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It Came From The Wastebasket #05: The Trouble With Troodon

Troodontids were small bird-like theropod dinosaurs, lightly built with slender legs and sickle-shaped “raptor” claws on the second toes of their feet. They had fairly big brains proportional to their body size, rather like modern birds, and their large forward-facing eyes had good depth perception. Owl-like asymmetrical ears in some species gave them a very keen sense of hearing, suggesting they may have been nocturnal hunters using sound to pinpoint the location of small prey.

The original specimen of the namesake of the group, Troodon formosus, was a serrated tooth discovered in the 1850s, about 77 million years old and originating from the Late Cretaceous Judith River Formation fossil beds in Montana, USA. It was so little to work with that it was initially mistaken for a lizard tooth, then during the 20^th century it was recognized as belonging to a dinosaur and spent time classified as a megalosaurid, then a pachycephalosaur, then finally as a small theropod similar to the Mongolian Saurornithoides.

In the late 1980s it was merged together with multiple other troodontids (including Stenonychosaurus of speculative “dinosauroid” fame), and since Troodon had been the first of all of them to be named it took priority as the genus name.

And then for a while every single Late Cretaceous troodontid specimen from North America was also lumped into Troodon, turning it into a wastebasket taxon.

An illustration of five troodontids standing in a circle facing each other. They're all very bird-like feathered dinosaurs with vaguely owl-like faces, reddish-orange and dark brown plumage, and white spots and bars on their longer wing and tail feathers. There are minor variations in their color shades and markings, but otherwise they all look incredibly similar to each other. The overall composition of the image is similar to that of the "Spider-man pointing at Spider-man" memes.

The problem was that all these troodontids came from locations separated by thousands of kilometers and millions of years of time, and it’s unlikely that they all actually represented just one single species. But they were only known from rare fragmentary remains, making distinguishing them from each other difficult, and the original Troodon tooth didn’t really have any distinctive features either – it turns out most troodontid teeth all look exactly the same!

It was becmoning increasingly dubious whether Troodon was even a valid name at all, and during the 2010s several paleontologists began trying to sort the mess out. The old names Pectinodon and Stenonychosaurus were revived, and some ‘Troodon’ fossils were also split off and given completely new names, becoming Albertavenator and Latenivenatrix*.

* Although Latenivenatrix might not actually be distinct enough from Stenonychosaurus to justify having a separate name.

As of 2022, Troodon itself is now in a sort of taxonomic limbo, with some paleontologists abandoning it as a dubious name while others are still arguing in favor of continuing to use it. The name could potentially be properly rescued if the original tooth can be clearly linked to better fossil material, letting Troodon take over priority again from one of the other better-established troodontids, or by defining a new type species similar to what happened with Iguanodon.

…But with how incredibly generic that tooth is, both of those options would be very difficult.

It Came From The Wastebasket #04: Breaking Up Bellerophon

Bellerophonts were small snail-like marine molluscs that were either early gastropods or very close relatives of them. They had symmetrically-coiled shells superficially shaped like those of nautiluses, with about half of the shell covered by their mantle similarly to some modern sea snails, and some fossil shells also preserve hints of banded color patterns.

First appearing in the late Cambrian (~500 million years ago), these molluscs existed all the way until the early Triassic, surviving the Great Dying mass extinction (~252 million years ago) only to go extinct just a short time later (~249 million years ago) – a phenomenon known as “dead clade walking”, when a group just barely scrapes through a mass extinction event but doesn’t manage to actually recover afterwards.

The whole group is something of a wastebasket of similar-looking shells, and might actually be more of an “evolutionary grade” made up of various early gastropods and gastropod-relatives than a single defined lineage.

But there’s also another wastebasket nestled inside this wastebasket: the namesake of them all, the genus Bellerophon.

An illustration of Bellerophon, an extinct sea-snail-like mollusc. It has a banded shell that coils vertically like a nautilus, with a ridge along the midline. It has a wide flat foot, its mantle covers about halfway up the sides of its shell, and it has a pair of snail-like head tentacles and a siphon. — *Bellerophon tenuifascia*

Originally named in 1808, this genus has had a huge number of species assigned to it over the last couple of centuries. This gives a false impression that Bellerophon-like molluscs didn’t change for hundreds of millions of years, and it makes figuring out their actual long-term patterns of evolution and extinction much more difficult.

In the last few decades some mollusc paleontologists have been gradually chipping away at Bellerophon, and multiple new genera have been broken off from it. But even today it remains a very bloated mess – there are still well over a hundred named species spanning about 230 million years of geologic time.

Studies do indicate the whole genus is highly polyphyletic, made up of a tangle of multiple different lineages that all really need to be revised and renamed – but there’s a lot of work still needing to be done to clean up this particular wastebasket.

It Came From The Wastebasket #03: The Gomphothere In The Room

The three living species of elephants are the last surviving members of the proboscidean lineage – but up until the end of the last ice age about 11,000 years ago their relatives were much more numerous and widespread, found on every continent except Australia and Antarctica. Mammoths are probably the most famous of these recently-extinct proboscideans, closely related to modern Asian elephants, but there were also the more distantly-related stegodonts and mastodons…

…And also the gomphotheres.

A cladogram showing the traditional classification of gomphotheres as a poorly-defined group evolutionarily between mastodons and modern elephants.

Traditionally any proboscideans that fell into the evolutionary grade between mastodons and elephants-and-stegodonts were all labelled as gomphotheres. As a result by the late 20^th century this group ended up as a wastebasket full of elephant-like forms that didn’t easily fit anywhere else, defined more by what they weren’t rather than by any features they all had in common.

This big collection of gomphotheres was highly diverse. Some species independently evolved similar convergent features, and there was also considerable individual physical variation within species, making the actual taxonomy of these animals very difficult to figure out. But over the last few decades there’s been a lot of revision of proboscidean evolutionary relationships, and gradually the gomphothere wastebasket has been clearing up. Groups like the choerolophodontids, amebelodontids, and anancids have been split off, leaving a more defined lineage of gomphotheres that do have shared anatomical characteristics – distinctive three-lobed trefoil-shaped wear patterns on their molar teeth.

A cladogram showing a more modern classification of gomphotheres, with the gomphotheriids as just one of several different lineages originating between mastodons and modern elephants.

These gomphotheriids were widespread, found across Africa, Europe, Asia, and the Americas – and they were especially successful in the latter. They arrived in North America during the Miocene (~16 million years ago) via the Beringia land bridge, and rapidly spread across the continent and down into Central America. They went on to become the only proboscideans to disperse into South America during the Great American Biotic Interchange, with two different lineages arriving at separate times – Notiomastodon around 2.5 million years ago, and Cuvieronius around 750,000 years ago.

An illustration of Cuvieronius, an extinct elephant-like gomphothere. It has a longer flatter head than modern elephants, and its tusks have a spiral twist to them. — *Cuvieronius hyodon*, a 2.3m tall (7’7″) South American gomphotheriid with distinctive spiraled tusks.

The exact relationships of the gomphotheriids to other elephant-like proboscideans are still a little uncertain. Both protein sequences and mitochondrial DNA have recently been recovered from 35,000-13,000-year old Notiomastodon specimens, but these studies have given different taxonomic conclusions – with the protein results suggesting gomphotheriids were most closely related to mastodons, and the DNA results suggesting they were much closer to true elephants.

It Came From The Wastebasket #02: What Makes A Monoclonius?

The first fossil remains of Monoclonius crassus were discovered in the Late Cretaceous Judith River fossil beds (~75 million years old) in 1876 in Montana, USA. It was one of the many dinosaur species hurriedly named as part of the Bone Wars, and was described based on a mixture of bones from several different sites.

At first much of this dinosaur’s anatomy was poorly understood, and at first it was misidentified as a hadrosaur. The skull remains were fragmentary and ceratopsians hadn’t yet been identified as a group, so Monoclonius‘ horns weren’t even recognized as being horns and a piece of the frill was initially misinterpreted as part of a breastbone.

Once the much better-preserved Triceratops was discovered in 1889, and the existence of ceratopsians was recognized, Monoclonius was re-examined and identified as a similar dinosaur – and three more species were quickly described within the genus, also based on very fragmentary fossils.

An illustration of Monoclonius, a dubious species of horned ceratopsian dinosaur. It has a parrot-like beak, a long straight nose horn, and a pair of small stubby brow horns. Its large bony neck frill is rimmed with small spikes, with a pair at the very top being longer and curling sharply downwards against the front of the frill. Its body is bulks and quadrupedal with a a thick tapering tail, and there are bumpy scales and sparse short quill-like spines on its back. It's colored mottled orange-and-brown, and there are hints of bright blue on its frill. — *Monoclonius crassus*

Then for a while afterwards every ceratopsid fossil that wasn’t clearly a Triceratops was then just dumped into Monoclonius, quickly turning the genus into a wastebasket full of dubious indistinct remains.

But then…

The new challenger screen from Super Smash Bros Ultimate, with the character silhouette replaced by that of Centrosaurus, a horned ceratopsian dinosaur. Text on the image reads "A new foe has appeared! Challenger approaching!"

Centrosaurus apertus was named in 1904, from the similarly-aged Dinosaur Park Formation in southern Alberta, Canada. It had originally been one of the various species of Monoclonius, but was now claimed to be different enough to deserve its own separate genus name – and this started a decades-long controversy between several paleontologists.

Over the new few decades arguments went back and forth over whether Centrosaurus was actually valid or if it was just a junior synonym of Monoclonius. As more and better ceratopsid fossil material was discovered several other Monoclonius species were eventually split off into their own separate genera, too, creating Styracosaurus, Chasmosaurus, and the somewhat dubious Brachyceratops. But other new species also continued to be lumped into Monoclonius up until 1990, meaning that over its century of existence this wastebasket taxon had at one point or another contained at least 16 different species.

During the 1990s opinion began to turn against Monoclonius, increasingly regarding it as a dubious name. Its original type specimen was a chimera of multiple different individuals (and possibly multiple different species), and it just didn’t have any distinct enough anatomical features to distinguish it from other ceratopsids.

Centrosaurus, meanwhile, was further validated by the discovery of huge bonebeds containing thousands of individuals, making it into one of the best-known of all ceratopsians.

Today Monoclonius‘ name remains attached to a few fossil specimens, but only the ones that are too indistinct to classify as anything else. Some “Monoclonius” have also turned out to actually be juveniles and subadults of other ceratopsians – it seems many young centrosaurines had a Monoclonius-like stage in their growth, before they went on to develop their own species’ distinctive horn and frill shapes.

So Monoclonius may never have been a distinct genus at all – it was just a bunch of different ceratopsian teenagers!

It Came From The Wastebasket #01: Is This An Insectivore?

Most of the wastebasket taxa featured this month are completely extinct and known only from fossils, but to start things off let’s take a look at a major example of how even groups with living members could have their classification muddled up for centuries.

The name Insectivora first came into use in the early 1820s, and was used to refer to various “primitive-looking” small insect-eating mammals, with modern shrews, moles, hedgehogs, tenrecs, and golden moles as the original core members.

An illustration showing the animals that originally made up "Insectivora". From left to right it pictures a shrew, a tenrec, and a hedgehog on the top row, and a mole and a golden mole on the bottom row. Text at top of the the image reads "Insectivora". — Insectivora

Then over the next few decades solenodons, treeshrews, sengis, and colugos all got lumped in with them too.

By the early 20^th century insectivorans were considered to represent the “primitive” ancestral stock that all other placental mammals had ultimately descended from, and any vaguely similar fossil species also got dumped under the label. Extinct groups like leptictids, cimolestans, adapisoriculids, and apatemyids all went into the increasingly bloated Insectivora, too, making the situation even more of a wastebasket as time went on.

An illustration showing the animals that made up the expanded historical version of "Insectivora". From left to right it pictures a leptictidan, a shrew, a tenrec, a hedgehog, and a sengi on the top row, an apatemyid, a mole, a golden mole, and a solenodon in the middle row, and a cimolestan, a colugo, and a treeshrew on the bottom row. Text at top of the the image reads "…Insectivora?", styled like a typewritten label that has been stuck over the previous image's text. — …Insectivora?

The problem was that the only characteristics that really united these various animals were very generic “early placental mammal” traits – small body size, five clawed digits on the hands and feet, relatively unspecialized teeth, and mostly-insectivorous diets – and attempts at making sense of their evolutionary relationships were increasingly convoluted.

An image of a diagram from a 1967 academic paper, showing a complicated attempt to figure out the evolutionary relationships of "insectivores", with many different group names linked by arrows. For comparison next to it is the "Pepe Silvia" conspiracy wall meme. — …They’re the same image.

(Image sources: http://hdl.handle.net/2246/358 & https://knowyourmeme.com/memes/pepe-silvia)

The rise of cladistic methods from the 1970s onwards resulted in a lot of “insectivores” finally being recognized as unrelated to each other, removing them from the group and paring things back down closer to the name’s original definition. The idea that insectivorans were ancestral to all other placentals was abandoned, instead reclassifying them as being related to carnivorans, and the remaining members were recognized as just retaining a superficially “primitive” mammalian body plan.

Just shrews, moles, hedgehogs, solenodons, tenrecs, and golden moles were left, and to disassociate from the massive mess that had been Insectivora this version of the group was instead now called Lipotyphla.

An illustration showing the animals that made up "Lipotyphla". From left to right it pictures a solenodon, a tenrec, and a hedgehog on the top row, and a shrew, a mole, and a golden mole on the bottom row. Text at top of the the image reads "Lipotylpha", styled like an embossed label-maker sticker that has been stuck over the previous images' text. — Lipotyphla

But there were still no unique anatomical links between the remaining lipotyphlans. And then once genetic methods became available in the late 1990s, something unexpected happened.

Studies began to suggest that tenrecs and golden moles were actually part of a completely different lineage of placental mammals, the newly-recognized afrotheres, with their closest relatives being sengis and aardvarks. Meanwhile the rest of the lipotyphlans were laurasiatheres, closely related to bats, ungulates, and carnivorans.

Lipotyphla was suddenly split in half. For a while it was unclear if even the remaining shrew-mole-hedgehog-solenodon group was still valid – hedgehogs’ relationships were especially unstable in some studies – but by the mid-2000s things began to settle down into their current state.

Finally, after almost 200 years of confusion, the insectivore wastebasket has (hopefully) now been cleaned up. The remaining “true lipotyphlans” do seem to all be part of a single lineage, united by their genetics rather than by anatomical features, and are now known as Eulipotyphla.

A few fossil groups like nyctitheriids and amphilemurids are generally also still included, but since this classification is based just on their anatomy it isn’t entirely certain. The only exception to this are the nesophontids, which went extinct recently enough that we’ve actually recovered ancient DNA from them and confirmed they were eulipotyphlans closely related to solenodons.

An illustration showing the animals that now make up Euipotyphla. From left to right it pictures a solenodon, and a hedgehog on the top row, and a shrew, a mole, and an amphilemurid on the bottom row. Text at top of the the image reads "Eulipotylpha", with the letters "E" and "U" hastily scribbled onto the front of the previous image's text. — Eulipotyphla

And a bonus image with species IDs:

It Came From The Wastebasket

Taxonomy – the naming, description, and classification of living things – is one of the foundations of biology and ecology. We need to know what things are in order to properly understand them and their evolutionary relationships, and without that we can’t build up an accurate picture of the true diversity of life on Earth.

Taxonomy of living species is also vital for conservation efforts, recognizing unique species that would otherwise go unnoticed. Accidentally using the same name for multiple things can easily mask the decline and potential extinction of critically endangered populations – for example, if we’d just assumed all Galápagos giant tortoises were exactly the same we’d never have realized that Lonesome George was the last known individual of the Pinta Island subspecies, or made efforts to find living hybrid descendants of his kind.

Meanwhile the paleontological taxonomy of fossils helps us to understand where things came from, and to identify long-term trends of evolution, diversity, and extinction over time. The history of life shows us how different types of organisms coped with changing conditions in the past, so we can try to predict how current climate change will affect the biosphere in the present and future.

But sometimes species don’t neatly fit into our classification system. Maybe they’re rather “generic” or “primitive” examples of that type of organism and don’t really have many unique or specialized features, or maybe the scientists describing them just weren’t able to classify them more specifically at the time, but either way they often end up with the same fate: dumped into a wastebasket taxon.

A pencil sketch of a wire mesh waste-paper basket, tipped over on its side with crumpled pieces of paper spilling out. A whale's tail and a trilobite are poking out of the trash, while a bird-like feathered dinosaur and a shrew-like mammal peer around the sides of the toppled basket.

Wastebaskets aren’t natural lineages, just a default label for things that don’t seem to fit anywhere else, and they’re basically somebody else’s problem to sort out later. Sometimes they can even end up containing things that superficially look very similar to each other but later turn out to not even be closely related at all.

This can be especially bad in paleontology, where there’s often only poorly-preserved and fragmentary fossils to work with and usually no way to verify evolutionary relationships with modern genetic analysis. This can result in wastebaskets getting especially bad if left unchecked – like how for a while in the 19^th and 20^th centuries many fragmentary theropod dinosaurs were just dumped into Megalosaurus, resulting in over 50 dubious species that eventually needed to be carefully reevaluated, renamed, and reclassified.

Every weekday this October we’ll be looking at a different example of these sort of taxonomic tangles – so I’ll see you all on Monday with one the worst historical wastebaskets…

Spectember 2022 #04: Aquatic Brontotheres

Squeezing in one last bonus #Spectember post this year!

This one isn’t based on a specific prompt, but instead is a companion piece to a previous one.

While North American brontotheres were adapting to the spread of grasslands, some of their Asian cousins took a very different evolutionary path through the rest of the Cenozoic.

So today’s concept comes from an anonymous submitter, who requested an arboreal ornithopod dinosaur:

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