Fossils of cambroernids were first discovered in the early 1900s, but these Paleozoic animals were so confusing that for a long time their evolutionary relationships were a mystery.
Discophyllum peltatum lived during the late Ordovician, about 458-448 million years ago, in what is now New York, USA. Up to around 11cm in diameter (~4.3″), it was one of the disc-shaped cambroernids – a lineage known as eldoniids – with a shallow domed body containing a clockwise-coiling sac and delicate feeding tentacles around its mouth.
Its disc would have been tough but flexible, containing numerous supporting radial structures that were probably part of a fluid-filled hydrostatic skeleton, giving it an almost-radially-symmetric body plan superficially resembling a jellyfish.
The lifestyle of eldoniids is still uncertain, but they seem to have mostly sat on the seafloor, possibly extending their tentacles out from under their discs to grab nearby food.
Early echinoderms seem to have gone through an asymmetrical phase before starting to evolving their characteristic radial symmetry. The first truly radial forms had three-way symmetry, but soon a group called the edrioasteroids upped that count to five.
First appearing in the fossil record around 525 million years ago in the early Cambrian, edrioasteroids were mostly shaped like discs or domes, and were immobile filter-feeders that lived permanently attached onto surfaces like the seafloor or the shells of other animals. Unlike most modern echinoderms their pentaradial symmetry was actually created by taking a tri-radial body plan and forking two of their arms near the bases to create a total of five.
Thresherodiscus ramosus
Thresherodiscus ramosus was an unusal edrioasteroid that lived in the shallow seas of what is now central Canada during the late Ordovician, around 460-450 million years ago. Up to about 4cm in diameter (~1.6″), its arms split additional times at irregular intervals, creating a complex asymmetrical branching pattern across its upper surface.
The tips of its arms protruded slightly over the rim of its body, and along with the erratic extra branching this may have been an adaptation to increase its food-gathering surface area.
Another group of early pentaradial echinoderms known as the blastozoans were characterized by erect feeding appendages called brachioles. But some blastozoans abandoned their five-way symmetry in favor of much stranger arrangements, sometimes having as few as two arms – and, in some cases, two mouths.
Amygdalocystites radiatus
Known from the same general area and time period as Thresherodiscus, Amygdalocystites radiatus was part of an Ordovician-to-early-Silurian lineage known as paracrinoids, which attached their irregularly-shaped bodies to the seafloor via a stem.
About 5cm long (~2″) Amygdalocystites had just two asymmetric arms forming “food grooves” along its upper edge, each lined with numerous long brachioles along just one of their sides. It probably orientated itself so its body was facing down-current, which would have created eddies that brought suspended food particles within easier reach of its brachioles.
Represented today by starfish, brittle stars, sea urchins, sea cucumbers, and crinoids, the echinoderms have a characteristic five-way radial symmetry that makes them barely even recognizable as bilaterians. Their true ancestry is only revealed by their genetics and their larvae, which still retain bilateral symmetry – and the way they metamorphose into adults is bizarre, essentially growing a whole new radial body from within the left side of their larval body.
(Sea cucumbers and sand dollars are superficially bilateral as adults, but evolved this secondarily on top of their existing radial symmetry. And some adult echinoderms like starfish also seem to retain a little bit of “behavioral bilaterism”, generally preferring to move with a specific arm always acting as their “front” end.)
The first known echinoderms appeared in the fossil record during the early Cambrian, about 525 million years ago, but the common ancestor of the whole group probably actually originated a few tens of millions of years earlier in the mid-to-late Ediacaran. Early echinoderms seem to have started off as flattened animals that sat on the seafloor filter-feeding, and with this largely immobile way of life their bodies started to shift into asymmetry, no longer constrained by the locomotory advantages of being bilaterally symmetric.
In fact, for these early sedentary filter-feeders being radial was actually much more advantageous, able to distribute sense organs all around their bodies and grab food from any direction without having to reposition themselves, converging on the lifestyle of non-bilaterian cnidarian polyps. The evolutionary transition from bilateral to asymmetrical to pentaradial seems to have happened incredibly quickly during the Cambrian Explosion, and all modern echinoderms probably evolved from a group called the edrioasteroids, maintaining their new base body plan even when they later began taking up more mobile lifestyles again.
But during the process of all that some very alien-looking lineages split off at various stages of anatomical weirdness.
Stylophorans had asymmetrical bodies with a single feeding arm at the front, and varied from irregular boot-like shapes to almost bilateral heart shapes depending on their specific ecologies. The highly asymmetrical forms were probably spreading their weight out over soft soupy mud in quiet waters, while the more bilateral forms may have been more streamlined to deal with stronger water currents.
Sokkaejaecystis serrata
Sokkaejaecystis serrata was a stylophoran that lived during the late Cambrian, about 501-488 million years ago, in what is now South Korea. It was tiny, only about 1cm long (~0.4″), and its boot-shaped body was surrounded by spines and flanges that spread out its surface area and probably also made it much more awkward for small predators to attempt to eat.
Meanwhile the solutes started off as immobile animals living attached to the seafloor via a stalk-like appendage. But fairly early in their evolution they switched to a more active mode of life, modifying their stems into tail-like “steles” that were used to push themselves along.
Maennilia estonica
Maennilia estonica lived in what is now Estonia during the late Ordovician, about 450 million years ago. It was quite large for a solute at about 12cm long (~4.7″), with a sort of vaguely-trapezoidal body, a short feeding arm, and a long thin stele.
Both of these strange early echinoderm lineages were surprisingly successful, surviving for a good chunk of the Paleozoic Era alongside their more familiar radial relatives. The solutes lasted until the early Devonian about 400 million years ago, and the stylophorans continued all the way into the late Carboniferous about 310 million years ago.
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.
Cambroernids were a bizarre group with branching feeding tentacles and a gut enclosed in a coiled sac. They came in a range of forms from worm-like to cup-like to disc-shaped, and despite their fossils being known since the early 1900s their evolutionary affinities were a longstanding problem. Various species had been interpreted in the past as sea cucumbers, jellyfish, tunicates, gnathiferans, or lophophorates, but in recent years they’ve been recognized as all being related, and linked to the ambulacrarians.
And it’s still not entirely clear where in that group they actually belong. They were probably a weird early stem lineage, but they might also be early stem-hemichordates or stem-echinoderms.
It seems like echinoderms became five-way symmetric incredibly quickly following the group’s first appearances in the early Cambrian. We don’t really know why this secondary radial symmetry evolved in the group – but we do know that the common ancestors of all modern pentaradial echinoderms were suspension-feeding animals that lived attached to the sea floor.
And those ancestors were probably a group called the edrioasteroids.
While many of the earliest echinoderms had bizarre asymmetrical forms, at some point members of their lineage adopted radial symmetry instead – a development that would eventually lead to the familiar five-way symmetry of most modern species.
And they may have transitioned to that via three-way symmetry.
During their early evolution, echinoderms started developing unusual asymmetric body plans – and some of them were so strange-looking that for a while it wasn’t clear if they even were echinoderms.
Modern echinoderms typically have five-way radial symmetry as adults, and don’t at all resemble other deuterostomes – but their larvae give away their ancestry, still retaining bilateral traits and only developing radial symmetry when they mature and metamorphose.
The earliest definite echinoderms are known from the early Cambrian, about 525 million years ago, which seems to be around the point when early members of the group first developed biomineralized skeletons and became much more likely to fossilize. However, they must have an evolutionary history going back further than that, and molecular clock estimates suggest their last common ancestry with their closest relatives the hemichordates was in the Ediacaran about 570 million years ago.
For a long time the transition from bilateral to radial symmetry was a mystery, but various fossil discoveries are starting to reveal how this unique group of animals evolved.
Enteropneusts, commonly known as acorn worms, are the most numerous group of modern hemichordates with over 100 known species. Most of them burrow in sediment eating organic detritus, but a few are filter-feeders and some deep-sea species crawl and drift around over the sea floor.
Their fossil record is poor due to their soft bodies, but the transitional form Gyaltsenglossus has recently given us a glimpse at acorn worms’ ancestral links with their cousins the tube-dwelling pterobranchs.
But that’s not the only fossil hemichordate with surprising traits from both lineages. It turns out the characteristic tubes of pterobranchs may actually have been ancestral to all modern hemichordates – with the acorn worms later secondarily losing the ability to make them.
