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Almost-Living Fossils Month #23 – Enamel-Armored Lizards

The glyptosaurines were a group of lizards that first appeared in the Late Cretaceous, about 85 million years ago. They were an early branch of the anguid lineage, originating in North America, and had heavily armored bodies covered in bony osteoderms – superficially similar to those of modern beaded lizards, but structurally much more complex with the outermost layer formed from a unique enamel-like substance called osteodermine.

They were some of the few lizards to survived through the end-Cretaceous extinction 66 million years ago (which killed off over 80% of the lizard species known at the time) and went on to become quite successful in the warm climates of the early Cenozoic.

They spread across to Europe and Asia and developed much larger body sizes, going from small 10cm-long (4″) forms in the Early Paleocene (~65 mya) to over 60cm long (2′) by the mid-Eocene (~40 mya). In North America and Europe they became common enough that they were probably important parts of the local ecosystems, and their widespread distribution suggests they were able to adapt to a variety of different habitats and environmental niches.

Their teeth resembled those of modern omnivorous lizards like blue-tongued skinks, suggesting they had a similar generalist diet – although their strong jaws have also been proposed to be specializations for crushing hard-shelled invertebrates such as snails.

Helodermoides tuberculatus here was one of the largest glyptosaurines, about 65cm long (2′2″). It lived during the Late Eocene and Early Oligocene (~34-33 mya) in the northwestern and midwestern United States, with fossils known from Montana, Wyoming, and Nebraska.

One fossil shows evidence of having lost part of its tail, probably dropping it in a self-defense behavior to escape a predator. However, unlike the regenerating tails of many other lizards, the osteoderms of Helodermoides instead seem to have formed a thick rounded bony cap over the wound, preventing any significant regrowth and leaving its tail permanently stumpy.

During the Late Eocene and Early Oligocene the glyptosaurines began to disappear, probably struggling to cope with cooling and drying climates, and their last definite fossils date to about 30 million years ago. Possible fragmentary remains from as late as the Early Miocene of Central Europe (~16 mya) may indicate that a few isolated late-surviving members of the group persisted on for a while longer, but if they did hang on that long they were probably finished off by further sharp temperature drops in the mid-Miocene.

Almost-Living Fossils Month #22 – Some Marine Crocs

First appearing in the Middle Jurassic, about 175 million years ago, the tethysuchians were a group of neosuchian crocodilians – part of the same lineage that includes all living crocs, although they were probably more closely related to the highly marine thalattosuchians than to modern forms.

Their fossil remains have been found almost globally, except for in Antarctica and Australia, and they appear to have been highly aquatic animals living in both freshwater and marine environments. Most members of the group had very long and slender gharial-like snouts, indicating they were specialized for fish-eating, but some (like the enormous Sarcosuchus) developed broader or shorter snout shapes that suggest more generalized diets of whatever they could catch.

One lineage of tethysuchians known as the dyrosaurids evolved around the mid-Cretaceous (~100-90 mya) and quickly spread around most of the world. These crocs mainly inhabited coastal marine waters, with a few species also living full-time in estuaries or rivers.

They had tall vertebrae around their shoulders, giving them a slightly hump-backed appearance and anchoring large neck muscles that allowed them to quickly whip their jaws around to catch fast-moving fish. Their deep vertically-flattened tails were capable of an even more powerful swimming stroke than those of modern crocs, and their well-muscled limbs probably made them strong walkers when on land.

The dyrosaurids were some of the few marine reptiles to survive through the end-Cretaceous extinction (~66 mya) relatively unscathed, and several species are known from both sides of the K-Pg boundary. This may be because the marine dyrosaurids are thought to have seasonally migrated inland to breed in freshwater environments, with juveniles spending their early lives in rivers and only returning to the coasts as adults – and since freshwater ecosystems were much less affected by the mass extinction than marine ones, this allowed them to continue on while groups like the mosasaurs and plesiosaurs died out.

Dyrosaurus maghribensis here lived during the Late Paleocene and Early Eocene of Morocco (~56-48 mya). It was similar in size to the largest living crocs, around 6m long (19′8″), but had thinner and less extensive bony osteoderm armor.

During the Eocene the dyrosaurids began to disappear, and by the Late Eocene (~37 mya) the last known species were found only in northern Africa. It’s not entirely clear why these once-successful tethysuchians began to decline, but they may have been struggling to deal with the cooling climate trends at the time. If they managed to persist until the end of the Eocene, sudden temperature and sea level drops during the Eocene-Oligocene extinction (~33 mya) probably finished them off entirely.

Almost-Living Fossils Month #21 – More Sharks

First appearing in the Early Permian, about 290 million years ago, the synechodontiformes were an early branch of the neoselachian lineage of cartilaginous fish, slightly closer related to living sharks and rays than to the hybodontiformes featured earlier this month.

They originated in the Paleo-Tethys Ocean and survived through the devastating end-Permian “Great Dying” mass extinction (~252 mya), then went on to quickly spread around most of the world and also survive through the Triassic-Jurassic extinction (~201 mya). During the Jurassic and Cretaceous they became quite common and diverse, taking over some of the niches previously occupied by the hybodontiformes and adapting to a range of marine environments from shallow coastal waters to open ocean.

Most known synechodontiform fossil remains are just their teeth, since cartilage skeletons don’t preserve very often, but there are a few rare body fossils that show they were varied in appearance with differing arrangements of dorsal fins and spines.

Paraorthacodus jurensis here was one of the species known from the Late Jurassic of Germany (~155-150 mya). Reaching lengths of at least 1.3m (4′2″), it had only one dorsal fin far back on its body, along with large pectoral fins and a low asymmetrical tail that gave it a superficial resemblance to the modern sixgill sharks.

Its teeth were close in shape to those of sand tiger sharks, and it may have had a similar lifestyle opportunistically hunting prey just above the sea floor in the waters around the continental shelf and slope. Remains of a chimaera in the mouth and gut contents of a couple of Paraorthacodus jurensis fossils suggest that smaller cartilaginous fish were fairly common elements of its diet.

A few synechodontiformes managed to survive the end-Cretaceous extinction 66 million years ago – but while the ancestors of moderns sharks thrived in the Cenozoic, the synechodontiformes never recovered anything close to their Mesozoic levels of success and instead began to decline.

The last known synechodontiforme was a currently-unnamed member of the Paraorthacodus genus, hanging on in the waters around Antarctica in the Late Eocene (~37 mya). If they managed to survive past that time it probably wasn’t for very much longer, and it’s likely they finally disappeared during another extinction event at the Eocene-Oligocene boundary.

Almost-Living Fossils Month #20 – Some Very Spiky Turtles

The meiolaniformes were a group of terrestrial turtles that first appeared in the fossil record in the Early Cretaceous, around 125 million years ago. Although they were originally thought to be cryptodirans, more recent studies suggest they weren’t actually quite true turtles at all, instead being close evolutionary cousins to them in a much older and more “primitive” lineage that may go back as far as the Triassic.

They’re known mainly from South America and Oceania, but they may have had a more global distribution during the Cretaceous, with some fossils from the northern continents sometimes being classified as members of the group. However, only the South American meiolaniformes seem to have actually survived through the end-Cretaceous extinction.

The most distinctive meiolaniformes were the heavily armored meiolaniids, which first appeared in Patagonia during the Early Eocene (~48 mya). With large horns on their heads and thorn-like spikes along their long tails, they seem to have convergently evolved to fill the same sort of large-herbivore-tank niche as ankylosaurs and glyptodonts.

They also had fairly large nasal cavities, which might indicate a well-developed sense of smell – or may have been an adaptation for regulating the heat and moisture content of each breath, similar to the complex noses of ankylosaurs.

The South American meiolaniformes all went extinct around the end of the Eocene (~33 mya), but some meiolaniids had already dispersed across to Australia via Antarctica (before the continents had fully separated, and before Antarctica had frozen over) and they continued to survive there for most of the rest of the Cenozoic. They even went on to spread to various islands around Oceania, suggesting they were able to float and swim like modern giant tortoises.

The largest Australian meiolaniids reached sizes of around 2.5m long (8′2″), making them some of the biggest of all known terrestrial turtles. These giant forms went extinct in the Late Pleistocene, around 50,000 years ago, alongside much of the other Australian megafauna.

A few smaller varieties hung on in smaller islands to the east, with one of the latest-surviving species being Meiolania platyceps on Lord Howe island. It was only about half the size of its biggest Australian relatives – an example of insular dwarfism – and lived into the Late Holocene just 3000-2000 years ago.

Meiolania species on other islands seem to have gone extinct after the arrival of humans. But Lord Howe Island appears to have never been inhabited prior to European settlement in the late 1700s, so it’s unclear why this last of the meiolaniformes disappeared.

[Edit: A 2018 study of Meiolania platyceps’ anatomy suggests it may have been more aquatic than previously thought. It might have been something like a giant herbivorous snapping turtle or an armored reptilian hippo, bottom-walking around in coastal lagoons, with its big nasal cavity housing salt glands.]

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 true 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 it’s 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.

These last polydolopimorphs survived until at least the end of the Pliocene, around 2.5 million years ago. Their disappearance coincides with the time of the Great American Interchange – when South America became connected to Central and North America – and they may have been some of the victims of the extinction caused by the influx of placentals from the north.

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.

The herpetotheriid’s extinction seems to have happened soon after the Middle Miocene disruption, a sudden shift towards cooler temperatures that may have altered their local ecosystems too quickly for them to adapt. The peradectids hung on until at least 11 million years ago, with some of the last members known from Southeast Asia, but they may have likewise fallen victim to further episodes of global cooling towards the end of the Miocene and into the Pliocene.

Almost-Living Fossils Month #17 – Flowering Before It Was Cool

The Bennettitales were a group of seed-bearing plants found all around the world during the Mesozoic. They were an incredibly important part of ancient ecosystems, dominating the mid-level vegetation during the Jurassic and Early Cretaceous and making up over a third of the known plant species in some places. They came in two main varieties: the stocky cycad-like Cycadeoidaceae, mostly found in the northern continents; and the more slender branching tree-like Williamsoniaceae, which were more globally distributed. Most seem to have been around 2m tall (6′6″), but some may have reached much larger maximum sizes, perhaps as much as 15-25m (~49′-82′).

They date back to at least the Late Triassic (~230 mya), but they were already so diverse and numerous at that time that they probably originated much earlier – and very similar-looking leaves from the Permian and the Carboniferous hint that these plants may have actually first appeared at least 300 million years ago.

Their exact evolutionary relationships are still under dispute, with different paleobotanists classifying them as different types of seed-bearing plant. They’ve been traditionally placed close to the cycads due to their leaf shapes and growth patterns, but these similarities might be convergent since their complex flower-like reproductive structures and stomata instead suggest a possible evolutionary link to angiosperms or gnetophytes or “seed-ferns”.

The “flowers” of the bennettitaleans were round cup-like structures (sometimes resembling artichokes) which didn’t actually ever open up. While this would have made it very easy for them to self-pollinate, they may also have been some of the first plants to experiment with a mutualistic relationship with insects for pollination, possibly even partnering with early cupedid beetles in a similar manner to modern figs and their wasps.

Towards the end of the Cretaceous the rise of the true flowering plants seems to have started to send the bennettitaleans into decline, and the Cretaceous-Paleogene mass extinction at first appeared to have finished them off entirely. While there are a few Paleocene and Eocene-aged fossils that might be bennettitalean leaves, they’re not preserved in enough detail to tell for certain and they could actually be cycads.

But a few specimens from the Early Oligocene (~28 mya) of southeastern Australia and Tasmania have actually been identified as the leaves of a late-surviving species named Ptilophyllum muelleri. Since there were several other types of “relict” Mesozoic plants known to have still been present in the region around the same time (including horsetails and ginkgoes) it shouldn’t be much of a a surprise that the bennettitaleans had hung on there too – but sadly it’s not clear how much longer past this time they would have managed to survive. A drying and cooling climate towards the end of the Oligocene and into the Miocene, along with continuing competition from diversifying flowering plants, may have been enough to finally overwhelm these last few members of a once-impressive lineage.

Almost-Living Fossils Month #16 – Fancy Triangle Clams

Trigonia was a genus of bivalve mollusc that first appeared in the Middle Triassic, around 245 million years ago. Part of a much older lineage (the trigoniidans) that originated over 400 million years ago in the Late Silurian or Early Devonian, and distantly related to modern freshwater mussels, these bivalves have been found in marine deposits all around the world.

Their triangular shells had complex internal hinges, and often featured elaborate patterns of ribs and tubercules (which may have been adaptations to increase burrowing efficiency) that made them very visually distinctive. They lived mainly in shallow coastal environments, and in some places their fossils are so common that they must have been very numerous animals in their ecosystems.

Trigonia costata was a species living in Europe during the Early-to-Middle Jurassic (~174-166 mya), around the time when the trigoniidans were exploding in diversity. Usually around 5-7.5cm in length (2-3″), it was one of the longest-lasting individual species of Trigonia and one of the most common at the time.

Along with their other trigoniidan relatives, various Trigonia species continued to evolve throughout the entire rest of the Mesozoic, and while almost all of them went extinct at the end of the Cretaceous a few did manage to hang on into the Cenozoic.

The last record of an actual Trigonia comes from Argentina at the very end of the Paleocene, about 56 million years ago. After a nearly 200-million-year run, this long-lived genus finally disappeared – but although Trigonia itself was gone, that wasn’t quite the end of the trigoniidans altogether.

A single remaining lineage quietly continued on all the way into modern day, either descended from one of the Trigonia species or very closely related to the genus, living in waters off the coast of Australia and Tasmania. Known as Neotrigonia, they’re not quite as elaborately ornamented as some of their ancient relatives, but their complex shell hinges give them away as the only living trigoniidans – and their anatomy can give us some hints about what Trigonia’s soft tissue parts may have looked like, such as the presence of an unusual boot-shaped muscular foot that helps them burrow rapidly into the seafloor.

Almost-Living Fossils Month #15 – Digging Dryolestoids

First appearing in the mid-Jurassic, about 168 million years ago, a group of mammals called dryolestoids were some of the closest known relatives to the therians (the group that contains modern marsupials and placentals). They were found throughout North America, Eurasia, and North Africa up until the Early Cretaceous (~125 mya), but then mostly disappeared from the northern continents and instead migrated into South America – where they went on to flourish and became some of the most common mammals in the continent by the Late Cretaceous.

Although mostly known from only teeth and jaw fragments, the dryolestoids seem to have been a pretty diverse group of mammals during the Mesozoic, adapting to a variety of different diets and lifestyles ranging from small insectivores to relatively large herbivores.

Most of them died out in the end-Cretaceous mass extinction, except for the dog-sized herbivorous Peligrotherium which survived until 59 million years ago in the Early Paleocene… and one other known example from much much later into the Cenozoic.

Despite being absent from the fossil record for over 40 million years, the dryolestoids reappeared again in the Early Miocene of Patagonia (~21-17.5 mya) with a single late-surviving member: Necrolestes patagonensis.

The known Necrolestes fossils are surprisingly well-preserved compared to most other dryolestoids, with about a third of its skeleton represented. It was a small mole-like burrowing animal, about 10-15cm long (4-6″), with large canine teeth and an upturned snout. The cartilage in its nose was ossified into bone, strengthening it and probably supporting a pad of thick toughened skin – and also suggesting that it was a “head-lift digger”, using its snout like a shovel to dig through the soil.

While it superficially resembled the earlier mole-like dryolestoid Paurodon, it was actually much closer related to more generalist Mesozoic forms like the sabertoothed Cronopio.

After Necrolestes there’s no further evidence of dryolestoids living any closer to modern day. Much like the late-surviving gondwanatheres they lived alongside, these last dryolestoids may have specialized themselves so much that they couldn’t cope with sudden environmental changes, and the Middle Miocene extinction could have finished them off entirely.

Almost-Living Fossils Month #14 – Ancient Snakes

While the evolutionary origin of snakes is still rather poorly understood, one very early branch of their lineage – known as the madtsoiids – were a particularly long-lived group.

Originating back in the mid-Cretaceous (~100 mya), these “primitive” snakes were found mostly in the southern continents of Gondwana (known from South America, Africa, India, and Australia), but a few also spread into Europe. They were either some of the earliest true snakes or perhaps ophidians very closely related to them, and may have retained small hindlimbs that were slightly more well-developed than the vestigal ones of some modern snakes.

They ranged in length from under 1m (3′3″) to at least 7m (23′), with biggest of them rivaling some of the very largest living snakes in size.

They would have been similar to pythons, non-venomous and relying on constriction to kill their prey, although they had less flexible skulls than their modern relatives and couldn’t easily swallow animals much larger than their own heads. At least some of the Cretaceous species would have preyed on smaller dinosaurs, with one fossil even preserving a mid-sized madtsoiid in a sauropod nest alongside a hatchling.

Although the madtsoiids survived the end-Cretaceous extinction quite well and kept going throughout most of their range for the first half of the Cenozoic, most of them eventually disappeared in the Eocene-Oligocene extinction about 33 million years ago.

Aside from a single possible Late Olgiocene/Early Miocene record from South America (~29-21 mya), after that point the madtsoiids were found only in Australia, where they persisted almost into modern times.

Wonambi naracoortensis was one of the last of the Australian madtsoiids, living from the mid-Miocene (~11 mya) to at least the Late Pleistocene (~40,000 years ago). It was also one of the larger members of the group, 5-6m long (16′5″-19′8″), and seems to have been an ambush predator that lurked around waterholes to catch drinking animals.

The last madtsoiids went extinct at the same time as many of the other Australian megafauna, and it’s not clear exactly what caused them to die out. Humans had arrived in Australia about 20,000 years earlier, and hunting – either directly targeting the large snakes, or simply gradually reducing their available prey – combined with a changing climate may have been too much for them to handle.