toothed whale – Nix Illustration

Nihohae

Nihohae matakoi was a dolphin that lived in the coastal waters around what is now Aotearoa New Zealand during the late Oligocene, about 25 million years ago. Part of a group known as waipatiids, it was much closer related to modern South Asian river dolphins than to modern oceanic dolphins.

Around 2m long (6’6″), it had unusually long tusk-like teeth at the front of its jaws, splaying out almost horizontally forwards and to the sides.

These teeth lay too flat to effectively interlock as a “fish trap”, and their fairly delicate structure and lack of wear marks suggests they also weren’t used for piercing large prey, sifting through gritty sediment, defending against predators, or for fighting each other. But Nihohae did have a highly flexible neck and the ability to quickly snap its jaws from side to side – although with a relatively weak bite force, suggesting it was primarily tackling small soft-bodied prey that could be easily swallowed whole.

Overall its feeding ecology seems to have been similar to modern sawfish, stunning prey such as squid with rapid slashing swipes of its jaws.

Strange Symmetries #22: The Whalerus And The Twisted Tusks

Mammalian tusks usually grow in symmetrical pairs with only minor developmental asymmetry, but a few species have evolved much more uneven arrangements.

A colored line drawing of the extinct toothed whale Odobenocetops. Its body is beluga-like but it has a face more like a walrus than a whale, with a big fleshy bristly upper lip and a pair of protruding tusks. The right side tusk is much longer than the left. It's depicted with a mottled brown and white color scheme. — *Odobenocetops peruvianus*

Odobenocetops peruvianus was a small toothed whale that lived during the Miocene, about 7-3 million years ago, in shallow coastal waters around what is now Peru. Around 3m long (~10′), it was a highly unusual cetacean with binocular vision, a vestigial melon, muscular lips, and a pair of tusks – features convergent with walruses that suggest it had a similar lifestyle suction-feeding on seafloor molluscs and crustaceans.

In males the right tusk was much more elongated than the left, measuring around 50cm long (~1’8″) in this species and up to 1.35m (4’5″) in the closely related Odobenocetops leptodon. Since these teeth were quite fragile they probably weren’t used for any sort of combat, and they may have instead served more of a visual display function.

And despite being closer related to modern narwhals and belugas than to other toothed whales, Odobenocetops’ long right-sided asymmetric tusks actually seem to have evolved completely independently from the iconic left-sided asymmetric spiral tusks of narwhals.

An edited meme image using screenshots of Dr. Doofenshmirtz from "Phineas and Ferb". The text reads: "If I had a nickel for every time whales evolved asymmetric tusks, I'd have two nickels. Which isn't a lot, but it's weird that it happened twice."

A colored line drawing of an extinct woolly mammoth. It's an elephant-like animal covered in a thick coat of brownish hair, with a high domed forehead, small ears, and long curving tusks. The tusks are noticeably asymmetrical, one curving more downwards than the other. — Woolly Mammoth (*Mammuthus primigenius*)

The woolly mammoth (Mammuthus primigenius) lived across Eurasia and North America during the last ice age, mostly from the Pleistocene about 400,000 years ago to the early Holocene about 10,000 years ago – altohugh a few relict populations survived until around 4,000 years ago in isolated areas of Alaska, Siberia, and eastern Russia.

Around 3m tall at the shoulder (~10ft), these hairy proboscideans had very long curving tusks that were used for digging out vegetation from under snow and ice, scraping bark from trees, and for fighting.

The tusks showed a lot of variation in their curvature, and were often rather asymmetrical, a condition also seen in the closely related Columbian mammoth. Like modern elephants mammoths may have also favored using one side over the other for certain tasks, which over their lifetimes could result in uneven wear exaggerating the natural asymmetry even more.

Strange Symmetries #19: Wonky Whales

Toothed whales – the branch of cetaceans that includes modern dolphins, porpoises, beaked whales, and sperm whales – have surprisingly asymmetrical skulls, with some of the bones skewed to one side and just the left nostril forming their blowhole.

Some of the most obvious external manifestation of this lopsidedness can be seen in sperm whales, which have their blowhole at the front left side of their head, and in male narwhals, which usually have a single left-side tusk.

This sort of asymmetry first appeared in the skulls of early toothed whales around 30 million years ago. And since the highest amounts of wonkiness have gone on to develop in lineages that hunt in dark, cluttered, or murky waters, this suggests that the trait is somehow linked to the evolution of complex echolocation.

Some ancient members of the river dolphin lineage also had some additional unusual asymmetry, sometimes having slightly sideways-bending snouts.

Ensidelphis riveroi was one of the weirdest of these, living around the coasts of what is now Peru during the Miocene, about 19 million years ago. Around 3m long (~10′), it had a very long narrow toothy snout that curved distinctly off to the right along its length.

A sketch showing Ensidelphis' bizarre side-curving snout. A hypothetical straight snout is shown outlined in blue, while the actual curvature is overlaid in red. — Expectation vs reality

It’s not clear what the function of this bend was, or even if the only known skull actually represents the normal condition for this species. But Ensidelphis’ bendy snoot might have been used to probe around in muddy seafloor sediment or to extract prey from crevices, possibly like an underwater version of the modern wrybill.

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.

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.

Weird Heads Month #19: Sword-Snouted Whales

Cetaceans are just weird animals in general. Fully aquatic mammals best described as “fat screaming torpedoes“, with bizarre head anatomy and their nostrils pulled up to the top of their heads behind their eyes. Some of them are among the largest animals to ever exist, some of them can live to over 200 years old, and some can dive to incredible depths below the ocean surface.

And they’re all descended from tiny deer-like creatures, with their closest living relatives being hippos and other even-toed ungulates.

Some ancient cetaceans were particularly odd-looking, evolving walrus-like tusks or elongated chins – or in the case of Eurhinodelphis longirostris here, an incredibly long swordfish-like snout.

Living during the mid to late Miocene, about 14-7 million years ago, Eurhinodelphis ranged across the Mediterranean and the North Atlantic, with fossil remains known from Western Europe, Turkey, and the East Coast of the United States. It was a fairly small dolphin-like cetacean about 2m long (6’6″), and was part of a lineage of early toothed whales called eurhinodelphinids.

Its upper jaw was around five times longer than the rest of its skull, and toothless past the point where the lower jaw ended. Much like the modern billfish it resembled, it probably used its snout to slash at fast-moving fish, stunning them and making them easier to catch.

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.