Island Weirdness #60 — Hippopotamus minor

Cyprus is one of the most isolated islands in the Mediterranean, having had no close connections to the mainland for the last 5.3 million years and being uplifted to close to its modern size during the Pleistocene. As a result it had very few land mammals, all of which arrived by swimming or rafting: rodents, shrews, a genet, dwarf elephants, and a dwarf hippopotamus.

Hippopotamus minor (sometimes called Phanourios minor) seems to have been descended from the common hippo, which probably swam across to Cyprus from the Levant region sometime in the mid-to-late Pleistocene, around 400,000 years ago. Isolated with no predators and limited space it rapidly became dwarfed compared to its ancestors, reaching at most 75cm tall at the shoulder (2’6″) — making it the tiniest known island hippo, and slightly smaller than the modern pygmy hippo.

It became much more terrestrial, with more digitgrade feet adapted for walking and climbing over rugged rocky terrain. Its teeth suggest a diet of pig-like browsing on forest vegetation — and much like pigs (and other hippos) they may have been opportunistic omnivores occasionally also eating small animals and carrion.

Despite being so small for a hippo, it was still one of the largest animals living on Cyprus, weighing about the same as the dwarf elephants it lived alongside. It also seems to have been the most common of the mammals on the island, with remains of thousands of individuals having been found.

While larger dwarf hippos are known from several other Mediterranean islands, the Cypriot species is the only one that seems to have survived into the early Holocene.

The earliest known evidence of humans in Cyprus comes from a rock shelter on the southern coast, dating to about 12,000 years ago, consisting of stone tools and a massive concentration of burned animal bones — with over 200,000 of them coming just from Hippopotamus minor. It’s possible that in addition to being so abundant on the island, the dwarf hippos’ evolution in the absence of predators meant they had no fear of humans and were much less aggressive than their larger relatives, making them particularly easy to hunt and kill.

…Or they were just especially tasty.

Later deposits from about 2000 years later show no sign of the hippos at all, with their role in the Cypriot ecosystem completely replaced by introduced species like deer, sheep, and goats.

Inticetus

While most modern toothed whales have jaws full of teeth that are all the same simple pointed shape – an adaptation for better holding onto slippery prey – their ancient ancestors had teeth much more like other mammals, with differentiated incisors, canines, and molars.

In-between them were whales like Inticetus vertizi, which lived off the coast of southwestern Peru during the Early Miocene, about 18 million years ago.

At over 3.5m long (11′6″) it was one of the larger known toothed whales around at the time, but it wasn’t the direct ancestor of any living whales. Instead it was more of an evolutionary “cousin” to them, part of an older offshoot lineage that lived alongside the early members of modern toothed whale groups.

Inticetus had a long and unusually wide-based snout, somewhat croc-like in appearance, with sharp pointed teeth at the front and multi-lobed cheek teeth further back. A lack of obvious wear on its back teeth suggests it wasn’t using them to chew up its food, and it may have had a fairly specialized diet – possibly using those back teeth to sieve small prey out of the water in a similar manner to modern lobodontine seals.

An close-up view of Inticetus' jaws, showing the differences in tooth shape from front to back.
Closeup of Inticetus‘ jaws

Inticetus-like teeth have also been found in Miocene-aged deposits in the eastern USA, the Atlantic coast of France, and southeast Italy, indicating that this ancient whale lineage was quite widespread.

Island Weirdness #21 – The Malagasy Mini-Hippos

Much like elephants, hippos have frequently made their way onto islands and developed dwarfed forms. These mini-hippos are mostly known from the Mediterranean, but further south they also occurred on Madagascar.

Hippopotamus madagascariensis (also sometimes called Hexaprotodon madagascariensis or Choeropsis madagascariensis; its exact classification is uncertain) was similar in size and appearance to the modern West African pygmy hippo – which it might have been closely related to, or may have just ended up resembling through parallel evolution.

Standing under 1m tall at the shoulder (3′3″) and measuring about 1.8m long (5′11″) it lived in the forested highlands and was much more terrestrial than its larger cousins. Its eyes were further down on the sides of its head, and it was better adapted for walking and running around on land, with proportionally longer legs and a more digitigrade posture.

It also had an unusually small brain for its size – about 30% smaller than expected – which may have been an energy-saving adaptation.

Two other species of dwarf hippo have been identified on Madagascar – the similarly-sized but more aquatic Hippopotamus lemerlei in the west of the island, and the larger and poorly-known Hippopotamus laloumena in the east. It’s not clear when exactly the ancestors of these various hippos first arrived on the island, and they may even represent three independent colonization events.

The last known subfossils date to about 1000 years ago, but surprisingly accurate accounts of similar creatures in Malagasy folklore raise the possibility that small isolated populations of these hippos may have survived into more recent times. One of the most intriguing examples is the kilopilopitsofy, described as having large floppy ears and supposedly being sighted as recently as the 1970s.

Rayanistes

Remingtonocetids were an early branch of the whale evolutionary family tree, known from about 49-41 million years ago and splitting off somewhere between the famous “walking whale” Ambulocetus and the more oceanic protocetids. With otter-like bodies, tiny eyes, and long gharial-like snouts, they lived in near-shore shallow marine habitats and probably swam using a combination of their hind feet and tails.

They were initially found only in Pakistan and India, but then Rayanistes afer here was discovered all the way over in Egypt – suggesting that these early whales were much more widespread than previously thought, dispersing through the Tethys Sea at about the same time as their protocetid cousins.

Dating to the Middle Eocene (~45-41 mya), Rayanistes was probably about 2.5m long (8′2”). It had powerful hindlimb musculature that would have given it a very strong kicking swimming stroke, but it probably couldn’t actually support its own weight on land since its femur wasn’t very well anchored into its pelvis.

Orcinus citoniensis

Despite commonly being called “killer whales” modern orcas are actually the largest living members of the oceanic dolphin family. Their ancestors are thought to have diverged from other dolphins between 10 and 5 million years ago – and surprisingly their closest relatives are the much smaller snubfin dolphins found in Australasia.

Living during the Pliocene (5-2 mya) in the Mediterranean, Orcinus citoniensis was an early member of the orca lineage, and was probably a transitional form between their early dolphin ancestors and the modern Orcinus orca.

It was half the size of modern orcas, at about 4m long (~13′). While it had a higher tooth count than its living relatives its teeth were also proportionally smaller, suggesting it wasn’t specialized for tackling large prey and probably fed mainly on fish and squid.

Waharoa

Waharoa ruwhenua, a whale from the Late Oligocene of New Zealand (~27-25 mya). Part of an early branch of the baleen whale lineage, it’s known from partial remains of an adult and a couple of juveniles and would have reached a full size of about 6m long (19′8″).

It had an unusually long flattened snout, with its nostrils further forward than modern whales, and only had baleen in the back half of its mouth – an interesting comparison to the intermixed teeth-and-baleen of some other early mysticetes. It’s not clear whether it had any vestigial teeth in the front of its jaws, although a single possible tooth has been found associated with its close relative Tokarahia.

The rather delicate nature of Waharoa’s jawbones suggests it wasn’t capable of rapid lunges at swarms of its small prey, instead probably using slow-cruising surface skim-feeding similar to modern right whales.

Inermorostrum

Inermorostrum xenops, a recently-named ancient cetacean!

Living about 30 million years ago in shallow coastal waters around the southeast USA, in what is now South Carolina, it was a member of one of the very earliest groups of toothed whales known as the xenorophids. Although only very distantly related to modern forms, xenorophids show evidence of being able to echolocate, suggesting the ability was probably ancestral to all toothed whales.

Estimated to have measured about 1m long (3′3″), Inermorostrum had a very short downturned snout and was completely toothless – specialized adaptations for suction feeding on small soft-bodied creatures on the seafloor.

Unusually for a toothed whale it also had proportionally large infraorbital foramina, openings in the bones of its snout for blood vessels and nerves to pass through. This suggests the presence of well-developed fleshy lips and possibly whiskers (as illustrated here), or maybe even an electroreceptive sense similar to some modern dolphins.

Ambulocetus

Ambulocetus natans, the Eocene “walking whale” – who might not actually have been able to walk at all!

A study published in 2016 suggests this early cetacean was actually fully aquatic and unable to support its own weight on land. So here’s an updated version compared to the Ambulocetus I did a couple of years ago.

Globicetus

Globicetus hiberus, a 5m long (16′4″) beaked whale from the Atlantic coast of Portugal and Spain. Its fossils can’t be easily dated since they were fished up from the seafloor, but it was probably around Early-to-Mid Miocene in age (~20-14 mya).

Its skull sported an odd bony sphere at the base of its snout, just in front of the melon, which appears to have been larger and more prominent in males than in females. Many modern beaked whales also have sexually dimorphic crests, ridges, and domes in their skulls, and these structures may function as sort of “internal antlers” – a display structure the whales can “see” via echolocation that signals their size, strength, and health to each other.