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 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 #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.

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.

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