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 ArgentinePatagonia 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.
The mancallines were a lineage of flightless semi-aquatic birds closely related to auks. Known from the Pacific coasts of what are now California and Mexico, between about 7.5 and 0.5 million years ago, they convergently evolved a close resemblance and similar lifestyle to both the recently-extinct North Atlantic great auk and the southern penguins.
Miomancalla howardi here lived in offshore waters around southern California during the late Miocene (~7-5 million years ago). The largest of the mancallines, it just slightly beat out the great auk in size – standing around 90cm tall (~3′) and weighing an estimated 5kg (11lbs).
Like great auks and penguins it would have been a specialized wing-propelled diver, swimming using “underwater flight” to feed on small bait fish. It probably spent much of its life out at sea, probably only returning to land to molt and breed.
September might have ended, but guess what? I am not remotely done with this yet so we’re continuing on.
Now it’s Spectober.
(Also just a reminder: I am not currently taking new requests. I’ve got far too many existing ones that I’m still working through!)
Someone who identified themself only as “Adam” asked for “eagles evolving into terrestrial predators to pursue larger prey”:
A flightless eagle occupying an apex predator niche in the same island chain as the giant herbivorous tegu, Terraetus adamii is descended from a species similar to the modern harpy eagle. It isn’t substantially larger than the biggest modern eagles – standing about 1.2m tall (~4′) – but it’s certainly much more massive, weighing around 25kg (~55lbs).
In the absence of other large terrestrial predators its ancestors originally took up a caracara-like lifestyle, preferring hunting on foot over flying, before gradually becoming totally flightless and converging on terror birds with large heavy skulls, reduced wings, and powerful legs.
Terraetus’ head is only very sparsely feathered, an adaptation for feeding inside the carcasses of large prey, which it dispatches using a combination of kicking and blows from its large hooked beak. It’s usually a solitary hunter that can tackle prey up to two or three times its own weight – preferencing the juveniles of the herbivorous tegu – but during the breeding season pairs will occasionally hunt cooperatively to take down larger targets.
Despite possessing sharp beaks and talons, these weapons aren’t actually used in fights between individuals of this species. Instead they bodily shove each other back and forth, battering at each other with large bony knobs that grow from the hand bones of each wing.
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.
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).
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.
Flightlessness is very rare in the long-legged fly family (Dolichopodidae), however, with only about 12 out of over 5000 species known to have lost functional wings — and eight of those are endemic to the Hawaiian islands.
The Koʻolau spurwing long-legged fly (Emperoptera mirabilis, sometimes classified as Campsicnemus mirabilis) was found only on Mount Tantalus in the southern Koʻolau Range of Oʻahu, close to Honolulu. About 2mm long (>0.1″), its wings were reduced to thin stiff spines, and it moved around by walking and hopping in leaf litter in the moist cool forest at elevations of about 300m (~1000ft).
Like most other long-legged flies it would have been predatory, hunting other tiny invertebrates.
The Koʻolau spurwing was actually still common on Tantalus as recently as the early 1900s, but multiple searches since the 1980s have failed to find any more of them at all. The species is most likely completely extinct, probably due to a combination of predation from invasive ants and habitat destruction from feral wild boar rooting up the forest floor.
Of the other flightless Hawaiian long-legged flies several other species are now possibly extinct — only one out of the five known Emperoptera species still definitely survives on the highest slopes of Mount Kaʻala, and one of the three Campsicnemus is either very rare or also extinct.
The Hawaiian islands also have three endemic species of flightless crane fly in the genus Dicranomyia, all of which are incredibly rare.
With the lack of large terrestrial mammals in New Zealand, birds were free to exploit the “big herbivore” niches in the ecosystem — and the giant moa were the ultimate result.
Closely related to modern South American tinamous, the ancestors of moa were small flying birds that arrived in New Zealand sometime in the early Cenozoic. At the time of the MioceneSt Bathans fossil deposits they were already large and flightless, and by the Holocene they had grown truly enormous.
Uniquely they completely lost their wings, not even having the tiny vestigial bones seen in other large flightless birds.
Dinornis robustus
The South Island giant moa (Dinornis robustus) was the largest of them all, with females standing almost 2m tall at the back (6’6″) and able to reach their heads up to heights of around 3.6m (11’10”). It had a long neck, a relatively tiny head with a curved beak, and large powerful legs — and preserved hair-like body feathers show that it was reddish-brown in color.
It also had some of the most extreme sexual dimorphism seen in any bird species, with the males being significantly smaller than the females at only about 1.2m tall (3’11”). This seems to have been the result of scaling-up smaller differences in body size from their ancestors.
Aptornis defossor
The adzebills were another odd group of big flightless birds whose ancestors also date back to sometime before the St Bathans deposits. They had large downward-curving beaks, short strong legs, and highly reduced wings that were smaller proportionally than those of the dodo.
They were less common than the moa, found only the drier forested parts of the lowlands, and based on isotope analysis of their bones they seem to have been predators hunting invertebrates, reptiles, and smaller birds.
When their remains were first discovered they were even thought to be a type of moa, but later studies (including recovered ancient DNA) have shown they were actually gruiformes, with their closest living relatives being either the South American trumpeters or the African flufftails.
The South Island adzebill (Aptornis defossor) was the slightly larger of the two species known from the Holocene, reaching sizes of about 0.8-1m tall (2’8″-3’3″).
The ancestors of the Māori people arrived in New Zealand around the year 1300, and sadly within about a century a combination of human hunting pressure and predation by introduced mammals sent both the moa and adzebills into total extinction.
New Zealand is probably one of the most famous modern examples of a unique island ecosystem, having been isolated for the last 80 million years. Lacking any living terrestrial mammals, birds instead became the dominant land animals, adapting to a wide range of niches and evolving unusual flightless forms like the modern kiwi and kākāpō.
The St Bathans fossil site on the South Island dates to the early Miocene, about 19 to 16 million years ago, just after a period known as the “Oligocene drowning” when large portions of the landmass were underwater. It gives us a glimpse of an ancient version of New Zealand when the ecosystem was recovering and rapidly diversifying in a then-subtropical climate, and has produced a wide range of new fossil species (including a mysterious mammal).
One of the most recent discoveries from the site is Heracles inexpectatus, a close relative of the modern New Zealand parrots — but significantly larger than any living species, estimated to have stood almost 1m tall (3’3″). Known from just a few leg bones, the fossils were so big that they initially weren’t even recognized as belonging to a parrot, instead being mistaken for a large eagle before their true nature was realized.
At such a size it would likely have been flightless, although it may have still been capable of climbing trees and gliding. Much like the modern kea it was probably an omnivore, using its large curved beak and powerful crunching bite to eat pretty much whatever it wanted.
By the late Miocene, around 13 million years ago, New Zealand’s climate rapidly shifted cooler and drier, and the tropical forests were quickly replaced with temperate ones. This changing habitat may have been too much too fast for the giant parrots to deal with, and they went extinct alongside many of the other St Bathans species.
Out of all the extinct island species we’ll be covering in this theme, there’s probably none more famous than the dodo (Raphus cucullatus) – a large flightless pigeon about 1m tall (3′3″).
The island of Mauritius was formed from a volcanic hotspot in the Indian Ocean about 10-7 million years ago, in a location roughly 1000km east of Madagascar (~620 miles). The dodo’s smaller flying ancestors must have arrived there sometime after that point via island-hopping from Southeast Asia – the area where its closest living relative the Nicobar pigeon is still found today – and finding themselves in an ecosystem completely lacking terrestrial mammals they quickly evolved to fill a large herbivore niche.
Although frequently depicted as blue-grey, the dodo’s actual life appearance is unknown for certain. No complete preserved specimens have survived into the present day, and contemporary accounts and drawings are somewhat inconsistent – but common elements among them suggest it was more of an earthy brown, with cream-colored primary feathers, yellow legs, a naked pale face, and a green-and-yellow streaked beak. The large white ostrich-like tail plumes shown in many images have also probably been highly exaggerated, since older images depict the dodo with only a tiny tufted tail at best.
Its appearance probably also varied based on the time of year, molting its feathers at the end of summer and being fattest during the breeding season in early spring.
And despite often being stereotyped as a slow dim-witted animal, the proportions of the dodo’s leg bones suggest it was actually quite fast and agile. Its brain-to-body size ratio was also typical compared to other pigeons – which are known to be highly intelligent birds – and it had a well-developed sense of smell.
Sadly this fascinating bird disappeared within only a century of being discovered by humans in the late 1500s. Its “island tameness” due to its lack of natural predators made it easy prey, its forest habitat was rapidly destroyed, and introduced mammal species (such as dogs, pigs, cats, rats, and macaques) competed for its food sources and ate its eggs and young – pressures that its population simply couldn’t hope to recover from all at one.
Its loss wasn’t even properly recognized until much later in the 1800s, but since then it has ironically become immortalized as a icon of human-caused extinction.
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