Island Weirdness #14 – Hoplitomeryx

During the mid-Miocene, about 15 million years ago, a region of central and southeast Italy around Gargano and Scontrone was cut off from the mainland by rising sea levels.

For the next 7-10 million years this island (or perhaps a cluster of islands) was left isolated, and an unusual ecosystem developed known as the “Mikrotia fauna”. With the island starting off lacking large predators, small herbivorous animals like rodents, pikas, and waterfowl became huge – and then small predators like gymnures and carnivorous birds also grew to keep up with the increasing size of their prey.

One of the strangest residents of the island(s) was Hoplitomeryx, an early type of ruminant that resembled a deer or pronghorn. Nicknamed the “prongdeer”, it had a total of five horns on its head and large protruding fangs similar to some modern deer.

Multiple species of Hoplitomeryx have been identified, representing four different size classes ranging from huge down to tiny insular dwarfs. The largest is estimated to have been similarly sized to modern moose, standing around 2m tall at the shoulder (6′6″), while the smallest would have been under 50cm (1′8″).

Each of these size classes was specialized for slightly different ecological niches, eating different types of vegetation to avoid directly competing with each other for the limited amount of food on the island.

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.

Island Weirdness #12 – Anatoliadelphys maasae

The Pontide island 43 million years ago had no placental mammalian carnivores, and in their absence a different type of mammal took up the main predatory niches – the metatherians.

Metatherians today are represented solely by marsupials, but there were once many other branches of this mammal lineage. One group present in Europe for much of the Cenozoic were the herpetotheriids, and these rat-sized animals seem to have also occupied the small predator and opportunistic omnivore niches on the Pontide island.

And then there were the bigger metatherians.

Anatoliadelphys was a specialized carnivore, with strong jaws and bone-crushing teeth similar to the modern Tasmanian devil. It had a body size similar to a domestic cat, giving it a total length of up to 1m long (3′3″), and had opossum-like grasping digits and a prehensile tail indicating it was capable of climbing trees.

It was initially thought to be descended from another type of metatherian found in Europe at the time, the peradectids, but a later study came up with a more surprising result for its evolutionary relationships. It instead may be closer related to the polydolopimorphs, a group known mainly from South America. Quite how its ancestors got as far as Turkey is a bit of a mystery, but it’s possible they’d previously dispersed into Africa at a time when the continents were still much closer to each other than they are today, and then spread northwards until they reached the Pontide island.

The main problem with this hypothesis is the current lack of any polydolopimorphans in the African fossil record – but the African Eocene is still poorly understood, so there may be some metatherian fossils yet to be discovered that could fill in this gap.

Island Weirdness #11 – Hilalia

During the mid-Eocene, about 43 million years ago, the Pontide region of modern day north-central Turkey was an island located between the Paratethys Sea to the north and the Tethys Ocean to the south.

This island had an ecosystem of native mammals unlike anything existing today, with a unique mixture of species whose ancestors had arrived over several million years via island-hopping or temporary land connections with Europe, Africa, and Asia. There were large rhino-like embrithopods, cat-sized metatherian predators, early primates, early bats, and “enigmatic” small insectivores – but the island completely lacked any of the rodents, carnivorans, creodonts, perissodactyls, or artiodactyls which were dominating the rest of Eurasia.

But one of the most surprising inhabitants of the island were the small herbivores – a genus called Hilalia that represented a type of early ungulate known as pleuraspidotheriids. Resembling deer-dogs, this group had originated in Europe during the Paleocene and had gone extinct there around 13 million years earlier than the date of the Pontide remains, making them living fossils at the time.

These survivors of an archaic lineage thrived on their isolated island refuge and there were at least four or five species of Hilalia at different sizes, suggesting they’d diversified to each occupy a slightly different niche in their ecosystem. It’s hard to accurately measure their body size from their very fragmentary remains, but they may have ranged from around 25cm to 60cm in total length (~10″- 2′).

Island Weirdness #10 – The Kogaionids

Multituberculates were a group of rodent-like mammals that originated back in the Early Jurassic, at a point on the mammalian family tree between the origin of monotremes and the earliest therians (represented today by marsupials and placentals).

While they were an incredibly successful group, found around the world in large numbers, in Late Cretaceous Europe multis had become incredibly rare and restricted to just a single place: Hațeg Island.

Isolated there, they evolved into a unique family known as the kogaionids, diverging from their ancestral mostly-herbivorous diet to instead become specialized insectivores with distinctly red iron-pigmented teeth and huge blade-like lower premolars.

Skull of Barbatodon, from fig 2 in Smith T, Codrea V (2015) Red iron-pigmented tooth enamel in a multituberculate mammal from the Late Cretaceous Transylvanian “Haţeg Island.” PLoS ONE 10(7): e0132550. doi: 10.1371/journal.pone.0132550  | CC-BY-4.0

Some of them also had oddly domed skulls and proportionally tiny brains, along with highly acute senses of smell, eyesight, balance, and motor control.

Kogaionon ungureanui was one of the first kogaionids to be discovered, and gives its name to the group as a whole. Although known only from a skull, it was probably rat-sized, around 30cm long (~12″).

Unusually for island species, which are often ecologically fragile and vulnerable, the kogaionids’ insectivorous habits allowed them to successfully survive through the end-Cretaceous mass extinction 66 million years ago while the Hațeg dinosaurs and pterosaurs perished. And when conditions changed and their island home became reconnected to the rest of Europe they rapidly spread out and became common across the entire region for a further 10 million years, only finally disappearing in the early Eocene about 56 million years ago.

Island Weirdness #09 – Hatzegopteryx thambema

There are no big theropod dinosaurs known from the end-Cretaceous Hațeg Island ecosystem, but that doesn’t mean there weren’t any large carnivores at all.

The apex predator niche here instead seems to have been occupied by Hatzegopteryx, an enormous azhdarchid pterosaur. Standing up to 4.5m tall (14’9″) when on the ground, and with an estimated wingspan rivaling Quetzalcoatlus (~11m / 36′), it was one of the largest animals to ever fly – although like other azhdarchids it probably actually spent most of its time stalking around quadrupedally on the ground eating whatever it could fit into its mouth.

Its neck was shorter and chunkier than most other azhdarchids, and its skull was wider and more massively built. The walls of its hollow bones were also unusually thick and reinforced for a pterosaur, so much so that they were initially mistaken for those of a theropod instead.

Fossils of Hatzegopteryx are very fragmentary, however, so its full appearance and the specifics of its diet are still uncertain. But it would have probably been able to tackle much larger prey than other azhdarchids, possibly capable of using its sturdy beak to bludgeon or stab anything too big to pick up and swallow whole in a similar manner to modern marabou storks.

Island Weirdness #08 – Balaur bondoc

When Balaur was described in 2010 it was initially thought to be a dromaesaurid closely related to Asian forms like Velociraptor. With its particularly stocky legs built for strength rather than speed, two-fingered hands, and two large sickle claws on each foot, it was interpreted as a weird highly specialized predator terrorizing the other Hațeg Island species at the end of the Cretaceous. Although only 1.8m long (5’10”), it was hypothesized to have taken down prey much larger than itself with powerful slashing kicks.

But later analyses cast doubt on this interpretation.

A lot of the anatomical features of Balaur’s skeleton were odd for a dromaeosaurid, but matched those of avialans – a group of close evolutionary “cousins” to the dromaeosaurids, containing Archaeopteryx and the common ancestors of all modern birds. And, by 2015, multiple studies had confirmed Balaur wasn’t really a “raptor” but instead a little further along on the bird lineage.

So now our picture of this dinosaur is very different: a chunky-bodied island bird, grown large and secondarily flightless sort of like a Cretaceous equivalent to the dodo. Its double sickle claws were probably adaptations for climbing and perching in trees, and based on similar avialans it was likely a herbivore rather than a hypercarnivore.

Island Weirdness #07 – Telmatosaurus transsylvanicus

The hadrosauroid dinosaur Telmatosaurus was another resident of Hațeg Island, and while it wasn’t quite as small or specialized as its cousin Tethyshadros it was still dwarfed compared to their other relatives, only growing to about 5m long (16’4″).

It was also the first dinosaur fossil found with a specific type of non-cancerous tumor known as an ameloblastoma on its lower jaw – a surprising discovery, since ameloblastomas were previously only known to occur in mammals and a single snake species. Various other types of abnormal tissue growth have been identified in other hadrosauroids and hadrosaurs, however, suggesting that this particular lineage of dinosaurs may have been unusually susceptible to developing tumors.

Island Weirdness #06 – Zalmoxes robustus

Hațeg Island was home to quite a few unique species right at the end of the Cretaceous, so we’ll be focusing on it for a few more days.

Zalmoxes robusutus was a member of the rhabdodontids, a group of herbivorous ornithopod dinosaurs related to well-known names like Iguanodon, Tenontosaurus, and the hadrosaurs.

It had a chunkier build than its closest relatives, with a deep skull, a large beak, and a rotund body. Like other rhabdodontids it would have had powerful jaw muscles and ridged cheek teeth specialized for scissoring, adaptations for cutting up particularly tough plant matter.

It was also quite small, about 2.4m long (7’10”), although since the largest known fossils represent subadults this may not have been its full size. A second species in the same genus (Zalmoxes shqiperorum) lived on the same island and was actually slightly bigger, suggesting that Z. robustus represented a minor case of insular dwarfism.

Island Weirdness #05 – Magyarosaurus dacus

At the very end of the Cretaceous period, between about 72 and 66 million years ago, tectonic uplift from the start of the formation of the Alps created an island in the area corresponding to modern-day Romania.

Known as Hațeg Island, after the region where fossils of the native species were found, it was similar in size to Hispaniola and was surrounded by deeper waters than most of the other European archipelago islands.

Magyarosaurus here was was a titanosaur living on this island, and was one of the smallest known of all sauropod dinosaurs at just 6m long (19’8″). Much of that length would have been in its neck and tail, and its body was only actually about the size of a horse.

Like some other titanosaurs it had bony osteoderm armor along its back, although since only one isolated piece has been found the exact arrangement isn’t known.

Discovered by Franz Nopsca (the gay Transylvanian baron-paleobiologist-spy), in the early 1900s, it was one of the first dinosaurs proposed as an example of insular dwarfism. Later researchers disagreed with this hypothesis, suggesting instead that the Magyarosaurus fossils were just juveniles – and it wasn’t until 2010 that studies of bone microstructure proved that these miniature sauropods really were fully grown adults.