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''Morrosaurus'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of herbivorous
elasmaria Elasmaria is a clade of Ornithopoda, ornithopods known from Cretaceous deposits in the former Gondwana (South America, Antarctica, Australia, and possibly Africa) that contains many bipedal ornithopods that were previously considered Hypsilophodo ...
n
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
that lived in the late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
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 (also known as the Antarctic Ocean), it contains the geographic South Pole. ...
. The only known species is the
type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * ...
''Morrosaurus antarcticus''.


Discovery

In 2002, the Argentine paleontologist Fernando Novas 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–so ...
, where the remains of the species were found. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
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 International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
age. The remains consists in a right hind leg, including the top of the
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
, the lower end of the femur, the upper part of the
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
, 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 a genus of ornithopod dinosaur that lived during the late Campanian stage of the Late Cretaceous, Upper Cretaceous, around 73 to 72 million years ago in what is now James Ross Island off the coast of northern Antarctica near Pa ...
'', 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, 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'', '' 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 spe ...
'' 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 elasmarian ornithopod dinosaur found in Late Cretaceous rocks of South Africa. It is known from a tooth and possibly some postcranial remains dating between the middle-Campanian to ...
'', 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 has a triangular shape from the front and a concave surface from the same angle.


Phylogeny

''Morrosaurus'' was classified in the group
Iguanodontia Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively sm ...
, as a basal member of Euiguanodontia. This in turn formed a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
with other ornithopods of
Patagonia Patagonia () is a geographical region that includes parts of Argentina and Chile at the southern end of South America. The region includes the southern section of the Andes mountain chain with lakes, fjords, temperate rainforests, and glaciers ...
and Antarctica, particularly ''
Trinisaura ''Trinisaura'' is a genus of ornithopod dinosaur that lived during the late Campanian stage of the Late Cretaceous, Upper Cretaceous, around 73 to 72 million years ago in what is now James Ross Island off the coast of northern Antarctica near Pa ...
'', ''
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 spe ...
'', '' Anabisetia'', '' Notohypsilophodon'', ''
Talenkauen ''Talenkauen'' is a genus of Basal (phylogenetics), 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 Ma ...
'' and ''
Macrogryphosaurus ''Macrogryphosaurus'' (meaning "big enigmatic lizard") is a genus of elasmarian dinosaur from the Coniacian age Upper Cretaceous Sierra Barrosa Formation (Neuquén Group) of Argentina in Patagonia. It was described by Jorge Calvo and colleagues ...
'' in a group called
Elasmaria Elasmaria is a clade of Ornithopoda, ornithopods known from Cretaceous deposits in the former Gondwana (South America, Antarctica, Australia, and possibly Africa) that contains many bipedal ornithopods that were previously considered Hypsilophodo ...
, 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, 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 The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
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 language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
based in the
phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
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 ...
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 "high-crested 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 this ...
''. In 2020 a study was conducted on the growth rate of ''Morrosaurus'', and its relative ''
Trinisaura ''Trinisaura'' is a genus of ornithopod dinosaur that lived during the late Campanian stage of the Late Cretaceous, Upper Cretaceous, around 73 to 72 million years ago in what is now James Ross Island off the coast of northern Antarctica near Pa ...
'', through use of
histological Histology, also known as microscopic anatomy or microanatomy, is the branch of biology that studies the microscopic anatomy of biological tissue (biology), tissues. Histology is the microscopic counterpart to gross anatomy, which looks at large ...
study of thin samples from the holotype and fourth . Ornithischian dinosaurs living in polar regions in the Northern Hemisphere (such as ''
Edmontosaurus ''Edmontosaurus'' ( ) (meaning "lizard from Edmonton"), with the second species often colloquially and historically known as ''Anatosaurus'' or ''Anatotitan'' (meaning "duck lizard" and "giant duck"), is a genus of hadrosaurid (duck-billed) din ...
'' and '' Pachyrhinosaurus'') 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 spe ...
'' 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''.


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 Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
. The distinction between fauna of this part of the world as compared with the northern hemisphere equivalent
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
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 () is a geographical region that includes parts of Argentina and Chile at the southern end of South America. The region includes the southern section of the Andes mountain chain with lakes, fjords, temperate rainforests, and glaciers ...
, 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 elasmarian ornithopod dinosaur found in Late Cretaceous rocks of South Africa. It is known from a tooth and possibly some postcranial remains dating between the middle-Campanian to ...
'' from Africa. Similar biogeographic patterns have been observed in
megaraptora Megaraptora is a clade of carnivorous theropod dinosaurs. Its derived members, the Megaraptoridae are noted for their large hand claws and powerfully-built forelimbs, which are usually reduced in size in other large theropods. Although undoubt ...
n theropods, diamantinasaurian sauropods, and
parankylosauria Parankylosauria is a group of Basal (phylogenetics), 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 ...
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, crown-group medusozoan by Han ''et al.'' (2016). * A study on the fossil corals fro ...


References

{{Taxonbar, from=Q21122031 Elasmaria Dinosaur genera Maastrichtian dinosaurs Dinosaurs of Antarctica Fossil taxa described in 2016 Taxa named by Fernando Novas