Morrosaurus
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''Morrosaurus'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of herbivorous
elasmaria Elasmaria is a clade of ornithopods known from Cretaceous deposits in South America, Antarctica, and Australia that contains many bipedal ornithopods that were previously considered "hypsilophodonts".Madzia, Daniel; Boyd, Clint A.; Mazuch, Marti ...
n
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
that lived in the late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
in
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest cont ...
. The only known species is the type ''Morrosaurus antarcticus''.


Discovery

In 2002, the Argentine paleontologist
Fernando Novas Fernando Emilio Novas (born 1960) is an Argentine paleontologist working for the Comparative Anatomy Department of the Bernardino Rivadavia Natural Sciences Museum in Buenos Aires, Argentina.
reported the discovery of a partial skeleton of a euornithopod in Antarctica. In 2016 these remains were the basis for naming the type species ''Morrosaurus antarcticus'', named and described by Sebastian Rozadilla, Federico Lisandro Agnolin, Fernando Emilio Novas, Alexis Rolando Aranciaga Mauro, Matthew J. Motta, Juan Manuel Lirio Marcelo, and Pablo Isasi. The genus name refers to the site of El Morro on
James Ross Island James Ross Island is a large island off the southeast side and near the northeastern extremity of the Antarctic Peninsula, from which it is separated by Prince Gustav Channel. Rising to , it is irregularly shaped and extends in a north–south ...
, where the remains of the species were found. The specific name refers to Antarctica. The holotype specimen MACN Pv 197, was found in a layer of the
Snow Hill Island Formation The Snow Hill Island Formation is an Maastrichtian, Early Maastrichtian geologic Formation (geology), formation found on James Ross Island, James Ross Island group, Antarctica. Remains of a Paraves, paravian Theropoda, theropod ''Imperobator anta ...
(Cape Lambe, previously assigned to the Lopez de Bertodano Formation), dating to the
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval ...
age. The remains consists in a right hind leg, including the top of the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
, the lower end of the femur, the upper part of the
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
, the underside of the tibia, the upper half of the foot, the bottom of the midfoot and the top of the first joint of the third toe.


Description

The only known specimen, thought to represent an adult, is estimated to have belong to an individual in length. This makes it larger and more robust in skeletal build than many of its relatives, including the fellow Antarctic genus ''
Trinisaura ''Trinisaura'' is an extinct genus of ornithopod dinosaur known from the lower levels of the Late Cretaceous Snow Hill Island Formation (early Maastrichtian stage) of James Ross Island, Antarctica. It contains a single species, ''Trinisaura sant ...
'', only in length. The fragmentary nature of the material precludes saying much about the preserved regions. It is united anatomically with other elasmarians by the strongly compressed outer front edge of the , lacking the distinct cleft and thick edge seen in other types of ornithopod. Other uniting traits of elasmarians seen in the and cannot be evaluated in ''Morrosaurus'' due to their lack of preservation in the genus. The seems to have been rather gracile, though more stout than in ''Trinisaura'', as was the elongate foot, with the central third being larger than the others. A series of distinctive features characterizes the genus. Two of these are
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
, completely unique traits acquired by the taxon. In bottom view, the greater trochanter near the top of the femur forms an S-shape, with the back edge thick across and the front edge thin, similar to the condition seen in other elasmarians ''
Anabisetia ''Anabisetia'' ( ) is a genus of ornithopod dinosaur from the Late Cretaceous Period of Patagonia, South America. It was a small bipedal herbivore, around long. Discovery Argentine paleontologists Rodolfo Coria and Jorge Orlando Calvo nam ...
'', '' Notohypsilophodon'', and ''
Gasparinisaura ''Gasparinisaura'' (meaning "Gasparini's lizard") is a genus of herbivorous ornithopod dinosaur from the Late Cretaceous. The first fossils of ''Gasparinisaura'' were found in 1992 near Cinco Saltos in Río Negro Province, Argentina. The type sp ...
'' but displayed to a more extreme extent. The fourth metatarsal bone has a prominent rearward projection that wraps around the third metatarsal, likewise similar but more developed to the condition seen in ''Gasparinisaura'' as well as ''
Kangnasaurus ''Kangnasaurus'' (meaning "Farm Kangnas lizard") is a genus of iguanodontian ornithopod dinosaur found in supposedly Early Cretaceous rocks of South Africa. It is known from a tooth and possibly some postcranial remains found in the early- Aptia ...
'', a genus with overall similar morphology and proportions to ''Morrosaurus''. In addition, there is a unique combination of two features that by themselves are not unique features. In the femur, the is positioned forward and to the outside of the greater trochanter. In the tibia, medial
malleolus A malleolus is the bony prominence on each side of the human ankle. Each leg is supported by two bones, the tibia on the inner side (medial) of the leg and the fibula on the outer side (lateral) of the leg. The medial malleolus is the promine ...
has a triangular shape from the front and a concave surface from the same angle.


Phylogeny

''Morrosaurus'' was classified in the group
Iguanodontia Iguanodontia (the iguanodonts) is a clade of herbivorous dinosaurs that lived from the Middle Jurassic to Late Cretaceous. Some members include ''Camptosaurus'', ''Dryosaurus'', ''Iguanodon'', '' Tenontosaurus'', and the hadrosaurids or "duck-bil ...
, as a basal member of Euiguanodontia. This in turn formed a clade with other ornithopods of
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and g ...
and Antarctica, particularly ''
Trinisaura ''Trinisaura'' is an extinct genus of ornithopod dinosaur known from the lower levels of the Late Cretaceous Snow Hill Island Formation (early Maastrichtian stage) of James Ross Island, Antarctica. It contains a single species, ''Trinisaura sant ...
'', ''
Gasparinisaura ''Gasparinisaura'' (meaning "Gasparini's lizard") is a genus of herbivorous ornithopod dinosaur from the Late Cretaceous. The first fossils of ''Gasparinisaura'' were found in 1992 near Cinco Saltos in Río Negro Province, Argentina. The type sp ...
'', ''
Anabisetia ''Anabisetia'' ( ) is a genus of ornithopod dinosaur from the Late Cretaceous Period of Patagonia, South America. It was a small bipedal herbivore, around long. Discovery Argentine paleontologists Rodolfo Coria and Jorge Orlando Calvo nam ...
'', '' Notohypsilophodon'', ''
Talenkauen ''Talenkauen'' is a genus of basal iguanodont dinosaur from the Campanian or Maastrichtian age of the Late Cretaceous Cerro Fortaleza Formation, formerly known as the Pari Aike Formation of Patagonian Lake Viedma, in the Austral Basin of Sa ...
'' and '' Macrogryphosaurus'' in a group called
Elasmaria Elasmaria is a clade of ornithopods known from Cretaceous deposits in South America, Antarctica, and Australia that contains many bipedal ornithopods that were previously considered "hypsilophodonts".Madzia, Daniel; Boyd, Clint A.; Mazuch, Marti ...
, whose members are distinguished by their adaptation to a running lifestyle which would be reflected by the narrow foot with a thin fourth metatarsal which indicates a high speed running; subsequently expanded
chevrons Chevron (often relating to V-shaped patterns) may refer to: Science and technology * Chevron (aerospace), sawtooth patterns on some jet engines * Chevron (anatomy), a bone * '' Eulithis testata'', a moth * Chevron (geology), a fold in rock l ...
, a feature that is associated with a greater surface area for attachment of the lateral muscles of the tail, which would give good control of the movements of this; and a curved humerus which demonstrates the absence of a deltopectoral ridge and therefore that the front leg was not used for walking. It cannot be determined, however, if ''Morrosaurus'' itself possessed these characteristics due to their limited remains. The existence of this clade may indicate that Patagonia, Antarctica and Australia shared the same type of fauna. The exact phylogenetic relationships within this clade could not be identified, except for ''Gasparinisaura'', which proved to be the most basal member of group.
Cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
based in the
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
of Rozadilla ''et al''., 2016:


Palaeobiology

Based on the hindlimb anatomy of ''Morrosaurus'' and other elasmarians, notably the slim , it's thought they were very capable runners, distinguishing them from other herbivorous dinosaurs they lived alongside. Elements of relatives not preserved in ''Morrosaurus'' such as expanded and distinctive bony intercostal plates along the torso would've further aided their
cursorial A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often us ...
abilities by improving tail balance and musculature control as well as breathing efficiency. This developed tail musculature and running ability of elasmarians is noted as more developed in elasmarian taxa than in other ornithopods such as ''
Hypsilophodon ''Hypsilophodon'' (; meaning "''Hypsilophus''-tooth") is a neornithischian dinosaur genus from the Early Cretaceous period of England. It has traditionally been considered an early member of the group Ornithopoda, but recent research has put thi ...
''. In 2020 a study was conducted on the growth rate of ''Morrosaurus'', and its relative ''
Trinisaura ''Trinisaura'' is an extinct genus of ornithopod dinosaur known from the lower levels of the Late Cretaceous Snow Hill Island Formation (early Maastrichtian stage) of James Ross Island, Antarctica. It contains a single species, ''Trinisaura sant ...
'', through use of
histological Histology, also known as microscopic anatomy or microanatomy, is the branch of biology which studies the microscopic anatomy of biological tissues. Histology is the microscopic counterpart to gross anatomy, which looks at larger structures vis ...
study of thin samples from the holotype and fourth . Ornithischian dinosaurs living in polar regions in the Northern Hemisphere (such as '' Edmontosaurus'' and ''
Pachyrhinosaurus ''Pachyrhinosaurus'' (meaning in Greek "thick-nosed lizard", from ' (), thick; ' (), nose; and (), lizard) is an extinct genus of centrosaurine ceratopsid dinosaur from the Late Cretaceous period of North America. The first examples were discove ...
'' typically show different growth rates than relatives from lower latitutes, owing to the extreme temperatures of their environments, but studies have failed to find the same signs in Antarctic animals. The study confirmed this pattern in elasmarians, with comparison with South American elasmarian ''
Gasparinisaura ''Gasparinisaura'' (meaning "Gasparini's lizard") is a genus of herbivorous ornithopod dinosaur from the Late Cretaceous. The first fossils of ''Gasparinisaura'' were found in 1992 near Cinco Saltos in Río Negro Province, Argentina. The type sp ...
'' finding incredibly similar histology to the Antarctic genera. All three show rapid growth with periodic interruptions, variation in growth marks, and changes in growth rate across their life cycle, growing slower as they aged. Similar growth strategies are reported in other ornithopods from much lower latitudes. This indicates that rather than adapting their growth physiology in response to the cold, southern environments they lived in, elasmarians were able to succeed in such ecosystems due to the existing possession of necessary adaptations to thrive there. Higher growth rate was noted in ''Morrosaurus'' as compared to ''Trinisaura'', consistent with being a larger species. Additionally, it was found the animals reached their sexual maturity well before their skeletal maturity, and ceased growth at a later point than did, consistent with other dinosaurs. Neonatal fossils of elasmarians from Australia, which offer strong evidence of elasmarians living year round in high-latitude conditions rather than being migratory in nature, were similarly found to display growth rates very similar to those of other ornithopods such as ''
Orodromeus ''Orodromeus'' (meaning "Mountain Runner") is a genus of herbivorous orodromine thescelosaurid dinosaur from the Late Cretaceous of North America. Only one species is known, the type species ''Orodromeus makelai''. Discovery and naming The rem ...
''.


Palaeobiogeography

The discovery of ''Morrosaurus'' was concurrent with the first evidence of a previously hypothesized uniting clade of ornithopod dinosaurs from the continents that had comprised Gondwana. The distinction between fauna of this part of the world as compared with the northern hemisphere equivalent Laurasia had long been noted, but evidence was traditionally lacking for a biogeographic link for dinosaurs between the different Gondwanan regions. Brazil and Northern Africa shared a fauna distinct from that of
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and g ...
, and each of these distinct from that of India and Madagascar; likewise, Australia, New Zealand, and Antarctic seemed connected to the exclusion of the others. This contrasted with existing recognition of links between the flora and marine invertebrate fauna between Patagonia, Antarctica, Australia, and New Zealand, termed the Weddellian Bioprovince. The recognition of ''Morrosaurus'' and ''Trinisaura'' as Antarctic elasmarians closely related to animals from South America constituted important evidence of such ties between these continents existing in dinosaurs as well. This would be bolstered by later studies producing firmer evidence of Australian ornithopods as also belonging to the group, as well as possibly ''
Kangnasaurus ''Kangnasaurus'' (meaning "Farm Kangnas lizard") is a genus of iguanodontian ornithopod dinosaur found in supposedly Early Cretaceous rocks of South Africa. It is known from a tooth and possibly some postcranial remains found in the early- Aptia ...
'' from Africa. Similar biogeographic patterns have been observed in
megaraptora Megaraptora is a clade of carnivorous Tetanurae, tetanuran theropod dinosaurs with controversial relations to other theropods. Its Derived (phylogenetics), derived members, the Megaraptoridae are noted for their elongated hand claws and proporti ...
n theropods, diamantinasaurian sauropods, and
parankylosauria Parankylosauria is a group of basal ankylosaurian dinosaurs known from the Cretaceous of South America, Antarctica, and Australia. It is thought the group split from other ankylosaurs during the mid-Jurassic period, despite this being unpreserved ...
n ankylosaurs. These discoveries of connected endemic Gondwanan ecosystems have overturned the traditional view of the southern continents, including the fauna seen in Late Cretaceous Antarctic, acting as a refugium for animals more successful elsewhere earlier in the Cretaceous.


See also

* South Polar region of the Cretaceous *
2016 in paleontology Flora Plants Fungi Cnidarians Research * '' Yunnanoascus haikouensis'', previously thought to be a member of Ctenophora, is reinterpreted as a crown-group medusozoan by Han ''et al.'' (2016). * A study on the fossil corals from the Late Tr ...


References

{{Taxonbar, from=Q21122031 Ornithopods Late Cretaceous dinosaurs Maastrichtian life Dinosaurs of Antarctica Fossil taxa described in 2015 Taxa named by Fernando Novas Ornithischian genera