Wednesday, December 18, 2013

Basilosauridae, the so-called zeuglodonts, referring to their complex, many-cusped teeth (the Greek zugotos means yoked or joined, and odous, of course, tooth). The most primitive archaeocete identified to date was Nalacetus, known mainly from isolated teeth. Pakicetus, another small, very early archaeocete, had eyes on top of its head, drank only fresh water (confirmed from oxygen isotope ratios in its tooth enamel), and was predominantly wolf- or hyena-like in appearance. The other families of archaeocetes had been largely supplanted by the zeuglodonts during the late Eocene. Probably the best-known zeuglodont was Basilosaurus, or the "king lizard" (from the Greek basileus for king and sauros for lizard). This animal could be almost 70 ft (21 m) long and weighed at least 11,000 lb (5,000 kg). Its small head in relation to the long body made it appear truly serpentine. The front appendages had been modified into short, broad paddles, but were still hinged at the elbow; and the rear appendages had atrophied to nothing more than stumps. 

Basilosaurids may have had dorsal fins and horizontal tail flukes, and they were likely hairless, or nearly so. In short, Basilosaurus was well along the path to becoming what cetologists now think of as a whale. The archaeocetes are replaced in the fossil record by odontocetes and mysticetes beginning in the Oligocene, about 38 mya. By approximately the middle of that epoch, the archaeocetes appear to have died out completely. The oldest known cetacean in the mysticete clade is Llanocetus denticrenatus, found in late Eocene rocks on the Antarctic Peninsula. 

This species' most characteristic feature was its series of lobed, widely spaced teeth, which were somewhat reminiscent of the teeth of the crabeater seal (Lobodon carcinophagus). Like the crabeater seal, L. denticrenatus was probably a filter feeder on krill-like invertebrates or possibly small schooling fish. At least four families of tooth-bearing mysticetes have been described from the Oligocene (24-38 mya). The transition leading to rudimentary baleen plates in the spaces between teeth probably occurred about 30 mya with the emergence of the Cetotheriidae, or primitive baleen-bearing mysticetes. It is a slight misconception to say that the presence of teeth is a diagnostic feature of Odontoceti, the so-called toothed whales, because all archaeocetes and some of the primitive fossil mysticetes also had teeth. Further, all of the modern baleenbearing mysticetes have teeth in the early fetal stages of their development. 

Odontocetes also radiated rapidly and widely during the Oligocene, by the end of which there were more than 13 families and 50 species of cetaceans in the world's oceans. This diversity was probably driven by changes in foraging opportunities related to breakup of the southern supercontinent of Gondwana, opening of the Southern Ocean, and the consequent polar cooling and sharpening of latitudinal temperature gradients. Several of the early odontocete lineages failed to survive beyond the Miocene (5-23 mya). The shark-toothed dolphins (Squalodontidae), with their sharp, triangular, serrated teeth, were likely active carnivores, while the very longbeaked Eurhinodelphinidae, with their overhanging upper jaws and many small, conical teeth, were more like the dolphins that cetologists know today. Both of these groups had vanished from the fossil record, and others had dwindled to mere remnants, by the end of the Miocene. 

The cetotheres radiated further during the Miocene (5-23 mya), with more than 20 genera in which the blowholes were positioned about as far back on the top of the head as they are in living mysticetes. Also, by the early Miocene, the two main branches of cetotheres were evident, one leading to the modern right whales (Balaenidae) and the other to the rorquals (Balaenopteridae) and gray whale (Eschrichtiidae). Gray whales do not appear in the fossil record until only about 100,000 years ago, and their ancestry is therefore particularly problematic. For their part, the odontocetes also experienced a major Miocene radiation. Beaked whale (Ziphiidae) fossils are common in marine sediments worldwide by 5-10 mya, and these include animals belonging to the modern genus Mesoplodon. Sperm whales in the family Physeteridae, similar in some important ways to the living species, were present by 22 mya.


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