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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 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

― 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.

― 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[]

― 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[]

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.

― 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[]

― 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.

― 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.

― 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[]
- ↑ 1.0 1.1 1.2 The Princeton Field Guide to Dinosaurs
- ↑ 2.0 2.1 Bone histology in the Ornithopods from the Maastrichtian of Hațeg Basin – were these dinosaurs really dwarfs?
- ↑ 3.0 3.1 Phylogeny, Histology and Inferred Body Size Evolution in a New Rhabdodontid Dinosaur from the Late Cretaceous of Hungary
- ↑ 4.0 4.1 Osteology and phylogeny of Zalmoxes (n. g.), an unusual Euornithopod dinosaur from the latest Cretaceous of Romania
- ↑ 5.0 5.1 The Rhabdodontidae (Dinosauria, Ornithischia), an enigmatic dinosaur group endemic to the Late Cretaceous European Archipelago
- ↑ Phylogeny of iguanodontian dinosaurs and the evolution of quadrupedality
- ↑ Histogenesis and growth dynamics of the tiny Vegagete rhabdodontomorph hindlimb (Ornithischia, Ornithopoda): Paleoecological and evolutionary implications
- ↑ An early-diverging iguanodontian (Dinosauria: Rhabdodontomorpha) from the Late Cretaceous of North America
- ↑ Dinosaur extinction battle flares
Dr. Darren Naish[]
- ↑ It has been suggested that the small size of Zalmoxes was an adaptation for life on an island, but it may have simply inherited its small size from its ancestors.
- ↑ 2.0 2.1 Rhabdodontids diverged relatively early on from other iguanodontians, probably around the Middle Jurassic Period, making them more primitive compared to other iguanodontians, especially hadrosaurs, which came long after they split off. It is therefore proposed that rhabdodontids were "living fossils" during the Late Cretaceous Period, standing out as they heavily reflected the appearance of their Jurassic ancestors. This idea, however, is up for debate.
- ↑ Hatzegopteryx was likely an island-hopping pterosaur, an idea backed up by the remains of similar azhdarchids found across Europe.
- ↑ The mosasaur in the first segment of Islands is Prognathodon.
Prehistoric Planet[]
- ↑ 1.0 1.1 1.2 I Know Dino: The Big Dinosaur Podcast, Episode 443: Dinosaurs on Islands. Featuring Prehistoric Planet 2
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