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.

Auroralumina

Cnidarians – a group of animals that includes modern corals, sea anemones, sea pens, jellyfish, hydra, and a couple of parasitic forms – are one of the most ancient animal lineages, originating at least 580 million years ago in the Ediacaran period.

Actual identifiable fossils of cnidarians that old are incredibly rare, however, and until now there was only one example – the small polyp-like Haootia from Canada.

But a second definite Ediacaran cnidarian has now been described: Auroralumina attenboroughii.

It was discovered in Charnwood Forest, England, in the very same site where the first recognized Precambrian fossils were found in the 1950s. About 20cm tall (~8″) it dates to around 560 million years ago and was made up of a pair of forking stiff-walled tubes which expanded into wide four-sided goblet-like shapes full of stubby tentacles. These densely-tentacled crowns would have been used to capture tiny planktonic organisms from the water around it, making it the current earliest known example of a predatory animal.

The one known fossil specimen has an incomplete base, so it’s uncertain if this was actually the full life appearance of Auroralumina or if it was even larger with more branches and goblets. And although it was preserved in deep-water sediments, it appears to have originated from much shallower waters, being swept down into the depths during a volcanic eruption.

While it superficially resembled a sea anemone, details of its anatomy suggest it was actually much closer related to medusozoans, having similar traits to the immobile polyp stage of the jellyfish life cycle. Its four-way symmetry and boxy shape may also link it to the enigmatic conulariids.

It’s not clear if it was able to bud off swimming medusa stages like its modern relatives – that might be an evolutionary innovation that came along later – but it at least shows that a basic medusozoan body plan was already in place around 20 million years earlier than previously thought.

Stanleycaris

Radiodonts were early arthropods with specialized frontal appendages, disc-like mouths, complex compound eyes, and swimming flaps along the sides of their bodies. Once considered to be bizarre “weird wonders” of the Cambrian Explosion that represented a failed evolutionary experiment, we now know that they were actually a highly diverse and successful lineage that lasted for at least 120 million years.

While some radiodonts were the largest animals of their time periods, Stanleycaris hirpex here was one of the smallest known members of the group – although at around 10cm long (~4″) it was still respectably big compared to most other Cambrian animals.

Discovered in the Canadian Burgess Shale deposits (~508 million years ago), it was originally known only from isolated frontal appendages and mouthparts, and had been assumed to be a fairly typical member of the hurdiid family. But the recent discovery of over 200 new fossils, including some exceptionally well-preserved full body specimens, has catapulted it directly from being poorly-known into now being one of the most completely known of all radiodonts.

And it had a very big surprise for us, right in the middle of its face.

It turns out that Stanleycaris had a huge third eye, unlike anything ever seen in a radiodont before. A large unpaired eye was also part of the five-eyed arrangement in opabiniids and Kylinxia, and finding a similar example in radiodonts too raises the possibility that this sort of well-developed “median eye” may have been more widespread in early arthropods than previously thought.

Along with the third eye, some of the Stanleycaris specimens preserve fine internal details of its nervous system and show that its brain was made up of two segments instead of the three seen in modern arthropods. It also had gills positioned on its underside, unlike most other radiodonts which had them on their backs.

Rhenopyrgus

Despite looking more like some sort of scaly tubeworm, Rhenopyrgus viviani here was actually an echinoderm, distantly related to modern starfish, brittle stars, sea urchins, crinoids, and sea cucumbers.

It was part of an extinct Paleozoic echinoderm lineage known as edrioasteroids, which lived attached to the seabed or on hard surfaces like the shells of other marine animals, using the tube feet on their five arms to catch food particles from the water around them.

Living during the Silurian, about 435 million years ago, in what is now Quebec, Canada, it stood around 3-4cm tall (1.2-1.6″), firmly anchored into the seafloor sediment by a bulbous sac-shaped base. Its long stalked body was somewhat flexible, and it was able to partially contract the top feeding region down under a “collar” of large scale-like armor plates.

Antarcticarcinus

Euthycarcinoids were a group of arthropods that lived between the mid-Cambrian and the mid-Triassic – but despite existing for over 250 million years their fossil record is incredibly sparse, and it’s only within the last decade that they’ve been recognized as being close relatives of modern centipedes and millipedes.

The earliest members of this group were marine, living in shallow tidal waters, but they quickly specialized into brackish and freshwater habitats and were even some of the very first animals to walk on land. Fossil trackways show they were amphibious, venturing out onto mudflats to feed on microbial mats, avoid aquatic predators, and possibly lay their eggs in a similar manner to modern horseshoe crabs.

Most euthycarcinoid species are known from tropical and subtropical climates, but Antarcticarcinus pagoda here hints that these arthropods were much more widespread and diverse than previously thought. Discovered in fossil deposits in the Central Transantarctic Mountains of Antarctica, it lived in freshwater lakes during the Early Permian (~299-293 million years ago), at a time when the region was in similar polar latitudes to today with a cold icy subarctic climate.

About 8.5cm long (3.3″), it would have had a similar three-part body plan to other euthycarcinoids – with a head, a limb-bearing thorax, and a limbless abdomen ending in a tail spine – but its most distinctive feature was a pair of large wing-shaped projections on the sides of its carapace. These may have helped to stabilize its body when resting on soft muddy surfaces, spreading out its weight, or they might even have functioned as a hydrofoil generating lift while swimming.

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 19th century, although they continued to be reconstructed as surface-floaters. Bottom-dwelling ammonite interpretations were also popular for a while in the early 20th 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 20th 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 21st 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.

Utaurora

Ever since the bizarre anatomy of Opabinia was first recognized in the 1970s, it’s been a persistently unique “weird wonder” of the Cambrian period. Over the decades we’ve figured out that it was an early type of arthropod in an evolutionary position between lobopodians and radiodonts, but this whole time it’s still been sitting there alone as the only known representative of a weird stem-lineage with no other known close relatives.

…Until now!

A fossil from the Wheeler Shale in Utah, USA (~507 million years ago) that was originally thought to be a tiny radiodont has been re-studied, and now we finally have another member of the opabiniid family: Utaurora comosa.

Only about 3cm long (1.2″), Utaurora had 15 pairs of swimming flaps along the sides of its body, and a tail region with a 7-part fan and a pair of serrated spines. Hair-like gill blades covered both its back and the bases of its swimming flaps, and although its head region was poorly preserved it probably had an arrangement of 5 eyes and a long flexible claw-tipped proboscis similar to that of Opabinia.

Its discovery extends both the geographical and temporal known range of opabiniids, and suggests that their continued scarcity in other Cambrian fossil sites compared to other soft-bodied arthropods may simply be because they were just incredibly rare animals in those habitats at the time.

Austrolimulus

Horseshoe crabs are famous examples of “living fossils“, having changed their external appearance very little over hundreds of millions of years. But some fossil species were much more varied in shape than their morphologically conservative modern relatives, such as Austrolimulus fletcheri here.

Living in freshwater environments in what is now New South Wales, Australia, during the Middle Triassic (~247-242 million years ago), Austrolimulus had incredibly long spines on each side of its head, reaching a span of around 18cm (7″) – wider than its total body length!

The function of these spines is unclear, but they may have acted like a hydrofoil in fast-moving currents, or they may have served a defensive purpose by making Austrolimulus‘ carapace too wide and unwieldy for some predators to deal with.

Eons Roundup 10

Time for some more PBS Eons commission work!

The radiodonts Lyrarapax and Tamisiocaris, from “How Plankton Created A Bizarre Giant of the Seas”
https://www.youtube.com/watch?v=G0oKBPZODhM


The rhynchocephalians Sphenotitan, Clevosaurus, and Kawasphenodon, from “When Lizards Took Over the World”
https://www.youtube.com/watch?v=peeX3PKOE_w

April Fools 2021: Time for T. rex

Did you know that in all the time I’ve been posting paleoart here, I’ve never actually done a proper full illustration of the most popular name in all of paleontology?

I think it’s finally time to fix this glaring oversight.

It’s time for T. rex.

Continue reading “April Fools 2021: Time for T. rex”