the menagerie

April Fools 2022: The Aquatic Dinosaur That Wasn’t

So, Spinosaurus wasn’t technically the first known aquatic non-avian dinosaur.

That title instead temporarily went to Compsognathus corallestris.

While the idea that hadrosaurs and sauropods were wallowing swamp-dwellers had been completely abandoned at the start of the Dinosaur Renaissance, the new view of dinosaurs as active sophisticated animals led to a surprising aquatic hypothesis during the early days of this paleontological revolution.

A specimen of the small theropod Compsognathus discovered in southeastern France in the early 1970s was only the second skeleton ever found of this dinosaur, and came over a century after the first. It was initially thought to represent a new species since it was about 50% larger than the German specimen of Compsognathus longipes, and it seemed to have something very unusual going on with its hands – its forelimbs were somewhat poorly-preserved and distorted, and had traces of some sort of large fleshy structure around the hands that was interpreted as representing elongated three-fingered flippers used for swimming.

This wasn’t necessarily as ridiculous of an idea as it might sound. Compsognathus lived during the Late Jurassic, about 150 million years ago, at a time when Europe was a group of islands in a shallow tropical sea. A semiaquatic dinosaur specialized to swim and dive, hunting the abundant aquatic prey in its environment, and easily able to island-hop all around the European archipelago seemed at least somewhat plausible, and reconstructions of fin-handed C. corallestris even appeared in several popular dinosaur books of the time.

But it didn’t last.

Within just a few years doubt was being cast on this idea, and further studies of both known Compsognathus skeletons in the late 1970s and early 1980s concluded that C. corallestris was actually a fully-grown adult individual of the juvenile C. longipes. The French Compsognathus had normal-looking hands for its kind after all, with two large clawed fingers and a vestigial third finger, and the “flipper” impressions had just been ripples in the fossil slab.

For a long time after that the general view became that there just weren’t any aquatic non-avian dinosaurs at all – but more recent discoveries like the new Spinosaurus material and Halszkaraptor are starting to suggest that some of these animals were much more at home in the water than previously thought.

Something resembling Compsognathus corallestris might still surprise us in the future.

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Retro vs Modern #14: Therizinosaurus cheloniformis

Therizinosaurs were some of of the most unique theropod dinosaurs. It’s only in the last few decades that we’ve started to understand much about them, and they’re still somewhat enigmatic even today.

1950s

The first known therizinosaur fossil discovery was Therizinosaurus cheloniformis, found in Southern Mongolia during the late 1940s and described and named in the mid-1950s based on a few fragments that included some unusually large and elongated claws.

These remains were interpreted as belonging to a giant turtle-like reptile that used its scythe-like claws to harvest aquatic plants – inspiring both parts of its scientific name, with Therizinosaurus meaning “reaper lizard” and cheloniformis meaning “turtle-shaped”.

1990s

The turtle interpretation began to be questioned during the 1970s, and the discovery of some slightly better (but still fragmentary) specimens reclassified Therizinosaurus as an unusual theropod dinosaur.

In the late 1970s and early 1980s fossils of another group of dinosaurs known as “segnosaurs” were also starting to be discovered, with a confusing mixture of anatomical features that seemed to link them to multiple different dinosaur lineages. As a result opinions about their evolutionary relationships varied during the 1980s, sometimes considering them to be theropods, sometimes late-surviving “relic” prosauropods, and sometimes a whole new major lineage of rare and weird saurischians.

Similarities between segnosaurs and the known material of Therizinosaurus were soon noted, and the discovery of the fairly complete Alxasaurus in the early 1990s confirmed that they were all part of the same group of bizarre herbivorous theropods – and the name “segnosaurs” was dropped in favor of “therizinosaurs”, since older names usually get priority in taxonomy.

Reconstructions of Therizinosaurus during this time tended to look rather weird and awkward. As with the majority of dinosaurs during this period it was depicted as entirely scaly and reptilian, and was often shown with a stiff hunched downcurving neck and an oddly tiny-looking tail.

Further discoveries during the 1990s finally began to clarify therizinosaurs’ evolutionary affinities, eventually placing them as an early branch of bird-like maniraptoran theropods, closely related to both oviraptorosaurs and the alvarezsaurs – and in 1999 the discovery of the small early therizinosaur Beipiaosaurus helped to confirm this relationship, revealing impressions of an extensive coat of filamentous feathers and longer stiffer quill-like structures.

2020s

Over the next couple of decades more and more therizinosaur fossils were found in both Asia and North America, and details about these dinosaurs’ appearance, biology, and ecology gradually became better understood. While there’s still a lot we don’t know about them, we do now at least have some fairly complete examples like Nothronychus, fossilized therizinosaur footprints, more feathers, an idea of Beipiaosaurus’ coloration (it was brown!), and even eggs and potential colonial nesting sites.

And while Therizinosaurus itself is still only represented by fragmentary and incomplete material, we now have a much better idea of what it was probably like. It lived in what is now the Gobi Desert during the Late Cretaceous, about 70 million years ago, and would have been both the largest known therizinosaur and the largest known maniraptoran dinosaur, estimated to have been as much as 10m long (33’) based on the proportions of its relatives.

It would have had a tiny head with a toothless beak at the front of its jaws, a long neck, and a wide bulky “pot-bellied” body housing its huge plant-fermenting gut. With its especially large body size it probably wasn’t as extensively feathered as its smaller relatives, but it may have still been sparsely fuzzy across parts of its body.

Unlike most other theropods it walked on all four toes of its feet, with the dewclaw enlarged into an extra weight-bearing digit. Recent analysis of footprints has also suggested that the larger therizinosaurs like Therizinosaurus may actually have been plantigrade, walking with their feet completely flat on the ground. This might turn out to just be an artifact of how the tracks were preserved, but therizinosaurs are certainly already weird enough that it could be a plausible interpretation.

But Therizinosaurus’ most distinctive feature was its hands, with extremely long narrow claws each at least 50cm long (1'8"). Unlike the strongly curved claws seen in other therizinosaurs, these ones were fairly straight for most of their length, only curving more sharply towards their tips.

While in the past these claws have been proposed as being weapons or digging adaptations, they were actually relatively delicate and were probably mainly used for pulling clumps of vegetation closer in a convergently similar manner to the later mammalian chalicotheres and ground sloths.

Therizinosaurus would have been a heavy slow-moving animal, and probably spent a lot of time sitting on its haunches supported by its especially robust hip bones while it browsed on large amounts of vegetation. It likely relied on its pure bulk and intimidation to deter potential predators, possibly even making aggressive displays with its claws when threatened – essentially it may have been a giant goose-sloth.

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Retro vs Modern #13: Stegosaurus stenops

The first known stegosaur fossils were found in England and South Africa between the 1840s and 1870s, but these dinosaurs weren’t properly recognized as a highly distinctive group until the discovery of Stegosaurus itself in North America during the late 1870s.

1880s

The first Stegosaurus reconstructions were based on fragmentary and disarticulated fossil material, and its life appearance was very poorly understood. Initially it was depicted as a bipedal long-necked animal, with its plates laying flat against its back like a turtle shell, numerous spikes across its back, and more plates running along its tail.

1890s-1970s

Better skeletons of the species Stegosaurus stenops were discovered in the late 1880s, and by the 1890s stegosaur anatomy was becoming clearer. Reconstructions quickly adopted an arch-backed body shape with a tiny head and drooping tail, short semi-sprawling forelimbs and long hindlimbs, and with the plates now properly upright on the back and the spikes at the end of the tail.

Stegosaurus’ unique appearance rapidly made it one of the most famous and recognizable dinosaurs to the general public. Its comical-seeming tiny head and even tinier brain also unfortunately ended up contributing to the prevailing early 20^th century attitude that dinosaurs were sluggish and unintelligent, with the myth that it needed a “second brain” in its hips to control its huge body becoming a popular notion for quite some time.

The exact arrangement of the iconic back plates and tail spikes was uncertain for several decades, with early versions in the 1890s having up to eight tail spikes and a single row of plates. This was then updated in the 1900s to a double row of symmetrical plate pairs, and by the 1920s the standard arrangment had soon become an alternating two-row pattern with the tail spikes reduced to four – a layout that’s still considered correct today.

2020s

In the second half of the 20^th century a combination of numerous new stegosaur species from China and the Dinosaur Renaissance began to revise the way Stegosaurus was understood, bringing it into a fully upright posture with its head and tail held high, and recognizing the convergently sauropod-like anatomy of its hands and feet.

But something still wasn’t right.

Compared to other known stegosaurs, Stegosaurus itself was starting to seem… rather weird. Its short neck, short forelegs, giant plates, sloping back and high rump were much more exaggeratedly proportioned than any of its relatives.

This was finally resolved in the 2010s when a near-complete specimen nicknamed “Sophie” was thoroughly described – and revealed that Stegosaurus’ proportions had been wrong the whole time. All previous skeletal reconstructions had been composites, put together from remains of multiple individuals that had all been different ages and sizes, and in the process had heavily distorted our idea of what this animal actually looked like.

Our modern view of Stegosaurus is now a much more typical stegosaur than before. It lived during the Late Jurassic, about 155-145 million years ago, across the Western and South Central United States (with a possible additional occurence in Portugal), alongside several other iconic dinosaurs of the “Jurassic savanna” like Brontosaurus, Diplodocus, and Allosaurus.

It grew up to around 9m long (~30ft), and had a small head with a long narrow snout, with a toothless beak at the front of its jaws and small peg-shaped teeth further back. Bony ossicles lined the underside of its neck, possibly providing chainmail-like protection to its throat, and its skin was covered in tiny pebbly scales interspersed with “rosettes” formed around slightly larger oval scales. Its neck was longer than previously thought, more in line with other stegosaurs, and its torso and hind legs were a bit shorter, making its posture more horizontal and its back less arched.

The actual function of the large back plates is still uncertain. Ideas about them being defensive armor (and speculation about them even being moveable!) have mostly been discounted at this point, since they were actually relatively fragile – although their keratinous covering may have had a fairly sharp edge. Thermoregulation has been a popular explanation for many decades, with blood vessel impressions in the plates being proposed as evidence they were used as “radiators” to prevent overheating like the ears of modern African elephants.

But currently the most likely primary plate function is thought to be visual display, with the large plates increasing the perceived size of Stegosaurus either to intimidate predators and rivals or to impress potential mates. If this was the case then they may have also been strikingly colored and patterned in life.

Meanwhile the “thagomizer” on its tail actually does seem to have been a weapon, with injuries to that area of the body being fairly common, and several Allosaurus fossils have been found with puncture wounds the exact size and shape of Stegosaurus spikes. Articulated specimens have also shown that the tail curved downwards at the tip, holding the thagomizer with the spikes pointing horizontally outwards and backwards.

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