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.

Eons Roundup 8

Once again it’s a PBS Eons commission roundup day!

An unnamed Cerro Ballena rorqual whale and the long-necked seal Acrophoca, from “How the Andes Mountains Might Have Killed a Bunch of Whales”

The poposauroid pseudosuchians Shuvosaurus (life restoration) and Effigia (skeletal) from “When Dinosaur Look-Alikes Ruled the Earth”


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.

Expectation vs reality

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.


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 #27: The Weirdest Wildebeest

Earlier in this series we saw some ruminants with bizarre-looking headgear, but there was another species in that group that evolved a completely different type of strange head.

Rusingoryx atopocranion was a close relative of modern wildebeest that lived during the late Pleistocene, around 100,000 to 50,000 years ago. Its fossil remains are known from the Kenyan part of Lake Victoria, on Rusinga Island – an area which wasn’t actually an island at the time due to lower lake levels, and was instead part of a hot dry grassland environment.

Standing about 1.2m at the shoulder (~4′), it had an oddly-shaped skull with a pointed snout and a highly domed forehead. But this wasn’t the thick bony dome of a headbutting animal – this structure was narrow and fairly fragile, and had looping nasal passages running through it.

Instead it was something never before seen in any mammal: a bony nasal crest convergently similar to those of hadrosaurid dinosaurs.

Juveniles had less developed crests, developing them as they matured, and one skull that may represent an adult female also has a smaller crest, suggesting that this feature was sexually dimorphic.

Based on just the anatomy of the nasal passages Rusingoryx may have honked at a frequency similar to a vuvuzela, but the added length of its vocal tract could have lowered this pitch even further, closer to infrasound ranges – so more like a tuba! Such low frequencies can travel very long distances and are also below the hearing range of many carnivores, and would have effectively allowed Rusingoryx to shout at each other in “stealth mode”.

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.

Weird Heads Month #09: Butterfly Faces

The nose-forks and head-crests we saw last time weren’t the only unusual headgear in ancient ruminants.

The giraffoids are represented today by just pronghorns, giraffes, and okapi, but in the past they were much more diverse, modifying their prongs and ossicones into multiple sets of horns, or into deer-like and moose-like antler shapes.  

And Prolibytherium was probably the most striking of the lot.

Two different species have been identified, with Prolibytherium magnieri here living in North Africa during the early-to-mid Miocene, about 17-16 million years ago. Its exact evolutionary relationships are uncertain but it was probably part of a group called climacoceratids, deer-like giraffoids which often had thorny branching ossicones that resembled antlers.

It stood around 1.2m tall at the shoulder (~4′), and exhibited dramatic sexual dimorphism – females had slender forked horn-like ossicones, while those of the males flared out into large wide flat shapes that resembled butterfly wings.

Heavy reinforcement in the bones of the back of the males’ skulls helped to support all the extra weight of those huge ossicones, and if they actually used the structures to fight with each other then this may have also provided some protection or shock absorption.

Weird Heads Month #08: Nose-Forks and Handlebar Heads

Modern ruminants are the only living mammals with bony headgear, with four different  lineages each sporting a slightly different type: deer antlers, bovid horns, giraffid ossicones, and the prongs of pronghorns.

We still don’t actually know much about the evolutionary origins of ruminant headgear, although a recent genetic study suggests they’re all derived from a single common ancestral structure (and that deer antlers started off as controlled bone cancer).

And some extinct species were even stranger.

The protoceratids were an early group of North American ruminants whose relationships are uncertain, but may have been related to modern chevrotains. They were convergently deer-like in appearance, with teeth adapted for grazing on tough grasses – and along with having a pair of horns in the usual position on their heads, males also sported an additional pair of ossicone-like growths on their noses.

Synthetoceras tricornatus lived during the Late Miocene, around 10-5 million years ago, and was one of the largest protoceratids, standing about 1.1m tall at the shoulder (3’7″). Its two nose-horns were partially fused into a single long structure with a forked tip, which may have been used for sparring in a similar manner to the antlers of modern deer.

A colored line drawing of an extinct deer-like animal. It has a pair of horns on its head, along with a long horn on its nose that has a two-pronged forked tip.
Synthetoceras tricornatus

Meanwhile on a different branch of the ruminant family tree, closer related to deer and giraffes, a group known as the palaeomerycids independently developed a similar sort of extra head appendage – but at the opposite end of their skulls.

These ruminants were a little more heavily built than the protoceratids, and specialized in feeding on soft vegetation in humid forest environments. They were a highly successful group, existing for almost 30 million years, ranging across Eurasia, Africa, and North America, and even ventured into South America during the early phases of the Great American Interchange.

Males had two giraffe-like ossicones above their eyes, along with a third crest-like one at the very back of their heads. In some species this formed a single central “horn” shape, while in others it forked out to each side. They also often had long saber-like canine teeth similar to modern water deer and musk deer, which were probably used for fighting while their elaborate headgear was purely for visual display.

Xenokeryx amidalae lived in Spain during the mid Miocene, about 16 million years ago. It stood around 0.8-1m tall at the shoulder (2’7″-3’3″) and had a unique T-shaped “handlebar” crest which ended up inspiring its genus name – a reference to the similar shape of one of Queen Amidala’s headpieces in Star Wars, which was itself based on Mongolian imperial fashion.

A colored line drawing of an extinct deer-like animal. It has fang-like tusks protruding from the sides of its mouth, a pair of giraffe-like ossicones above its eyes, and a T-shaped handlebar-like crast on the back of its head.
Xenokeryx amidalae


The earliest baleen whales didn’t actually have any baleen plates in their mouths, and the evolutionary origin of these unique filter-feeding structures is still poorly understood.

It was thought to have been a fairly simple linear process from toothed ancestors to a mix of teeth and baleen and then to fully toothless with just baleen, but more recent discoveries have begun to cast doubt on that idea. The teeth of ancestral baleen whales weren’t suited to filter-feeding at all, instead still being adapted for predatory piercing and chewing – actions which would have been constantly interfering with and damaging any proto-baleen forming alongside them, and making it seem much more unlikely that there would have ever been a transitional form that had both teeth and baleen at the same time.

But then how did baleen whales get their baleen?

Maiabalaena nesbittae here provides a possible solution. Discovered in Oregon, USA, this early baleen whale dates to the early Oligocene, around 33 million years ago, and compared to most of its modern relatives it was comparatively tiny, only about 4.6m long (15′).

And it had no teeth at all, but possibly also no baleen.

Baleen rarely fossilizes, so it’s unclear whether Maiabalaena actually had any or not, but the shape of its skull suggests it probably didn’t – it lacked the broad thickened upper jaw associated with supporting racks of baleen plates. It instead seems to have been adapted for suction feeding similar to modern belugas and beaked whales, using muscular cheeks and tongue to manipulate water pressure and pull small prey like fish and squid straight into its mouth.

Since it lived at a time when the Antarctic Circumpolar Current was forming and cooling the oceans, changing ecosystems and prey availability, it may represent a previously unknown stage in baleen whale evolution – a point when they’d moved towards specializing for suction feeding and lost their teeth entirely, before transitioning again over to filter-feeding with baleen in a completely separate evolutionary development a few million years later.

Eons Roundup 4

Some more recent commission work for PBS Eons!

The entelodonts Eoentelodon and Brachyhyops, from “The Hellacious Lives of the Hell Pigs”

The early ichthyosaur Tholodus and the mosasaurPluridens, from “When Ichthyosaurs Led a Revolution in the Seas”

The early bats Onychonycteris and Icaronycteris, from “When Bats Took Flight”