Earlier in this series we saw some ruminants with bizarre-looking headgear, but there was another species in that group that evolved a completely different type of strange head.
Rusingoryx atopocranion was a close relative of modern wildebeest that lived during the late Pleistocene, around 100,000 to 50,000 years ago. Its fossil remains are known from the Kenyan part of Lake Victoria, on Rusinga Island – an area which wasn’t actually an island at the time due to lower lake levels, and was instead part of a hot dry grassland environment.
Standing about 1.2m at the shoulder (~4′), it had an oddly-shaped skull with a pointed snout and a highly domed forehead. But this wasn’t the thick bony dome of a headbutting animal – this structure was narrow and fairly fragile, and had looping nasal passages running through it.
Instead it was something never before seen in any mammal: a bony nasal crest convergently similar to those of hadrosaurid dinosaurs.
Juveniles had less developed crests, developing them as they matured, and one skull that may represent an adult female also has a smaller crest, suggesting that this feature was sexually dimorphic.
Based on just the anatomy of the nasal passages Rusingoryx may have honked at a frequency similar to a vuvuzela, but the added length of its vocal tract could have lowered this pitch even further, closer to infrasound ranges – so more like a tuba! Such low frequencies can travel very long distances and are also below the hearing range of many carnivores, and would have effectively allowed Rusingoryx to shout at each other in “stealth mode”.
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.
The Mediterranean island of Crete had very few predators during the Pleistocene, with most being birds of prey. And with the terrestrial carnivore niches in the ecosystem left vacant, it was a semi-aquatic mammal and an owl that ended up taking advantage of that opportunity.
Neither were large enough to threaten the dwarf elephants and hippos, and don’t even seem to have habitually eaten even the smallest of the miniature giant deer. Instead these Cretan predators focused much more on the smaller land vertebrates on the island, preying on birds, shrews, rodents, amphibians, and reptiles.
Lutrogale cretensis
Lutrogale cretensis (previously known as Isolalutra cretensis) was a close relative of the modern smooth-coated otter. It was about the same size as its living cousin, around 1m long (3’3″), but had stronger jaws and chunkier limbs.
Its skeleton shows features associated with walking and running more than swimming, and it seems that this was something of a “land otter” — still able to swim, but spending most of its time on land similar to the modern small-clawed otter.
Shellfish were likely still the main part of its diet, indicated by its crushing teeth. But it probably also regularly ate whatever small terrestrial vertebrates it could catch, since more aquatic otters are already known to prey on those types on animals when they can.
Athene cretensis
Athene cretensis was yet another weird island owl, but this time not a descendant of a Strix or Tyto species. Instead this owl was descended from the Eurasian little owl — except it had become much much larger.
It stood around 60cm tall (2′), over three times bigger than its living relative. Its legs weren’t quite as long as those of the modern burrowing owl, but they were still proportionally much longer than those of little owls and show adaptations for terrestrial movement. Little owls already sometimes chase down prey on foot, and Athene cretensis was probably even more of a ground-based hunter, convergently similar to the Hawaiian stilt-owls and the Cuban terror owls.
Preserved pellets show that it ate small mammals and birds, mainly large mice.
Its wings were still quite large, and it was probably also a good flier — and may even have spread over to some of the Dodecanese islands to the east of Crete, since a wing bone closely resembling that of Athene cretensis has been found on Armathia.
Both of these predators seem to have disappeared around the end of the Pleistocene, at the same time as many of the other native Cretan species about 21,500 years ago. Much like the situation with Candiacervus, this may have been a result of a combination of a rapidly shifting climate and the presence of humans disrupting the already fragile island ecosystem.
The island of Crete has been isolated since about 5.3 million years ago, when the dried-out Mediterranean Searefilled — but at that time it started off as several much smaller islands, and only gained its larger modern shape thanks to tectonic uplift in the Pleistocene.
It only had a small number of endemic land mammals during the Pleistocene, whose ancestors all seem to have reached the island by swimming or rafting from southern Greece: dwarf elephants, a small hippo, an otter, a shrew, large mice, and several deer.
Deer are surprisingly good swimmers, and seem to have colonized Crete by the mid-to-late Pleistocene 300,000 years ago. They were by far the most diverse mammals on the island, with eight species in six size classes, each living in different types of habitat and specializing in their own ecological niche in a similar situation to the older Italian Hoplitomeryx. Their anatomy was modified so much that it’s unclear what their original ancestors actually were, or even if they were all descended from a single colonization or multiple arrivals, but they seem to have been close relatives of the huge Megaloceros.
All eight species are usually classified in the genus Candiacervus, and the smallest and weirdest of them all was Candiacervus ropalophorus.
Ironically for a cousin of the giant deer it was tiny, just 40-50cm tall at the shoulder (1’4″-1’8″), with proportionally short stocky legs more like a goat. It seems to have convergently evolved to occupy the same niche as wild goats do elsewhere, clambering over steep rocky mountainous terrain and eating tough prickly vegetation.
The antlers of the males were huge for their body size, around 77cm long (2’6″), and they were simplified into a long straight beam with only a single small spike at the base. The far ends were wider and rounded, described as club-like or spatula-like, and their odd shape suggests they probably weren’t much use for fighting and wrestling like in other deer. Instead they seem to have been more just for show and visual display.
Meanwhile a second dwarf species, Candiacervus reumeri, had more standard-looking antlers and probably still fought each other.
The largest species, Candiacervus major, was as big as a modern wapiti, with a shoulder height of around 1.65m (5’5″) and body proportions much more like a normal long-legged deer. Its antler shape isn’t actually known yet, but since it lived in thickly forested areas of Crete the stags may have had more streamlined antlers to avoid getting snagged on low branches.
The various Candiacervus species went extinct towards the end of the Pleistocene, around the start of the Last Glacial Maximum 21,500 years ago. Originally this was thought to be long before humans ever reached the island, but more recent discoveries have brought that into question.
Humans do actually seem to have encountered living Candiacervus ropalophorus, since petroglyphs in Asphendou Cave appear to depict the dwarf deer and so must be at least 21,500 years old. Additionally, even older stone tools on the southern coast of Crete from at least 130,000 years ago match those made by archaic humans (probably Homo erectus) who may have arrived over sea from northern Africa.
So it’s possible the weird Cretan deer survived alongside humans for some time, but then their habitat started to degrade as the climate shifted rapidly colder and drier. Some remains show that many individuals were suffering from secondary hyperparathyroidism and metabolic bone disease, signs of severe nutritional deficiencies, and their weakening population may have ultimately been unable to deal with both the malnutrition and the additional pressures of human hunting.
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:
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.
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”.
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.
Located on the Pacific Ring of Fire, the islands of the Philippines were formed by volcanic activity at the junction between several tectonic plates. Most of the 7461 islands that make up the archipelago have never been connected to any other landmass, leading to a huge number of unique endemic species evolving from whatever managed to arrive via ocean rafting events.
Celebochoerus cagayanensis
Celebochoerus cagayanensis was a giant species of pig, known from the island of Luzon. Living around 800,000 years ago in the mid-Pleistocene, it had enormous tusks and stood around 1m tall at the shoulder (3’3″) — similar in size to the very largest modern pigs, the African giant forest hogs.
The giant forest hogs are also some of its closest living relatives, along with the river pigs, and back in the Miocene and Pliocene similar pigs were present in Asia. Celebochoerus‘ ancestors probably arrived in the Philippines from Taiwan, and eventually spread onwards to the Indonesian island of Sulawesi to the south, where another species of Celebochoerus existed.
Bubalus cebuenesis
In contrast to the huge pigs of Luzon, the Cebu tamaraw (Bubalus cebuensis) was a particularly tiny species of wild cattle related to modern water buffalo. Just 75cm at the shoulder (2’6″), it was an example of insular dwarfism even smaller than the modern tamaraw which still survives on the island of Mindoro.
The spotty fossil record of these animals makes it difficult to determine when they disappeared, but it’s likely that they went extinct sometime around the arrival of early humans about 700,000 years ago.
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.
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.
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.
Standing at an adult height of about 1.1m tall (3′7″), they were smaller than any population of modern humans and are thought to represent an unusual case of insular dwarfism.
They also had much smaller brains than would be expected for their size, similarly to the miniature hippos of Madagascar, which was probably an energy-saving adaptation in an island environment with limited resources, since brains are metabolically expensive organs. An area of their brains associated with higher cognition was about the same size as in modern humans, however, so they weren’t necessarily less intelligent – stone tools and butchery marks on dwarf elephant bones suggest they were cooperatively hunting, and there’s also possible evidence of fire use for cooking.
It’s not clear exactly where they belong on the human family tree, and attempts at extracting DNA from the known remains have so far failed. They might be descendants of a population of Homo erectus who arrived on Flores about 1 million years ago, or they may even have been part of a much older unknown lineage that dispersed from Africa over 2 million years ago.
Although they were initially thought to have lived on Flores from 190,000 years ago up to about 12,000 years ago, more accurate dating of the cave where their skeletal remains were discovered suggests they actually disappeared about 50,000 years ago – about the same time that modern humans arrived on the island.
