avialae – Nix Illustration

Avisaurus

Avisaurus darwini here lived at the very end of the Cretaceous, about 66 million years ago, in what is now the Hell Creek fossil beds in Montana, USA.

It was a member of a diverse group of Mesozoic birds known as enantiornitheans, which retained claws on their wings and often still had toothed snouts instead of beaks – and being part of the avisaurid family it was also one of the larger known examples of these birds, similar in size to a modern hawk at around 60cm long (~2′).

Although this species is only known from isolated foot bones, the remains have distinct enough anatomical features to show that Avisaurus had powerful gripping talons similar to those of modern hawks and owls, suggesting it had a similar lifestyle hunting small vertebrate prey in the ancient swampy Hell Creek ecosystem.

Patagopteryx

While birds are one of the few animal groups to have achieved powered flight, they’re also very prone to losing their aerial abilities. Many times over their evolutionary history, multiple different bird lineages have convergently become secondarily flightless – and Patagopteryx deferrariisi was one of the earliest known examples of this.

Living during the Late Cretaceous, about 86-84 million years ago, in what is now the northern part of Argentine Patagonia in South America, Patagopteryx was roughly the size of a modern chicken at around 50cm long.

When it was first discovered it was classified as a ratite, but soon after it was recognized as actually being a much earlier type of bird, an early ornithuromorph only distantly related to any modern groups.

It had small wings, little-to-no keel, and no wishbone, indicating it lacked the large powerful musculature required for flight. Its legs were quite long, with large feet with all four toes facing forward – proportions that suggest it was built more for walking than for high-speed running.

Growth rings in its bones also show that it had a much slower growth rate than modern birds, taking several years to reach adult size.

Falcatakely

Modern birds’ upper beaks are made up mostly from skull bones called the premaxilla, but the snouts of their earlier non-avian dinosaur ancestors were instead formed by large maxilla bones.

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.

Retro vs Modern #04: Archaeopteryx lithographica

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.

1860s-1970s

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 20^th 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.

2020s

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.

Qinornis

66 million years ago, the end-Cretaceous mass extinction wiped out all dinosaurs except for the avian bird lineage.

…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.

Elsornis

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.

Island Weirdness #08 – Balaur bondoc

When Balaur was described in 2010 it was initially thought to be a dromaesaurid closely related to Asian forms like Velociraptor. With its particularly stocky legs built for strength rather than speed, two-fingered hands, and two large sickle claws on each foot, it was interpreted as a weird highly specialized predator terrorizing the other Hațeg Island species at the end of the Cretaceous. Although only 1.8m long (5’10”), it was hypothesized to have taken down prey much larger than itself with powerful slashing kicks.

But later analyses cast doubt on this interpretation.

A lot of the anatomical features of Balaur’s skeleton were odd for a dromaeosaurid, but matched those of avialans – a group of close evolutionary “cousins” to the dromaeosaurids, containing Archaeopteryx and the common ancestors of all modern birds. And, by 2015, multiple studies had confirmed Balaur wasn’t really a “raptor” but instead a little further along on the bird lineage.

So now our picture of this dinosaur is very different: a chunky-bodied island bird, grown large and secondarily flightless sort of like a Cretaceous equivalent to the dodo. Its double sickle claws were probably adaptations for climbing and perching in trees, and based on similar avialans it was likely a herbivore rather than a hypercarnivore.

Island Weirdness #04 – Gargantuavis philoinos

By far the biggest island in the Late Cretaceous European archipelago, the Ibero-Armorican island (sometimes also known as the Ibero-Occitan island) was made up of most of the Iberian Peninsula and France and was larger than modern-day Madagascar.

Around 73-71 million years ago one of the residents of this island was the aptly-named Gargantuavis – the largest known Mesozoic bird, and probably an example of island gigantism.

Although only known from a few isolated bones, it’s estimated to have been slightly larger than a modern cassowary, somewhere in the region of 2m tall (6′6″). At that size it would have also been secondarily flightless, which is surprising for a bird that was living alongside larger fast-moving theropods like abelisaurs.

Not much else is known about it due to the scarce remains, but it seems to have had a long slender neck and probably had a small head. Its hips were fairly broad, suggesting it wasn’t capable of running very fast, and it was likely a slow-moving herbivore that was a fairly rare member of its ecosystem.

Exactly where it belongs in the bird evolutionary tree is also unclear, with the best current guess being “some sort of euornithean”.

Enantiophoenix

Enantiophoenix electrophyla, an enantiornithean bird from the Late Cretaceous of Lebanon (~95 mya).

It was similar in size to a modern starling, around 20cm (8″) long, and although only known from a fragmentary fossil it had fairly chunky leg bones with large claws. It was probably a strong percher like most other avisaurid enantiornitheans.

Several tiny pieces of amber were also found within the fossil, which have been suggested to be stomach contents. This could perhaps be evidence of Enantiophoenix feeding on tree sap like modern sapsuckers, but without a known skull it’s hard to tell for certain whether it was specialized for that sort of diet or not.

Longipteryx

Longipteryx chaoyangensis, an enantiornithine from the Early Cretaceous of China, about 120 million years ago. With a body length of only around 15cm (6″), it had a long snout tipped with a few hooked teeth and feet capable of perching – features that indicate it may have lived very similarly to modern kingfishers, feeding on fish and small invertebrates in its swampy forest habitat.

The enantiornithines were a sort of “cousin” lineage to modern birds. Most had toothy jaws and clawed wings, and the wide variety in their skull shapes suggests that they were specialized for many different dietary niches. The entire group went extinct during the K-Pg mass extinction and left no living descendants, but during the Cretaceous they were the most widespread and diverse group of birds*, with fossils currently known from every continent except Antarctica.

* Depending on how you define “bird”.