#theriiformes

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".

[ 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 ]

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Almost-Living Fossils Month #24 -- Sabertoothed Sparassodonts

Along with the marsupials and the polydolopimorphs, the sparassodonts were one of the lineages of metatherian mammals that inhabited South America during its “great isolation” for most of the Cenozoic. And despite having to share the large carnivore niches with both the terror birds and the sebecosuchian crocs, they still managed to become the main mammalian predators of the region.

Their first definite fossils come from the start of the Paleocene (~65 mya), but they probably actually originated sometime in the Late Cretaceous before the mass extinction. A currently-unnamed skull from Mongolia (~70 mya) appears to be either an early sparassodont or a very close relative, and a North American metatherian called Varalphadon (~90 mya) may also be linked to the group. It’s possible that, like the marsupials, they may have first evolved in North America and later spread into South America before it became isolated.

Like their marsupial relatives they would have given birth to tiny undeveloped young, although we don’t know for certain if they actually had pouches or not. Their epipubic bones were highly reduced, so it’s possible they didn’t have pouches -- but they also might have had mostly cartilaginous epipubics (like thylacines) that just didn’t fossilize.

Over the course of the Cenozoic the sparassodonts convergently evolved many similar features to placental carnivorans, with carnassial teeth for shearing through flesh and a wide variety of body shapes ranging from small weasel-like forms to long-snouted ambush hunters to large hyaena-like bone-crushers.

But by far the most famous members of the group were the thylacosmilids. First appearing in the Early Miocene, about 20-15 million years ago, these sparassodonts developed huge elongated canine teeth that resembled those of sabertoothed cats. Unlike the felid sabertooths, however, thylacosmilids’ fangs grew continuously and their lower jaws had long bony flanges that supported and protected their teeth when theirs jaws were closed.

Thylacosmilus atrox was the last and most highly specialized of the thylacosmilids, living from the Late Miocene to the Late Pliocene, around 9-3 million years ago. About 1.2-1.5m long (~4-5′) and standing 60cm tall at the shoulder (2′) it was similar in size to a modern jaguar -- not huge compared to some placental predators, but still one of the largest of all known carnivorous metatherians.

Despite its huge fangs it actually had a fairly weak bite force, instead relying on its strong forelimbs to immobilize its prey before delivering precise deep stabs into soft body parts using powerful neck muscles. The structure of its limbs also suggests it wasn’t a fast runner, and it probably had to stalk or ambush its targets.

It’s finally time for “Almost-Living” Fossils Month!

There are various modern species that we call “living fossils” because they’ve survived for very long periods of geological time, but there are also plenty of long-lived and successful groups that didn’t quite manage to make it to the present day.

For all of August I’ll be doing six posts on this topic a week (taking a break on Sundays), and the basic criteria for entries are that a featured organism has to be part of a group that:

originated in the Mesozoic or earlier

existed for at least 50 million years

and survived through the end-Cretaceous mass extinction, only to go extinct later in the Cenozoic

So, let’s begin with...

Almost-Living Fossils Month #01 -- The Saint Bathans Mammal

Dating to the mid-Miocene, about 19-15 million years ago, this little mammal was discovered in the Saint Bathans fossil deposits in the South Island of New Zealand, and was probably the size of a large shrew, around 10cm long (4″). It doesn’t have an official scientific name yet, but its fragmentary remains might represent something very special.

Not only was it a terrestrial mammal living at a time when none were thought to inhabit New Zealand at all, but it also doesn’t seem to fit into any known group of mammals. It’s definitely not a placental or marsupial or monotreme, and it doesn’t match any of the other “archaic” groups -- so it could represent an entirely new lineage we didn’t even know existed.

At best it’s been classified as a theriiform, belonging somewhere between eutriconodonts and multituberculates in the mammal family tree. This means it would have to be part of a group with a very long ghost lineage going all the way back to the Late Triassic or Early Jurassic (~220-190 mya). Whatever it actually was, its ancestors were probably already present on New Zealand when the islands split away from Gondwana in the Late Cretaceous (~85 mya), and must have remained isolated there for tens of millions of years more.

It’s unknown how much closer to present day these mysterious archaic mammals actually survived, but it’s possible they were one of the casualties during the Middle Miocene disruption extinction about 14 million years ago, when the relatively warm climate suddenly cooled. And then land mammals were completely absent from New Zealand until the arrival of humans ~750 years ago.

Almost-Living Fossils Month #18 -- Metatherians Everywhere

While modern marsupials are native only to Australasia and the Americas, they’re part of a larger grouping of mammals known as metatherians that were once found around most of the world. Thought to have originated in Asia, perhaps as far back as the Late Jurassic (~160 mya), the ancestral metatherians spread to Europe and the Americas during the Cretaceous and gave rise to multiple different lineages, ranging from small sabertoothed carnivores to otter-like swimmers to the main mammalian predators of Cenozoic South America.

True marsupial fossils didn’t appear in the fossil record until the start of the Paleocene (~65 mya), but some of their closest evolutionary “cousins” -- the herpetotheriids -- originated a little earlier in the Late Cretaceous, around 70-66 million years ago.

Herpetotheriids were small opossum-like animals found in North America, Asia, Europe, and Africa. They were more terrestrial than the opossums they resembled, adapted for running on the ground rather than climbing, but were probably similarly opportunistic omnivores eating a wide variety of foods.

They survived through the end-Cretaceous extinction and seem to have actually been quite common in the northern continents for most of the Cenozoic. They also weren’t the only metatherians in those regions, living alongside another group called the peradectids which were true marsupials closely related to opossums.

In North America the native metatherians all went extinct sometime around the early Miocene (~20 mya) -- although the continent would later be recolonized by opossums from South America during the Great American Interchange. Meanwhile the African herpetotheriids disappeared around the same time, but the European and Asian metatherians continued on into a little longer into the mid-Miocene.

Amphiperatherium frequens was one of the last known members of the herpetotheriids, living in Europe during the early-to-mid Miocene (~20-13 mya). It was about the size of a small rat, around 15-20cm long (6-8″), and seems to have mainly inhabited warm humid environments.

Although the disappearance of the northern metatherians has traditionally been blamed on them being out-competed by placental mammals, the fact that they survived alongside placentals just fine for over 45 million years suggests that something else pushed them over the brink. Their extinction may in fact be more linked to the ongoing drying and cooling climate trends in the latter half of the Cenozoic.

Almost-Living Fossils Month #19 -- Even More Metatherians

While the opossum-like herpetotheriids and peradectids survived in the northern continents for most of the Cenozoic, a wider variety of metatherian mammals were found in the south. Alongside the true marsupials and the sparassodonts, a group known as the polydolopimorphs existed in South America for over 60 million years. Although most of the their fossil remains consist only of isolated teeth and jaw fragments, they seem to have been a very diverse group that adapted to a wide range of ecological niches including insectivores, herbivores, and fruit-eating and seed-eating specialists.

Their exact evolutionary position within the metatherians is still rather unclear and under dispute, with different studies giving different results. They were probably marsupialiformes, slightly less closely related to marsupials than the herpetotheriids, but some paleontologists instead consider them to have been true marsupials related to either the shrew opossums or the microbiotheres. (And some go with both options, proposing that they weren’t even a natural group but were polyphyletic, with some being marsupialiformes and others being marsupials.)

The earliest definite polydolopimorph fossils come form the very start of the Paleocene in South America (~66 mya), but their lineage likely goes further back into the Late Cretaceous -- possible remains from North America suggest they may have originated there at least 70 million years ago, with their ancestors migrating into South America shortly before the end-Cretaceous extinction. A few also reached Antarctica by the Late Eocene (~40-33 mya), before the continent had fully separated from its neighbors and frozen over, but its unclear whether any ever made it as far as Australia alongside their marsupial relatives.

They were most diverse during the first half of the Cenozoic, and in the latter half they were represented mainly by a highly specialized lineage called the argyrolagids. Known from western and southern South America (Peru, Bolivia, Argentina, and Chile) from the Early Oligocene onwards, these polydolopimorphs were convergently rodent-like desert herbivores with short forelimbs and long hopping hindlimbs that gave them a resemblance to jerboas or springhares.

Argyrolagus palmeri here lived during the Early Pliocene of Argentina (~5-3.5 mya). About 40cm long (1′4″), it had only two toes on its feet, a long pointed snout, and large eyes and ears that indicate it was probably nocturnal.

Month of Mesozoic Mammals #26: Crunch Time Didelphodon

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 #23: Toothy Jaws Cronopio

Cronopio was one of the earliest South American dryolestoids, living during the start of the Late Cretaceous of Argentina (100-94 mya). Known only from partial skull material, it had a long snout and enlarged canine teeth which gave it a superficial resemblance to the fictional “sabertoothed squirrel” Scrat from the Ice Age film series.

(from fig 4 in Rougier GW, Apesteguía S, Gaetano LC (2011) Highly specialized mammalian skulls from the Late Cretaceous of South America. Nature 479(7371):98–102 doi: 10.1038/nature10591)

Although its full size and appearance is unknown, it’s estimated to have measured about 15cm long (6″). It was probably an insectivore or an omnivore, but the rather delicate nature of its snout and fangs suggest it had a weak bite and relied on strong jaw muscles to chew up its food with a specialized rotating motion. It’s also not clear whether its saberteeth had a function for feeding, or if they were used for display and fighting like in some modern deer.

And while I’ve reconstructed Cronopio here with ankle spurs, it’s actually unknown whether dryolestoids had this feature. They occupied an evolutionary position between mammals that definitely had spurs (symmetrodonts) and ones that definitely didn’t (therians), but fossil remains of dryolestoid ankles are poorly-preserved and incomplete.

(I also wanted to do a depiction that wasn’t so blatantly Scrat-like, because that’s become sort of a paleoart meme. Soft-tissue and long fur can really change the outward appearance of mammals, so enjoy this weird tiny pig-rat version of Cronopio.)

Month of Mesozoic Mammals #28: Hop To It Zalambdalestes

Living during the Late Cretaceous of Mongolia (85-70 mya), Zalambdalestes was part of an early branch of the eutherian evolutionary tree -- a highly specialized group of mammals that it lends it name to, the zalambdalestids.

About 20-25cm in length (8-10″), it had relatively long limbs with especially strong hindlegs that show adaptations for rabbit-like hopping. Its long narrow snout may have ended in a flexible proboscis similar to those of modern sengi, and sharp interlocking teeth indicate a carnivorous or insectivorous diet.

Its long rodent-like incisors grew continuously throughout its life, suggesting it was gnawing on something tough enough to constantly wear down its front teeth.

(from fig 51 in Wible JR, Novacek MJ, Rougier GW (2004) New data on the skull and dentition in the Mongolian late Cretaceous eutherian mammal Zalambdalestes. Bulletin Of The American Museum Of Natural History 281:1-144 uri: http://hdl.handle.net/2246/449)

Some studies have proposed that zalambdalestids were actually very basal members of placental mammal groups such as rodents or rabbits, but the presence of epipubic bones in front of their pelvises (bones not found in placentals) shows they were a much earlier type of eutherian that still reproduced more like marsupials. Any anatomical similarities to later placentals were probably just the result of convergent evolution.