera
Aegyptocetus Protocetinae Artiocetus Crenatocetus Dhedacetus Gaviacetus Indocetus Kharodacetus Maiacetus Protocetus Qaisracetus Rodhocetus Takracetus Makaracetinae Makaracetus Georgiacetinae Babiacetus Carolinacetus Eocetus Georgiacetus Natchitochia Pappocetus Pontobasileus

The protocetids form a diverse and heterogeneous group of cetaceans known from Asia, Europe, Africa, and North America.

Description

There were many genera, and some of these are very well known (e.g., Rodhocetus). Known protocetids had large fore- and hindlimbs that could support the body on land, and it is likely that they lived amphibiously: in the sea and on land. It is unclear at present whether protocetids had flukes (the horizontal tail fin of modern cetaceans). However, what is clear is that they are adapted even further to an aquatic life-style. In Rodhocetus, for example, the sacrum – a bone that in land-mammals is a fusion of five vertebrae that connects the pelvis with the rest of the vertebral column – was divided into loose vertebrae. However, the pelvis was still connected to one of the sacral vertebrae. Furthermore, the nasal openings are now halfway up the snout; a first step towards the telescoped condition in modern whales. Their supposed amphibious nature is supported by the discovery of a pregnant Maiacetus,^[1] in which the fossilised fetus was positioned for a head-first delivery, suggesting that Maiacetus gave birth on land. The ungulate ancestry of these early whales is still underlined by characteristics like the presence of hooves at the ends of toes in Rodhocetus.

Taxonomy

The protocetid subfamilies were proposed by Gingerich et al. 2005. They placed Makaracetus in its own subfamily (Makaracetinae) based on its unique adaptations for feeding (including only two incisors in each premaxilla). They then erected two subfamilies for the rest of the protocetids based on their degree of aquatic adaptation:^[2]

Protocetinae- Protocetines are Lutetian protocetids with generalized skulls retaining three incisors in the premaxilla and three molars in the maxilla. To the extent postcrania are known, they also have a pelvis similar to those in land-living mammals, with a sacrum composed of several fused vertebrae, articulated to the ilia and innominates, and large hindlimbs used for foot-powered propulsion.^[2]

Georgiacetinae- Georgiacetines are Bartonian protocetids transitional to the basilosaurids. Their skulls and dentition is similar to those of protocetines, but their pelvis is usually reduced without substantial articulations between the sacrum and the ilia and innominates. No hindlimbs have been found, but the reduced pelvis indicates that they probably swam using their tail rather than their feet.^[2]

Notes

1. Gingerich et al. 2009
2. Gingerich et al. 2005, Table 1, pp. 207–8

References

• Gingerich, Philip D; ul-Haq, Munir; von Koenigswald, Wighart; Sanders, William J; Smith, B Holly; Zalmout, Iyad S (2009). "New Protocetid Whale from the Middle Eocene of Pakistan: Birth on Land, Precocial Development, and Sexual Dimorphism". PLOS ONE. 4 (2): e4366. doi:10.1371/journal.pone.0004366. PMC 2629576. PMID 19194487.
• Gingerich, Philip D.; Zalmout, Iyad S.; Ul-Haq, Munir; Bhatti, M. Akram (2005). "Makaracetus bidens, a new protocetid archaeocete (Mammalia, Cetacea) from the early middle Eocene of Balochistan (Pakistan)" (PDF). Contributions from the Museum of Paleontology. 31 (9): 197–210. OCLC 742723177. Retrieved March 2013. Check date values in: |access-date= (help)
• Stromer, Ernst (1908). Die Archaeoceti des ägyptischen Eozäns: Beiträge zur paläontologie und geologie Österreich-Ungarns und des Orients (PDF). W. Braumüller. OCLC 21174007. Retrieved July 2013. Check date values in: |access-date= (help)