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Zalmoxes (named after the Thracian deity "Zalmoxis", whose supposed resurrection is likened to the excavation of this dinosaur's remains) is a genus of rhabdodontid ornithopod dinosaur that lived on Hațeg Island, an area which would eventually become a basin within modern-day Romania, 70 - 66 million years ago, during the Maastrichtian stage of the Late Cretaceous Period.

Paleobiology[]

Size and Physique[]

Zalmoxes trying to stay afloat on a small raft
Zalmoxes trying to stay afloat on a small raft

Zalmoxes was a fairly small animal, with the type species Z. robustus growing to a maximum length of 2.5 meters (8 feet, 3 inches), and weighed no more than 45 kilograms (100 pounds).[1] As indicated by its species name, "robustus", Zalmoxes has a stout body that is more heavily-built than its relatives. Like most of the inhabitants of Hațeg Island, Zalmoxes is believed to be subjected to insular dwarfism. However, as brought up by Dr. Darren Naish, lead consultant of Prehistoric Planet, it is possible that Zalmoxes simply inherited the standard diminutive size of its ancestors,[DN 1] with its status as a dwarf being questioned since at least 2005; instead of evolving a shrunken body size in response to its island habitat, Zalmoxes may have instead relied on growing slowly for an extended period of time, as indicated by analysis of its bones.[2] A 2012 study also concluded that rhabdodontids really were small to begin with, and that some species like Rhabdodon itself actually attained their larger sizes due to gigantism.[3]

Dentition[]

Zalmoxes is known from a skull that is 80% complete, indicating that it had a deep, triangular head that was large for its body size, its beak filled with the characteristic spade-shaped teeth of other rhabdodontids. A study in 2003 indicates that Zalmoxes fed on tough fibrous vegetation from angiosperms and ferns to horsetails and the soft shoots of various plants.[4]

Prehistoric Planet's Portrayal
[]

Prehistoric Planet depicts Zalmoxes with a black color scheme streaked with teal stripes, with a creamy white underside to serve as countershading to help further camouflage the animal in the shadowy undergrowth of the forests of Hațeg Island.

Paleoecology[]

Paleoenvironment[]

Main: Hațeg Island

Main: Hațeg Island

Five juvenile Zalmoxes cautiously about to cross an open area of a forest in Hațeg Island
Five juvenile Zalmoxes cautiously about to cross an open area of a forest in Hațeg Island
“ Even by day, it's still dark in the understory of the dense forests, such as this one in Europe. Animals of any kind are difficult to distinguish. Yet, they're everywhere. ”

David Attenborough, Forests

During the time of Zalmoxes, 70 - 66 million years ago, most of Europe was a series of islands scattered across the prehistoric Tethys Ocean. Hațeg Island, which would one day become a town in Hunedoara County, Romania, was 200 kilometers (120 miles) from the nearest landmass, and is estimated to be 80,000 square kilometers (31,000 square miles) worth of tropical woodland, sustained by braided rivers, lakes, and seasonal monsoons. Due to limited resources on the island, most of the inhabitants grew smaller due to an evolutionary process known as insular dwarfism, though whether Zalmoxes was also subject to this same process remains to be confirmed given a lack of better support and evidence of the contrary.

Confined to the islands of Europe, the only way for rhabdodontids to proliferate in other places is to raft (above), swim short distances, hop across islands, and traverse land bridges
Confined to the islands of Europe, the only way for rhabdodontids to proliferate in other places is to raft (above), swim short distances, hop across islands, and traverse land bridges
“ Telmatosaurus seldom breaks cover. And equally inconspicuous, Zalmoxes, the last of a very ancient dinosaur lineage. This forest is one of their few remaining strongholds. ”

David Attenborough, Forests

By the end of the Cretaceous, one superfamily of iguanodontian dinosaur, hadrosauroids from Telmatosaurus to Edmontosaurus, became one of the most widespread animals on land. Their other relatives, however, were not as successful. There is the hypothesis that, because rhabdodontids may have split away from the iguanodontian clade approximately 160 million years ago during the Middle Jurassic Period (long before the hadrosauroids, which first appeared around 130 million years ago in the Early Cretaceous Period), they are a primitive group of "living fossils" that heavily resembled their early ancestors. Determining the validity of this view is difficult given how their existence throughout the Mesozoic remains enigmatic because, despite having been known since the describing of Rhabdodon in 1869, to date, only nine species within six rhabdodontid genera have been found, isolated from the rest of the world and confined to their "few remaining strongholds", the European islands of Spain and France (including - but not only - Ibero-Armorica, which is comprised of southern France and northeastern Spain), western Hungary, eastern Austria, and Hațeg (Romania) during the Late Cretaceous Period (around 85 - 66 million years ago), giving them an exceptionally long ghost lineage spanning at least 75 million years between the first known rhabdodontid and their Middle Jurassic ancestor before they supposedly split off from the rest of the iguanodontians.[4][3][5][DN 2] However, although he stated that the idea influenced the depiction of Zalmoxes as the last of an ancient lineage on the show, Dr. Darren Naish did consider the hypothesis debatable, likely in part due to some studies classifying some ornithopods from the Early Cretaceous and the start of the Late Cretaceous (e.g. an unnamed ornithopod from the Vegagete dig site in the Castrillo de la Reina Formation of Spain, Muttaburrasaurus of Australia, Tenontosaurus and Iani smithi of North America) as rhabdodontomorphs, closely related to (but not actual members of) the rhabdodontid family, which may or may not imply that true rhabdodontids did not appear until their supposed cousins came into existence, overall making rhabdodontid ancestry more mysterious, bringing about more questions regarding when they really did diverge from the other iguanodontian clades.[6][7][8][5][DN 2]

Paleofauna
[]

Juveniles Zalmoxes look around, checking for incoming threats
Juveniles Zalmoxes look around, checking for incoming threats
“ Seven inches tall, baby Zalmoxes are no more than snacks for many predators... but snacks are nonetheless worth eating, so they have to be cautious. ”

David Attenborough, Forests

Regardless of whether or not it underwent insular dwarfism like most of the other animals in the same place and time, Zalmoxes is one of the smallest residents of Hațeg Island. It coexisted with other herbivores like the hadrosauromorph Telmatosaurus, titanosaurs like Magyarosaurus and Paludititan, the nodosaur Struthiosaurus (also found in Austria, France, and Hungary), the alvarezsaurid Bradycneme (as well as its possible junior synonym, Heptasteornis, if considered valid), and Transylvanosaurus, another rhabdodontid. These different herbivores likely occupy different niches, feeding on different food and otherwise not competing for the exact same resources. Hațeg Island is also home to various multituberculate mammals, lizards, turtles, anurans (frogs and toads), and birds from the small Balaur bondoc (once interpreted as a dromaeosaur, possibly a junior synonym of Elopteryx) to the massive Gargantuavis, both of which appear to be flightless. Given its small size and lack of armor, Zalmoxes is considered a light, easy snack by various predators like the madtsoiid snake Nidophis and crocodylomorphs (e.g. Allodaposuchus, Aprosuchus, and Sabresuchus). Threats can also come from the sky in the form of azhdarchid pterosaurs, with at least three (as well as a fourth, unnamed medium-sized azhdarchid with a robust neck) prowled Hațeg Island, Albadraco, Eurazhdarcho, and the giraffe-sized Hatzegopteryx, apex predator of the region (and possibly also the top hunter of other European islands).[DN 3] Due to its extreme vulnerability, this rhabdodontid needed to be very wary of its surroundings, preferring to mostly stay under the cover of vegetation, quickly crossing open areas to minimize the chances of being caught. Zalmoxes also relies on herding behavior, for safety in numbers protects the many at the expense of the few who are targeted by predators.

Since Europe was mostly a sea during the Cretaceous Period, and land bridges did not seem common by that time, the only way that the non-flying inhabitants of the European islands can spread is via island hopping and rafting, traveling intentionally or accidentally on floating masses of land. Given its small size, Zalmoxes can easily take advantage of the latter method, but this does put it at risk of being caught by mosasaurs like Prognathodon in the open sea.

As one of the last non-avian dinosaurs, Zalmoxes, along with most of the animals it coexisted with, would have bore witness to the Cretaceous-Paleogene Extinction Event 66.043 million years ago,[9] a catastrophe that ultimately wiped out three-quarters of all life on Earth, including all non-avian dinosaurs.

Appearances[]

Forests[]

Juveniles Zalmoxes rushing down a fallen tree, hoping to make it to the other side without getting spotted by hunters
Juveniles Zalmoxes rushing down a fallen tree, hoping to make it to the other side without getting spotted by hunters

The seventh, final segment of Forests takes place in the dark understory of a forest in Europe. As a Telmatosaurus keeps a low profile amongst the lush vegetation, five juvenile Zalmoxes warily look around, as they are about to cross a log over the river, out in a more open area of the forest. As they rush down a log, the last one to cross is suddenly snatched up and swallowed alive by a Hatzegopteryx. Gulping his snack down, the majestic pterosaur slowly weaves through the forest, at one point stopping by to check out one moss-covered tree before continuing his casual but regal walk, eventually making it to the beach, where he spreads his large wings, flapping them to keep them warmed up before picking at them, grooming himself.

A group of Zalmoxes, visiting the sea spray-coated plants by the beaches of Hațeg Island
A group of Zalmoxes, visiting the sea spray-coated plants by the beaches of Hațeg Island
“ Many of the forest's permanent residents regularly come down here, because the plants are coated with sea spray, and are a welcome source of salt. Here, sauropods can meet one another, renewing family bonds, and creating new ones. ”

David Attenborough, Forests

Some Zalmoxes are seen amongst the foliage, looking around as Telmatosauruses feed on the salted vegetation, with one even licking a propped-up piece of wood on the beach. Two titanosaurs make their way down to the beach, affectionately rubbing their heads and necks against one another. And as the sun sets, Hatzegopteryx spreads his wings once more and finally leaves the island, flying off into the horizon.

Islands[]

Zalmoxes sees a larger rafting island up ahead
Zalmoxes sees a larger rafting island up ahead
“ The mouth of a river in southern Europe, 66 million years ago. In the aftermath of a tropical storm, debris is drifting downstream. And amongst the wreckage, rafts of wreckage ripped from the land. For a weary pterosaur, this one may be a welcome place to rest. But it's far from safe... ”

David Attenborough, Islands

In the first segment of Islands, a weary adult Alcione rests on a collection of vegetation in southern Europe, a raft drifting out of a river into the sea. For a moment, it feels safe, flapping its wings as the raft drifts on peacefully. But suddenly, sensing that something is wrong, the Alcione takes to the skies just as a large marine predator lunges at its raft.

Zalmoxes prepares to jump off his raft, aware of the Prognathodon closing in on him
Zalmoxes prepares to jump off his raft, aware of the Prognathodon closing in on him
“ A huge mosasaur, a deadly underwater hunter, is looking for an easy meal... such as this little dinosaur, Zalmoxes. Staying here is too risky. He needs a larger raft, and quickly. There's little choice but to swim for it. ”

David Attenborough, Islands

Seeing yet another easy meal, the predator, a Prognathodon,[DN 4] approaches another raft, where a Zalmoxes begins to panic. Barely able to keep his balance on the drifting log, the ornithopod jumps into the water as the Prognathodon closes in on him. Paddling frantically, he manages to clamber up on a larger drifting island, retreating into the safety of the trees. However, he senses that he is not the first on the large raft, and looks around, sniffing.

Zalmoxes bonding with each other
Zalmoxes bonding with each other
“ He is not the first here. Another castaway, a female. Their raft is drifting out to sea. Sometimes, castaways on rafts like these are washed up on the shores of distant islands. If they're very lucky, this pair could become pioneers... and may establish a new population, which eventually becomes a new species, unlike anything seen before. ”

David Attenborough, Islands

Emerging from the vegetation is a female Zalmoxes, which turns towards him in surprise. Together, the pair look out to the open seas, then turn to each other, nuzzling. The segment ends with the two Zalmoxes continuing to look ahead of them, with the waters stretching out for miles around them, and the afternoon sun shining down on their raft.

Gallery[]

References[]

General[]

Dr. Darren Naish[]

Prehistoric Planet[]

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