Many decapod crustaceans have slightly asymmetrical pincers, often with one claw being chunkier and specialized for “crushing” while the other is more slender and used for “cutting”.
But fiddler crabs take this sort of asymmetry to the extreme as part of their sexual dimorphism – males have one massively oversized claw, which is used for both visual display to potential mates and for physical fights against rivals.
Some of the earliest fiddler crabs are known from the Miocene of what is now northern Brazil. Although the fossils have been given several different taxonomic names since their discovery in the 1970s (including Uca maracoani antiqua, Uca antiqua, and Uca inaciobritoi) they’re currently considered to be indistinguishable from the modern Brazilian fiddler crab, Uca maracoani, meaning that these crabs have remained externally unchanged for the last 16 million years.
Up to about 4cm in carapace width (~1.6″), modern Uca maracoani are found in coastal mangrove swamps and tidal mudflats around the northern and eastern coasts of South America – and some of these environments have also undergone little change since the Miocene. Males of the species can develop their enlarged pincer on either side of their bodies, with lefties and righties seeming to occur in equal numbers.
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.
Palaeopagurus vandenengeli lived in what is now northern England during the Early Cretaceous, about 130 million years ago. Around 4-5cm long (~1.6-2″), it was found preserved inside the shell of the ammonite species Simbirskites gottschei.
Its left claw was much larger than its right, and together they would have been used to block the shell opening when it was hiding away inside. And while the exact shape of its abdomen isn’t known, it probably asymmetrically coiled to the side to accomodate the spiralling shape of the host shell.
Hermit crabs seem to have switched over to using gastropod shells by the Late Cretaceous, around 90-80 million years ago, possibly due to marine snails developing much stronger sturdier shells during this period in response to the increasing prevalence of specialized shell-crushing predators. The more upright snail shells would also have been much easier to drag around the seafloor than ammonite shells – and meant that they were ultimately less affected by the total disappearance of ammonites during end-Cretaceous mass extinction.
Brachiopods (also known as “lamp shells”) superficially look very much like bivalves, but these two groups aren’t very closely related to each other – although they’re both lophotrochozoans, their last common ancestor probably lived sometime in the Ediacaran at least 560 million years ago, and their similarities in appearance are due to convergent evolution.
The two valves of their shells are also arranged differently. Bivalve shells grow on their left and right sides and are usually symmetrical, but brachiopods form their shells from the upper and lower surfaces of their bodies.
As a result brachiopod shells are usually unequal in size and shape but have their own plane of bilateral symmetry down the center – but some of them still managed to become asymmetrical anyway.
Torquirhynchia inconstans lived during the Late Jurassic, about 161-145 million years ago, in the warm shallow seas that covered what is now Europe and Iran. Around 3cm across (~1.2″) it had a strongly ridged shell with an asymmetrical closing edge, positioned high on one side and low on the other.
This unusual uneven arrangment is thought to be an adaptation to living on soft sediments. Asymmetrical brachiopods like Torquirhynchia may have lived with one side of their body mostly buried into the seafloor, and twisted their shell edges so the still-exposed half was raised up to better function for water circulation and filter-feeding.
But some bivalves break that arrangement, developing asymmetrical valves that can be massively different in size and shape.
Gryphaea arcuata was an oyster that lived during the Early Jurassic, about 200-174 million years ago, in the warm shallow seas that covered what is now Europe and eastern Greenland. Around 6cm long (~2.4″), its left valve was thick and strongly convex and curled, while the right valve was relatively thin and slightly concave forming a “lid”.
The gnarled curled claw-like shape of Gryphaea fossils led to them being colloquially known as “devil’s toenails” in some of the regions where they’re commonly found, with folk beliefs that they had the power to prevent joint pain.
Their shape was actually an adaptation to living on very soft seafloor sediments. The larger curled valve acted sort of like a boat on the soupy mud, supporting the Gryphaea‘s weight and preventing it from sinking.
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.)
First discovered in the early 1970s, in the mid-CarboniferousBear Gulch Limestone deposits (~324 million years ago) of Montana, USA, it was initially mistaken for the long-sought-after “conodont animal” due to the presence of numerous conodont teeth inside its body. But just a few years later well-preserved eel-like conodont animals were found elsewhere, and it became apparent that the conodont teeth inside Typhloesus had actually just been part of its last meal.
But if it wasn’t a conodont… then what was it?
Up to about 10cm long (4″), Typhloesus had a streamlined body with a vertical tail fin and paired “keels” along its sides. It had a mouth and a gut cavity, but no apparent anus, and it also didn’t seem to have any eyes or other sensory structures. And in the middle of its body there was something very weird – a pair of “ferrodiscus” organs, disc-shaped structures which contained high concentrations of iron but whose function was completely unknown.
This anatomy just didn’t match any other known animals, so much so that it gained the nickname of “alien goldfish”.
For the next few decades it remained a bizarre enigma, at best tentatively considered to represent an unknown lineage of some sort of metazoan that left almost no other fossil record due to being entirely soft-bodied.
But now, 50 years after its initial discovery, we might just finally have a clue about Typhloesus’ true identity.
It’s not a definite identification yet, and even if it was a mollusc it was an incredibly strange one, with features like the ferrodiscus still lacking any explanation. But this discovery at least shows that there are still new details waiting to be found in the “alien goldfish” fossils, and gives us a start towards bringing its classification back down to earth.
The Ptychopariida were some of the earliest known trilobites, first appearing in the early Cambrian about 521 million years ago and surviving until the end of the Ordovician about 444 million years ago. They included some of the most numerous and common trilobite species, and were probably ancestral to multiple other major lineages – including the very last trilobites at the end of the Permian – making them incredibly important in understanding the overall evolution of trilobites as a whole.
…But this group is also one of the biggest wastebaskets in paleontology.
First established in the early 20th century, the ptychopariids seemed to have some fairly good defining characteristics based on their facial sutures, large thoraxes, and relatively small pygidia. But the group quickly became a dumping ground for a massive amount of Cambrian trilobites, eventually containing numerous different families, hundreds of genera, and many more individual species.
Actually figuring out their internal evolutionary relationships also turned out to be extremely difficult – so much so that some paleontologists working on them just gave up trying and arranged the genera names alphabetically instead!
Even cladistic studies from the 1970s onward struggled to make sense of these highly “problematic” trilobites, and any larger-scale analysis was a daunting task due to how huge and diverse the ptychopariid wastebasket had become over the years. Worse, some of the anatomical features the group had been based around were starting to look more like the result of a lot of convergent evolution across multiple lineages than any actual shared ancestry.
Efforts were still made at breaking up the mess, however, with better-understood sub-groups like the Proetida, Harpida, Asaphida, Trinucleida, and Olenida being gradually split off into their own separate orders over the course of the last few decades.
Ptychoparia striata
But even by the early 2010s what remained of the Ptychopariida was still paraphyletic at best, more of an “evolutionary grade” of early trilobites than a single lineage, with most of its constituent families also rather poorly defined. There was even a proposal to abandon the group entirely, stating that “it serves no scientific purpose” and that its orphaned contents should be considered “order uncertain” until their actual relationships can be untangled.
Today the “ptychopariids” are in dire need of a full revision – since they were the ancestors of many other major groups they’re still crucial for understanding early trilobite evolution. There may be a salvageable single lineage somewhere in the remains of this wastebasket, even if it’s restricted to just close relatives of the genus Ptychoparia, but until somebody tackles them properly they’re stuck in taxonomic limbo with their name only being used in a loose sense.
Protorthoptera was a group of fossil insects created in the early 20th century to categorize “primitive” neopterans – some of the earliest insects to have evolved the ability to fold their wings down over their backs. Known mostly from just fossilized forewings, they first appeared around 320 million years ago in the late Carboniferous, and after heavy losses during the Great Dying mass extinction they eventually disappeared in the mid-Triassic about 240 million years ago.
As early as the mid-20th century the protorthopterans were recognized as being a general taxonomic dumping ground, containing a mixture of early members of multiple different “orthopteroid” insect lineages. But invertebrate paleontologists at the time considered this collection of “primitive” insects to lack enough distinctive features to confidently separate them out from each other, and so the highly paraphyletic grouping continued to be used well into the 1990s.
Ctenoptilus elongatus
But in the early 2000s this situation finally changed. Proper cladistic analysis of protorthopteran fossils identified defining features of the wing vein patterns, and many species were reclassified into various lineages within the Archaeorthoptera – which includes modern grasshoppers, crickets, and locusts along with several closely related fossil groups like the titanopterans and caloneurodeans.
“Protorthoptera” is still sometimes used in a loose sense for fossil neopteran insects that still can’t be confidently classified anywhere else, so the wastebasket isn’t entirely cleared here.
And there are some alternate classification systems (mainly proposed by Russian paleontologists) that instead consider many protorthopterans to be notopterans closely related to modern ice-crawlers, and place others as part of other modern neopteran lineages such as webspinners and true bugs.
Hopefully better fossil discoveries and future studies will eventually help clear things up, and give us a better overall picture of the evolution of these insects.
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.
Hamites attenuatus
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 severalotherheteromorphammonitelineages, 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.
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