During the mid-Pleistocene, between about 900,000 and 500,000 years ago, the Mediterranean islands of Malta and Sicily were connected and shared a unique ecosystem made of up a mix of weird endemic species. While the tiny elephants and giant swans are probably the most famous, there were also several other unusual animals such as dwarf hippos, huge owls, large cranes, giant tortoises, and big lizards.
And also massive rodents.
Leithia melitensis, the Maltese giant dormouse, was descended from garden dormice, but thanks to the lack of large land predators on Siculo-Malta it was able to evolve a much much larger body size – about 60-70cm long (2′-2’4″), almost the size of a cat.
Recent reconstructions of its skull have shown it was also proportioned differently compared to its tiny modern relatives, more chunkily built with a shorter and wider snout, bigger teeth, and thicker cheekbones that must have anchored some incredibly powerful muscles for chewing. It may have been eating a much more herbivorous diet than other dormice, processing a lot of tough fibrous vegetation.
The Mediterranean island of Crete had very few predators during the Pleistocene, with most being birds of prey. And with the terrestrial carnivore niches in the ecosystem left vacant, it was a semi-aquatic mammal and an owl that ended up taking advantage of that opportunity.
Neither were large enough to threaten the dwarf elephants and hippos, and don’t even seem to have habitually eaten even the smallest of the miniature giant deer. Instead these Cretan predators focused much more on the smaller land vertebrates on the island, preying on birds, shrews, rodents, amphibians, and reptiles.
Lutrogale cretensis (previously known as Isolalutra cretensis) was a close relative of the modern smooth-coated otter. It was about the same size as its living cousin, around 1m long (3’3″), but had stronger jaws and chunkier limbs.
Its skeleton shows features associated with walking and running more than swimming, and it seems that this was something of a “land otter” — still able to swim, but spending most of its time on land similar to the modern small-clawed otter.
Shellfish were likely still the main part of its diet, indicated by its crushing teeth. But it probably also regularly ate whatever small terrestrial vertebrates it could catch, since more aquatic otters are already known to prey on those types on animals when they can.
Athene cretensis was yet another weird island owl, but this time not a descendant of a Strix or Tyto species. Instead this owl was descended from the Eurasian little owl — except it had become much much larger.
It stood around 60cm tall (2′), over three times bigger than its living relative. Its legs weren’t quite as long as those of the modern burrowing owl, but they were still proportionally much longer than those of little owls and show adaptations for terrestrial movement. Little owls already sometimes chase down prey on foot, and Athene cretensis was probably even more of a ground-based hunter, convergently similar to the Hawaiian stilt-owls and the Cuban terror owls.
Preserved pellets show that it ate small mammals and birds, mainly large mice.
Its wings were still quite large, and it was probably also a good flier — and may even have spread over to some of the Dodecanese islands to the east of Crete, since a wing bone closely resembling that of Athene cretensis has been found on Armathia.
Both of these predators seem to have disappeared around the end of the Pleistocene, at the same time as many of the other native Cretan species about 21,500 years ago. Much like the situation with Candiacervus, this may have been a result of a combination of a rapidly shifting climate and the presence of humans disrupting the already fragile island ecosystem.
The island of Sicily was isolated about 5.3 million years ago when the Mediterranean rapidly refilled. During the next few million years changes in sea level and tectonic uplift allowed repeated colonizations by mainlaind species via the sea strait separating Sicily from Italy, and opened up occasional connections with nearby Malta, resulting in a series of different ecosystems over time.
During the mid-Pleistocene, between about 900,000 and 500,000 years ago, a lack of large land predators on Sicily and Malta allowed a weird mix of endemic species to evolve. Most famous are the tiniest elephants (Palaeoloxodon falconeri), but there were also a couple of giant owls, a small long-legged owl, a giant crane, a big lizard, a giant tortoise, an otter, and giant dormice.
And then there were the swans.
Cygnus falconeri was enormous, at least a third larger than the biggest living swans, at least 1.5m tall (4’11”) — taller than the native elephants, although not nearly as heavy. Its wings were large, with a span of around 3m (9’10”), but at such a hefty size it would have been either a very poor and reluctant flier or functionally flightless.
Its legs were better adapted for walking around on land than for swimming, with shorter toes and possibly reduced webbing. It would have been one of the biggest terrestrial herbivores on Siculo-Malta, probably mainly a grazer but also capable of reaching much higher vegetation than the elephants or tortoises.
It lived alongside another unique swan species, the goose-like dwarf swan Cygnus equitum. Both the giant and dwarf swans probably evolved from the same whooper swan-like ancestor species, but each resulted from separate colonization events — otherwise interbreeding would have probably prevented them from developing such a huge difference in size.
Or an alternative scale comparison to highlight the utter ridiculousness of this island:
Between 500,000 and 200,000 years ago multiple sea level fluctuations allowed new species to colonize Siculo-Malta from the mainland, including various large mammalian herbivores and carnivores. With new competition and predators, Cygnus falconeri probably disappeared around the same time as the tiny elephants and most of the other mid-Pleistocene endemic animals.
The dwarf swan, smaller and still a strong flier, may have survived the altered ecosystem for a bit longer, but would have gone extinct during the rapid climate changes at the start of the last glacial period 115,000 years ago.
Megalenhydris barbaricina was the largest of the Sardinian-Corsican otters, reaching lengths of over 2m (6’6″), slightly bigger than the modern South American giant river otter. Its crushing teeth indicate it specialized in crunching through hard-shelled invertebrates like molluscs and crustaceans, and its highly flexible backbone and flattened tail suggest it was a strong swimmer that may have been even more aquatic than most other otters.
It was possibly an equivalent of the modern sea otter, spending most of its time in the water, although it’s not clear whether it was a river or marine species.
Dating on the one known partial skeleton of Megalenhydris is uncertain, but it may be late Pleistocene to early Holocene in age, between about about 70,000 and 10,000 years old. Unfortunately this puts it within the same age range as the arrival of humans on Sardinia and Corsica, and its extinction may have been a direct result of being hunted for its meat and pelt.
5.3 million years ago the Strait of Gibraltar formed, re-establishing the connection with the Atlantic, and the Mediterranean refilled incredibly rapidly, possibly in as little as two years. As a result, various species that had colonized across the dried-out Mediterranean from the continental mainland were left stranded out on islands that had been re-formed throughout the sea.
And on what is now Menorca a population of rabbits found themselves isolated, with little competition and no large terrestrial predators.
Nuralagus rex was the evolutionary result, an enormous rabbit 50cm tall at the shoulder (1’8″). It was heavily built with a stiff spine — making it unable to hop — and had weaker senses than its ancestors, with small eyes and stubby ears. It would have been a slow-moving animal ambling around the scrublands of Menorca, digging for its main foods of roots and tubers.
It’s unclear what happened to this big bunny, but it seems to have disappeared around the end of the Pliocene, about 3-2.5 million years ago. Possibly the onset of global cooling at the beginning of the Pleistocene ice ages changed the climate too quickly for it to adapt to, or the dropping sea levels that connected Menorca with nearby Mallorca introduced new competition from the other island that Nuralagus couldn’t cope with.
And one of the animals that spread into Menorca from Mallorca was Myotragus.
About the same size as the giant rabbits, at 50cm tall (1’8″), Myotragus balearicus was a close relative of modern sheep that had undergone dwarfing on Mallorca since its ancestors’ isolation 5.3 million years ago.
It was possibly one of the most unusual mammals ever, with a combination of features not seen anywhere else. Its snout was relatively short and rabbit-like, with ever-growing front teeth in its lower jaw, and its eyes faced directly forward, giving it stereoscopic vision more like a primate than a herbivore. Its brain and sense organs were highly reduced compared to its ancestors, its legs were shorter and stockier, and its feet had lost much of their flexibility, making it unable to run or jump.
But strangest of all was its metabolism, as indicated by growth lines in its bones. It was essentially cold-blooded, functioning more like a reptile than a mammal, growing at a slow rate that varied or even stopped entirely depending on the conditions of its environment. It would have taken around 12 years for it to reach maturity, an incredibly long time for its size, and so it would have reproduced very very slowly — but this metabolic strategy also allowed it to conserve a lot of energy and survive long periods of scarce food availability.
Myotragus‘ extreme weirdness obviously worked to its advantage, because it was quite common on Mallorca-Menorca and unlike many of the other native species it survived through the much colder drier conditions of the Pleistocene.
It was still around in the Holocene when humans reached the islands in about 3000 BCE, and this slow-moving sluggish goat-like animal with a rather uncanny face must have been a very strange sight to them.
There was a hypothesis that the early settlers actually attempted to domesticate Myotragus, based on remains found in caves with what appeared to be trimmed horns. But in recent years this has been disputed, since some of the “trimmed” horns pre-date human arrival and may be better explained as the result of Myotragus individuals chewing on the bones of others for the mineral content.
Shortly after the arrival of humans Myotragus and the other remaining endemic mammals on the islands (a giant dormouse and a giant shrew) disappeared entirely, suggesting that a combination of hunting, competition from introduced livestock, and predation from dogs was finally too much for these strange sheep to handle.
The Saint Helena giant earwig (Labidura herculeana) was, as its name suggests, an enormous species of earwig. Growing as large as 8.4cm long (3.3″) it was the biggest of its kind in the world, and was completely flightless with no hind wings.
It lived in deep burrows in the arid plain and gumwood forest regions of the island, and only came out at night during the summer rains. Since it was probably descended from the shore earwig (Labidura riparia) it was likely a similar sort of opportunistic carnivore, eating smaller invertebrates and carrion.
Humans didn’t reach Saint Helena until the early 1500s, and it wasn’t until 1798 that the earwig was noticed by naturalists and given its scientific name. Then it was more or less forgotten about, and scientific interest in it didn’t start to resume until the 1960s.
But by then it was just very slightly too late.
Extensive habitat destruction and predation by invasive cats, rodents, and centipedes had taken a huge toll on the earwig, and it had become incredibly rare. The last sighting of a live individual was in 1967, and attempts to locate more for potential captive breeding programs in the 1980s and 1990s failed to find any at all.
Like much of the Caribbean, Puerto Rico originated as part of a Mesozoic volcanic island arc. While in theory this could have resulted in the region retaining some relict populations of weird Cretaceous species, there’s a major reason this didn’t actually happen: the asteroid impact event 66 million years ago. The proximity to Chicxulub would have been devastating to the proto-Caribbean at the time, with tsunamis as high as 500m (1640′) battering the islands and destroying practically everything that lived there at the time.
Caviomorph rodents arrived from South America around the start of the Oligocene, about 33 million years ago, and became very successful due to being some of the only land mammals present on many of the islands. The only modern survivors of this lineage are a handful of hutias, but in the past they were much more diverse — and some of them were giants.
Tainotherium valei lived in Puerto Rico during the Late Pleistocene and early Holocene, and while it wasn’t quite the largest of the giant hutias it was still enormous. It’s only known from a single partial leg bone, so its full size is difficult to estimate, but it was probably somewhere around 80cm tall at the shoulder (2’7″) and weighed about 100kg (220lbs) – similar in size to a modern black bear.
And that leg bone is especially unusual, showing anatomical adaptions associated with tree-climbing. This is strange for such a huge heavy mammal, but it may have been an ecological equivalent of something like the giant Malagasy lemurs, a slow-moving animal that climbed up trees to feed and regularly traveled over the ground from one feeding site to another.
During the Pleistocene lower sea levels meant most of the Puerto Rican archipelago was part of a much larger landmass that also included most of the nearby Virgin Islands, and the predominate climate at the time was a dry savanna. As sea levels rose at the start of the Holocene the climate shifted wetter and Puerto Rico became densely forested. This would have been advantageous for a tree-climbing animal like Tainotherium, so it was likely still around when the first humans arrived in around 3000-2000 BCE.
Unfortunately these early settlers appear to have burned large amounts of the forests for the first few centuries of their presence, and such widespread habitat destruction would have driven this giant rodent extinct very quickly.
Like many other isolated islands ancient Cuba lacked any large land predators, allowing some birds to exploit more terrestrial lifestyles.
The Cuban flightless crane (Grus cubensis, or possibly Antigone cubensis) lived during the Late Pleistocene and Holocene. It was probably a descendant of the sandhill crane — and although an endemic variety of sandhill crane still exists in Cuba today, the two aren’t directly related to each other and instead are the result of two different colonization events.
It was about the same size as modern sandhill cranes, around 60cm tall at the back (2′) with a full height of about 1m (3’3″), but it was much more heavily built. It had stockier legs and a thicker beak, suggesting it may have been specialized for a different ecological niche than its ancestors, and its wings were reduced enough that it was probably completely flightless.
And, once again, there was also a weird owl on this island.
But it was almost twice the size of its Hawaiian cousins, measuring about 1.1m tall (3’7″) and potentially being the largest owl to ever exist. Its remains were so big, in fact, that they were initially mistaken for those of a terror bird.
It was powerfully built and was probably a good runner, mainly preying on giant rodents and dwarf ground sloths. While its wings and flight muscles were reduced it might not have been entirely flightless, and it may have been still been capable of turkey-like short bursts of flight.
Three other species of Ornimegalonyx also stalked ancient Cuba at the same time, varying slightly in size from each other and probably each specializing in a different size class of prey.
Remains of both of these birds have been found in natural petroleum seeps on the northern coast of Cuba that date to as recently as about 6000 years ago, around the same time that humans first arrived. After that point they probably both went extinct very quickly — the flightless cranes were probably actively hunted and eaten into extinction, and the terror owls would have disappeared as their prey species dwindled away due to the same hunting pressures.
It probably evolved in Cuba, and colonized the Bahamas shortly after the hutias did, sometime in the last 400,000 years during a glacial period when a particularly low seal level meant the islands were only about 30km apart.
It was the main nocturnal predator in the Bahamas, and much like its older Italian relative Tyto gigantea it also had a giant hawk counterpart in the daytime: the huge Titanohierax.
Although many popular online sources refer to Tyto pollens as being flightless, it actually had large robust wings and could probably fly quite well. This might be due to some confusion between it and a completely different giant Caribbean owl, Ornimegalonyx.
The Lucayan people probably reached Andros Island sometime around 1000 CE, and coexisted with the giant owls for several centuries. It was only after European arrival in the 1500s and the felling of their forest habitat that they seem to have vanished.
Local legends of an owl-like creature called the chickcharney may have been inspired by historical encounters with Tyto pollens, and suggest that they were aggressively territorial.
The complete lack of land mammals on the Hawaiian islands left the terrestrial predator niches available to birds — and it was owls who ended up taking advantage of that role in the ecosystem.
The Grallistrix owls were found on Kauaʻi, Oʻahu, Molokaʻi, and Maui, with each island having its own endemic species. They were a type of true owl, probably descended from a species of Strix, and developed especially long legs that led to their nickname of “stilt-owls”.
The Molokaʻi stilt-owl (Grallistrix geleches) was the largest of the four known species, about 60cm tall (2′). Although it had proportionally short wings it was still capable of flying, and probably specialized in stalking and ambushing smaller birds like Hawaiian honeycreepers in the dense forests.
(These owls were also the inspiration for the pokémon Decidueye!)
Meanwhile the forest floor insectivore niche was occupied by Talpanas on Kauaʻi, but on the other main islands a different type of bird took up the same role.
The Moloka’i flightless ibis (Apteribis glenos) was a typical example of the genus, about 50cm long (1’8″). It was flightless, with reduced wings, and had unusually short stocky legs that gave it proportions much more like a kiwi than an ibis. And it probably lived very much like a kiwi, too, probing around in the forest litter with its beak searching for snails and other invertebrates.
We even have an idea of the coloration of these birds, thanks to subfossil remains with preserved feathers. They seem to have been brown and beige, similar to the juvenile plumage of their close relatives.
The arrival of humans at least 1000 years ago would have unfortunately been devastating to both the stilt-owls and the flightless ibises. The combination of hunting, habitat destruction, and invasive dogs, pigs, and rats attacking them and their ground-based nests probably drove them all into extinction very quickly.