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Casatia

Modern beluga whales and narwhals are the only living representatives of the monodontid lineage, found only in cold Arctic and sub-Arctic waters. But this whale family actually first evolved in much warmer climates – and some of them were downright tropical.

Casatia thermophila lived about 5 million years ago during the early Pliocene, in the Mediterranean Sea around Tuscany, Italy. Although known only from a couple of partial skulls and a few vertebrae it was probably similar in size to its modern relatives, around 5m long (16’4″).

It seems to have had a larger number of functional teeth than modern monodontids, and probably didn’t suction feed like its modern close relatives. Instead it may have fed more like most porpoises and dolphins, relying more on speed and snapping jaws to capture prey.

It inhabited the Mediterranean at a time not long after the sea there had mostly dried up and then been rapidly refilled. The presence of warm-water marine species such as bull sharks, tiger sharks, and dugongs in the same fossil beds as Casatia indicates the local climate at the time was hotter than it is today, with tropical temperatures – and suggests that this whale’s ancestors must have originally moved into the replenishing Mediterranean from lower latitudes alongside these other warmth-adapted animals.

This tropical monodontid was also much closer related to modern belugas than modern narwhals are, which raises the possibility that the two living monodontid species actually specialized for colder conditions completely independently of each other rather than descending from a cold-adapted common ancestor. Instead modern belugas and narwhals may have originated from separate warm-water monodontid ancestors who evolved similar cold-tolerant adaptations in parallel as the climate cooled during the onset of the Quaternary ice age, while the rest of their relatives all went extinct.

Kogiopsis

Kogiopsis floridana was a physeteroid whale that lived near the coast of the southeastern United States from the mid-Miocene to the early Pliocene, about 14-4 million years ago.

Known just from fossilized lower jaws and teeth, with some teeth up to nearly 13cm long (~5″), its full life appearance and size are uncertain – but it may have been slightly larger than a modern bottlenose dolphin at around 4.5m long (~14’9″). It’s traditionally been considered to be part of the kogiid family, closely related to modern pygmy and dwarf sperm whales, but some studies disagree with that classification and instead place it in the true sperm whale lineage.

It was probably a predator in a similar ecological role to modern orcas, adapted for hunting prey like squid, fish, and smaller marine mammals. But unlike orcas it wouldn’t have been the apex predator of its ecosystem, subject to predation pressure by even larger carnivores like macroraptorial sperm whales and everyone’s favorite ridiculously huge shark – and as a result it probably had a “live fast and die young” lifestyle similar to modern kogiids and other small-to-medium-sized Miocene physeteroids, rapidly maturing and only living to around 20 years old.

I’ve reconstructed Kogiopsis here as a kogiid-like animal, with a similar sort of shark-like head shape and “false gill” markings. In the background a second individual is depicted displaying “inking” behavior, releasing a defensive cloud of reddish-brown fluid from a specialized sac in its colon.

Phiomicetus

Named after the canine-headed Ancient Egyptian god, Phiomicetus anubis is the first fossil cetacean to discovered, described, and named entirely by a team of Arab paleontologists.

Living during the mid-Eocene, about 43 million years ago, in a shallow sea-covered region that is now part of Egypt‘s Western Desert, Phiomicetus was an early protocetid – an amphibious foot-powered swimmer, at a transitional point in the evolution of whales from deer-like terrestrial animals to fully aquatic screaming torpedoes.

About 3m long (~10′), it had large jaw muscles and sharp teeth with wear patterns that suggest it was a raptorial hunter grabbing and snapping at prey with powerful bites. It would have probably tackled fairly big prey compared to other protocetids, hunting things like large fish, turtles, and even smaller whales in an ecological role similar to that of modern orcas.

Along with the distantly-related long-snouted Rayanistes it’s one of the earliest known whales from Africa, giving us further glimpses at a time period when early cetaceans were first dispersing out from the South Asian subcontinent via the ancient Tethys Sea.

Borealodon

Modern mysticete whales all have baleen plates in their mouths, but before the evolution of these specialized filter-feeding structures the early members of their lineage still had toothy jaws.

Borealodon osedax here was one of those “toothed mysticetes”, living about 30-28 million years ago during the mid-Oligocene off the coast of Washington state, USA.

Unlike modern baleen whales it was small, about the size of a modern porpoise at around 2m long (6’6″), and the wear on its multi-cusped teeth suggest it was a predator taking slicing bites of fish – possibly using suction-assisted feeding like its close relatives the aetiocetids.

Its fossilized remains are also a rare example of an ancient whale fall, with characteristic bore holes in its bones from Osedax worms.

Aegicetus

The protocetids were some of the first oceanic cetaceans, occupying a transitional position in the evolution of whales, with four paddle-like limbs and nostrils only partway up their snouts.

Early members of this group swam like otters, using a combination of undulating their bodies and paddling with large hind limbs, but somewhere in the Late Eocene they switched over to propelling themselves entirely with their tails and gave rise to even more whale-like forms like the basilosaurids.

And Aegicetus gehennae was right in the middle of that switch.

Discovered in the Wadi Al-Hitan (“Valley of the Whales”) fossil site in Egypt, Aegicetus lived around 37-35 million years ago. It was similarly-sized to earlier protocetids like Georgiacetus, measuring about 3.5m long (11’6″), but its hind limbs were proportionally smaller. Its hips were also completely disconnected from its vertebrae, giving it much more flexibility to undulate its body and tail – and preventing it from supporting its weight on land, suggesting that it spent its entire life in the water.

It wasn’t a direct ancestor to more “advanced” cetaceans, since it lived alongside several species of basilosaurids. Instead it seems to represent a late-surviving example of what the earlier protocetid-basilosaurid transitional forms would have looked like.

Ensidelphis

Last week’s weird-snouted Furcacetus wasn’t the only recently-discovered ancient platanistoid dolphin that deserves some attention.

Ensidelphis riveroi was described in the same paper, and also lived in the coastal waters around Peru during the early Miocene, about 19 million years ago. It was a little less closely related to its modern river-dwelling cousins than Furcacetus, and was slightly larger, estimated to have measured about 3m long (9’10”).

But what made it weird was its incredibly long snout, lined with around 256 tiny sharp teeth, which also curved markedly to the right side along its 55cm (1’10”) length.

With only one known skull of Ensidelphis it’s impossible to tell if this was a natural condition for the species or if it was some sort of anomalous individual. It doesn’t seem to be a deformation of the fossil, at least.

Similar unusual right-side bending has been seen in the skulls of a few individuals of modern South Asian river dolphins, franciscanas, and Amazon river dolphins, possibly caused by injuries at a young age being exaggerated as the animals grew. However, many other platanistoid dolphins (especially squalodelphinids) are known to have naturally had similar bends in their snouts – but always to the opposite side, curving to the left instead of the right.

But naturally bent or not, what might Ensidelphis have been doing with that incredibly lengthy snoot?

Its long slender jaws would have had a fairly weak bite, so it probably wasn’t able to catch large prey, and it had a very flexible neck. Possibly it swam along near the seafloor using its snout to probe and sweep around in the sediment for buried small prey.

Modern South Asian river dolphins swim along on their sides while doing this – almost always on their right sides, interestingly enough – and if Ensidelphis did the same sort of thing then a snout bent in that direction might have been an advantage.

Furcacetus

The two living subspecies of the South Asian river dolphin are the last surviving members of a lineage known as the Platanistoidea, an early evolutionary branch of the toothed whales. This group was once much more diverse and widespread than their modern representatives, found in oceanic habitats around the world from the Oligocene to the mid-Miocene.

Many of them had forward-pointing protruding teeth at the tips of their snouts, resembling those of some plesiosaurs or pterosaurs, suggesting they were a convergent adaptation used for snagging hold of slippery soft-bodied aquatic prey.

Furcacetus flexirostrum is one the newest additions to this group, named and described in late March 2020. It lived in Pacific coastal waters around Peru during the early Miocene, about 19-18 million years ago, and was about the same size as modern South Asian river dolphins at around 2.3m long (7’7″).

And it had a uniquely-shaped snout for a cetacean, curving upwards for most of its length but then turning downwards right at the tip, which along with large forward-pointing teeth gave its jaws a vaguely crocodilian appearance.

A closeup view of the jaws of Furcacetus.

Much like slender-snouted crocodilians and spinosaurids, this arrangement would have allowed Furcacetus to make quick bites at small-fast-moving prey like fish and crustaceans.

Livyatan

The modern sperm whale is already an impressive animal, being by far the largest of the living toothed whales and famous for its ability to dive over 2km down (1.2 miles) to feed on deep-sea animals like giant squid.

But some of its ancient relatives were terrifying.

Livyatan melvillei here has an appropriately monstrous name, inspired by both the Hebrew name for the Leviathan and Herman Melville, the author of Moby-Dick. Known from the Pacific coast of South America during the late Miocene, around 10-9 million years ago, it’s estimated to have measured somewhere between 13.5m and 17.5m long (~44′-57′) – comparable in size to an adult male sperm whale.

Unlike the relatively slender mouth of its modern cousin, however, it instead had thick strong jaws full of enormous teeth.

It was part of a loose grouping of what are known as “macroraptorial sperm whales“, which all had similarly toothy jaws and occupied the same sort of ecological niche as modern orcas, specializing in hunting prey like large fish, squid, seals, and other whales.

Livyatan‘s main food source was probably smaller baleen whales about half its own size, and its only real competition for this prey was the equally huge megalodon shark that shared the same waters.

A huge fossil tooth found in Australia suggests that Livyatan or a very close relative of it survived at least into the early Pliocene, about 5 million years ago. Around this time a cooling climate and dwindling numbers of its preferred prey would have eventually made a population of such enormous apex predators unsustainable, and driven this “killer sperm whale” into extinction – probably around the same time megalodon disappeared, about 3.6 million years ago.

Inticetus

While most modern toothed whales have jaws full of teeth that are all the same simple pointed shape – an adaptation for better holding onto slippery prey – their ancient ancestors had teeth much more like other mammals, with differentiated incisors, canines, and molars.

In-between them were whales like Inticetus vertizi, which lived off the coast of southwestern Peru during the Early Miocene, about 18 million years ago.

At over 3.5m long (11′6″) it was one of the larger known toothed whales around at the time, but it wasn’t the direct ancestor of any living whales. Instead it was more of an evolutionary “cousin” to them, part of an older offshoot lineage that lived alongside the early members of modern toothed whale groups.

Inticetus had a long and unusually wide-based snout, somewhat croc-like in appearance, with sharp pointed teeth at the front and multi-lobed cheek teeth further back. A lack of obvious wear on its back teeth suggests it wasn’t using them to chew up its food, and it may have had a fairly specialized diet – possibly using those back teeth to sieve small prey out of the water in a similar manner to modern lobodontine seals.

An close-up view of Inticetus' jaws, showing the differences in tooth shape from front to back. — Closeup of *Inticetus*‘ jaws

Inticetus-like teeth have also been found in Miocene-aged deposits in the eastern USA, the Atlantic coast of France, and southeast Italy, indicating that this ancient whale lineage was quite widespread.

Rayanistes

Remingtonocetids were an early branch of the whale evolutionary family tree, known from about 49-41 million years ago and splitting off somewhere between the famous “walking whale” Ambulocetus and the more oceanic protocetids. With otter-like bodies, tiny eyes, and long gharial-like snouts, they lived in near-shore shallow marine habitats and probably swam using a combination of their hind feet and tails.

They were initially found only in Pakistan and India, but then Rayanistes afer here was discovered all the way over in Egypt – suggesting that these early whales were much more widespread than previously thought, dispersing through the Tethys Sea at about the same time as their protocetid cousins.

Dating to the Middle Eocene (~45-41 mya), Rayanistes was probably about 2.5m long (8′2”). It had powerful hindlimb musculature that would have given it a very strong kicking swimming stroke, but it probably couldn’t actually support its own weight on land since its femur wasn’t very well anchored into its pelvis.