Island Weirdness #57 — Cygnus falconeri

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:

A scale comparison between a dwarf elephant and a giant swan. The swan is the taller of the two.
Palaeoloxodon falconeri vs Cygnus falconeri

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.

Island Weirdness #56 — Cynotherium sardous

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.

So the existence of a unique canid on the Mediterranean islands of Sardinia and Corsica is quite unusual.

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.

Island Weirdness #55 — Megalenhydris barbaricina

Along with its miniature mammoths, the Mediterranean island of Sardinia (and neighboring Corsica) had an unusually large amount of endemic mustelids 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.

Island Weirdness #54 — Big Bunnies & Strange Sheep

At the end of the Miocene, about 6 million years ago, the movement of tectonic plates caused the narrow connection between the Mediterranean Sea and the Atlantic Ocean to close — and over the next several hundred thousand years the Mediterranean dried up almost completely.

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.

A stylized illustration of an extinct giant rabbit. It doesn't resemble a modern rabbut much at all, with a small head, tiny eyes and ears, a fat body and thick somewhat bear-like limbs.
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.


A stylized illustration of an extinct goat-like sheep. It has a short snout, oddly forward-facing eyes, short pointed horns, and a chunky body.
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.

Island Weirdness #53 — Tiny Elephants On Parade Part 5: Western Mediterranean

During the Pleistocene elephants colonized many of the islands in the Mediterranean Sea, with each island independently developing its own unique dwarfed form.

A stylized illustration of an extinct pygmy mammoth. It has long tightly-curved tusks, small ears, and a coat of hair over its body.
Mammuthus lamarmorai

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.


A stylized illustration of an extinct dwarf elephant. It has long thin tusks and tiny ears, and its body shape resembles a chubby baby elephant.
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.

A stylized illustration of an extinct dwarf elephant. It has short thick twisting mammoth-like tusks and small ears.
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”.

Island Weirdness #52 — Labidura herculeana

Located in the South Atlantic Ocean, Saint Helena is a small volcanic island that formed several million years ago on the Mid-Atlantic Ridge. It’s one of the most remote islands in the world, and with no native land mammals and no large predators it developed an ecosystem with several unique flightless birds, and hundreds of endemic invertebrates and plants.

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.

The last traces of the species were some isolated subfossil pincers found in the mid 1990s, and the giant earwig was officially declared extinct in 2014.

Island Weirdness #51 — Tainotherium valei

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.

Afterwards subsidence in the early Cenozoic submerged many of the islands and drowned whatever was left, so it wasn’t until later tectonic uplifting that land re-emerged and was able to be recolonized by species via rafting and island hopping (or possibly via a short-lived land bridge).

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.

Island Weirdness #50 — Caracara tellustris

Jamaica is unusual among the Caribbean islands for having had relatively few predatory birds.

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.

Island Weirdness #49 — Sloth-Monkeys & Fighting Ibises

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.

A stylized illustration of an extinct titi monkey. It has long soft fur and a very long tail.
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.


A stylized illustration of an extinct flightless ibis. It has stout legs, and has its wings raised as if threatening to hit something with its heavy club-like arm bones.
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.

Island Weirdness #48 — Chunky Cranes & Terror Owls

Like many other isolated islands ancient Cuba lacked any large land predators, allowing some birds to exploit more terrestrial lifestyles.

A stylized illustration of an extinct flightless sandhill crane. It has a somewhat chunkier beak than its modern relatives, along with smaller wings and thicker legs.
Grus cubensis

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.


A stylized illustration of an extinct giant owl. It has proportionally short wings and long stilt-like legs.
Ornimegalonyx oteroi

And, once again, there was also a weird owl on this island.

Ornimegalonyx oteroi was closely related to true owls in the genus Strix, and in a great example of convergent evolution did the exact same thing as the Grallistrix stilt-owls — it evolved into a long-legged short-winged ground-based apex predator.

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.