ecdysozoa – Nix Illustration

Retro vs Modern #09: Hallucigenia sparsa

If just one single species had to represent how our reconstructions of prehistoric animals can drastically change, it would have to be Hallucigenia sparsa.

1970s

First discovered in the 1910s in the Canadian Burgess Shale fossil deposits, specimens of Hallucigenia were initially categorized as being a species of the early polychaete worm Canadia. It wasn’t until the 1970s that they were recognized as being something else entirely, and the first reconstruction of this tiny animal was bizarre.

It was depicted as a long-bodied creature with a single row of tentacles along its back, and several pairs of long sharp spines that were interpreted as being stilt-like “legs” used to walk. The tentacles were thought to catch food from the water and pass it forwards to the bulbous “head” – and at one point it was even proposed that all the tentacles had their own additional “mouths” at their tips!

It’s easy to look back on this version now and laugh at how ridiculous and obviously wrong it was, but it’s important to remember the historical context here. This was coming from a point when the incredible animal diversity of the Cambrian Explosion was only just starting to be understood, revealing a range of poorly-understood bizarre and alien-looking forms like Opabinia – “weird wonders” that were considered to be representatives of previously unknown ancient branches of life.

At the time, Hallucigenia‘s utter weirdness and impractical body plan seemed to almost make sense as a unique evolutionary “failed experiment” that had left no living relatives.

1990s

Discoveries of legged-and-armored lobopodian “worms” in the Chinese Chengjiang fossil deposits during the 1980s prompted a re-interpretation of Hallucigenia in the early 1990s. Speculatively reconstructing it as a lobopodian with the spines on its back and with the tentacles as a set of paired clawed legs started to make it seem a lot less alien and a lot more like a real velvet-worm-like animal – and just a year later the “missing” other half of the leg pairs was confirmed to be present in some of the fossil specimens.

But it was still unclear which end was actually the head, and whether the large blob-like structure was a real part of Hallucigenia‘s anatomy or just an artifact of the fossilization process.

2020s

New research in the mid-2010s finally settled the head problem and clarified a lot of Hallugicenia‘s anatomy, discovering that the slender elongated end had a pair of simple eyes and a mouth with a throat ringed with tiny teeth.

We now know Hallucigenia sparsa lived all around the world during the mid-Cambrian, about 518-508 million years ago, with body fossils known from Canada and China and isolated spines found in numerous other similarly-aged locations. Instead of an evolutionary dead-end “weird wonder” it was actually an early member of the vast arthropod lineage, just one of a highly diverse collection of successful Cambrian lobopodians, and its closest living relatives are probably velvet worms and tardigrades.

It grew up to about 5cm long (2″) and had seven pairs of long sharp defensive spines along its back, covered with a microscopic surface texture of tiny triangular “scales”. It had seven pairs of clawed walking legs, with most of its feet tipped with two claws each but the final two pairs having just one, and its body ended right at the final pair of limbs – the “blob” structure in some fossils was actually just an artifact the whole time, formed by Halligenia‘s innards being forcefully squeezed out during its burial in the seafloor sediment.

Its neck region bore three pairs of long delicate tendril-like limbs, which may have been covered in feathery hair-like structures for filter-feeding similar to some other lobopodians. A small pair of velvet-worm-like antennae may also have been present on its head, and could have been a sexually dimorphic feature.

Cambrian Explosion #61: Crustacea – Little Wigglers

We’re finally at the end of this series, and to finish off let’s look at one of the few types of Cambrian true crustaceans that are known only from fully mature adults: the skaracarids.

These tiny soft-bodied meiofaunal animals are known from late Cambrian areas of “Orsten-type preservation” in Sweden and South China, with a possible additional fragmentary occurrence in Poland – suggesting that they had a global distribution.

Cambrian Explosion #60: Crustacea – Larvae Larvae Everywhere

One of the characteristic features of the crustacean lineage are their larval forms, passing through various tiny larval stages. They often look nothing like their eventual adult forms and historically weren’t even recognized as being the same species, with their complex lifecycles not being properly recognized until the late 1800s.

A lot of Cambrian crustaceans are only known from their larvae, preserved in exquisite microscopic detail in sites of “Orsten-type preservation”. Only disarticulated fragments of larger-bodied forms have been found in a few places, and it isn’t until much later in the Paleozoic that fossil crustaceans actually seem to become abundant in marine ecosystems.

It’s not clear why there’s such a bias in their early fossil record compared to most other arthropods, but possibly they were just very very rare animals early on. Adult forms may have mostly lived in places where they just didn’t fossilize, while their tiny larvae sometimes dispersed into different environments with a better chance of preservation.

Cambrian Explosion #59: Stem-Crustacea – Actual Ancient Aliens & Bivalved Buddies

The majority of known fossils of Cambrian crustaceans are in the form of minuscule microfossils with “Orsten-type preservation” – formed in oxygen-poor seafloor mud and exceptionally well-preserved in three-dimensional detail. They can only be discovered and studied after dissolving away the rock around them with acid and picking through the residue under a microscope, then they’re scanned with an electron microscope to see their fine details.

And it turns out some of these tiny early crustaceans looked really weird.

Cambrian Explosion #58: Hymenocarina

The pancrustaceans are a grouping of mandibulates that contains all of the crustaceans and hexapods (insects and their closest relatives) along with their various stem-relatives.

They’re critical components of most ecosystems on the planet, and are major parts of the nutrient cycle. In aquatic environments the crustaceans dominate, with modern copepods and krill being some of the most abundant living animals and making up enormous amounts of biomass providing vital food sources for larger animals. On the land springtails and ants are especially numerous, and the air is full of flying insects, the only invertebrates to ever develop powered flight. Some groups of insects have also co-evolved complex mutualistic partnerships with flowering plants and fungi.

Hexapods and insects don’t appear in the fossil record until the early Devonian, but they’re estimated to have first diverged from the crustaceans* in the early Silurian (~440 million years ago), around the same time that vascular plants were colonizing the land.

(* Hexapods are crustaceans in the same sort of way that birds are dinosaurs. They originated from within one of the major crustacean lineages with their closest living relatives possibly being the enigmatic remipedes.)

But crustaceans and their pancrustacean ancestors go back much further into the Cambrian, and we’ll be finishing off this month and this series with some of those early representatives.

Cambrian Explosion #57: Tuzoiida

What were tuzoiids?

We don’t know.*

Despite hundreds of specimens having been found, and around 20 different species being described, these arthropods are an ongoing puzzle.

They’re known from between about 518 and 505 million years ago, in deposits associated with tropical and subtropical regions all around the world. They had large spiny bivalved carapaces up to 18cm long (7″), shaped like an upside-down domed taco shell, with a distinctive reticulated net-like surface ornamentation – but the rest of their ecology and anatomy is very unclear.

Most fossils are just empty carapaces, which appear to have been made of unmineralized chitin. Rare examples of soft-part preservation show they had a pair of stalked eyes sticking out the front, and a pair of short simple antennae, but impressions of the rest of their bodies are fragmentary and indistinct enough to not be particularly helpful.

Cambrian Explosion #56: Euthycarcinoidea

The euthycarcinoids were a group of euarthropods known from the mid-Cambrian to the mid-Triassic (~500-254 million years ago), surviving through multiple mass extinctions including the devastating “Great Dying” at the end of the Permian that finished off the trilobites. But despite an evolutionary history spanning around 250 million years they have a very sparse fossil record, extremely rare and known from less than 20 species across their entire time range.

For a long time their affinities were uncertain, and they’ve been variously suggested to have been crustaceans, trilobites, or chelicerates, or even to have been a lineage of earlier stem-euarthropods. But since the early 2010s better understanding of their anatomy has placed them in the mandibulates, probably as the closest relatives of the myriapods and helping to close the gap between the aquatic ancestors of that group and their earliest known terrestrial forms.

Cambrian Explosion #55: Fuxianhuiida

In the final stretch of this month we finally come to the last of the major groupings of euarthropods: the mandibulates, which include the modern myriapods (centipedes and millipedes) and pancrustaceans (crustaceans and insects), along with several extinct groups.

Characterized by possessing mandible mouthparts, mandibulates are by far the largest lineage of arthropods and the most successful group of animals on Earth. Over a million living species are known (most of of which are insects, particularly beetles) and an estimated six-to-ten times more than that are probably still undiscovered.

Mandibulates probably diverged from their chelicerate cousins around the start of the Cambrian 540 million years ago. If the trilobites and their artiopodan relatives were early or stem-mandibulates then the earliest known fossils of the group are about 521 million years old, otherwise the first records come from a few million years later in the Chinese Chengjiang fossil deposits (~518 million years ago).

Cambrian Explosion #54: Trilobita – Transform and Roll Up

Most trilobites were able to roll themselves up into a protective ball – a behavior known as enrollment or volvation – exposing just their heavily armored backs to attackers. They’re often found fossilized curled up like this, and rare preservation of soft tissues shows that they had a complex system of muscles to help them quickly achieve this pose while simultaneously tucking their antennae and all their limbs safely inside their enrolled shells.

Some species also developed sharp defensive spines and spikes that jutted out when they enrolled, making themselves even more daunting to potential predators in one of the earliest known examples of an evolutionary “arms race”.

Cambrian Explosion #53: Trilobita – A Prolific Paleozoic Posse

The biggest stars of the Cambrian euarthropods, and most of the Paleozoic Era, were of course the trilobites. Known from literally tens of thousands of species spanning over 270 million years, they’re some of the most recognizable and popular fossils.

Trilobites’ exact evolutionary origins and transitional forms are unknown, but they’re thought to have originated in Siberia in the very early Cambrian and their leg anatomy indicates they were a part of the artiopodan lineage. They made a sudden and dramatic entrance to the fossil record about 521 million years ago, appearing fully-formed and rapidly diversifying and spreading all around the world within just a couple of million years.

Their hard calcified exoskeletons made them much more likely to fossilize than soft-bodied animals, with a distinctive three-part body plan consisting of a head shield, three-lobed thorax segments, and a tail shield. Each individual regularly molted their carapace throughout their life, meaning that most trilobite remains are actually empty discarded shells rather than actual carcasses.

Along with being heavily armored arthropod tanks, most species were also able to roll themselves up to defend against predators, and some developed additional elaborate spines and spikes.

…And some were just weird.