Strange Symmetries #06: Trilobite Tridents

The genus Walliserops was one of the weirdest-looking trilobites, covered in numerous pointy spines and sporting a large three-pronged “trident” on the front of its face.

They also had some degree of asymmetry in their bodies. Their tridents often didn’t fork evenly, and their long forehead spines curved off to one side – possibly so they could lift their heads up without stabbing themselves in the back.

Walliserops hammii lived in what is now Morocco during the early-to-mid Devonian, about 403-392 million years ago. Around 5cm long (~2″) It was one of the “short trident” species of Walliserops, and its chunky forehead spine curved particularly strongly to the right.

The function of these trilobites’ elaborate tridents is still poorly understood. But an unusual individual of the long-tridented species Walliserops trifurcatushas been found with a lopsided four-pronged trident, and since it was able to grow to full maturity the shape of the structure probably wasn’t absolutely vital for survival, suggesting it wasn’t used for feeding or sensory purposes.

The tridents may instead have been used for combat with each other similar to the horns of some modern beetles. However, these sorts of features are usually only seen in males, and there’s currently no definite evidence for any significant sexual dimorphism in trilobites.

(Although perhaps like ceratopsid dinosaurs their ornaments were just present in both males and females, being also useful for species recognition, visual display, and defense against predators.)

Strange Symmetries #05: Enigmatic Eldoniids

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.

They had coiled bodies and fractal-branching feeding tentacles, and they ranged in shape from worm-like to cup-like to disc-like. Historically various species were classified as sea cucumbers, jellyfish, tunicates, gnathiferans, or lophophorates, but more recently they’ve been recognized as a single united lineage of ambulacrarians, closely related to modern echinoderms and hemichordates.

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.

Strange Symmetries #04: Even More Echinoderms

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.

A colored line drawing of Thresherodiscus, an extinct early echinoderm. It's a domed disc-shaped creature that looks like it has a starfish merged onto its upper surface – but the "arms" branch many more times than five, and not totally symmetrically, creating an erratic forking pattern. It's depicted with a dark purplish body and brighter orange arms.
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.

A colored line drawing of Amygdalocystites, an extinct early echinoderm. It has an oval body with a curving stem coming out from its right side, with the stem ending in a circular holdfast. Two "food grooves" run along its top edge, roughly in line with each other on each side of where its not-visible mouth is located. Each food groove has a single row of long tendril-like feeding appendages growing from its left edge. It's depicted with a red and yellow color scheme, with a striped stem and irregular stripes on its body giving a sort of flame-like pattern, and purple tips on its feeding appendages.
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.

Strange Symmetries #03: Eerie Early Echinoderms

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.

A colored line drawing of Sokkaejaecystis, an extinct early echinoderm. It has a body shaped roughly like a flattened boot, with spikes and flanges growing from around its margin. What looks like a long tail-like appendage growing from the sole of the boot shape is actually a starfish-like feeding arm at the animals' "front" end. It's depicted with orange-brown coloration with brighter yellow on the spikes and flanges, and darker brown irregular stripes over its body.
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.

A colored line drawing of Maennilia, an extinct early echinoderm. It has a flattened body shaped like a lumpy trapezoid, with a single short starfish-like arm growing from its left side and a long thin segmented tail-like appendage growing from the right side of its back end. It's depicted with mottled red and yellow coloration like a camouflage pattern.
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.

Strange Symmetries #02: Oh Worm

Living during the Cambrian Period about 518 million years ago, Wufengella bengtsonii was discovered in the Chinese Chengjiang fossil deposits and was recently named and described in late 2022.

It was a small worm-like animal about 1.6cm long (~0.6″), with bundles of long bristles along its sides and flap-like structures on its underside. Its back was also covered with sclerite armor arranged in a strangely asymmetrical fashion, with larger overlapping plates in the middle and numerous smaller cap-like sclerites distributed unevenly along each side.

Although its bristles and appendages resemble those of annelid worms, the distinctive structure of the sclerites identifies Wufengella as being a member of the tommotiids – early relatives of modern lophophorates (bryozoans, brachiopods, and horseshoe worms). 

Its discovery actually confirms an old prediction that lophophorates probably originated from armored worm-like animals, representing an evolutionary link between earlier free-living annelid-like forms and later immobile filter-feeding tommotiids.

It’s not known why the armor on Wufengella‘s back was so unevenly organized – but some of the later tube-like tommotiids also had weird symmetry going on, with forms like Eccentrotheca having irregular sclerites arranged in a spiral around their bodies.

Strange Symmetries #01

Most animals are bilaterally symmetric, having body plans with mirrored left and right sides – which also allows them to have a defined head end, rear end, top side, and underside.

It’s not entirely clear what evolutionary advantage this type of symmetry gave to the first bilaterians, which would have been been small “simple” worm-like animals living sometime during the Ediacaran Period between 600 and 560 million years ago. The current generally accepted explanation is that it probably allowed for better active locomotion – clustering sense organs at the head end and directing body movement more efficiently towards food sources and away from threats.

However, this sort of symmetry is never completely perfect. Internal structures like organs are often arranged nonsymmetrically, and the realities of genetics, physical development, and environmental influences always result in external small deviations.

…But not every bilaterian has stayed roughly symmetrical.

Over the last half-billion years or so some bilaterians have abandoned their roughly-mirror-image body plans in favor of something distinctly wonkier. Asymmetry has evolved multiple times in various different lineages, and so every weekday this month we’ll be looking at some examples.

And we might as well start way back near the beginning:


Strange Symmetries #01: Almost Bilateral

Living in the Ediacaran between about 567 and 550 million years ago, the proarticulatans were flattened rounded organisms with two rows of soft “quilted” rib-like segments (known as isomers) and sometimes a larger fused “head” section at the front. The left and right isomers weren’t perfectly mirrored, instead being offset from each other in a glide reflection pattern – but the presence of a clear central body axis suggests these animals may have had some sort of relation to the earliest bilaterians, possibly even being a very early stem group that was experimenting with a not-quite-totally-bilateral body plan.

A colored line art drawing of Vendia, an extinct enigmatic Precambrian animal. It's shaped like a flattened oval and vaguely resembles an eyeless limbless gummy trilobite, with several rib-like segments on each side of its midline. The segments alternate instead of lining up symmetrically, and the larger front pair are fused into a head-like structure.
Vendia sokolovi

Discovered in what is now northwest Russia, and dating to around 555 million years ago, Vendia sokolovi was a small proarticulatan measuring about 1.1cm long (0.4″). It had a rather small number of isomers compared to some of its relatives, only 7 per side, and seems to have had a simple digestive tract that branched into each isomer.

(The superficial resemblance to trilobites was coincidental – while we might not be entirely sure what these things were, we do at least know they weren’t closely related to early arthropods.)

Very little overall is known about these animals’ lifestyles. Trace fossils suggest they were able to move around, feeding on microbial mats on the seafloor, and they may also have been able to firmly stick themselves onto the spots they were currently grazing.

Paucipodia

Lobopodians were some of the earliest known panarthropods, closely related to velvet worms, tardigrades, and the ancestors of all the true arthropods. They were small soft-bodied worm-like animals with multiple pairs of fleshy legs, and some species also bore elaborate spikes, armor plates, and fleshy bumps all over their bodies – with the spiny Hallucigenia being the most famous example.

But unlike its more charismatic relative Paucipodia inermis here didn’t seem to have any ornamentation at all.

Known from the Chinese Chengjiang fossil deposits, dating to about 518 million years ago, Paucipodia lived in what was then a shallow tropical sea. Its 13cm long (~5″) tubular body had nine pairs of legs, with each foot tipped with a pair of hooked claws, and the inside of its mouth was ringed with tiny sharp teeth.

Several specimens have been found preserved in association with the weird gummy-disc animal Eldonia, which may indicate Paucipodia either preyed on them or scavenged on their carcasses.

Some Paucipodia fossils also have enigmatic tiny “cup-like” organisms attached to their legs. It’s currently unknown what exactly these were, or whether they were parasitic in nature or simply opportunistically “hitching a ride” similar to the Inquicus found on armored palaeoscolecid worms in the same fossil beds.

Noripterus

Noripterus complicidens was a pterosaur that lived in what is now Mongolia and northwest China during the early Cretaceous, about 140 million years ago.

It had a wingspan of up to 4m (~13′), with a head-and-body length of around 1.2m (~4′), and like other dsungaripterids it had a distinctively reinforced skull, with a toothless beak at the front of its jaws and strong widely-spaced teeth further back – an arrangement that was probably used to catch and then powerfully crack open hard-shelled prey.

Unlike other pterosaurs, however, Noripterus also had some very unusual feet.

All other known pterosaurs seem to have had plantigrade hindlimbs, standing and walking with the whole foot on the ground. But Noripterus had toe joints that looked more like those of theropod dinosaurs than other pterosaurs, with a higher level of upward flexibility and potentially a more digitigrade posture standing on just its toes.

With dsungaripterids already having fairly stout body proportions that suggest they spent a lot of time walking around on the ground, Noripterus may have been even more agile and adept at terrestrial locomotion. Digitigrady is generally more efficient for moving at higher speeds, so this pterosaur might have been a runner behaving similarly to modern ground birds, preferring to sprint away from threats on foot rather than launch itself into the air.

Tropidosuchus

Proterochampsids were a group of Triassic archosauriformes, closely related to the true archosaurs (crocodilians, pterosaurs, and dinosaurs/birds).

Known only from South America between about 242 and 205 million years ago, these reptiles’ heads were wide at the back but very narrow along the snout, often with prominent bony bumps and ridges on their skulls, and they had less osteoderm armor on their bodies than other archosauriformes.

They’ve traditionally been interpreted as very crocodile-like and semi-aquatic, but their long slender limbs and presence in rather arid paleoenvironments suggest they may have been more terrestrial fast-running predators.

Tropidosuchus romeri here lived about 235 million years ago in what is now Argentina. It was one of the smaller proterochampsids, only about 50cm long (1’8″), with just a single row of osteoderms along its back, and had larger and lower-set eyes compared to its relatives.

CT scans of its braincase indicate it had a particularly good sense of smell, and it may have relied mainly on scent to locate prey.

Antaecetus

Antaecetus aithai was an early whale that lived during the late Eocene (~40 million years ago) in what is now Morocco, at a time when northern Africa was covered by a warm shallow sea.

It was part of the “basilosaurids“, some of the first fully aquatic cetaceans – traditionally considered to be a single defined group, but more recently found to be more of an “evolutionary grade” of multiple early whale lineages – and much like Basilosaurus it had elongated back vertebrae that would have given it a very long slender body shape.

Antaecetus also had a proportionally smaller head and smaller teeth than other basilosaurids, along with much denser bones and a stiffer spine that would have made it a rather slow swimmer with reduced maneuverability. It was also fairly small overall compared to most of its relatives, probably around 6m long (~20′).

It was probably a slow-moving coastal water animal somewhat like modern sirenians – except unlike manatees and dugongs it was carnivorous. Its relatively delicate teeth suggest it was feeding on soft-bodied prey like cephalodpods, and with its lack of speed it must have been some sort of ambush predator, waiting around for potential prey to come within striking range.