Almost-Living Fossils Month #09 – Horned Sharks

All modern species of sharks and rays are part of a single lineage of cartilaginous fish known as neoselachians, and the closest evolutionary “cousins” to all of them were the hybodontiformes.

First appearing way back during the Devonian, about 400 million years ago, these early sharks were widespread around the world and incredibly successful as a group, living in both marine and freshwater environments.

Although due to their cartilaginous skeletons hybodontiformes are mostly known from fossilized teeth, there are still some complete specimens known that show us their overall body shape. They had two dorsal fins, each with a long spine in front, and an asymmetrically-shaped tail. Some of them also had small horn-like spines on their heads – this seems to be a sexually dimorphic trait, since the ones with “horns” also have claspers which show they were males – and they generally had powerful jaws with teeth specialized for crushing.

They were probably fairly slow swimmers most of the time, but would have still been capable of occasional bursts of higher speed, and various species were adapted to a wide range of food sources. Some had wider flatter teeth for cracking open hard-shelled seafloor invertebrates, and others were more opportunistic hunters that would have crunched on pretty much anything they could fit in their mouths.

Hybodontiformes were the dominant type of shark around the world before the end-Permian “Great Dying” mass extinction (~252 mya), and then went on to recover and flourish once again up until the mid-Jurassic.

Hybodus hauffianus was one of the Early Jurassic species, living around 183 million years ago in Europe. About 2m long (6′6″), it had two different types of teeth in its mouth – sharper ones in the front and flatter ones in the back – suggesting it was a generalist predator eating whatever it could catch. We do know its diet at least included the fast-swimming squid-like belemnites, since some fossils preserve clusters of their internal hard skeletons in Hybodus’ stomach region.

Towards the end of the Jurassic neoselachians began to diversify and take over most of the marine shark ecological niches, and the hybodontiformes became increasingly restricted to freshwater. During the Cretaceous they continued to do fairly well in those environments, but most of them still disappeared around the time of the end-Cretaceous extinction (~66 mya). Since most other sharks weren’t actually particularly affected by the extinction event, it’s not clear whether the hybodontiformes were more vulnerable for some reason or whether it was the ongoing competition from neoselachians that drove the majority of them extinct at that time.

Still, a few of them did seem to make it through to the Cenozoic, although they were absent from the fossil record until the Miocene. Freshwater deposits in Sri Lanka have evidence of a late-surviving member of the group living perhaps as recently as 5 million years ago – so they would have only gone completely extinct sometime after that, and we probably missed seeing them alive by only a few million years at most.

Almost-Living Fossils Month #08 – A Lot Of Lobsters

Hoploparia was a type of clawed lobster that first appeared in the fossil record in the Early Cretaceous about 140 million years ago. Many many different species within this genus have been found all over the world – over 100 of them have been described! – with quite a lot of anatomical diversity between them, showing that these lobsters were very good at adapting to a wide range of habitats and climates.

Although the vast majority of Hoploparia species lived just in the Cretaceous period, a small number of them did survive the end-Cretaceous mass extinction 66 million years ago. Hoploparia stokesi here was one of them, known from both the Late Cretaceous and Early Paleocene of Antarctica (~70-61 mya) – and was actually one of the first fossils ever described from the continent.

Specimens of this species are usually about 13cm long (5″), and show an evolutionary shift over time, developing much stronger claws and jaws, suggesting they were adapting their diet towards hard-shelled prey.

Various species of Hoploparia persisted on in North America, Europe, and Antarctica for the first half of the Cenozoic, but they never recovered to anywhere close to their Cretaceous levels of diversity. By the Early Miocene (~23-16 mya) there was just one known species left hanging on in Antarctica, and then they were gone.

(However, some modern lobster genera may in fact have originated from somewhere within the huge Hoploparia lineage back in the Cretaceous, so they might at least still have some close living relatives!)

Almost-Living Fossils Month #07 – Scaly Amphibians

Although there are just three main types of modern amphibians alive today – the frogs, salamanders, and caecilians, collectively known as “lissamphibians” – they weren’t always the only ones.

A fourth major lineage called albanerpetontids originated in the Middle Jurassic, about 166 million years ago. They’re usually thought to be slightly closer related to frogs-and-salamanders than to caecilians, but they also might not quite be true lissamphibians and instead belong just outside the group as evolutionary “cousins”. It’s a little complicated since we’re still not actually sure which group of ancient amphibians the lissamphibians even evolved from.

Resembling tiny salamanders, usually only around 5cm long (2″), albanerpetontid fossils have been found in North America, Europe, Africa, and Asia. In some places their remains are actually quite common, suggesting they were one of the more numerous small vertebrates in their ecosystems. They’re thought to have lived mostly in leaf litter, similarly to some small modern lizards like ground skinks, wriggling and burrowing through the loose material and preying on small invertebrates.

Their most notable feature was their body being covered in a mosaic of small scales – although unlike reptile scales these were bony structures formed under a layer of skin, structurally much more like fish scales, and they probably weren’t particularly visible in life. They also had very flexible necks for amphibians, with a convergently mammal-like joint between their skull and vertebrae.

After surviving the end-Cretaceous extinction alongside their lissamphibian relatives the albanerpetontids ended up mostly restricted to just Europe, but they seem to have continued on there for pretty much the entire Cenozoic.

Albanerpeton pannonicus here was one of the very last known members of the group, living just 2.5 million years ago in the late Pliocene and early Pleistocene. Despite the albanerpetontids’ 160-million-year history and having made it through multiple mass extinctions, it seems to have been the cooling Ice Age climate that finally sent these scaly little amphibians into extinction.

Almost-Living Fossils Month #06 – Circle Fish

Pycnodonts were a group of fish that originated in the western Tethys Sea during the Late Triassic (~215 mya), and later spread to most of the rest of the world with the exception of Antarctica and Australia. Ranging in size from a few centimeters to around 2m (6′6″), they had deep vertically-flattened bodies and almost circular silhouettes. Although they somewhat resembled modern marine angelfish or butterflyfish, they weren’t actually very closely related, instead being part of a much older branch of neopterygian fish.

They inhabited a range of shallow coastal waters from marine to freshwater environments, and most of them had jaws full of round flat teeth used to crush hard-shelled prey – but some may have been herbivorous grazers similar to parrotfish, and one lineage even became sharp-toothed piranha-like predators.

Some also developed quite elaborate appearances, such as Hensodon spinosus here. Living during the peak of the pycnodonts’ diversity in the mid-to-late Cretaceous, its fossils are known from Lebanon and date to about 100-95 million years ago.

It was only about 7cm long (2.75″) but it was bristling with various small spines and large “horns”, with different specimens showing two distinct arrangements. One type had double-pronged forward-facing horns, while the other had two horns one after the other – this may be evidence of sexual dimorphism, with the “bull horned” form thought to be male and the “rhino horned” form thought to be female.

Hensodon was also probably stripy in life, since one fossil preserves faint evidence of a light-and-dark stripe pattern on its dorsal fin.

Only a few pycnodonts survived into the Cenozoic, and their last appearance in the fossil record was in the mid-Eocene (~40 mya). Since this was at about the same time that more modern types of reef fish began to evolve, it’s likely that a combination of new competition and changing climate conditions resulted in the last pycnodonts going extinct by the end of the Eocene around 33 million years ago.

Almost-Living Fossils Month #05 – Cryptic Choristoderes

The choristoderes were a group of aquatic reptiles that mostly inhabited freshwater environments. Known mainly from North America, Europe, and Asia, they first appeared in the fossil record in the Late Triassic (~205 mya) – although their lineage could potentially go back further than that – and they varied in appearance from large long-snouted croc-like creatures to more lizard-like and miniature plesiosaur-like forms.

Many them were fully aquatic and spent their entire lives in the water, with some developing the ability to give live birth and others returning to land only to lay eggs (with only females having well-developed enough limbs to be able to haul themselves out onto shore). In some places their fossils are incredibly common, with every life stage represented from babies to adults (even one with two heads!), and yet despite having such detailed knowledge of their lives we still don’t know exactly what type of reptile they actually were.

Their evolutionary origins and relationships are very unclear, with the only real certainly being that they’re at least diapsids. They’re often classified as either archosauromorphs (closer related to crocodilians and dinosaurs/birds) or as lepidosauromorphs (closer related to lizards), but they could also be a much earlier separate branch of the reptile family tree.

Some of the large crocodilian-like neochoristoderes survived into the Cenozoic and initially did quite well for themselves – even outcompeting actual crocodilians in the northern continents for a while – but then they seem to have fallen victim to the cooling and drying climate of the Eocene-Oligocene extinction about 33 million years ago.

But that wasn’t the end of the choristodere lineage just yet.

A small number of fossils of a little choristodere named Lazarussuchus have been found in a few different places around Europe, with the youngest specimens dating to as recently as the Early Miocene (~20-16 mya). Surprisingly it wasn’t closely related to the neochoristoderes at all, but instead seems to have been part of a much older and more “primitive” branch of the choristodere family tree that must have been surviving since at least the mid-Jurassic with very little presence in the fossil record.

At about 30cm long (1′) it was less aquatic than most other choristoderes, with large claws that would have given it good traction on land and a more generalized lizard-like body plan. One specimen even preserves soft tissue impressions, showing that its toes lacked webbing and that it had a low crest running along its tail.

Its lack of specialization may have been the reason for its longer survival, being able to adapt to a wider variety of environments compared to its more water-reliant cousins.

It’s unclear exactly how much closer to present day these rare last choristoderes survived. If they managed to make it through the mid-Miocene extinction then they might potentially have persisted until the onset of the Pleistocene Ice Age 2.5 million years ago – but their fossils are scarce enough that we’ll probably never know for certain.

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 #03 – The Crowned Starfish

Known mainly from around Europe – with a few records from the Atlantic coast of North America and the Caribbean – the stauranderasterids were a family of starfish that first appeared about 190 million years ago in the Early Jurassic.

Not much is known about their life ecology, although they seem to have inhabited shallow tropical seas and like many other starfish would probably have preyed on various slow-moving marine invertebrates. Complete specimens are very rare compared to just isolated elements, making their maximum size difficult to estimate, but they likely grew to at least 5-10cm across (2″-4″).

They had enlarged ossicles forming a bumpy “crown” over their central disc, with five arms that could be either narrow and elongated (such as in Stauranderaster coronatus here) or shorter and club-shaped (like Manfredaster bulbiferus). This gave them some visual similarities to modern starfish like Protoreaster, and since they were both part of a larger grouping of starfish called valvatids it’s unclear whether these features mean that they were very closely related to each other or if it was simply due to convergent evolution.

The stauranderasterids survived until at least the Late Paleocene/Early Eocene (~56 mya), but some possible remains from Cuba date to as recently as the Early Micoene (~23-16 mya).

Almost-Living Fossils Month #02 – The Saber-Toothed “Herring”

First appearing in the mid-Cretaceous, about 113 million years ago, Enchodus was a small-to-medium-sized genus of predatory fish. Different species ranged from a few centimeters to up to 1.5m in length (4′11″), with Enchodus gladiolus here being an averaged-sized example at about 60cmlong (2′).

The most distinctive feature of these fish were the enlarged fang-like teeth in both their upper and lower jaws, over 6cm (2.4″) long in the largest individuals, which may have been a specialization for feeding on soft-bodied cephalopods.

Despite having been nicknamed “saber-toothed herrings”, they weren’t actually closely related to herrings at all, instead being part of the aulopiformes – a group also containing modern lizardfish, lancetfish, and a different type of sabertooth fish.

Fossils of various Enchodus species have been found all over the world, and they seem to have been very common and important members of ancient marine ecosytems, occupying mid-level carnivore niches and in turn being eaten by other predators. Their remains have been identified within the preserved stomach contents of marine reptiles such as plesiosaurs and mosasaurs, as well as sharks and hesperornithean birds.

These toothy fish survived through the end-Cretaceous mass extinction and continued their success for almost 30 million years into the Cenozoic, with the last known fossils dating to just 37 million years ago in the Late Eocene. They probably didn’t survive much longer beyond that date, since there was an extinction event at the Eocene-Oligocene boundary (~34 mya), a period of sudden cooling that affected many marine animals at the time.

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.)

Month of Mesozoic Mammals #31: The Survivors


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.