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.


Litovoi tholocephalos, a multituberculate mammal from the Late Cretaceous of Romania (~70-66 mya). Living on what was at the time the large offshore Hațeg Island, this rat-sized animal (about 25cm /10″ long) was part of a lineage of insectivorous multis called the kogaionids, with the same sort of red-colored enamel on its teeth as other species like Barbatodon.

Its brain was surprisingly tiny proportional to its size – one of the smallest known brain-to-body ratios of any mammal, and more similar to those of non-mammalian cynodonts – but it also seems have been highly specialized for processing sensory input, with relatively enormous regions associated with smell, eyesight, balance, and motor control. The olfactory bulbs of its brain were so enlarged, in fact, that they caused its skull to bulge out into an unusually dome-shaped forehead.

Its reduced brain size may have been due to limited food availability on its isolated island home. Brains are very metabolically expensive organs, and some other extinct island mammals like hippos, hominids, and goats are also known to have evolved smaller brain sizes. Modern shrews even seasonally shrink their own brains during winter for similar energy-saving reasons.