Paleontology and science illustration, and feathering ALL the dinosaurs
It’s been a while, but let’s catch up with some more work I’ve done for PBS Eons:
The enigmatic Paleodictyon, from “Something Has Been Making This Mark For 500 Million Years”
The archaic ungulates Loxolophus, Arctocyon, and Eoconodon, from “How a Mass Extinction Changed Our Brains”
And the nautilid Aturia, from “When Nautiloids Met Their Match”
Hexameroceras panderi was a nautiloid cephalopod that lived during the late Silurian, about 425-423 million years ago, in what is now Czechia.
Around 5cm long (2″), it had a downwards-curving egg-shaped shell that preserved the original color pattern on one fossil specimen, showing closely-packed crisscrossing vertical and horizontal bands.
Like several of its close oncocerid and discosorid relatives, its shell also developed a highly constricted opening as it reached maturity. This eventually formed into a narrow visor-like shape with several lobes that probably correlated to the life positions of the eyes and arms, with a “spout” at the bottom for the siphon.
The function of this structure is still unclear. It may have been a defensive measure against predators – but it would have also severely limited the range of motion of the arms and the size of food that could be eaten through the mouth, suggesting that Hexameroceras may have specialized in very small prey, perhaps even filter-feeding.
Another possibility is that these visored nautiloids might represent brooding females, walling themselves into their shells to protect their eggs and dying after releasing the hatchlings through the tiny remaining gap.
Continue reading “Hexameroceras”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.
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.
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.
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.
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.)
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.
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.
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.
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.
Continue reading “Cambrian Explosion Month #26: Phylum Mollusca – Tentacle Time”Nautiloids are represented today by just two living genera (Nautilus and Allonautilus), but they have a lengthy evolutionary history going back almost 500 million years.
The peak of their diversity was during the first half of the Paleozoic, with many different shapes of shells from coiled to straight, then they began to decline when their relatives the ammonites and coleoids appeared and began to compete for similar ecological niches. Although a few groups of nautiloids survived through the end-Permian mass extinction, most of them had disappeared by the end of the Triassic, leaving just one major remaining lineage known as the Nautilina (or Nautilaceae).
During the mid-to-late Jurassic (~165 mya) two new groups split away from the ancestors of the modern nautiluses – the cymatoceratids and the hercoglossids.
Cymatoceratids such as Cymatoceras sakalavum here had shells with a ribbed texture. Living during the Early Cretaceous, about 112-109 million years ago, this particular species is known from Japan and Madagascar and could reach a shell diameter of over 15cm (6″).
Hercoglossids, meanwhile, were much more smooth in appearance, but both groups also had more complex undulating sutures between their internal chambers than modern nautiluses do.
These nautiluses made it through the end-Cretaceous mass extinction and had a brief period of renewed success, filling the ecological roles left vacant by the extinct ammonites. But by the end of the Oligocene (~23 mya) both the cymatoceratids and hercoglossids vanished, possibly unable to deal with cooling oceans and the evolution of new predators.
Some of the hercoglossids’ Cenozoic descendants, the aturiids, managed to last a little longer into the Early Pliocene (~5 mya) before another period of cooling seems to have finished them off. Past that point, all that was left of the once-massive nautiloid lineage were their cousins the nautilids, who gave rise to today’s few living representatives.
(It’s also worth noting that the classification of the cymatoceratids seems to be in flux right now. Some paleontologists currently don’t consider Cymatoceras itself to actually be part of the group, instead being a nautilid much closer related to modern nautiluses. If this is the case then the cymatoceratids may not have actually survived past the Late Cretaceous – but the Cymatoceras genus alone still counts as an “almost-living” fossil since its various species ranged from the Late Jurassic to the Late Oligocene.)
Ammonites (or “ammonoids” in technical terms) are one of the most recognizable types of fossil, found in such high abundance that they’re frequently used to precisely date rock layers. They’re absolutely everywhere in fossil collections, and are even made into jewelry.
So we must already know everything we possibly could about them, right?
Except… we really don’t know what their soft parts looked like.
The fossil record for ammonite soft tissue is surprisingly empty for a group that existed for over 300 million years. A possible ink sac and a few organs have been found, but nothing else.
Based on their other cephalopod relatives, they probably had at least ten arms (the two longer tentacles shown on this Collignoniceras are a little speculative), along with a siphon for propulsion – but until we find that elusive exceptional preservation we just don’t know for sure.
[Edit: As of 2022, a few more traces of soft tissue have been found!]
