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.
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.
Just 21km south of the Bahamas (13 miles), Cuba is the biggest island in the Caribbean and has a complex geological history. It originated as part of a volcanic island arc in the Late Cretaceous and during its existence in the Cenozoic it was colonized by a richer variety of mammals than the Bahamas ever had, including rodents, the enigmatic solenodons and nesophontes, and ground sloths.
The Caribbean ground sloths were all part of the megalonychid lineage — which eventually reached North America and produced giants — and they arrived on Cuba by the early Miocene, about 20 million years ago.
Megalocnus rodens lived during the Late Pleistocene and Holocene, between about 125,000 and 5000 years ago. It had unusually rodent-like front teeth, and while it was dwarfed compared to its mainland relatives it was actually one of the largest of the Cuban ground sloths — close to the size of a modern black bear, about 80cm tall at the shoulder (2’5″).
Subfossil remains show that Megalocnus survived well into the Holocene, and there’s been speculation that it may even have still been around in the highland forests as late as the time of European colonization in the 1500s.
However, radiocarbon dating of remains has given no dates younger than about 5000-4000 years ago, about 1000 years after the earliest arrival of humans in Cuba.
Isla Pedro González is part of the Pearl Islands in the Gulf of Panama, about 48km (30 miles) offshore. It was formed after the end of the last glacial period, as steadily rising sea levels cut it off completely from mainland Panama about 8500 years ago — and isolating the population of deer that lived there.
Although Pedro González is the third largest of the Pearl Islands it’s still fairly tiny, just 6.5km long (4 miles). As a result of this small space, limited available food supplies, and a lack of any large predators, these deer rapidly shrank down into a dwarfed form to survive. By 6000 years ago they were the size of a small dog, just 35-40cm tall at the shoulder (1’2″-1’4″) and weighing less than 10kg (22lbs).
The Pedro González dwarf deer haven’t been given any official scientific name just yet, but since they’re thought to be descendants of brocket deer they’d be a part of the genus Mazama, either as their own separate species or as a subspecies — similar to the larger native deer on nearby Isla San José, which are probably their closest living relatives.
Paleoindian settlers arrived on Pedro González just over 6000 years ago, and we know they hunted and ate the tiny deer because all the known remains come from a preserved trash heap and show signs of human butchering and chewing.
Younger deposits have gradually less and less deer bones, and although those particular settlers had left again by about 5500 years ago the damage was already done — layers from another group of people about 2300 years ago show no deer bones at all, so the dwarf deer had to be already extinct by that time.
In the cool-temperate climate of Pleistocene Japan one type of small elephant seems to have convergently become somewhat of a mammoth-mimic, with twisting tusks and possibly even a thick coat of hair.
Palaeoloxodon naumanni
Palaeoloxodon naumanni lived during the late Pleistocene, between about 500,000 and 16,000 years ago. At about 2-2.5m tall at the shoulder (6’6″-8’2″) it was still fairly large, similar in size to the smallest living elephant species — but it was a dwarf in comparison to its immediate ancestors, the absolutely enormous Asian straight-tusked elephant (Palaeoloxodon namadicus).
Actual woolly mammoths (Mammuthus primigenius) were also present in Japan, but the two similar-looking elephants inhabited different environments — Palaeoloxodon naumanni preferred the southern forests, while the true mammoths roamed the colder north.
Humans arrived in Japan around 40,000-30,000 years ago, so Palaeoloxodon naumanni actually coexisted with them for quite some time. Although it was hunted, it seems to have mainly been climate change towards the end of the last glacial maximum that led to its extinction.
Mammuthus exilis
Over on the other side of the Pacific Ocean, at least 60,000 years ago, some huge Columbian mammoths (Mammuthus columbi) swam the 6.5km (4 miles) distance to the ancient island of Santa Rosae — a landmass that today is mostly submerged, with its remaining peaks forming the modern California Channel Islands.
With a lack of large predators and then steadily rising sea levels reducing the available habitat on their new home, the mammoths shrank into a dwarfed species known as the Channel Islands pygmy mammoth (Mammuthus exilis). Standing around 1.75-2m at the shoulder (5’9″-6’6″), they were less than half the size of their ancestors and had only about 10% of the body mass.
The pygmy mammoths survived until about 13,000 years ago, around the same time that early Paleoindians arrived. While they may also have been hunted by humans, the warming post-glacial climate is currently thought to be the main factor in their extinction, changing the types of vegetation on their still-shrinking islands and reducing fresh water sources.
We left off last time with the dwarf stegodontids of Flores, but other Indonesian islands also had their own populations of unusually small elephant-relatives — so here’s a few more to start off this month!
Sinomastodon bumiajuensis
Sinomastodon bumiajuensis lived on the island of Java during the early Pleistocene, about 2-1.5 million years ago. It stood around 2m tall at the shoulder (6’6″), less than half the size of most other Sinomastodon species from mainland Asia. Although it looked convergently similar to modern elephants it was actually a member of the gomphotheres, much more closely related to the weird “shovel-tuskers” than to any living species.
Stegodon semedoensis
Stegodon semedoensis, also from the early Pleistocene of Java about 1.5 million years ago, is only known from a few isolated molar teeth — but the size of those teeth suggest it was one of the smallest known pygmy stegodontids. It was probably no more than 1.2m at the shoulder (3’11”), comparable in size to its close relative Stegodon sondaari over on Flores.
Elephas celebensis
Meanwhile on Sulawesi, Elephas celebensis (sometimes called Stegoloxodon celebensis) was an actual true elephant closely related to the modern Asian elephant. Living during the late Pliocene and early Pleistocene, between about 2.5 million and 800,000 years ago, it was only 1.5m tall (5′) and had a second set of tusks in its lower jaw, a “primitive” feature retained from the gomphothere-like ancestors of modern elephants.
Stegodon sompoensis
At the same time Sulawesi also had yet another small stegodontid, Stegodon sompoensis, also around 1.5m tall.
The cooling climate of the Pleistocene and dropping sea levels eventually connected the islands of western Indonesia to the Sundaland landmass of mainland Asia. Influxes of new predators and competitors — and early humans — probably drove these endemic small elephants to extinction.
In Africa during the Eocene and Oligocene, the main terrestrial herbivores were a different type of mammal entirely: hyraxes, the close relatives of elephants and manatees. Although their only modern representatives are small climbing rodent-like animals, hyraxes were once a much more diverse and widespread group, filling a variety of ecological niches and ranging from the size of rats up to the size of rhinos.
Antilohyrax pectidens was a mid-sized example of these diverse hyraxes, standing about 50cm tall at the shoulder (1′8″) and living around 34-28 million years ago in Egypt. It had a deer-like snout and long slender limbs adapted for running and leaping, with leg bones incredibly similar in size and proportion to modern springbok.
Its incisor teeth were comb-shaped and resembled those of colugos, so it was probably a similar sort of selective browser eating soft leaves and shoots.
