Cimolestans were one of the major mammal lineages that survived through the K-Pg mass extinction 66 million years ago. Closely related to early placentals, they had a burst of diversification during the first half of the Cenozoic and rapidly evolved into a wide range of specialized forms – some uniquely weird, and others convergently resembling more familiar modern animals like squirrels, bears, ground sloths, and hippos.

And one group known as the pantolestids were incredibly otter-like.

(Because synapsids love them some lutrinization.)

Palaeosinopa didelphoides here lived during the mid-Eocene, about 52 million years ago, in what is now the Mountain West region of the USA. It was similar in size to a small otter, about 1m long (3’3″), and had a streamlined body with a well-muscled neck, short powerful forelimbs, slightly longer hindlimbs, and a very long tail.

Inhabiting a subtropical lake ecosystem, it probably swam using both hindlimb paddling and otter-like tail undulations. Its strong jaws and teeth suggest it was specialized for crunching hard shellfish prey, but so far preserved gut contents have only shown fish bones and scales. Fairly large claws indicate it was also able to dig out burrows similarly to modern otters and beavers.

Although pantolestids were never particularly common animals they were quite widespread, expanding their range from their evolutionary origins in North America across to Europe and eventually into Asia. A cooling and drying climate at the end of the Eocene seems to have driven most of the group into extinction alongside all their other cimolestan relatives – but a few of the Asian species clung on slightly longer as the very last of their kind, with the last known fossils dating to about 28 million years ago in the early Oligocene.


If there’s any equivalent to carcinization in mammals, it’s turning into an otter-beaver-like semi-aquatic form.

Because it just keeps happening.

Modern examples alone include otters, beavers, muskrats, giant otter shrews, desmans, aquatic genets, yapoks, lutrine opossums, and platypuses – and in the fossil record there were early pinnipeds, remingtonocetids, pantolestids, stagodontids, and Liaoconodon going as far back as the early Cretaceous. Even outside of the true mammals there were also Castorocauda, Haldanodon, and Kayentatherium during the Jurassic, and much further back in the late Permian there was the early cynodont Procynosuchus.

So a non-cynodont synapsid doing the exact same thing really isn’t all that surprising.

Perplexisaurus foveatus was a member of the therocephalians, a group of synapsids that were close evolutionary “cousins” of the cynodonts-and-true-mammals lineage. Similar in size to a modern rat, about 20cm long (8″), it lived in Western Russia during the Late Permian about 268-265 million years ago.

At the time this region was a river plain with a tropical climate, experiencing seasonal floods that turned the whole area into what’s known as “viesses” (a name based on the abbreviation “V.S.S.” standing for “very shallow sea”), vast shallow lake-seas that persisted for weeks or months at a time.

So this little animal has been interpreted as being semi-aquatic, swimming around and feeding on aquatic invertebrates and tiny fish and amphibians. Its skull had numerous pits around the front of its face, suggesting that it had a highly sensitive snout – probably whiskery, allowing it to hunt entirely by touch in dark murky water, but it’s also been proposed to have possibly had an electroreceptive sense similar to modern platypuses.


The docodonts were a group of mammaliaformes (close relatives of the earliest true mammals) which lived across North America, Europe, and Asia from the Middle Jurassic to the Early Cretaceous. Originally only known from teeth and jaw fragments they were traditionally thought to be fairly generic shrew-like insectivores, but more recent discoveries of better fossils have revealed they were actually much more diverse, occupying ecological niches ranging from squirrel-like tree-climbers to mole-like diggers to beaver-otter-like swimmers.

Most of the more complete fossil material of these animals comes from the mid-Jurassic of China, but one species from elsewhere is also known from a partial skeleton.

Haldanodon exspectatus here lived in central Portugal during the Late Jurassic, about 155 million years ago. Around 15-20cm long (6-8″), it had small eyes and short chunky well-muscled limbs with the front paws adapted for digging. Since it inhabited a very swampy environment it probably wasn’t a pure mole-like burrower – extensive tunnels would have constantly flooded – but it may have instead been a similar sort of semi-aquatic animal to modern platypuses and desmans, foraging for invertebrates in the water and excavating burrows in the banks.

Roughened areas of bone on its snout may also have supported a patch of tough keratinous skin, which would have helped protect its face while digging.


We have a fairly good picture of the evolutionary origins of most groups of aquatic mammals – except for the pinnipeds. The fossil record of early seals is still rather sparse, and for a long time the earliest known species was Enaliarctos, an animal that was already very seal-like and didn’t help much in figuring out whether seals’ closest living relatives are bears or musteloids.

But then Puijila darwini was found in the late 2000s, a transitional form with a near-complete skeleton, filling in a gap in our understanding so conveniently it almost seems too good to be true.

This is the equivalent of Archaeopteryx for seals.

Discovered in Nunavut, Canada, Puijila dates to the early Miocene, about 23-20 million years ago. It was a small freshwater otter-like animal, about 1m long (3’3″), with a long tail and webbed feet adapted for paddling with all four of its limbs.

It lived at around the same time as the more specialized Enaliarctos, so it wasn’t a direct ancestor of modern seals, instead being part of an early offshoot lineage that retained more basal characteristics – but it does gives us a clue as to what the earliest pinnipeds looked like. Along with genetic studies it also helped to clarify that seals’ closest relatives are indeed the musteloids, although they’re estimated to have last shared a common ancestor around 45 million years ago so there’s still a lot of time unaccounted for in the proto-seal fossil record.

Several other fossil species that were previously thought to be musteloids have now also been recognized as close relatives of Puijila, and it seems that they were a fairly widespread group basically filling the ecological niche of otters at a time before true otters existed.

Most surprising and frustrating of all, however, is that some of these other otter-seals actually survived all the way into the Pleistocene, only going completely extinct sometime in the last 2 million years.

We barely missed having them still alive today!


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.


Procynosuchus delaharpeae, a synapsid from the Late Permian (~259-252 mya). Measuring about 60cm long (2′), it was one of the earliest members of the cynodonts, the lineage that would eventually lead to mammals.

Its fossils are mostly known from southern Africa, but similar remains have also been found in Europe and Russia, suggesting it was actually quite widespread across the supercontinent of Pangaea that existed at the time.

It had a long vertically-flattened tail, strong leg muscles, and paddle-like feet – all adaptations that suggest it was a semi-aquatic otter-like animal capable of agile swimming. It also had forward-facing eyes, giving it good binocular vision and depth perception while pursuing fish underwater.

Month of Mesozoic Mammals #26: Crunch Time


While true marsupials didn’t appear in the fossil record until the early Cenozoic, some of their closest relatives in the Mesozoic were a group called stagodontids. These metatherians evolved in North America during the Late Cretaceous, and much like the deltatheroideans seem to have taken over some of the ecological niches left vacant after the extinction of most eutriconodonts.

Their strong jaws and large blunt premolar teeth were adapted for crushing hard-shelled food, and they seem to have been semi-aquatic swimming animals (somewhat similar to modern water opossums) specialized for eating freshwater invertebrates such as snails and crustaceans.

Most stagodontids went extinct during the end-Cretaceous mass extinction, but if the early Cenozoic genus Eobrasilia is a member of the group then at least some survived in South America up until about 52 million years ago.

Other metatherians persisted in Europe, Africa, and Asia for most of the Cenozoic, with some fossils dating to just 11 million years ago, and the sparassodonts were successful in South America until about 3 million years ago – but today the only living members of this branch of the therian lineage are the marsupials in the Americas and Australasia.

Didelphodon was a stagodontid living during the Late Cretaceous of North America (70-66 mya), and is known from skull and jaw remains, along with a fairly complete skeleton that hasn’t been officially described yet.

It had a long otter-like body and flexible feet, adaptations for efficient swimming, and was one of the largest known mammals at the time, measuring around 1m in length (3′3″).

Its sturdy jaws had an especially powerful bite force, one of the strongest relative to body size known for any mammal, and some of its molars were blade-like and similar in appearance to carnassials. Along with crunching on shellfish it would also have been capable of crushing bones and tough plants, and its diet was probably an omnivorous mixture of invertebrates, vegetation, carrion, and small vertebrates – potentially including other mammals and small dinosaurs.

Month of Mesozoic Mammals #14: Otter-Mimics


The last eutriconodont featured this month specialized for a semi-aquatic lifestyle very similar to modern otters.

Known from the Early Cretaceous of China (125-112 mya), Liaoconodon was about 35cm long (1′2″) and had a long streamlined body and paddle-like limbs. Like other eutriconodonts it was carnivorous, likely feeding on fish and aquatic invertebrates in its wetland habitat.

Its ears show a transitional state between those of earlier mammaliaformes and modern mammals, with the inner ear bones almost fully separated from the jaw aside from a thin rod of cartilage. While this cartilage disappears during embryonic development in modern mammals, in Liaoconodon it was ossified (turned to bone) and appears to have helped to support the eardrum – although it’s not clear whether this was the ancestral state for Mammalia and fully separated ear bones convergently evolved multiple times in different lineages, or whether this was an evolutionary reversal within the eutriconodonts.

Month of Mesozoic Mammals #06: Going for a Swim


While some docodonts like Agilodocodon were going up into the trees, another branch of the group was specializing into semi-aquatic habits instead.

Castorocauda is known from the Middle Jurassic of China (165-161 mya), represented by an exceptionally preserved fossil showing soft tissue and hair impressions. About 40cm long (1′4″), it would have lived in a wetland environment and was well-adapted for swimming, with a flattened scaly beaver-like tail, webbed toes, and a coat of dense fur very similar to that of modern mammals, made up of both guard hairs and underfur.

Its strong forelimbs suggest it was capable of digging burrows, like modern platypus, and its sharp backwards-pointing teeth indicate a diet of slippery prey such as fish and worms.

It was also one of the earliest known mammals with (possibly venomous) spurs on its ankles. This feature is only seen today in monotremes, but seems to have been an ancestral trait common to all early mammals that was later lost in the lineage leading to marsupials and placentals.

Month of Mesozoic Mammals #02: Swimming Cousins


Known from the Early Jurassic of Arizona (196-183 mya), Kayentatherium was part of a group of cynodonts called tritylodontids – very close cousins of the true mammals, specialized for herbivory. They had strong jaw muscles, large incisors, and grinding cheek teeth, an arrangement convergently similar to modern rodents, and were some of the latest-surviving non-mammalian synapsids, persisting into the Early Cretaceous.

Kayentatherium wellesi skull by 5of7 || CC BY 2.0

Kayentatherium was one of the larger tritylodontids at just over 1m long (3′3″), and appears to have been semi-aquatic, with oar-shaped hindlimbs and a flattened beaver-like tail. Although not the first non-mammalian synapsid to be interpreted as a swimmer, it was the earliest close relative of the true mammals to develop these sorts of adaptations.