Around 11 million years ago, during the late Miocene, much of what is now northern Honshu in Japan was submerged under fairly deep ocean waters. This offshore environment was inhabited by a variety of ancient sea-going tetrapods such as turtles, desmostylians, seal-like allodesmines, archaic baleen whales, and early oceanic dolphins… and also one very unexpected bird.
Meet the flightless marine swan.
Annakacygna hajimei, also known as the Annaka short-winged swan, was the same size as a modern black swan at about 1.2m long (~4′), but had a combination of features unlike any of of its living close relatives. Its head was proportionally large, and it had a long spoon-shaped bill like a shoveler duck, lined with comb-like structures for filter-feeding on plankton. It also had widened hips that would have helped keep it stable floating in rough waters, its tail was highly mobile and muscular, and its feet resembled those of diving birds like loons.
With thickened heavy bones and shortened forearms it was clearly completely unable to fly, but its reduced wings appear to have been highly specialized rather than just vestigial. Its shoulders were extra flexible while its wrists had a more limited range of motion, allowing it to fold its wings into a distinctive half-raised position similar to modern mute swans.
It probably used its wings and tail to perform elaborate “busking” visual displays, and also to carry and protect its young on its back while out at sea – basically making itself into a living swan boat.
And Falcatakely forsterae here had a very unusual combination of these features.
Living in Madagascar during the Late Cretaceous, about 70-66 million years ago, it was around 40cm long (1’4″) and was part of a diverse lineage of Mesozoic birds known as enantiornitheans. These birds had claws on their wings and usually had toothy snouts instead of beaks, and many species also had ribbon-like display feathers on their tails instead of lift-generating fans.
Falcatakely had a long tall snout very similar in shape to a modern toucan, unlike any other known Mesozoic bird, with the surface texture of the bones indicating it was also covered by a keratinous beak. But despite this very “modern” face shape the bone arrangement was still much more similar to other enantiornitheans – there was a huge toothless maxilla making up the majority of the beak, with a small tooth-bearing premaxilla at the tip.
This suggests that there was more than one potential way for early birds to evolve modern-style beaks, and there may have been much more diversity in these animals’ facial structures than previously thought.
Archaeopteryx lithographica was first discovered in the 1860s, still in the early days of our understanding of dinosaurs, and was a timely example of the sort of transitional form first proposed by Charles Darwin only a couple of years earlier. For over a century it was a famous icon of evolution, and has been part of a lot of weird drama over the years – it’s been central to arguments about bird origins, was accused of being a fake, and one specimen even vanished under mysterious circumstances.
At the time of its discovery Archaeopteryx was actually fairly quickly accepted as demonstrating an evolutionary link between dinosaurs and birds… but sadly this view wasn’t to last.
In the early 20th century opinion shifted towards birds not being dinosaurs but instead descended from “thecodont” reptiles (what we’d now call early archosaurs and pseudosuchians). And so for a long time Archaeopteryx ended up being depicted as simply the “first bird”, a half-reptile half-avian curiosity.
Reconstructions of it from this time period varied from very good to kind of awkward depending on how much the artist was trying to emphasize its reptilian ancestry, commonly featuring wonky-fingered wings and a scaly lizard-like face. It was also frequently depicted with bright gaudy parrot-like coloration, with a specific yellow-and-blue color scheme becoming a “paleoart meme” so prolific that it would eventually inspire the design of a Pokémon.
After decades of stagnation the dinosaur-bird link was resurrected in the early 1970s, with the discovery of the bird-like Deinonychus kicking off the Dinosaur Renaissance. Along with the explosion of spectacularly feathered dinosaur fossils from China in the mid-1990s, Archaeopteryx finally began to be properly presented as a feathered dinosaur again.
Continued study of the known Archaeopteryx specimens in the last couple of decades has vastly improved our knowledge of what this animal would have looked like, revealing previously unknown features like the exact plumage arrangement on its wings and legs, and even potentially some details about its coloration.
Living in southern Germany during the Late Jurassic, about 150-148 million years ago, Archaeopteryx inhabited what was then an island archipelago in a shallow tropical sea. It grew to around 50cm long (~1’8″) and was almost entirely covered with pennaceous feathers, externally probably just looking like a long-tailed bird.
It had broad wings, with asymmetrical flight feathers similar to those of modern birds but with more extensive coverts, some of which were probably a matte black color. Its legs also sported long “feather trousers” and a “raptor“-like hyperextensible second toe, and there was a slight forked shape to the tip of its tail.
Arguments have gone back and forth about how well it was actually able to fly, with current thinking being that it made short bursts of active flapping flight a little like a modern pheasant – but since its shoulder joints were less mobile than those of modern birds it must have used a different sort of flight stroke to generate lift.
It’s no longer always considered to have been the “first bird”, or even to have been the direct ancestor of any modern birds. Instead it represents an offshoot lineage of early birds (or very-bird-like dinosaurs) that was just one part of a still-expanding flock of feathery fossil discoveries.
…Or did it?
But I’m not talking about the dubious claims of non-avian dinosaur fossils found in places they shouldn’t be. This is about something else entirely: an unassuming little bird known as Qinornis paleocenica.
Living in Northwest China during the mid-Paleocene, about 61 million years ago, Qinornis was roughly pigeon-sized at around 30cm long (12″). It’s known only from a few bones from its legs and feet, but those bones are unusual enough to hint that it might have been something very special.
Uniquely for a Cenozoic bird, some of its foot bones weren’t fully fused together. This sort of incomplete fusion is seen in both juvenile modern birds and in adults of non-avian ornithurine birds from the Cretaceous – and the Qinornis specimen seems to have come from an adult animal.
If it was fully grown with unfused feet, then that would suggest it was actually part of a “relic” lineage living 5 million years after the mass extinction, surviving for quite some time longer than previously thought.
The last known non-avian dinosaur.
Brontornis burmeisteri was one of the largest flightless birds known to have ever existed, standing around 2.8m tall (9’2″) and estimated to have weighed 400kg (~880lbs).
Known from the early and mid-Miocene of Argentina, between about 17 and 11 million years ago, it’s traditionally considered to be one of the carnivorous terror birds that dominated predatory roles in South American ecosystems during the long Cenozoic isolation of the continent.
But Brontornis might not actually have been a terror bird at all – it may have instead been a giant cousin of ducks and geese.
The known fossil material is fragmentary enough that it’s still hard to tell for certain, but there’s some evidence that links it to the gastornithiformes, a group of huge herbivorous birds related to modern waterfowl.
If it was a gastornithiform, that would mean it represents a previously completely unknown lineage of South American giant flightless galloanserans. And, along with the gastornithids and the mihirungs, it would represent a third time that group of birds convergently evolved this sort of body plan and ecological role on entirely different continents during the Cenozoic.
The enantiornitheans (“opposite-birds”) were the most diverse and widespread group of Mesozoic birds, existing all around the world throughout the Cretaceous period. They retained claws on their wings and had toothy snouts instead of beaks, and while most of them lacked the lift-generating tail fans of modern birds they appear to have still been very adept fliers.
But Elsornis keni here was doing something different.
Known from the Late Cretaceous of Mongolia, about 80 million years ago, this opposite-bird lived alongside famous dinosaurs like Velociraptor and Protoceratops in what is now the Gobi Desert. Only a single partial specimen has ever been found, so its full life appearance is unknown and this reconstruction is somewhat speculative, but it would have been around the size of a pigeon at 25cm long (10″) – not including any decorative tail feathers it may have had, similar to other enantiornitheans.
It wing and shoulder bones were very odd for an opposite-bird, with proportions that don’t match anything capable of competent flight. Instead Elsornis appears to have been a flightless enantiornithean, a representative of a previously unknown terrestrial lineage.
It’s another PBS Eons commission roundup day!
Seriemas are part of a lineage known as cariamiformes, highly terrestrial birds that were widespread across most of the world but are today represented today by only two living species in South America. During the Cenozoic this group repeatedly evolved into large predatory flightless forms like the the phorusrhacids and bathornithids, and were probably the closest avians ever got to recreating the “carnivorous theropod” body plan and ecological niche.
And yet none of them ever seem to have experimented with more dromaeosaurid-like claws.
…With one known exception.
Qianshanornis rapax here lived in East China during the mid-Paleocene, about 63 million years ago. It was a small cariamiform, probably around 30cm tall (1″), and is only known from fragmentary fossil material – but part of those fragments was a fairly well-preserved foot. And the bones of its second toe were unlike any other known Cenozoic bird, shaped incredibly similarly to those of dromaeosaurids and suggesting it may have had the same sort of big hyperextendible “sickle claw”.
While it had sturdy legs and short wings, and probably spent a lot of time walking on the ground like other cariamiformes, it was probably also still a fairly strong flier based on the known anatomy of its arms and shoulders.
Unfortunately, though, its head and claws were entirely missing, so without more fossil discoveries it’s hard to say anything definite about its ecology. I’ve restored it here based on other predatory cariamiformes, but since it was also closely related to a herbivorous species it’s not clear whether Qianshanornis was truly a dromaeosaur-mimic or if something else was going on with that unique second toe.