Huge, flightless, and carnivorous, the phorusrhacids (or terror birds) were some of the largest apex predators in South America during its Cenozoic “splendid isolation” as an island continent – and they were possibly the closest that birds ever came to reclaiming the ecological roles of their extinct non-avian theropod dinosaur relatives.
And for a while in the late 1990s and early 2000s there was a hypothesis that they’d even re-evolved clawed hands.
Living during the Pliocene and Pleistocene in Florida and Texas, between about 5 and 1.8 million years ago, Titanis stood around 1.5-1.8m tall (~5-6′) and was heavily built, with long strong legs and a massive hooked beak. Remains of its small wings were incomplete and fragmentary but had seemingly unusual joints, with what looked like a stiffer wrist and more flexible “fingers” than other birds, which led paleontologist Robert Chandler to propose in 1994 that this terror bird species had modified its wings into clawed grasping arms similar to those of dromaeosaurs, used to restrain prey animals while its beak tore them apart.
But the idea of a giant murder-bird with added meathook-hands only lasted about a decade. Further investigation in 2005 showed that Titanis‘ arms weren’t that weird after all – the same sort of joints are found in terror birds’ closest living relatives, the seriemas, and so Titanis really had the same sort of small vestigial wings as many other large flightless birds.
…However, there still could have been some claws on there. Many modern birds actually have one or two small claws on their hands that aren’t visible under their feathers, and terror birds like Titanis having something like that going on is completely plausible – they just wouldn’t have been using them for any sort of specalized predatory function.
Mammalian tusks usually grow in symmetrical pairs with only minor developmental asymmetry, but a few species have evolved much more uneven arrangements.
Odobenocetops peruvianus
Odobenocetops peruvianus was a small toothed whale that lived during the Miocene, about 7-3 million years ago, in shallow coastal waters around what is now Peru. Around 3m long (~10′), it was a highly unusual cetacean with binocular vision, a vestigial melon, muscular lips, and a pair of tusks – features convergent with walruses that suggest it had a similar lifestyle suction-feeding on seafloor molluscs and crustaceans.
In males the right tusk was much more elongated than the left, measuring around 50cm long (~1’8″) in this species and up to 1.35m (4’5″) in the closely related Odobenocetops leptodon. Since these teeth were quite fragile they probably weren’t used for any sort of combat, and they may have instead served more of a visual display function.
The woolly mammoth (Mammuthus primigenius) lived across Eurasia and North America during the last ice age, mostly from the Pleistocene about 400,000 years ago to the early Holocene about 10,000 years ago – altohugh a few relict populations survived until around 4,000 years ago in isolated areas of Alaska, Siberia, and eastern Russia.
Around 3m tall at the shoulder (~10ft), these hairy proboscideans had very long curving tusks that were used for digging out vegetation from under snow and ice, scraping bark from trees, and for fighting.
The tusks showed a lot of variation in their curvature, and were often rather asymmetrical, a condition also seen in the closely related Columbian mammoth. Like modern elephants mammoths may have also favored using one side over the other for certain tasks, which over their lifetimes could result in uneven wear exaggerating the natural asymmetry even more.
It was smaller than modern pronghorns, around 70cm tall at the shoulder (~2’4″), and males had long antler-like horns with three tines. Bizarrely, one of these horns was always at least twice the size of the other, with “left-horned” and “right-horned” individuals seeming to occur in equal numbers.
Meanwhile in China another Miocene ungulate known as Tsaidamotherium hedini also had strange headgear, with an enlarged right “horn” forming a helmet-like dome on top of its head. This species was featured here on the blog just year, so check out that post for more details about it.
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.
Toothed whales – the branch of cetaceans that includes modern dolphins, porpoises, beaked whales, and sperm whales – have surprisingly asymmetrical skulls, with some of the bones skewed to one side and just the left nostril forming their blowhole.
Some of the most obvious external manifestation of this lopsidedness can be seen in sperm whales, which have their blowhole at the front left side of their head, and in male narwhals, which usually have a single left-side tusk.
This sort of asymmetry first appeared in the skulls of early toothed whales around 30 million years ago. And since the highest amounts of wonkiness have gone on to develop in lineages that hunt in dark, cluttered, or murky waters, this suggests that the trait is somehow linked to the evolution of complex echolocation.
Some ancient members of the river dolphin lineage also had some additional unusual asymmetry, sometimes having slightly sideways-bending snouts.
Ensidelphis riveroi was one of the weirdest of these, living around the coasts of what is now Peru during the Miocene, about 19 million years ago. Around 3m long (~10′), it had a very long narrow toothy snout that curved distinctly off to the right along its length.
Expectation vs reality
It’s not clear what the function of this bend was, or even if the only known skull actually represents the normal condition for this species. But Ensidelphis’ bendy snoot might have been used to probe around in muddy seafloor sediment or to extract prey from crevices, possibly like an underwater version of the modern wrybill.
Modern flatfish are characterized by their highly asymmetrical skulls, with both eyes positioned on just one side of their head. They aren’t actually born this way, but instead they undergo “eye migration” as juveniles, twisting up their skulls to bring one eye across the top of the head.
Progressive eye migration in a developing Summer Flounder, Paralicthys dentatus. From Helfman et al (2009). The diversity of fishes. 2nd ed., Wiley-Blackwell.
This bizarre arrangement is the result of flatfish adapting to life laying flat on the seafloor, but instead of slowly widening and flattening themselves out they took an evolutionary “shortcut” by simply tipping their tall narrow bodies over onto one side. Initially this would have left one of their eyes unusable, but random mutations causing slightly asymmetrical skulls would have rapidly become highly advantageous to the earliest members of this lineage – and over time they just got wonkier and wonkier.
We’ve even found fossils of early flatfish in the “halfway there” stage of their evolution!
Amphistium paradoxum lived in what is now northern Italy during the Eocene, around 50-48 million years ago. About 20cm long (~8″), it had one eye partially migrated towards the top of its head, but not all the way around yet, showing a transitional state between its bilaterally symmetric ancestors and its more twisted-skulled modern relatives.
Unlike most modern flatfish Amphistium came in both “right-eyed” and “left-eyed” forms in equal numbers, suggesting that a genetic preference for a specific side also hadn’t developed yet.
Styracosaurus albertensis was a ceratopsid dinosaur living during the late Cretaceous about 75 million years ago, in what is now Alberta, Canada. Around 5m long (~16′), it was one of the most elaborately ornamented horned dinosaurs, with a long nose horn and multiple elongated spikes on its frill.
There was actually quite a lot of variation in the frills of Styracosaurus, with varying numbers of long spikes and extra hook-like projections present on some individuals. But one recently-discovered specimen nicknamed “Hannah” is especially surprising – it had a noticeable amount of asymmetry in its skull. The left and right sides show different numbers and arrangements of spikes, so much so that if the two halves had been discovered separately they might have been identified as belonging to two completely different species.
Frill arrangements are often used to define different ceratopsids, so if this level of individual variation and asymmetry existed in other species, too, then we may need to reevaluate some of them.
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.
This repeated “pristification” suggests that saws are just incredibly useful and relatively “easy to evolve” structures for these types of fish, being both highly sensitive to bioelectric fields and able to physically slash and stab to kill prey.
Onchopristis numida was a sawskate known from what is now Northern and Western Africa during the mid-Cretaceous, about 95 million years ago. Up to about 3m long (~10′), it lived in both saltwater and freshwater, and was probably a bottom-dwelling ambush predator similar to modern angelsharks.
Whenever a denticle was lost from its saw, a larger one would grow to replace it, and over the life of an Onchopristis this resulted in an increasingly extreme amount of saw asymmetry.
Modern pristified fish also have rather asymmetrical saws. Sawfish are commonly born with a different number of denticles on each side, while sawsharks add extra denticles of varying sizes as they age, with the ongoing replacement of lost denticles resulting in more uneven arrangements over their course of their lives.
It’s not clear if the asymmetry gives any sort of advantage to these fish – but if nothing else it probably doesn’t cause them any disadvantage, so there’s no evolutionary pressure to stay more symmetrical.