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.
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.
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.
Uniquely they completely lost their wings, not even having the tiny vestigial bones seen in other large flightless birds.
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.
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.
Despite being located so close to mainland Africa, these enormous flightless ratites weren’t the closest relatives of ostriches as might be expected. Instead their closest living relatives are the kiwis of New Zealand, and they must have descended from flying ancestors that reached Madagascar across the Indian Ocean sometime during the early-to-mid Cenozoic.
Aepyornis maximus was one of the biggest of these big birds, standing around 3m tall (9′10″) and weighing over 500kg (1,100 lbs). Its eggs were equally massive, up to 34cm long (1’1″) and with a circumference of over 1m (3′3), making them the largest known eggs laid by any vertebrate.
Recent studies of the shape of its brain within its skull show that it had a good sense of smell but very poor eyesight – possibly being near-blind – suggesting that much like its kiwi relatives it was highly specialized for a nocturnal lifestyle.
There were several other species of elephant bird throughout Madagascar, and at least some of them appear to have successfully survived alongside humans for quite some time. Carbon dating of eggshells suggests they were still alive around 1000 years ago, and based on historical mentions they may have persisted as late as the 1600s before finally disappearing.
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”.
Known from around the North Pacific rim from about 33-15 million years ago, plotopterids were flightless diving birds which used their small but powerful wings to propel themselves through the water. They were convergently similar to penguins in body shape and lifestyle, but not actually closely related to them – instead being relatives of gannets, cormorants, and anhingas.
Smaller plotopterids were about the size of modern cormorants, around 70cm long (2′4″), but the larger known genera like Copepteryx rivalled the southern giant penguins at around 1.8m (6′).
And a second species of Copepteryx known only from a single leg bone (Copepteryx titan) may have been ever bigger. Estimated at over 2m in length (6′6″), it was possibly one of the largest diving birds to have ever lived.