Almost-Living Fossils Month #04 – The Last Multis

Known almost exclusively from the southern continents of Gondwana – hence their name – the gondwanatheres were part of a widespread and very long-lived group of mammals known as multituberculates.

Although multis resembled placental rodents, and gave birth to tiny undeveloped young in a similar manner to marsupials, they originated much further back in the mammal evolutionary tree. They existed by at least the Early Jurassic (~183 mya), and their ancestry may go even further back into the Late Triassic (~220 mya) if they were descended from haramiyidans.

Multis survived the end-Cretaceous extinction and became very diverse through the first half of the Cenozoic, until a combination of factors such as climate shifts, new types of vegetation, and the evolution of new mammalian predators (and possibly also competition from placental rodents) resulted in most of them going extinct by the Early Oligocene (~33 mya) – with only the gondwanatheres surviving past that point in the then-isolated continent of South America.

Patagonia peregrina was the very last known gondwanathere in the fossil record, living just 21-17.5 million years ago in the Early Miocene of Argentina. Although only teeth and jaw fragments have been found so far, it was probably about 15cm long (6″) and would have been a burrowing herbivore similar to modern gophers or tuco-tuco. Its ever-growing rodent-like teeth were adapted for grazing on tough grasses in its savanna-like habitat, and it would have lived alongside several other now-extinct types of mammal – but we’ll be getting to those ones later in the month.

Since it seems like these last gondwanatheres had survived by retreating into a rather specialized ecological niche, they sadly probably didn’t persist for very long beyond the time of Patagonia. A wave of extinctions associated with sudden climate cooling about 14 million years ago may well have been the final blow to the once-successful lineage of the multituberculates.

Almost-Living Fossils Month #01

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…


An illustration of an extinct small mammal. It resembles a shrew, with a pointed whiskery snout, small rounded ears, and a long furred tail. It also has pointed spurs on its ankles.

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.

(It’s also worth noting that the fossils of the Saint Bathans mammal were described over ten years ago, and our knowledge of ancient mammals and their evolutionary tree has changed a lot in that time. New studies or more fossil material might reveal the Saint Bathans mammal to actually be one of the very last survivors of an already-known group, but that would still make it an amazing and unique discovery.)

Eucladoceros

Eucladoceros dicranios, a deer from the Pliocene and Pleistocene of Europe (~3.5-1 mya). Close in size to a modern moose, standing about 1.8m tall at the shoulder (5′10″), the males of this species had a set of particularly large antlers – measuring up to 1.7 meters across (5′6″) and bristling with at least twelve prongs each – giving it the nickname of “bush-antlered deer”.

The more famous “Irish elk” (Megaloceros giganteus) would later develop even bigger antlers, but Eucladoceros was the earliest known deer to evolve this sort of extremely elaborate headgear.

Orcinus citoniensis

Despite commonly being called “killer whales” modern orcas are actually the largest living members of the oceanic dolphin family. Their ancestors are thought to have diverged from other dolphins between 10 and 5 million years ago – and surprisingly their closest relatives are the much smaller snubfin dolphins found in Australasia.

Living during the Pliocene (5-2 mya) in the Mediterranean, Orcinus citoniensis was an early member of the orca lineage, and was probably a transitional form between their early dolphin ancestors and the modern Orcinus orca.

It was half the size of modern orcas, at about 4m long (~13′). While it had a higher tooth count than its living relatives its teeth were also proportionally smaller, suggesting it wasn’t specialized for tackling large prey and probably fed mainly on fish and squid.

Month of Mesozoic Mammals #31: The Survivors

Purgatorius

Quite a few groups of Mezozoic mammals actually made it into the Cenozoic – including multituberculates, dryolestoids, various different metatherians, cimolestans, leptictidans, and possibly another unknown lineage in New Zealand – but most of them eventually declined and died out, and only monotremes, marsupials, and placentals still remain alive today.

We don’t know exactly when placentals originated. The first definitive fossils come from the start of the Cenozoic, but a few early ancestral forms probably already existed during the Late Cretaceous (estimated up to 90-75 mya) and only got their chance to rapidly diversify immediately after the mass extinction event.

One of the closest fossils we have to the earliest placentals is Purgatorius. Known mainly from teeth from the Early Paleocene of North America (66-63 mya), it’s not entirely clear whether it actually existed in the Mesozoic, but its remains have been found very close to the K-Pg boundary and one fossil might actually be from the end-Cretaceous.

A few foot bones have been associated with some of the fossil teeth, and if they do belong to Purgatorius then they show that it had very flexible ankles, a characteristic typical of tree-climbing animals. It would likely have been a squirrel-like creature, about 15-20cm long (6-8″), eating an omnivorous mixture of insects, seeds, and fruit. It may also have been capable of burrowing similar to modern chipmunks.

It’s often been interpreted as a placental mammal, specifically a very early type of primate, but more recent studies suggest it might not even be a true placental at all  – although it was probably still a very close relative of the common ancestor of all living placentals.

Month of Mesozoic Mammals #30: Strange Relations

Gypsonictops

A group known as the leptictidans were probably some of the weirdest early eutherians. With their tiny forelegs, big hindlegs, and long counterbalancing tails, they somewhat resembled jerboas or small kangaroos – except they also had long slender snouts that probably ended in sengi-like proboscises, and their feet were structured more like those of running animals than jumping ones. They’re also thought to have been mainly bipedal, convergently evolving a similar posture and movement style to non-avian theropod dinosaurs.

Leptictidium (Eocene, 50-35 mya) by Tim Bertelink || CC BY-SA 4.0

First appearing in the Late Cretaceous, they made it through the end-Cretaceous extinction and survived up until the mid-Cenozoic across the northern hemisphere, going extinct around 33 million years ago. They were probably omnivores, eating a mixture of insects, small vertebrates, and soft plant matter such as fruit and leaves.

Their mix of “primitive” skull features and highly specialized skeletons makes classifying them particularly difficult. They’ve been proposed to be placentals related to primates and rodents or afrotheres, a very early branch of the eutherians, or close to placentals but not quite true members themselves. The latter interpretation currently seems most likely, but they could also be a paraphyletic group at the base of placentals (suggesting that they could even be ancestral to placentals, and therefore all placentals would technically be leptictidans).

Gypsonictops was one of the earliest leptictidans, living during the Late Cretaceous of North America (70-66 mya). Known only from teeth and jaw fragments, we don’t know much about its appearance or full size – although it was probably smaller than its later relatives, perhaps about 35cm long (1′2″).

Any reconstruction of such fragmentary remains is going to be very speculative, but I’ve restored it here as a sort of transitional form, not yet quite as specialized. A more sengi-like animal, mainly quadrupedal but able to run and hop on its hind legs to flee from danger or chase after small fast-moving prey.

Month of Mesozoic Mammals #29: Rooting Around

Schowalteria

First appearing in the Late Cretaceous, a widespread and diverse group of mammals known as cimolestans were once thought to be early members of placental groups like pangolins and carnivorans. But more recent studies have shown them to be part of a different branch of the eutherian family tree altogether, more like cousins to the earliest placentals and leaving no living descendants.

However, they did make it through the end-Cretaceous mass extinction and were quite successful during the early Cenozoic, evolving into forms ranging from giant herbivores to fanged squirrel-like climbers to otter-like swimmers, with the latter surviving until about 33 million years ago.

One group of North American cimolestans, the taeniodonts, were specialized for digging up tough roots and tubers, with large claws, strong blunt jaws, and big front teeth that became ever-growing in some species.

Schowalteria was the earliest known member of this group, living during the Late Cretaceous of Canada (70-66 mya). Only represented by partial skull material, its full size is unknown – some estimates put it at a similar size to giants like Repenomamus, but it was likely closer to half that size at around 50cm in length (1′8″). Still one of the larger Mesozoic mammals around, but not nearly as big as some of the Cenozoic taeniodonts would later become.

Month of Mesozoic Mammals #28: Hop To It

Zalambdalestes

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

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.

Skull of Zalambdalestes || 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.

Month of Mesozoic Mammals #27: Treetop Origins

Juramaia

The other major branch of the therian mammals are the eutherians – represented today by only the placentals. But although the first eutherians split from their common ancestor with the metatherians all the way back in the mid-Jurassic, more then 160 million years ago, true placentals don’t seem to have actually appeared until right after the end-Cretaceous mass extinction.

There were many other side-branches of the eutherian lineage during the Mesozoic, however, and these last few days of the month will cover some of them.

(Some of these relationships are still under dispute, such as the exact position of leptictidans and Purgatorius and whether they’re actually placentals or not. Eutherian phylogeny is a bit of a mess.)

Juramaia was the earliest known eutherian, living during the Late Jurassic of China (161-156 mya). About 15cm long (6″), it was a shrew-like insectivore with limb anatomy that would have allowed it to climb up trees in a similar manner to modern rats.

Something very similar to it would have been the common ancestor of all later eutherians – suggesting that the earliest members of the group may have started out as tree-climbers before diversifying into different niches later on.

But despite it being closer related to living placentals than to marsupials, placental-style reproduction hadn’t actually evolved yet and it would have still given birth to tiny undeveloped young.

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.