It Came From The Wastebasket #20: Colossally Convoluted Condylarths

“Insectivora” was a wastebasket taxon so bad it had to be revised multiple times, but there’s another particularly infamous case in mammal taxonomy that’s still in the process of being resolved – the “condylarths.

This group was first created in the early 1880s, during the Bone Wars, and initially was just a subgroup of odd-toed ungulates containing only the phenacodontids. But just a few years later Condylarthra was promoted up to its own order, and groups like the periptychids and hyopsodontids were added in too.

Then over the next few daceds century various groups were added and removed from the condylarths, most notably with the mesonychids and arctocyonids being brought in from their previous position with the creodonts.

By the mid-20th century the condylarths had become a big convenient dumping ground for any and all “primitive” ungulate-like mammals that didn’t easily fit into any modern groups, ranging in age from the early Paleocene through to the early Oligocene. But it soon became apparent that they had the same problem as the “insectivores” – there weren’t really any unique anatomical features that united all these animals together.

They generally had rounded-cusped molar teeth and hoof-like toes, but they also had rather generalized “primitive mammal” features and a diverse range of ecologies. Some were small herbivores, but others were coati-like or dog-like omnivores, and some were even bear-sized carnivores.

An illustration showing five different "condylarths". On the top row is Hyopsodus, a small guinea-pig-like animal with a long horse-like head; then Meniscotherium, a slightly larger animal that somewhat resembles a capybara; then the much larger Arctocyon, which has a slightly bear-like body, hoofed towes, and a dog-like head. On the bottom row is Ectoconus, a small animal with a tapir-like body and a blunt rectangular snout; then Mesonyx, a much larger dog-like animal with hoofed toes and a long tail.
From left to right, top row: Hyopsodus lepidus (hyopsodontid), Meniscotherium chamense (phenacodontid), Arctocyon primaevus (“arctocyonid”).
Bottom row: Ectoconus ditrigonus (periptychid), Mesonyx obtusidens (mesonychid)

It wasn’t even clear how the various different condylarth groups were actually related to each other. The best guess was that arctocyonids had arisen from within the “insectivores”, with a Protungulatum-like form as the common ancestor of all the other condylarths. Where exactly modern ungulates had then evolved from within the condylarths was also still uncertain.

Cladistic analysis in the 1980s began to tackle the confusing pile of assorted condylarths, and showed that they weren’t the single ancestral source of all modern ungulates, but instead a loose collection of several unrelated groups from all over the ungulate evolutionary tree. Arctocyonids, periptychids, and hyopsodontids were placed as early “primitive” lineages, phenacodontids were loosely linked with the ancestors of odd-toed ungulates once again, and mesonychids were considered to be the ancestors of whales.

An image of a diagram from a 1994 academic paper, showing the proposed evolutionary relationships of various "condylarths" with main ungulate groups. It differs majorly from modern understanding by its inclusion of elephants, sirenians, hyraxes, and their extinct relatives, along with showing whales as descending from mesonychids instead of artiodactyls.
…And if you know modern mammal phylogeny you’ll probably see some big problems here. 🐘
(Image source:

And, once again paralleling the mess of the “insectivores”, it wasn’t until genetic methods became available in the late 1990s that larger-scale ungulate relationships began to be properly resolved. The paenungulates (elephants, hyraxes, and sirenians), which had been traditionally considered to be a major branch of ungulates, were removed entirely and reclassified as afrotheres. And, along with some new fossil discoveries, whales were recognized as having actually evolved from within the even-toed ungulates instead of from mesonychids.

This shake-up threw the still-problematic “condylarth” classifications back into question – with some “condylarths” turning out to also be afrotheres instead of true ungulates.

Today the actual relationships of the main “condylarth” ungulate families are still in the process of being figured out, and there’s a lot of remaining uncertainty and disagreement about them.

Phenacodontids seem to have mostly maintained their traditional position as early odd-toed ungulates, and hyopsodontids may potentially be part of this group too – possibly as members of the hippomorph lineage, closely related to horses and brontotheres. Arctocyonids might be a wastebasket themselves, with some studies finding them to be a mix of several different archaic ungulate lineages. Periptychids may have links to the even-toed ungulates. The mesonychids, meanwhile, are now generally considered to be a separate order from the traditional “condylarths”, and may be either an early branch of the even-toed ungulates or much more basal ungulates closely related to the “arctocyonids”.

Since the term “condylarth” no longer has any real taxonomic meaning some paleontologists have proposed replacing it with “archaic ungulate” to distance from the historical messiness of the old name. But this hasn’t really caught on, and many papers still use “condylarth” in a very loose sense to refer to an “evolutionary grade” of early ungulates of unclear evolutionary affinities.

A cladogram showing the modern classification of several different "condylarth" families. They're shown as potentially occupying positions throughout the branches of the even-toed and odd-toed ungulates.

And while that’s the last main entry for this month, we’re not quite done yet. There’s still one weekday left in October, and after digging through so many taxonomic garbage cans there’s only one place we can go now.

…See you in the trash heap.

It Came From The Wastebasket #11: A Cetothere Change

Cetotheres were a group of small baleen whales, one of three major lineages of these cetaceans alongside the rorquals and the right whales. They first appeared in the fossil record in the mid-Miocene, about 14 million years ago (but are estimated to have actually originated 10-15 million years earlier), and disappeared during the Pleistocene about 2 million years ago.

First recognized in the mid-19th century, for a long time the cetotheres were used as a wastebasket for all fossil baleen whales that didn’t clearly fit into any modern whale families. By the start of the 21st century nearly 30 different genera representing numerous different species were all lumped into the group – and the genus Cetotherium was another wastebasket in itself with at least 12 assigned species, many of which were based on fragmentary or dubious remains.

An illustration of Ciuciulea, an extinct cetothere baleen whale. It looks similar to modern rorquals, with a streamlined and relatively slender body, but lacking the typical throat grooves of that type of whale. Its jaws are long and narrow, with a paired blowhole at the top of its head and small eyes set near the corners of its mouth. It also has fairly small flippers, a dorsal fin set about two-thirds of the way down its back, and a horizontal tail fluke. It's coloration is countershaded, mostly grey on top and white underneath.
Ciuciulea davidi

This was finally cleaned up in the 2000s, when a revision of the cetotheres cut the group down to just 6 genera. Since then a handful of additional new genera and species have been named, and while a few polyphyletic Cetotherium species may still need tidying up the cetotheres have overall gone from being a total taxonomic mess to actually being one of the best studied groups of fossil baleen whales.

Their exact evolutionary relationships with each other are still in flux, but the most surprising discovery from the improved understanding of these ancient whales is that they might not be extinct after all.

A set of three screenshots from the animated movie "Ice Age: Dawn of the Dinosaurs". The first image shows the two opossum characters Crash and Eddie asking, "Were you killed?!" The second image shows the weasel buck looking away and replying, "Sadly, yes…" The third image shows Buck leaning closer and adding "but I lived!" Buck's face has been photoshopped into that of a Cetotherium whale in both shots.

A study in the early 2010s suggested that the pygmy right whale may actually be a living cetothere. This classification was initially controversial, but further discoveries of fossil relatives of this enigmatic modern whale and comparisons of their distinctive inner ear anatomy have provided stronger evidence for an evolutionary link.

At this point it seems fairly likely that the pygmy right whale really is either the last surviving representative of the cetothere lineage, or at least is a very close evolutionary “cousin” (a “cetotherioid”) closer related to them than to any other modern baleen whales.


Tsaidamotherium hedini was a ruminant ungulate living around 11 million years ago during the late Miocene, in the northeastern part of the Tibetan Plateau in what is now Northwestern China. Although it’s known only from partial skull remains it was probably similar in body size to a large sheep, about 80cm tall at the shoulder (2’7″).

Since its discovery in the 1930s it’s traditionally been classified as part of the muskox lineage, but in 2022 it was proposed to actually be a giraffoid very closely related to the newly-discovered Discokeryx.

Tsaidamotherium had some extremely unusual headgear, with highly asymmetrical “horns” (actually ossicones if was a giraffoid). The left one was small and positioned above the eye, while the right one was shifted back and towards the middle of the forehead, and was expanded out into a wide bony disk that would have supported a large helmet-like domed keratin covering.

Its skull also had a very large nasal cavity resembling that of the modern saiga antelope, suggesting it may have convergently evolved a similar sort of complex air-filtering snout to deal with dry cold air in its mountainous habitat.


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 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.


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.


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.


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.


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”