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.

Bathyergoides

Blesmols, or African mole-rats, are a group of rodents adapted for mole-like burrowing. Closely related to the more famous naked mole-rat, these little mammals have reduced eyes and ears along with incisors that protrude out even when their mouths are closed, allowing them to excavate tunnels primarily using their teeth.

One of the earliest known fossil blesmols is Bathyergoides neotertiarius here, from the early Miocene of Namibia about 20 million years ago. For almost a century this species was known only from teeth and partial skull remains, but recently a partial skeleton was described giving us a better idea of its overall appearance and lifestyle.

Bathyergoides was a fairly large blesmol, around 25cm long (~10″), and was already specialized for tooth-digging with a skull very similar to modern forms. It had powerful muscular forelimbs that would have been used to push back loose soil while burrowing, but unlike its living relatives it also had a long tail and relatively slender hindlimb bones – with anatomy suggesting its legs were used more for stabilizing its posture than for actively digging.

It may have had a less subterranean lifestyle than modern blesmols, digging out extensive burrows but still foraging for food above ground in a similar manner to modern semi-fossorial rodents like giant pouched rats.

Retro vs Modern #16: Uintatherium anceps

Discovered in the Western United States during the early 1870s, Uintatherium anceps was part of one of the earlier major conflicts in the the Bone Wars. Nearly 30 different scientific names were applied to various fossil specimens of this mammal in under two decades, and the taxonomic tangle wasn’t properly sorted out until nearly a century later in the 1960s when they were recognized as actually all being the same species.


1870s

Paleontologist Edward Cope considered Uintatherium (under the name “Loxolophodon”) and its close relatives to be proboscideans – part of the elephant lineage – due to some of the similarities in their anatomy. The first reconstruction of these animals showed this version, depicting elephant-like animals with downward-pointing tusks, short tapir-like trunks, and the multiple bony projections on their skulls speculatively shown as attachment points for large antler-like horns.

Cope’s rival Othniel Marsh heavily criticised that interpretation of Uintatherium, arguing that these huge mammals were instead a separate group within the ungulates named dinoceratans – although this wasn’t really as huge of a classification difference as it seems today, since at the time proboscideans were also considered to be ungulates!

The dinoceratan ungulate interpretation quickly won out, and for a while in the 20th century Uintatherium actually became a fairly popular and well-known prehistoric mega-mammal, commonly included in collections of cheap plastic “dinosaurs” and usually depicted as more of a knobbly-headed sabertoothed rhino.


2020s

In recent years the dinoceratans seem to have fallen into obscurity and some degree of paleontological neglect, with little modern work on the group and no major studies for the last couple of decades – although this might be starting to change.

Despite the early ideas about them being ungulates, the evolutionary relationships of dinoceratans have become much more murky over the last century or so. Due to different elements of their anatomy being highly convergent with various other mammals it’s easy to find “false positives” in morphological comparisons, and they’ve been proposed as being connected to a wide variety of groups including “condylarths“, “insectivores“, rodents, and cimolestans. But some mid-2010s research suggests they were in fact ungulates after all, closely related to early South American forms like Carodnia – a lineage whose own evolutionary relationships are murky, but may have close affinities with modern horses, rhinos, and tapirs.

We now know Uintatherium anceps lived across the Western and South Central USA during the mid-Eocene, about 46-40 million years ago, at a time when warm wet climates extended up into the Arctic and lush tropical-style rainforests covered much of the continent.

It was similar in size and build to a modern white rhino, about 4m long (13′) and stood around 1.7m tall at the shoulder (5’7″). It had three distinctive pairs of “horns” on its forehead, snout, and nose, that were similar in structure to the ossicones of giraffids, probably covered in skin and hair rather than keratin. Its elongated canine teeth were protected by bony flanges on its lower jaw, and seem to have been a sexually dimorphic feature that was much more prominent in males.

It also had an oddly concave skull, with its forehead dipping inwards, and an unusually tiny braincase for its size. It probably wasn’t a particularly intelligent animal, but it didn’t really need to be – as one of the first types of herbivorous mammal to get truly huge in the early Cenozoic, a fully-grown Uintatherium probably had no natural predators at all.

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.

Elasmotherium

Elasmotherium sibiricum was a giant rhinoceros that lived during the mid-to-late Pleistocene epoch, between about 800,000 and 39,000 years ago. Found across much of the Eurasian steppe dry grassland environments, it stood around 2.5m tall (8’2″) at the top of its humped shoulders and weighed about 4 tonnes (4.4 US tons), making it close in size and mass to a modern elephant.

It was the last known representative of a particularly ancient lineage of rhinos, last sharing a common ancestor with modern forms over 40 million years ago.

A large bony dome on its forehead is traditionally thought to have supported an enormous keratinous horn like the distantly-related woolly rhino, but a 2021 study has recently challenged that interpretation. The dome structure was actually rather thin-walled and wouldn’t have been able to support the weight of a giant horn, instead probably being covered by a much stumpier backwards-pointing nub – while an enlarged nasal cavity inside the dome also suggests it may have actually functioned as a resonating chamber, similar to the crests of hadrosaurs or the extinct wildebeest Rusingoryx.

It also had a smaller toughened “pad” on its nose that may have been used along with a prehensile upper lip to dig around in the soil for plant roots and tubers.

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.

Allodesmus

Desmatophocids were a group of seal-like pinnipeds that appeared very early in the group’s evolution, around 23 million years ago. They were found across the northern Pacific from the west coast of North America to Japan, and were the first pinnipeds to get big, with some species reaching sizes comparable to modern northern elephant seals.

They had a mixture of anatomical features similar to true seals, sea lions, and walruses, but weren’t actually the ancestors of any of those modern groups. Instead they seem to have just been their own separate thing, a very early diverging “cousin” lineage of pinnipeds that convergently developed close resemblances to their later relatives.

Allodesmus demerei here was one of the last known desmatophocids, living in the late Miocene (~9 million years ago) in what is now southwest Washington, USA. 

It would have been a sea lion-like animal, able to walk on all fours when hauled out on land, and showed distinct sexual dimorphism, with males growing to sizes of around 4m long (13′) and females being somewhat smaller. It powered its swimming using its front flippers, and may have mostly foraged in deep dark waters, using both keen vision and sensitive whiskers to locate prey.

The nasal region of its skull also shows some similarities to modern elephant seals, and some reconstructions depict males with the same sort of large proboscis. 

Hyrachyus

What’s the most unexpected fossil you’d think could be found on the island of Jamaica?

How about an ancient rhino?

Hyrachyus here was an early member of the rhinocerotoids, a lineage of odd-toed ungulates that also includes the true rhinoceroses, the tapir-like and hippo-like amynodontids, the long-legged hyracodontids, and the giant indricotheriines.

This particular genus was very widespread for much of the Eocene, found across Europe, Asia, and North America, crossing back and forth between the continents via the North Atlantic land bridge.

The Jamaican Hyrachyus lived during the mid-Eocene, around 45 million years ago, and was very anatomically similar to the North American Hyrachyus affinis – with the known fossil material not being considered distinct enough to be assigned to a new species yet. It was also about 15-20% smaller than its mainland relative, standing only 25cm tall at the shoulder (10″), but it’s not yet clear if this was a case of insular dwarfism or not.

Its presence in ancient Jamaica suggests that there may have been some sort of land connection between the proto-island and Central America during the early Eocene, when a chunk of what would eventually become western Jamaica was located much closer to the coasts of Honduras and Nicaragua. It’s the only fossil ungulate known from the Caribbean, and one of only a few terrestrial mammals in the region with North American evolutionary roots (the others being the extinct rodents Caribeomys merzeraudi and Oryzomys antillarum, and modern solenodons).

Unfortunately these little rhinos didn’t get much time on their island home. Jamaica subsided fully underwater about 40 million years ago, drowning its unique Eocene ecosystem entirely, and wouldn’t re-emerge and be re-colonized until much later in the Cenozoic.

Heliosus

Just before the 2017 solar eclipse, some unusual fossils were discovered in Southern Wyoming, USA.

Consisting of a partial jawbone and a humerus, and dating to the mid-Eocene (~47 million years ago), the remains clearly belonged to an early even-toed ungulate – but one much bigger than the rabbit-sized herbivores known from that time. This was something closer in size and build to a large modern pig, standing at least 1m tall at the shoulder (3’3″).

It turned out to belong to a member of a somewhat obscure lineage known as the helohyids, a group whose evolutionary relationships are a bit uncertain but are generally considered to be part of the whale-and-hippo lineage. These pig-like animals were large opportunistic omnivores, possibly occupying a similar ecological role to the later entelodonts, with some Late Eocene forms reaching sizes comparable to black bears.

This new helohyid was named Heliosus apophis, inspired by the eclipse, with its genus name meaning “sun pig”, and its species name referencing a sun-devouring Ancient Egyptian deity.

It was one of the earliest known large-bodied members of the group, and shows that these animals must have increased in size very rapidly during their early evolution, going from rabbit-sized to pig-sized within just a couple of million years.

Adalatherium

Even for a fossil species from an isolated island, Adalatherium hui is very weird.

This mammal was part of an enigmatic group known as gondwanatheres, which were probably early members of the theriiform lineage – slightly closer related to modern marsupials and placentals than to monotremes. Found in the southern continents of Gondwana between the Late Cretaceous and the Miocene, these animals were adapted for herbivory with convergently rodent-like ever-growing front teeth that helped them chew through tough plant matter.

They were previously known mainly from isolated teeth and jaw fragments, with some rare full skull material, but Adalatherium is remarkable for being represented by a complete skeleton.

And it’s turned out to be far stranger than anyone expected.

Living in northwestern Madagascar during the Late Cretaceous, about 70-66 million years ago, Adalatherium was one of the larger known Mesozoic mammals at around 60cm long (2′) – although the one known specimen seems to have been a juvenile, so mature individuals were probably slightly larger.

(And based on its body proportions, its close relative Vintana may actually have been even bigger than previously thought. Whether this sort of large size was common in Cretaceous gondwanatheres or if this was just island gigantism is still unknown, though.)

It was probably a marmot-like digging animal, excavating burrows with its large claws and powerful limbs, and since it likely evolved from ancestors that had become isolated on Madagascar over 20 million years earlier it had developed a very unusual mixture of both “primitive” and highly specialized anatomical features. It had more back vertebrae than any other known Mesozoic mammal, upright forelimbs, sprawling hind legs with bowed-out tibias, strong back and leg musculature, and a therian-like pelvis with epipubic bones.

And then there’s the snoot.

The snout region of Adalatherium‘s skull was pockmarked with a large number of foramina, holes that allow the passage of nerves and blood vessels through the bone. It had more of these than any other known mammal, and their presence suggests that it probably had a very sensitive upper lip and whiskery snout. Most mammals with a lot of whiskers just have one very big foramina, but Adalatherium seems to have evolved a different solution to the same problem.

It also had one other bizarre feature – a hole in the top of its nose. A large “internasal vacuity” between its nasal bones is a unique feature not known in any other mammal, and its function is a total mystery.

Since this hole was also surrounded by many foramina it may have supported some sort of soft-tissue sensory structure on top of its nose. So I’ve speculatively depicted it here with a leathery horn-like “shield”.

Adalatherium skull
From fig 2 in Krause, D. W. et al (2020). Skeleton of a Cretaceous mammal from Madagascar reflects long-term insularity. Nature 581, 421–427. https://doi.org/10.1038/s41586-020-2234-8