Neolicaphrium recens here might look like some sort of early horse, but this little mammal was actually something else entirely.
Known from southern South America during the late Pleistocene to early Holocene, between about 1 million and 11,000 years ago, Neolicaphrium was the last known member of the proterotheriids, a group of South American native ungulates that were only very distantly related to horses, tapirs, and rhinos. Instead these animals evolved their remarkably horse-like body plan completely independently, adapting for high-speed running with a single weight-bearing hoof on each foot.
Neolicaphrium was a mid-sized proterotheriid, standing around 45cm tall at the shoulder (~1’6″), and unlike some of its more specialized relatives it still had two small vestigial toes on each foot along with its main hoof. Tooth microwear studies suggest it had a browsing diet, mainly feeding on soft leaves, stems, and buds in its savannah woodland habitat.
It was one of the few South American native ungulates to survive through the Great American Biotic Interchange, when the formation of the Isthmus of Panama allowed North and South American animals to disperse into each other’s native ranges. While many of its relatives had already gone extinct in the wake of the massive ecological changes this caused, Neolicaphrium seems to have been enough of a generalist to hold on, living alongside a fairly modern-looking selection of northern immigrant mammals such as deer, peccaries, tapirs, foxes, jaguars… and also actual horses.
Some of the earliest human inhabitants of South America would have seen Neolicaphirum alive before its extinction. We don’t know whether they had any direct impact on its disappearance – but since the horses it lived alongside were hunted by humans and also went extinct, it’s possible that a combination of shifting climate and hunting pressure pushed the last of the little not-horses over the edge, too.
Mammalian tusks usually grow in symmetrical pairs with only minor developmental asymmetry, but a few species have evolved much more uneven arrangements.
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.
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.
It was smaller than modern pronghorns, around 70cm tall at the shoulder (~2’4″), and males had long antler-like horns with three tines. Bizarrely, one of these horns was always at least twice the size of the other, with “left-horned” and “right-horned” individuals seeming to occur in equal numbers.
Meanwhile in China another Miocene ungulate known as Tsaidamotherium hedini also had strange headgear, with an enlarged right “horn” forming a helmet-like dome on top of its head. This species was featured here on the blog just year, so check out that post for more details about it.
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.
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.
Antaecetus aithai was an early whale that lived during the late Eocene (~40 million years ago) in what is now Morocco, at a time when northern Africa was covered by a warm shallow sea.
It was part of the “basilosaurids“, some of the first fully aquatic cetaceans – traditionally considered to be a single defined group, but more recently found to be more of an “evolutionary grade” of multiple early whale lineages – and much like Basilosaurus it had elongated back vertebrae that would have given it a very long slender body shape.
Antaecetus also had a proportionally smaller head and smaller teeth than other basilosaurids, along with much denser bones and a stiffer spine that would have made it a rather slow swimmer with reduced maneuverability. It was also fairly small overall compared to most of its relatives, probably around 6m long (~20′).
It was probably a slow-moving coastal water animal somewhat like modern sirenians – except unlike manatees and dugongs it was carnivorous. Its relatively delicate teeth suggest it was feeding on soft-bodied prey like cephalodpods, and with its lack of speed it must have been some sort of ambush predator, waiting around for potential prey to come within striking range.
Towards the end of the Cretaceous, about 69 million years ago, the most diverse and numerous mammals in the northern hemisphere were the metatherians, close relatives of modern marsupials.
About the size of a modern mouse, around 10-15cm long (4-6″), and with teeth that suggest it was a shrew-like insectivore, this little metatherian lived in northern Alaska in what’s known as the Paaŋaqtat Province – a region with a distinctive population of endemic polar animals. At the time this area was located at an even higher latitude than it is today, around 80-85ºN, but due to a greenhouse climate it was also warmer, with no permanent ice and the average temperatures staying above freezing.
Unnuakomys was by far the most common mammal species in the Paaŋaqtat Province, represented by numerous fossil teeth and a few jaw fragments, and it also seems to have been the only metatherian living in the whole region. This may just be a preservation bias in the fossil record, but it might also indicate that Unnuakomys was uniquely specialized to endure the several months of continuous darkness each winter in its polar woodland environment, while other North American metatherians were restricted to more southerly latitudes.
And yet despite the huge diversity of gliding mammals, and their particular prevalence in tropical forests, there’s an entire continent famous for its rainforests that’s somehow completely lacking any modern examples: South America.
It’s not clear why the gliding lifestyle never took off in South America, but the continent is surprisingly devoid of any other gliding vertebrates, too. The only exceptions are a few species of flying frogs in the northwestern tropical forests around Colombia.
But back in the early Eocene, about 53-50 million years ago, there was at least one South American gliding mammal. Some fossil limb bones found in the Itaboraí Formation in southeastern Brazil look very much like those of a gliding mammal – long and thin, with a locking elbow joint, knees adapted for jumping, and flexible ankles typical of tree-climbers.
These remains haven’t been given a new scientific name, however, because there’s a good chance they belong to an already-described species. Fossils from Itaboraí are found disarticulated, broken, and with bones of multiple different species jumbled together, so most fossil mammals named from the site have been based on their more easily distinguishable teeth and jaw fragments.
The problem is matching those teeth with these bones.
Currently the best identity guess based on size is Gaylordia macrocynodonta. This mammal would have been around 30cm long (1′), about the size of a modern rat, and had distinctive large canine teeth. It used to be classified as a marsupial related to opossums, but more recent studies have found it to have actually been a marsupialiform metatherian instead, much more closely related to Pucadelphys and sparassodonts than to any modern true marsupials.
Gaylordia‘s crushing molars suggest it was carnivorous, able to crunch through bones or hard-shelled invertebrate prey. This would be a very unusual diet for a gliding mammal, since most other mammalian gliders are herbivores or omnivores – the only other known predatory examples were the volaticotheres over 110 million years earlier.
“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.
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.
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 toalso 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.
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.
The rodent-like multituberculates were a major lineage of mammals that were only distantly related to modern marsupials and placentals. They originated around the time of the mid-Jurassic (~168 million years ago), survived through the end-Cretaceous mass extinction, and went on to become one of the most diverse and successful types of mammal in the Paleocene. After that point they began to decline, and after anw over-130-million-year-long run they went extinct* in the early Oligocene (~33 million years ago).
(* Except, possibly, in South America, where an enigmatic fossil known as Patagonia peregrina may represent a multi surviving as recently as about 18 million years ago in the early Miocene.)
First discovered in North America in the 1880s, Catopsalis foliatus was part of a group of multituberculates called taeniolabidoids. These multis got significantly larger than the rest of their kind – averaging beaver-sized but with some species getting up to at least capybara-sized – and were some of the first mammals to evolve into relatively big herbivores after the extinction of the non-avian dinosaurs.
Catopsalis foliatus
Catopsalis was named based on a partial jawbone and a few teeth, and over the next century or so various other similar-looking fossils from both North America and Asia were added into the genus as additional species. Eventually Catopsalis contained eight different species, ranging over about 10 million years from the late Cretaceous to the early Eocene – not especially big compared to some other wastebaskets we’ve looked at this month, but it was still a problem, muddying up attempts to understand the actual evolutionary relationships and biogeography of the taeniolabidoids.
Cladistic studies in the 1980s showed that Catopsalis was paraphyletic, made up of at least five separate lineages, and a few of them were subsequently renamed and reclassified. The Cretaceous Asian forms became Djadochtatherium and Catopsbaatar, and are now considered to be part of a different lineage of multis known as djadochtatherioids, while one of the remaining North American species then became Valenopsalis.
…But a couple of other new Catopsalis species have also been named in the meantime (one as recently as 2018), so there are still seven different species that need sorting out in this particular wastebasket.
