When owls find their way onto isolated islands lacking any terrestrial predators, they have a tendency to take up that role for themselves – evolving longer legs and shorter wings, and specializing more towards hunting on foot. From New Zealand to Hawaii to the Caribbean to the Mediterranean to Macaronesia, leggy island ground-owls have independently happened over and over again in the last few million years—
—And, unfortunately, they’ve all also become victims of the Holocene extinction, their fragile island ecosystems too easily disrupted by human activity and the arrival of invasive species.
Its wing proportions indicate it would have been a poor flyer, instead primarily hunting on foot in the dense laurisilva forests. Since there were no terrestrial mammals or reptiles on São Miguel at the time, its diet probably mainly consisted of insects and other invertebrates – and it would have in turn been the potential prey of larger predatory birds like buzzards and long-eared owls.
All currently known subfossil remains of the São Miguel scops owl date only from the Holocene, between about 50 BCE and 125 CE. It’s likely that it was extinct by the 1400s, following the settlement of humans in the Azores, destruction of its forest habitat, and the introduction of rodents, cats, and weasels.
Mammuthus creticus was originally thought to also be a palaeoloxodontine, but more recent studies of its anatomy and ancient DNA have confirmed it was indeed another tiny mammoth. It was probably descended from either the Southern mammoth or Mammuthus rumanus, which would have arrived on Crete during the late Pliocene or early Pleistocene between about 3.5 and 1 million years ago.
Isolated on Crete, with no predators and living at a time when the island was much smaller, it quickly dwarfed and became the tiniest known mammoth to ever exist, standing just 1.1m tall at the shoulder (3’7″). Not much is known about its ecology, but its teeth suggest it was a browser feeding on leaves and shrubs, possibly filling a similar niche to the mid-sized deer that came later.
This mini-mammoth seems to have gone extinct by the mid-Pleistocene, about 1 million years ago, around the time when rising sea levels during an interglacial phase may have submerged so much of the smaller proto-Crete that its population could no longer be supported.
Later in the mid-to-late Pleistocene, after the sea level dropped again and tectonic uplift brought Crete close to its modern dimensions, the small mammoths were replaced by both newly-arriving deer and Palaeoloxodon elephants, which evolved into the much more moderately dwarfed forms of Palaeoloxodon creutzburgi and Palaeoloxodon chaniensis.
Palaeoloxodon tiliensis
To the north and east of Crete the Cyclades and Dodecanese islands had endemic dwarf elephants on at least eight islands, with the best known being the species that lived on Tilos.
Palaeoloxodon tiliensis stood about 1.8m tall (5’11”), on the larger side for a dwarf Mediterranean elephant but still one of the smallest palaeoloxodontines in the Aegean region. Several thousand specimens have been found, and radiocarbon dating shows it was a fairly recent evolutionary development, appearing just 45,000 years ago in the late Pleistocene.
This dwarf elephant was also the very latest surviving of its entire kind, living well into the Holocene until at least 4000 BCE. This is several thousand years after humans first arrived on Tilos, suggesting it was a rare case of an island elephant that managed to endure the effects of a human presence for quite some time.
In fact there’s some speculation that Palaeoloxodon tiliensis (or a similar unknown species) may have survived for even longer than that, since one Ancient Egyptian tomb from around 1480-1400 BCE contains a painting depicting traders with exotic animals, including what appears to be a small hairy elephant with slender limbs and thin upward-curving tusks. We may never know for certain if this was actually a late-surviving dwarf, a mutant modern elephant, or just artistic license with scaling, but the possibility is still intriguing.
Palaeoloxodon cypriotes
Over on isolated Cyprus further to the east, the only native large mammals were the miniature hippos and an equally miniature elephant.
Palaeoloxodon cypriotes was smaller than the Aegean palaeoloxodontines, about 1.4m tall (4’7″), and much like its cousin on Tilos seems to have evolved very recently towards the end of the Pleistocene, sometime around 20,000 years ago.
It wasn’t the first dwarf elephant on Cyprus — there was a larger, earlier species known as Palaeoloxodon xylophagou at least 200,000 years ago — but it’s not clear whether these two species represent a single evolutionary line or two entirely different colonizations of the island.
Similarly to the hippos it lived alongside, Palaeoloxodon cypriotes disappeared shortly after humans arrived on Cyprus, between 12,000 and 10,000 years ago. Collections of its bones have been found in a rock shelter with evidence of having been burnt, suggesting that it was being actively hunted and cooked.
And that’s all for the Island Weirdness series! Even over two months there are still plenty of species I didn’t have time to feature, so this definitely won’t be the last we see of strange endemic species.
Thank you for following along — with a shoutout to my Patreon supporters! — and regular weekly art posts will resume here next Monday.
Cyprus is one of the most isolated islands in the Mediterranean, having had no close connections to the mainland for the last 5.3 million years and being uplifted to close to its modern size during the Pleistocene. As a result it had very few land mammals, all of which arrived by swimming or rafting: rodents, shrews, a genet, dwarf elephants, and a dwarf hippopotamus.
Hippopotamus minor (sometimes called Phanourios minor) seems to have been descended from the common hippo, which probably swam across to Cyprus from the Levant region sometime in the mid-to-late Pleistocene, around 400,000 years ago. Isolated with no predators and limited space it rapidly became dwarfed compared to its ancestors, reaching at most 75cm tall at the shoulder (2’6″) — making it the tiniest known island hippo, and slightly smaller than the modern pygmy hippo.
It became much more terrestrial, with more digitgrade feet adapted for walking and climbing over rugged rocky terrain. Its teeth suggest a diet of pig-like browsing on forest vegetation — and much like pigs (and other hippos) they may have been opportunistic omnivores occasionally also eating small animals and carrion.
Despite being so small for a hippo, it was still one of the largest animals living on Cyprus, weighing about the same as the dwarf elephants it lived alongside. It also seems to have been the most common of the mammals on the island, with remains of thousands of individuals having been found.
While larger dwarf hippos are known fromseveralother Mediterranean islands, the Cypriot species is the only one that seems to have survived into the early Holocene.
The earliest known evidence of humans in Cyprus comes from a rock shelter on the southern coast, dating to about 12,000 years ago, consisting of stone tools and a massive concentration of burned animal bones — with over 200,000 of them coming just from Hippopotamus minor. It’s possible that in addition to being so abundant on the island, the dwarf hippos’ evolution in the absence of predators meant they had no fear of humans and were much less aggressive than their larger relatives, making them particularly easy to hunt and kill.
…Or they were just especially tasty.
Later deposits from about 2000 years later show no sign of the hippos at all, with their role in the Cypriot ecosystem completely replaced by introduced species like deer, sheep, and goats.
Despite being decent swimmers, canids are surprisingly rare in island ecosystems, only seeming to end up there when able to move over land connections with larger landmasses (or when brought there by humans). Even the most remote species, the recently-extinct warrah, is thought to have crossed over a short stretch of sea ice during the last glacial period.
The Sardinian dhole (Cynotherium sardous) was a small fox-sized canid, just 50cm tall at the shoulder (1’8″), related to the modern African wild dog and dhole. It was probably descended from the much larger wolf-like Xenocyon, which would have been able to reach Sardinia-Corsica during the early-to-mid Pleistocene about 1.2 million years ago, at a time when lower sea levels connected the island to the European mainland via Tuscany.
Isolated with very little large prey, it instead evolved to specialize in hunting small fast-moving animals, flattening its body low to the ground while stalking in a similar manner to modern foxes or Ethiopian wolves. Powerful shoulder muscles allowed it to launch into sudden high-speed lunges, and it had an especially strong flexible neck that would have been used to grab at its zig-zagging targets and shake them to death.
Cynotherium went extinct sometime in the early Holocene, around 11,000 years ago, after the arrival of humans on Sardinia and Corsica.
The earliest definite human remains on Sardinia are at least 20,000 years old, and while it’s unclear if those were permanent settlers it still seems like Cynotherium was able to deal with the effects of a human presence for several thousand years, probably due to its main prey (the Sardinian pika) also surviving at the time. So its disappearance may have been caused by a combination of problems that slowly whittled away at its population, like the warming climate, gradual habitat destruction, and competition from introduced predators like feral dogs — or possibly even new diseases caught from them.
Along with its miniature mammoths, the Mediterranean island of Sardinia (and neighboring Corsica) had an unusually large amount of endemicmustelids during the Pleistocene and early Holocene. There were at least four different otters — probably all descended from a single species of Lutra — that occupied various ecological niches in both the rivers and the coasts, and also an enigmatic grison-like terrestrial species.
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
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.
Myotragus balearicus
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.
Mammuthus lamarmorai lived on the island of Sardinia during the middle and late Pleistocene, between about 450,000 and 40,000 years ago. Standing around 1.4m tall at the shoulder (4’7″), it was a dwarf form roughly one-third the size of its ancestor, the huge Eurasian steppe mammoth.
Its remains are known only from the west and south of the island. Strangely it appears to be have been absent from the nearby island of Corsica, despite the two being joined as a single landmass a few times during lower sea level periods in the Pleistocene.
It’s not clear why this mini-mammoth disappeared. The date of the earliest human settlement of Sardinia is controversial (ranging from 250,000 to 20,000 years ago), so Mammuthus lamarmorai might never have actually encountered them. Instead it may have struggled to cope with climate changes during the last glacial period towards the end of the Pleistocene, which rapidly turned Sardinia colder and drier.
Palaeoloxodon falconeri
Over on Sicily and Malta (which were also occasionally a single island when sea levels were lower), there were several different species of miniature elephant during the mid-to-late Pleistocene, each with a different body size and occupying its own ecological niche. They were all descendants of the massive straight-tusked elephant (Palaeoloxodon antiquus) but each resulted from independent colonization waves swimming over to the island(s).
Palaeoloxodon falconeri lived during the mid-Pleistocene, about 550,000 years ago, and was both the smallest of the Mediterranean dwarfs and possibly the smallest elephant to ever exist. Just 0.8-1m tall (2’7″-3’3″), it had adult body proportions resembling the juveniles of its ancestors, limbs adapted for running and clambering over rough terrain, and a proportionally enormous brain relative to its body size.
Around 200,000 years ago a sea level drop allowed new colonization from the Italian mainland. If Palaeoloxodon falconeri was still around at that time it likely didn’t survive long with new competition from large herbivores like bison and deer, and being preyed on by newly-arriving large carnivores like wolves, lions, and hyenas.
Another small elephant soon evolved to take its place, although due to the presence of predators it was never able to get nearly so tiny.
Palaeoloxodon mnaidriensis
Palaeoloxodon mnaidriensis was in fact one of the largest dwarf elephants in the Mediterranean, standing about 1.8-2m tall (5’11”-6’6″), but despite its larger size its limbs still show signs of adaptation for more fast and agile movement. Its tusks also show a lot of variation in shape, with some much more curved and twisted than others.
This elephant had disappeared by about 13,000 years ago, probably due to the climate significantly warming towards the end of the last ice age. Much like Sardinia, the earliest arrival of humans on Sicily and Malta is controversial, and it’s unclear whether they ever encountered Palaeoloxodon mnaidriensis — the earliest definite date for Sicily is about 16,000 years ago, so a human-induced extinction can’t be ruled out entirely.
The subfossil remains of the Siculo-Maltese elephants may have also ended up inspiring legends of the cyclops, as their skulls would have resembled large human ones with the nasal cavity forming a single big “eye socket”.
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.
One of the few known fossil species is Caracara tellustris, a close relative of the modern crested caracaras that probably evolved in the mid-Pleistocene between about 1.2 and 0.4 million years ago.
About 60-65cm long (2′-2’2″), it was similar in body size to larger individuals of its living relatives but was heftier built and had slightly longer legs, and its wings were reduced enough that it was either a very weak flier or entirely flightless.
It lived only in the dry scrubland around the southern coast of the island, and probably mainly preyed on reptiles, small rodents, and crabs using its strong legs — a lifestyle very similar to the modern secretarybird. Like other caracaras it would have also opportunistically scavenged on carrion, which there would have been little competition for.
The known remains of Caracara tellustris date to as recently as 100 CE, showing that it existed well into the Holocene and survived through the initial arrival of humans on its island home (about 4000 BCE). This is likely due to its inhospitable hot, arid, and thorny habitat, where it would have been left relatively undisturbed, and it may even have persisted until the time of European colonization in the 1500s.
Unfortunately the scrubland was also very limited in size and the Jamaican caracara would have always been quite a rare species. If it was still around by then it would have faced a combination of introduced predatory mammals and habitat destruction by agriculture, which would have driven it into extinction so quickly its existence was never even noticed by naturalists at the time.
Jamaica is the third largest island in the Caribbean, and much like Cuba it originated as part of a Late Cretaceous volcanic island arc. It began to subside during the Eocene and was completely submerged for a large portion of the Cenozoic, then was uplifted again in the early-to-mid Miocene, reaching close to its present-day size around 13 million years ago.
Few land mammals ever colonized the island prior to human influence, and most of the known remains are from rodents. But another group did make it onto Jamaica, and became something especially weird.
Xenothrix mcgregori
Xenothrix mcgregori is a primate only known from fragmentary remains, but what is known of its skeleton shows a unique combination of features for a New World monkey. It had a reduced number of teeth in its jaws, with enlarged molars, and oddly-shaped heavily-built leg bones that resemble those of slow quadrupedal climbers like lorises.
It was probably about 70cm long in total (2’4″), including the tail, and is thought to have lived a lot like a tree sloth, spending most of its time moving slowly around in the trees and possibly even feeding while hanging upside down.
Its anatomy was so ununsual that its evolutionary relationships were a mystery until ancient DNA was recovered from subfossil bones and confirmed it was actually a titi monkey very closely related to the genus Cheracebus. Its ancestors probably arrived on Jamaica in the late Miocene, around 11 million years ago, and it had some close relatives on a couple of other Caribbean islands — the terrestrial Paralouatta on Cuba, and Antillothrix and Insulacebus on Hispaniola — although they likely all independently colonized the Caribbean via different rafting events from South America.
Xenicibis xympithecus
Another inhabitant of Jamaica was an equally strange bird.
Xenicibis xympithecus was one of only two lineages of ibis ever known to have become completely flightless (the other being Apteribis from Hawaii).
Around 60cm tall (2′), it had some of the most unique wings of any bird. The hollow bones were thickened, its forearm was proportionally short, and the hand was modified into a large heavy “club” — and blunt-force injuries on some of these birds’ remains suggest that they used their wings as weapons when fighting, clobbering each other with powerful blows.
Radiocarbon dating suggests the Xenothrix monkeys survived well into the Holocene, until around 1100 CE. Since various groups of humans had been present on Jamaica since about 4000 BCE the sloth-monkeys must have coexisted with them for several millennia, and their extinction may have been caused by more of a “slow fuse” of gradual habitat destruction than direct exploitation.
Dating on Xenicibis‘ extinction is less precise, with the youngest known remains being somewhere between 10,000 and 2200 years old. It may have still been around when the earliest humans arrived, but unlike the native monkeys it seems like it didn’t last long beyond that point.
