mollusc – Nix Illustration

Glossoceras

Although the only nautiloids living today have characteristic tightly coiled shells, earlier in their evolutionary history these cephalopods were much more diverse.

And Glossoceras gracile here is an example of one of the more unusual groups of nautloids: the ascocerids.

Living during the Late Silurian, about 422 million years ago, in wheat is now Gotland, Sweden, Glossoceras was only around 5cm long as an adult (~2″). Like other ascocerids it started out its life looking like a fairly standard early nautiloid, with a long straight shell that curved slightly upwards, but as it approached maturity things got weird – the front part of the shell grew out into a much more bulbous flask-like shape, and the old juvenile section broke off entirely.

The gas-filled buoyancy chambers of its adult shell were positioned directly above its body chamber rather than behind like in other nautiloids, giving it very good stability in the water. The shell walls were also very thin and lightweight, which would have made it a much more maneuverable swimmer.

Strange Symmetries #16: Go Home Heteromorphs You’re Drunk

Most ammonites had spiral-coiling shells, but during the Cretaceous a group known as the heteromorphs evolved a much weirder range of forms. Some were straightened, some were hooked, some had helical snail-like shapes, and some even ended up bearing a strange resemblance to paperclips.

But one of the most bizarre of all was the genus Nipponites, whose ribbed shell looked like a bundle of tangled asymmetrical coils.

Nipponites bacchus lived in what is now Hokkaido, Japan, during the late Cretaceous about 90 million years ago. Around 10cm long (~4″), its shell was less tightly coiled up than its better-known relative Nipponites mirabilis, but these looser whorls were formed in the same way via a series of U-bends in different directions during its growth.

Despite their irregular and ungainly appearance, the unique shape of these ammonites seems to have actually been very hydrodynamically stable. They weren’t fast-moving, but they didn’t need to be, probably spending most of their time floating suspended in the water column catching small planktonic prey from around themselves.

Strange Symmetries #09: Not Toeday, Satan

The shells of bivalve molluscs are formed on the left and rights sides of their bodies, and so are usually roughly bilaterally symmetric.

But some bivalves break that arrangement, developing asymmetrical valves that can be massively different in size and shape.

Gryphaea arcuata was an oyster that lived during the Early Jurassic, about 200-174 million years ago, in the warm shallow seas that covered what is now Europe and eastern Greenland. Around 6cm long (~2.4″), its left valve was thick and strongly convex and curled, while the right valve was relatively thin and slightly concave forming a “lid”.

The gnarled curled claw-like shape of Gryphaea fossils led to them being colloquially known as “devil’s toenails” in some of the regions where they’re commonly found, with folk beliefs that they had the power to prevent joint pain.

Their shape was actually an adaptation to living on very soft seafloor sediments. The larger curled valve acted sort of like a boat on the soupy mud, supporting the Gryphaea‘s weight and preventing it from sinking.

Strange Symmetries #07: Gastropods Do The Twist

Gastropods – snails and slugs – are a group of molluscs that originated sometime in the Cambrian Period, with the earliest definite stem-gastropods known from around 510 million years ago and the first true gastropods turning up in the early Ordovician.

The spiral-coiled shells of snails are their most familiar feature, giving them obvious external asymmetry, but gastropods are also defined by a specific type of internal asymmetry known as torsion.

Torsion is an anatomical process that occurs during larval development, and involves rotating their internal organs, mantle, and shell a full 180° relative to their head and muscular foot. This twists their gut into a U-shape, knots up their nervous system, and brings their respiratory organs and anus up close to their head.

And we still don’t really know why they do it.

One idea (the “rotation hypothesis”) is that it originated as a defensive function after early gastropods began developing their spiral shells. The shell opening may have originally been positioned at early gastropods’ rears, meaning they retracted their bodies back-end-first leaving their heads and sensory structures still vulnerable – but twisting the shell around would allow them to pull their front end in faster instead.

A competing idea (the “asymmetry hypothesis“) instead proposes that the shape of the coiled shell restricted the gills of early gastropods, which may have originally been positioned in mantle cavities on each side of their bodies. In response to this they developed a single larger gill cavity on just one side of their body, and then gradually expanded and rotated this asymmetric feature around to the front for better aeration.

In either case this resulted in some of the rest of their anatomy “coming along for the ride”. And regardless of whatever the original evolutionary advantage of torsion actually was, it made gastropods incredibly successful – they’re a massively diverse group, second only to the insects in terms of sheer number of species, and today they’re found all over the world in almost every habitat from deep sea trenches to high mountain elevations.

A colored line drawing of Spinyplatyceras, an extinct marine snail. It has a low coiling shell covered in very long thin pointed spines, and there are two short tentacles on its head. It's depicted with orange and black striped coloration on its shell, and a purplish body. — *Spinyplatyceras arkonense*

Spinyplatyceras arkonense lived in what is now Ontario during the mid-Devonian, about 391-385 million years ago. Around 5cm long (2″), it was part of a group of Paleozoic marine snails known as platyceratids, which were probably related to either modern limpets or neritomorphs.

Platyceratids seem to have had a unique parasitic relationship with crinoids, attaching themselves to the top of the host’s body and using their radula to drill into them, either robbing food directly from the crinoid’s gut or feeding on its other internal organs.

The long spines on Spinyplatyceras‘ shell probably helped to deter predators. In an interesting case of coevolution the crinoid hosts of some platyceratids developed their own defensive spines, too – and it seems this wasn’t to prevent the snails from infesting them, but to also discourage the snails’ predators. These crinoids may have been frequently indirectly injured during snail-eating predators’ attacks, and it might have actually “cost” them less to keep enduring an infestation than to deal with the collateral damage of the snails being removed.

Typhloesus

Typhloesus wellsi has been a mystery for a long time.

First discovered in the early 1970s, in the mid-Carboniferous Bear Gulch Limestone deposits (~324 million years ago) of Montana, USA, it was initially mistaken for the long-sought-after “conodont animal” due to the presence of numerous conodont teeth inside its body. But just a few years later well-preserved eel-like conodont animals were found elsewhere, and it became apparent that the conodont teeth inside Typhloesus had actually just been part of its last meal.

But if it wasn’t a conodont… then what was it?

Up to about 10cm long (4″), Typhloesus had a streamlined body with a vertical tail fin and paired “keels” along its sides. It had a mouth and a gut cavity, but no apparent anus, and it also didn’t seem to have any eyes or other sensory structures. And in the middle of its body there was something very weird – a pair of “ferrodiscus” organs, disc-shaped structures which contained high concentrations of iron but whose function was completely unknown.

This anatomy just didn’t match any other known animals, so much so that it gained the nickname of “alien goldfish”.

For the next few decades it remained a bizarre enigma, at best tentatively considered to represent an unknown lineage of some sort of metazoan that left almost no other fossil record due to being entirely soft-bodied.

But now, 50 years after its initial discovery, we might just finally have a clue about Typhloesus’ true identity.

Recently something new was discovered in some Typhloesus specimens – a radula-like feeding structure that was probably part of an eversible proboscis. This would mean that Typhloesus was a mollusc, possibly a gastropod that convergently evolved a swimming predatory lifestyle similar to modern pterotracheoids.

It’s not a definite identification yet, and even if it was a mollusc it was an incredibly strange one, with features like the ferrodiscus still lacking any explanation. But this discovery at least shows that there are still new details waiting to be found in the “alien goldfish” fossils, and gives us a start towards bringing its classification back down to earth.

It Came From The Wastebasket #09: Hammering Away At Hamites

Ancyloceratines were a lineage of distinctive-looking ammonites, commonly known as “heteromorphs“, which had unusual uncoiled shells that ranged in shape from near-straight to hooked to helical to paperclip-like to “balls of string“.

Heteromorphs’ strange shells would have created a lot of drag in the water, and they may not have been especially agile swimmers, but they were very hydrodynamically stable and easily maintained neutral buoyancy. Their paleobiology has only just started to be properly understood in recent years, and now most species of these ammonites are thought to have floated suspended in the photic zone and twilight zone of the open ocean, catching small zooplankton from the water around themselves.

What these ammonites were doing obviously worked very well for them, because they were incredibly diverse and successful during the Cretaceous period. They were also the only type of ammonite to persist for a short time after the end-Cretaceous mass extinction, existing as a “dead clade walking” for another half a million years or so before finally disappearing entirely.

An illustration of Hamites attenuatus, an unusually-shaped extinct ammonite. It has a mostly-uncoiled shell shaped like a long sideways U, almost paperclip-like. At the bottom end of the shell it has a squid-like body, with large eyes and ten short webbed arms that are lined with speculative fringes for filter-feeding. It has a pinkish color scheme with a faint iridescent sheen. — *Hamites attenuatus*

The hamitids were a group of heteromorphs from the mid-Cretaceous (~110-90 million years ago), with their namesake genus Hamites traditionally being used as a wastebasket taxon for anything that that didn’t neatly fit into any other group of similar heteromorph ammonites.

By the late 1990s Hamites had become a mess of multiple different diverse lineages, with over 20 species all lumped together – and this was a problem because the hamitids were the ancestors of several other heteromorph ammonite lineages, and having the taxon in such disarray made studying the evolutionary origins of all those other groups very difficult.

So in the early 2000s attempts were made to clean this all up, figuring out the relationships between the different Hamites species and dividing the genus into multiple new genera.

There hasn’t been much more detailed research on the relationships of hamitids since then – and other groups of heteromorphs are still in need of revision – but it’s a start at clearing the wastebasket, at least.

It Came From The Wastebasket #04: Breaking Up Bellerophon

Bellerophonts were small snail-like marine molluscs that were either early gastropods or very close relatives of them. They had symmetrically-coiled shells superficially shaped like those of nautiluses, with about half of the shell covered by their mantle similarly to some modern sea snails, and some fossil shells also preserve hints of banded color patterns.

First appearing in the late Cambrian (~500 million years ago), these molluscs existed all the way until the early Triassic, surviving the Great Dying mass extinction (~252 million years ago) only to go extinct just a short time later (~249 million years ago) – a phenomenon known as “dead clade walking”, when a group just barely scrapes through a mass extinction event but doesn’t manage to actually recover afterwards.

The whole group is something of a wastebasket of similar-looking shells, and might actually be more of an “evolutionary grade” made up of various early gastropods and gastropod-relatives than a single defined lineage.

But there’s also another wastebasket nestled inside this wastebasket: the namesake of them all, the genus Bellerophon.

An illustration of Bellerophon, an extinct sea-snail-like mollusc. It has a banded shell that coils vertically like a nautilus, with a ridge along the midline. It has a wide flat foot, its mantle covers about halfway up the sides of its shell, and it has a pair of snail-like head tentacles and a siphon. — *Bellerophon tenuifascia*

Originally named in 1808, this genus has had a huge number of species assigned to it over the last couple of centuries. This gives a false impression that Bellerophon-like molluscs didn’t change for hundreds of millions of years, and it makes figuring out their actual long-term patterns of evolution and extinction much more difficult.

In the last few decades some mollusc paleontologists have been gradually chipping away at Bellerophon, and multiple new genera have been broken off from it. But even today it remains a very bloated mess – there are still well over a hundred named species spanning about 230 million years of geologic time.

Studies do indicate the whole genus is highly polyphyletic, made up of a tangle of multiple different lineages that all really need to be revised and renamed – but there’s a lot of work still needing to be done to clean up this particular wastebasket.

Retro vs Modern #17: Ammonites

Ammonites (or ammonoids) are highly distinctive and instantly recognizable fossils that have been found all around the world for thousands of years, and have been associated with a wide range of folkloric and mythologic interpretations – including snakestones, buffalo stones, shaligrams, and the horns of Ammon, with the latter eventually inspiring the scientific name for this group of ancient molluscs.

(Unlike the other entries in this series the reconstructions shown here are somewhat generalized ammonites. They’re not intended to depict a specific species, but the shell shape is mostly based on Asteroceras obtusum.)

1830s

It was only in the 1700s that ammonites began to be recognized as the remains of cephalopod shells, but the lack of soft part impressions made the rest of their anatomy a mystery. The very first known life reconstruction was part of the Duria Antiquior scene painted in 1830, but to modern eyes it probably isn’t immediately obvious as even being an ammonite, depicted as a strange little boat-like thing to the right of the battling ichthyosaur and plesiosaur.

The argonaut octopus, or “paper nautilus”, was considered to be the closest living model for ammonites at the time due to superficial similarities in its “shell” shape, but these modern animals were also rather poorly understood. They were commonly inaccurately illustrated as floating around on the ocean surface using the expanded surfaces on two of their tentacles as “sails” – and so ammonites were initially reconstructed in the same way.

1860s

While increasing scientific knowledge of the chambered nautilus led to it being proposed as a better model for ammonites in the mid-1830s, the argonaut-style depictions continued for several decades.

Interestingly the earliest known non-argonaut reconstruction of an ammonite, in the first edition of La Terre Avant Le Déluge in 1863, actually showed a very squid-like animal inside an ammonite shell, with eight arms and two longer tentacles. But this was quickly “corrected” in later editions to a much more nautilus-like version with numerous cirri-like tentacles and a large hood.

The nautilus model for ammonites eventually became the standard by the end of the 19^th century, although they continued to be reconstructed as surface-floaters. Bottom-dwelling ammonite interpretations were also popular for a while in the early 20^th century, being shown as creeping animals with nautilus-like anatomy and numerous octopus-like tentacles, before open water active swimmers eventually became the standard representation.

2020s

During the 20^th century opinions on the closest living relatives of ammonites began to shift away from nautiluses and towards the coleoids (squid, cuttlefish, and octopuses). The consensus by the 1990s was that both ammonites and coleoids had a common ancestry within the bactridids, and ammonites were considered to have likely had ten arms (at least ancestrally) and were probably much more squid-like after all.

Little was still actually known about these cephalopods’ soft parts, but some internal anatomy had at least been figured out by the early 21^st century. Enigmatic fossils known as aptychi had been found preserved in position within ammonite shell cavities, and were initially thought to be an operculum closing off the shell against predators – but are currently considered to instead be part of the jaw apparatus along with a radula.

Tentative ink sac traces were also found in some specimens (although these are now disputed), and what were thought to be poorly-preserved digestive organs, but the actual external life appearance of ammonites was still basically unknown. By the mid-2010s the best guess reconstructions were based on muscle attachment sites that suggested the presence of a large squid-like siphon.

Possible evidence of banded color patterns were also sometimes found preserved on shells, while others showed iridescent patterns that might have been visible on the surface in life.

In the late 2010s the continued scarcity of ammonite soft tissue was potentially explained as being the same reason true squid fossils are so incredibly rare – their biochemistry may have simply been incompatible with the vast majority of preservation conditions.

But then something amazing happened.

In early 2021 a “naked” ammonite missing its shell was described, preserving most of the body in exceptional detail – although frustratingly the arms were missing, giving no clarification to their possible number or arrangement. But then just a few months later another study focusing on mysterious hook-like structures in some ammonite fossils concluded that they came from the clubbed tips of a pair of long squid-like tentacles – the first direct evidence of any ammonite appendages!

A third soft-tissue study at the end of the year added in some further confirmation that ammonites were much more coleoid-like than nautilus-like, with more evidence of a squid-style siphon, along with evidece of powerful muscles that retracted the ammonite’s body deep inside its shell cavity for protection.

Since ammonites existed for over 340 million years in a wide range of habitats and ecological roles, and came in a massive variety of shapes and sizes, it’s extremely likely that their soft anatomy was just as diverse as their shells – so there’s no single “one reconstruction fits all” for their life appearances. Still, at least we now have something less speculative to work with for restorations, even if it’s a bit generalized and composite, and now that we’re finally starting to find that elusive soft tissue there’s the potential for us to discover so much more about these iconic fossil animals.

Cambrian Explosion Month #26: Phylum Mollusca – Tentacle Time

Cephalopods‘ highly distinctive body plan and incredible intelligence make them some of the most charismatic marine animals. Today they’re mainly represented by the soft-bodied coleoids (octopuses, squid, and cuttlefish), with the modern giant squid and colossal squid being the largest living invertebrates. In comparison the shelled nautiluses seem like weird oddballs, but they’re actually far more typical examples of the group than their squishier cousins – as part of the conchiferan lineage the ancestors of all modern cephalopods were also shell-bearing molluscs, and for much of their evolutionary history shelled forms like ammonites and orthoceridans were extremely abundant.

The exact evolutionary relationships of cephalopods within the conchiferan family tree aren’t clear, but their closest relatives might be modern monoplacophorans and they probably descended from limpet-like “monoplacophoran-grade” ancestors in the early Cambrian. The current oldest potential cephalopod fossils come from about 522 million years ago, but the first definite cephalopods in the fossil record come from much later in the period.

Cambrian Explosion Month #25: Phylum Mollusca – Shelling Out

The exact evolutionary relationships of the main groups of modern molluscs are rather debated, with several different proposed family trees. But one of the main possibilities is that there are two major lineages: the aculiferans and the conchiferans.

Modern conchiferans include slugs and snails, cephalopods, bivalves, tusk shells, and monoplacophorans – all groups that ancestrally have either a single-part shell or a two-part bivalved shell, with some lineages later becoming secondarily shell-less.

The ancestral conchiferans are thought to have been monoplacophoran-like molluscs, limpet-like with a cap-shaped shell, and likely diverged from a common ancestor with the aculiferans around the end of the Ediacaran. (But modern monoplacophorans probably aren’t “living fossil” descendants of early Cambrian conchiferans, and may instead be close relatives of cephalopods that have convergently become similar in appearance to their ancestors.)

Some of the earliest conchiferans were the helcionelloids, a lineage of superficially snail-like molluscs with coiled cone-shaped mineralized shells. They appeared in the fossil record at the start of the Cambrian (~540-530 million years ago) and lasted until the early Ordovician (~480 million years ago), and have been found all around the world as components of the “small shelly fauna“.

And while they’re usually tiny, only a couple of millimeters in size, they may actually represent juveniles or larvae – there’s evidence that at least some species grew up into much larger 2cm (0.8″) limpet-like adult forms.