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''Antarctopelta'' ( ; meaning 'Antarctic shield') was a
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 nomencla ...
of
ankylosauria Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. ...
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 ...
with one known
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
, ''A. oliveroi'', which lived 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 contine ...
during the Late
Cretaceous Period 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 th ...
. It was a medium-sized ankylosaur, reaching 4 meters (13 ft) in length or more, and showed characteristics of two different families, making more precise classification difficult. The single known
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
specimen was discovered 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 ...
in 1986, constituting the first dinosaur remains ever discovered on Antarctica, although it is the second dinosaur from the continent to be formally named.


Discovery and naming

The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
(specimen used as the basis for the taxon) is the only known example of this genus and species, and was the first dinosaur ever found in Antarctica. It consists of three isolated teeth, part of the
lower jaw In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
with another tooth ''in situ'', some other skull fragments, vertebrae (neck, back, hips and tail), some partial limb bones (
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...
, ilium and
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 with ...
), toe bones (five
metapodial Metapodials are long bones of the hand (metacarpals) and feet (metatarsal The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes ...
s and two
phalanges The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...
), and numerous pieces of armor, representing approximately 15% of the skeleton. This specimen was initially located in January 1986 on James Ross Island, off the
Antarctic Peninsula The Antarctic Peninsula, known as O'Higgins Land in Chile and Tierra de San Martín in Argentina, and originally as Graham Land in the United Kingdom and the Palmer Peninsula in the United States, is the northernmost part of mainland Antarctic ...
. It was discovered by
Argentine Argentines (mistakenly translated Argentineans in the past; in Spanish (masculine) or (feminine)) are people identified with the country of Argentina. This connection may be residential, legal, historical or cultural. For most Argentines, s ...
geologist A geologist is a scientist who studies the solid, liquid, and gaseous matter that constitutes Earth and other terrestrial planets, as well as the processes that shape them. Geologists usually study geology, earth science, or geophysics, alth ...
s Eduardo Olivero and Roberto Scasso, but excavation was not completed for almost a decade due to the frozen ground and harsh weather conditions. The material was collected from an area of 6 square meters (64.5 sq ft) over several field seasons but is assumed to have belonged to a single individual. Much of the skeleton is in poor condition, as many of the bones nearest to the surface were subjected to years of fragmentation by freeze-thaw weathering. Although the material had been known for decades and written about in three separate publications, ''Antarctopelta oliveroi'' was not named until 2006, by Argentine
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of foss ...
s
Leonardo Salgado Leonardo Salgado is an Argentinean palaeontologist with a special interest in dinosaurs of the Cretaceous period and other investigations of the palaeobiology of fossil bearing geological formations. Salgado is the leading or coauthor of several ta ...
and Zulma Gasparini. It was therefore the second named genus of dinosaur from Antarctica after ''
Cryolophosaurus ''Cryolophosaurus'' ( or ; "CRY-oh-loaf-oh-SAWR-us") is a genus of large theropod dinosaur known from only a single species ''Cryolophosaurus ellioti'', from the early Jurassic of Antarctica. It was one of the largest theropods of the Early Jur ...
'' in 1993, despite being discovered first. The genus name refers to its location on the continent of Antarctica and its
armored Armour (British English) or armor (American English; see spelling differences) is a covering used to protect an object, individual, or vehicle from physical injury or damage, especially direct contact weapons or projectiles during combat, or ...
nature. Antarctica is derived from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
words ''αντ''/''ant-'' ('opposite of') and ''αρκτος''/''arktos'' ('bear' referring to the constellation
Ursa Major Ursa Major (; also known as the Great Bear) is a constellation in the northern sky, whose associated mythology likely dates back into prehistory. Its Latin name means "greater (or larger) bear," referring to and contrasting it with nearby Ursa ...
, which points north). The Greek ''πελτη''/''pelte'' ('shield') is commonly used to name genera of ankylosaurs (''
Cedarpelta ''Cedarpelta'' is a extinct genus of basal ankylosaurid dinosaur from Utah that lived during the Late Cretaceous period (Cenomanian to lower Turonian stage, 98.2 to 93 Ma) in what is now the Mussentuchit Member of the Cedar Mountain Formation. Th ...
'' and '' Sauropelta'', for example). The single known species, ''A. oliveroi'', is named after Eduardo Olivero, who discovered the holotype, first mentioned it in print, and has worked in Antarctica for decades. Earlier work suggested that the James Ross Island ankylosaur was a juvenile. More recent research indicates that the different parts of the vertebrae are completely fused together, while a juvenile would be expected to have visible sutures between the
neural arch The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...
and
body Body may refer to: In science * Physical body, an object in physics that represents a large amount, has mass or takes up space * Body (biology), the physical material of an organism * Body plan, the physical features shared by a group of animal ...
(''centrum'') of the vertebrae. A preliminary
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 ...
analysis of several bones also indicates a level of
remodeling Renovation (also called remodeling) is the process of improving broken, damaged, or outdated structures. Renovations are typically done on either commercial or residential buildings. Additionally, renovation can refer to making something new, ...
that would not be seen in newly formed bone.


Description

Like other ankylosaurs, ''Antarctopelta oliveroi'' was a stocky,
herbivorous A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
quadruped Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor ...
protected by
armor plates Military vehicles are commonly armoured (or armored; see spelling differences) to withstand the impact of shrapnel, bullets, shells, rockets, and missiles, protecting the personnel inside from enemy fire. Such vehicles include armoured fightin ...
embedded in the skin. Although a complete
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
has not been found, the species is estimated to have reached a maximum length of 4 meters (13 ft) from snout to tail tip. In 2010
Gregory Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology, and more recently has examined sociology and theology. He is best known for his work and research on theropod dino ...
gave a higher estimation of 6 meters (20 ft) and 350 kg (772 lbs). Very little of the
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
is known, but all of the known skull fragments were heavily
ossified Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in th ...
for protection. One bone in particular, identified as a ''supraorbital'', included a short spike which would have projected outwards over the eye. The
leaf A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
-shaped
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tea ...
are
asymmetrical Asymmetry is the absence of, or a violation of, symmetry (the property of an object being invariant to a transformation, such as reflection). Symmetry is an important property of both physical and abstract systems and it may be displayed in pre ...
, with the majority of the denticles on the edge closest to the tip of the snout. These teeth are also proportionately large compared to those of other ankylosaurs, with the largest measuring 10 millimeters (0.4 in) across. This compares to the much larger
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the C ...
n ''
Euoplocephalus ''Euoplocephalus'' ( ) is a genus of very large, herbivorous ankylosaurid dinosaurs, living during the Late Cretaceous of Canada. It has only one named species, ''Euoplocephalus tutus''. The first fossil of ''Euoplocephalus'' was found in 1897 i ...
'', 6–7 m (20–23 ft) in body length, which had teeth averaging only 7.5 mm (0.3 in) across. Six different types of osteoderms were found along with the skeletal remains of ''Antarctopelta'', but very few were articulated with the skeleton, so their placement on the body is largely speculative. They included the base of what would have been a large spike. Flat oblong plates resembled the ones that guarded the neck of the
nodosaurid Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous period in what is now North America, South America, Europe, and Asia. Description Nodosaurids, like their close relatives the ankylosaurids, we ...
'' Edmontonia rugosidens''. Large circular plates were found associated with smaller,
polygon In geometry, a polygon () is a plane figure that is described by a finite number of straight line segments connected to form a closed ''polygonal chain'' (or ''polygonal circuit''). The bounded plane region, the bounding circuit, or the two to ...
al nodules, perhaps forming a shield over the hips as seen in '' Sauropelta''. Another type of osteoderm was oval-shaped with a keel running down the middle. A few examples of this fifth type were found ossified to the
rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ch ...
s, suggesting that they ran in rows along the flanks of the animal, a very typical pattern among ankylosaurs. The final group consisted mainly of small bony nodules which are often called ''ossicles'', and were probably scattered throughout the body. Several ribs were also found with these ossicles attached.
Vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
e from sections of the tail were found. Although the tip of the tail did not fossilize, some of the smaller vertebrae recovered would have been situated near the end of the tail in life, and these were associated with ossified
tendons A tendon or sinew is a tough, high-tensile-strength band of dense fibrous connective tissue that connects muscle to bone. It is able to transmit the mechanical forces of muscle contraction to the skeletal system without sacrificing its ability ...
on the upper and lower sides. In
ankylosaurid Ankylosauridae () is a family of armored dinosaurs within Ankylosauria, and is the sister group to Nodosauridae. The oldest known Ankylosaurids date to around 122 million years ago and went extinct 66 million years ago during the Cretaceous–Pal ...
s, these tendons help to stiffen the end of the tail in support of a large, bony tail club. However, large osteoderms and specialized tail vertebra indicate that ''Antarctopelta'' may have in fact had a flat, frond-like tail structure resembling a
macuahuitl A macuahuitl () is a weapon, a wooden club with several embedded obsidian blades. The name is derived from the Nahuatl language and means "hand-wood". Its sides are embedded with prismatic blades traditionally made from obsidian. Obsidian i ...
, akin to that of '' Stegouros'', its closest relative.


Classification

''Antarctopelta'' shares several features with the nodosaurids, mainly in the teeth and armor, while the possibly-clubbed tail is far more similar to those of ankylosaurids. This mosaic of characters makes assignment to a specific family difficult. It had been designated as
Ankylosauria Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. ...
''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertaint ...
'' before being subjected to a
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 ...
. Later, a phylogenetic analysis performed by Thompson ''et al.'', 2011 suggested that ''Antarctopelta'' was the
basalmost In phylogenetics, basal is the direction of the ''base'' (or root) of a rooted phylogenetic tree or cladogram. The term may be more strictly applied only to nodes adjacent to the root, or more loosely applied to nodes regarded as being close to th ...
known
nodosaurid Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous period in what is now North America, South America, Europe, and Asia. Description Nodosaurids, like their close relatives the ankylosaurids, we ...
. However, a 2021 study found ''Antarctopelta'' to belong to a distinctive basal lineage of Southern Hemisphere
ankylosaurs Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. ...
, the
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 unpreser ...
. The following
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 ...
is reproduced from 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 ...
in the 2021 study by Sergio Soto-Acuña and colleagues:


Paleoecology

The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
skeleton was collected about 90 m (300 ft) from the base of the Gamma Member of the Santa Marta Formation in Antarctica. This
member Member may refer to: * Military jury, referred to as "Members" in military jargon * Element (mathematics), an object that belongs to a mathematical set * In object-oriented programming, a member of a class ** Field (computer science), entries in ...
was deposited in a shallow marine environment and also preserves marine fossils such as
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimor ...
teeth, remains of the
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on the ...
'' Taniwhasaurus'',
ammonite Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttlefish ...
s,
bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, biva ...
s, and
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. The ...
s.
Index fossil Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictionary of Bi ...
s like ammonites suggest the rocks were deposited in the early
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 from ...
stage Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly British theatre newspaper * St ...
of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
Period, or about 71.2–70.8 Ma (
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
). Despite being found in marine sediment, ''Antarctopelta'', like all ankylosaurs, lived on land. Other ankylosaurs have also been found in marine sediments, likely as a result of carcasses washing out to sea. Although Antarctica in the Cretaceous was in the southern
polar region The polar regions, also called the frigid zones or polar zones, of Earth are the regions of the planet that surround its geographical poles (the North and South Poles), lying within the polar circles. These high latitudes are dominated by floa ...
, the Earth had a much warmer
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
during this time period, and the continent would have been ice-free. Animals like ''Antarctopelta oliveroi'' would have lived in forests of conifers and even
deciduous tree In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, afte ...
s. Despite the higher temperatures, darkness would still have descended for the winter, just as it does today at high
latitude In geography, latitude is a coordinate that specifies the north– south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north ...
s. The Antarctic Peninsula, including James Ross Island, was connected to South America throughout this time period, allowing interchange of fauna between both continents. In fact, the recent discovery of the Chilean Parankylosaurian '' Stegouros'' shows that these dinosaurs inhabited also South America.


See also

*
Timeline of ankylosaur research This timeline of ankylosaur research is a chronological listing of events in the history of paleontology focused on the ankylosaurs, quadrupedal herbivorous dinosaurs who were protected by a covering bony plates and spikes and sometimes by a club ...
*
South Polar dinosaurs The South Polar region of the Cretaceous comprised the continent of East Gondwana–modern day Australia and Antarctica–a product of the break-up of Gondwana in the Cretaceous Period. The southern region, during this time, was much warmer than i ...


References

{{Taxonbar, from=Q134176 Ankylosaurs Dinosaurs of Antarctica Late Cretaceous dinosaurs Maastrichtian life Fossil taxa described in 2006 James Ross Island Ornithischian genera