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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