Island Weirdness #61 — Tiny Elephants On Parade Part 6: Eastern Mediterranean

Alongside the weird deer, otters, and owls, the island of Crete also had dwarf elephants — and much like Sardinia to the west the Cretan elephants were actually descendants of mammoths rather than the Palaeoloxodon seen in the rest of the Mediterranean.

A stylized illustration of an extinct pygmy mammoth. It has gently curving tusks, small ears, and a body shape more like a baby elephant.
Mammuthus creticus

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.


A stylized illustration of an extinct dwarf elephant. It has long thin tusks and small ears.
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.

A stylized illustration of an extinct dwarf elephant. It has long gently curving tusks and small ears.
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.

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 #44 — Tiny Elephants On Parade Part 4: Mammoth-Mimics & Mini-Mammoths

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.

A stylized illustration of an extinct mammoth-like elephant. It has long twisting tusks, small ears, and a speculative coat of long 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.


A stylized illustration of an extinct pgymy mammoth. It has curving tusks, small ears, and a speculative coat of hair.
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. 

(And if you prefer your pygmy mammoths less speculatively hairy, there’s always the version I did for PBS Eons earlier this year.)

Island Weirdness #32 — Tiny Elephants On Parade Part 3: More Indonesia

It’s finally time for part 2 of the Island Weirdness series!

(If you haven’t seen the previous installments, I suggest starting back at the beginning here.)


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!

A stylized illustration of an extinct dwarf elephant. It has long straight tusks and small ears.
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.

A stylized illustration of an extinct dwarf elephant. It has long gently curving tusks and proportionalyl short legs.
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.


A stylized illustration of an extinct dwarf elephant. It has four straight tusks, two longer ones in its upper jaw and two shorter ones in its lower jaw.
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.

A stylized illustration of an extinct dwarf elephant. It has long curving tusks and small ears.
Stegodon sompoensis

At the same time Sulawesi also had yet another small stegodontid, Stegodon sompoensis, also around 1.5m tall.

Both of these dwarfs lived alongside a larger Stegodon species, as well as giant tortoises and large-tusked pigs.

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.

Antilohyrax

This is not a deer.

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.

Eons Roundup 2

And speaking of island elephants, I recently illustrated some for PBS Eons’ episode on the California Channel Island pygmy mammoths:

https://www.youtube.com/watch?v=dXwoKEDtdlM

Island Weirdness #31 – Tiny Elephants On Parade Part 2: Flores

Much like Japan, ancient Flores had a succession of dwarf stegodontids – close relatives of modern elephants that were capable of island-hopping through Indonesia by swimming.

A stylized illustration of an extinct dwarf elephant. It has curved tusks, small ears, and very proportionaly short stumpy legs.
Stegodon sondaari

Stegodon sondaari lived on Flores during the Early Pleistocene, about 900,000 years ago, and was the size of a small water buffalo at just 1.2m (3′11″) tall at the shoulder. It was probably descended from the larger Stegodon trigonocephalus, known from Java, and it had proportionally short legs which may have been an adaptation to clambering over rough terrain and steep inclines.

Around 850,000 years ago Stegodon sondaari disappeared from Flores, probably due to a large volcanic eruption, but a new wave of stegodontids quickly recolonized the island. The mid-sized Stegodon florensis probably originated from either Java to the west or Sulawesi to the north, and eventually evolved into a new dwarfed subspecies.

A stylized illustration of an extinct dwarf elephant. It has long gently curving tusks and small ears.
Stegodon florensis insularis

Stegodon florensis insularis wasn’t quite as small as its predecessor, standing around 1.8m tall (5′10″). It probably didn’t shrink quite so much due to the existing presence of various predators on Flores, since it was likely the main prey of large Komodo dragons, it was hunted by Homo floresiensis, and it may also have been occasionally targeted by giant storks.

It seems to have disappeared around the same time as several other unique endemic species, between 50,000 and 20,000 years ago, due to either climate change, another volcanic eruption, or the arrival of modern humans – or perhaps a combination of all of those factors.

And that’s all for this month… but Island Weirdness will be back later for part 2, with more giants, more dwarfs, and so many elephants.

Island Weirdness #13 – Tiny Elephants On Parade Part 1: Japan

Part of the “island rule” is that large animals often become smaller – and no group seems to exemplify this more than the elephants.

Although they’re the largest living land animals today, and in the past included some of the largest known land mammals ever, ancient elephants also frequently ended up on islands thanks to their ability to swim long distances. They produced many different dwarfed forms around much of the world, and a few of them will be featured intermittently throughout both months of this theme.

The earliest known examples were the stegodontids of Japan in the early Miocene. These animals weren’t quite true elephants, instead being close evolutionary cousins to them, and had two small additional tusks in their lower jaws similar to the related gomphotheres.

A stylized illustration of an extinct dwarf elephant. It has four tusks – two short straight ones in its upper jaw, and another two smaller ones in its lower jaw.
Stegolophodon pseudolatidens

Stegolophodon pseudolatidens first arrived in Japan about 18 million years ago, and within just 2 million years they’d developed into insular dwarfs that were probably around 2m tall at the shoulder (6′6″)– still reasonably large, but only about 60% the size of their mainland relatives.

A stylized illustration of an extinct dwarf elephant. It has long curving tusks and small ears.
Stegodon aurorae

Much later in the Early Pleistocene another small almost-elephant appeared in Japan. Living between about 2 million years ago and 700,000 years ago, Stegodon aurorae was about the same size as the then-extinct Stegolophodon but probably wasn’t descended from them. Instead it was probably the result of a separate arrival and dwarfing of a larger Stegodon species from mainland Asia.