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''Glacialisaurus'' is a
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 sauropodomorph
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 ...
from the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic� ...
period of
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. ...
. It is known from two specimens; the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
(name-bearing specimen), a partial tarsus (ankle) and metatarsus, and a partial left
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 ...
(upper thigh bone). The fossils were collected by a team led by paleontologist William R. Hammer during a 1990–91 field expedition to the central region of the
Transantarctic Mountains The Transantarctic Mountains (abbreviated TAM) comprise a mountain range of uplifted rock (primarily sedimentary) in Antarctica which extends, with some interruptions, across the continent from Cape Adare in northern Victoria Land to Coats L ...
. They come from sedimentary rocks of the
Hanson Formation The Hanson Formation (also known as the Shafer Peak Formation) is a geologic formation on Mount Kirkpatrick and north Victoria Land, Ross Dependency, Antarctica. It is one of the two major dinosaur-bearing rock groups found on Antarctica to date; ...
and date to the Pliensbachian stage of the Early Jurassic, around 186 to 182 million years ago. The fossils were described in 2007, and made the basis of the new
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 ...
and
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
''Glacialisaurus hammeri''. The genus name translates as “icy” or "frozen lizard”, and the specific name honors Hammer. This dinosaur has been classified as a
massospondylid Massospondylidae is a family (biology), family of early massopod dinosaurs that existed in Asia, Africa, North America, South America and AntarcticaHellert, Spencer M. "A New Basal Sauropodomorph from The Early Jurassic Hanson Formation of Antarc ...
, a group of medium-sized, basal (early diverging or "primitive") sauropodomorphs that existed during the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
and Early Jurassic on every continent except Australia. Its length has been estimated at . ''Glacialisaurus'' was a large herbivorous dinosaur, though it was average sized for a massospondylid. ''Glacialisaurus'' was distinct from other sauropodomorphs in features such as having a robust medial epicondylar ridge on the lower femur, a robust adductor ridge extending from the upper end of the femoral medial condyle, and a second metatarsal with a front border that is weakly convex in the upper end.


Discovery and naming

Fossils of a sauropodomorph dinosaur were discovered by a field team from Augustana College led by paleontologist William R. Hammer during 1990–91 fieldwork in the lower
Hanson Formation The Hanson Formation (also known as the Shafer Peak Formation) is a geologic formation on Mount Kirkpatrick and north Victoria Land, Ross Dependency, Antarctica. It is one of the two major dinosaur-bearing rock groups found on Antarctica to date; ...
of Mount Kirkpatrick in the Central Transantarctic Mountains of
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. ...
, dating to the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic� ...
.Hammer, W. R., & Hickerson, W. J. (1996). Implications of an Early Jurassic vertebrate fauna from Antarctica. ''The Continental Jurassic'', 215–218. The fossils were from two different individuals: elements of the right ankle and metatarsus such as the
astragalus Astragalus may refer to: * ''Astragalus'' (plant), a large genus of herbs and small shrubs *Astragalus (bone) The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...
, two tarsals and four metatarsals preserved in articulation (specimen FMNH PR1823), and the lower part of a left
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 ...
(FMNH PR1822, a thigh bone), ending just after the dissipation of the medial epicondylar crest. Several other fossils were collected from the same site, including fossils of the carnivorous
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
dinosaur '' Cryolophosaurus'', a
pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
humerus (upper arm bone), and a large tooth of a tritylodont,Smith, N. D., Makovicky, P. J., Hammer, W. R., & Currie, P. J. (2007). Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. ''Zoological Journal of the Linnean Society'', ''151''(2), 377–421. all found at an elevation of about . The right ankle and tarsus were preserved in a thick layer of
strata In geology and related fields, a stratum (: strata) is a layer of Rock (geology), rock or sediment characterized by certain Lithology, lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by v ...
, while the femur was preserved at the surface weathering next to the ''Cryolophosaurus'' specimen. The fossils were sent to the
Field Museum of Natural History The Field Museum of Natural History (FMNH), also known as The Field Museum, is a natural history museum in Chicago, Illinois, and is one of the largest such museums in the world. The museum is popular for the size and quality of its educationa ...
in Chicago, USA, and were first reported in 1994. This report speculated that cervical vertebrae from ''Cryolophosaurus'' found nearby also belonged to the sauropodomorph, but this has since been disproven. The fossils were described by the paleontologists Nathan Smith and Diego Pol, who named the new genus and species ''Glacialisaurus hammeri'', with FMNH PR1823 as
holotype specimen A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was Species description, formally described. It is either the single such physical example (or illus ...
. The generic name is derived from the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
root meaning after its discovery in the
Beardmore Glacier The Beardmore Glacier in Antarctica is one of the largest valley glaciers in the world, being long and having a width of . It descends about from the Antarctic Plateau to the Ross Ice Shelf and is bordered by the Commonwealth Range of the Queen ...
region in the Central Transantarctic Mountains and the word meaning . The specific name honors Hammer for his contributions to Antarctic paleontology.


Description

While few remains are known of ''Glacialisaurus'', its leg bones show it was a robust basal (early diverging or "primitive") sauropodomorph. The femur fragment is the larger of the two known specimens, measuring as preserved, with an estimated total length when intact of . ''Glacialisaurus'' is estimated to have been about long. As a basal sauropodomorph, ''Glacialisaurus'' would have had a long neck and a proportionally small head with leaf-shaped teeth. The hand would have been short, wide, and robust with a large claw on the thumb.


Leg bones

The
cross-section Cross section may refer to: * Cross section (geometry) ** Cross-sectional views in architecture and engineering 3D * Cross section (geology) * Cross section (electronics) * Radar cross section, measure of detectability * Cross section (physics) ...
of the robust femoral shaft is slightly wider from side to side than from front to back, though not as extreme as in eusauropods. The medial epicondylar crest extends from the medial surface of the lower femoral shaft and is distinct from all other sauropodomorphs in that it is robust, a trait convergently evolved in basal theropods. The front surface of the femur is flat instead of convex from side to side, a feature shared with other basal sauropodomorphs. The top surface of the upper femur lacks any anterior extensor groove. At the lower end, the lateral and medial condyles are separated by a craniocaudal groove that ends abruptly with a popliteal fossa (opening in the bone). ''Glacialisaurus'' is distinguished by its robust adductor ridge extending from the upper end of the femoral medial condyle. This ridge starts at the end of the medial condyle and is kidney-shaped with a long axis spanning proximolateral−distomedially. The astragalus is low and elongate from across side to side and the medial portion lacks the craniocaudally broadening compared to the lateral portion, a trait found in most non-eusauropods. The astragalus is weakly convex at the lower end, though this is not as extreme as in '' Blikanasaurus'' and '' Lessemsaurus''. The upper surface of the astragalus is softly convex because it is where the lower end 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 ...
(shin bone) articulates with the astragalus. This surface is pierced by two foramina (small openings in bone) that have been interpreted as vascular foramina. The ascending process (protrusion of bone) is mound shaped and its upper articular surface faces proximomedially. The distal tarsals have a laterally elongated triangular shape in when seen from their top ends. The corners of the tarsals are rounded and bulbous, especially in the posteromedial corner. The medial distal tarsal is not confined solely to metatarsal III, but also barely contacts the proximal end of metatarsal II, like in ''
Saturnalia Saturnalia is an Roman festivals, ancient Roman festival and holiday in honour of the List of Roman deities, god Saturn (mythology), Saturn, held on 17 December in the Julian calendar and later expanded with festivities until 19 December. By t ...
''. The lateral distal tarsal has a quadrangular shape and was likely longer mediolaterally than proximodistally. Metatarsal I is roughly 3/4 the length of metatarsal II, as in most basal sauropodomorphs. Metatarsal I has a broad and short shaft that is ellipse shaped, more so than in most other basal sauropodomorphs. The upper portion of the small posterior groove separating the two distal condyles is similar to that of ''Plateosaurus''. The medial condyle is less robust and more proximally positioned than the lateral one. This would cause a medial displacement of the first digit, a characteristic in most saurischian dinosaurs (the group that includes sauropodomorphs and theropods). The upper end of metatarsal II is hourglass-shaped and has concave medial-lateral ends to articulate with the other metatarsals. The medial concavity is well developed, but the lateral concavity is less so. The diagnostic traits (characteristics that distinguish a taxon from others) of the second metatarsals include: a front border that is weakly convex in proximal aspect; a hypertrophied lateral plantar flange on the proximal end (present, but less developed in many basal sauropodomorphs, e.g., ''Saturnalia'', ''Plateosaurus''); and a medial distal condyle that is more robust and well−developed than the lateral distal condyle. The third metatarsal lacks much preserved detail, but has a trapezoidal upper end with a straight to concave front border and a slightly convex medial border for articulation with metatarsal II. The hind edge is narrower from side toside than the front one, but is not acute or rounded, causing the upper outline of metatarsal III to be almost trapezoidal, as in ''Lufengosaurus'', '' Gyposaurus'', and ''Coloradisaurus''. On the contrary, most non-eusauropod sauropodomorphs have almost triangular upper outlines. Only the upper portion of the metatarsal IV is preserved, but preserves an upper outline akin to that of ''Lufengosaurus'' that has a broad anterior face and a finger−like posteromedial projection. This finger-like process is slightly convex and would have articulated with metatarsal III.


Classification

The
phylogenetic 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 dat ...
position of ''Glacialisaurus'' is unstable due to its fragmentary nature, but it is frequently found to be a member of the family
Massospondylidae Massospondylidae is a family (biology), family of early massopod dinosaurs that existed in Asia, Africa, North America, South America and AntarcticaHellert, Spencer M. "A New Basal Sauropodomorph from The Early Jurassic Hanson Formation of Antarc ...
. Massospondylids are a group of non- eusauropod sauropodomorphs that existed during the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
to Early Jurassic in Africa, Antarctica, Asia, and the Americas. Massospondylids have been recovered as the
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
(most closely related group) to more derived sauropodomorphs, including
Sauropoda Sauropoda (), whose members are known as sauropods (; from ''wikt:sauro-, sauro-'' + ''wikt:-pod, -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative t ...
itself, and are more derived than groups like Plateosauridae. In their 2007 phylogenetic analysis of the relationships of ''Glacialisaurus'', Smith and Pol found that it was a massospondylid. Features of its foot are similar to ''Lufengosaurus'' (from the Early Jurassic of China), and the phylogenetic study suggested that these dinosaurs were close relatives, whereas ''Massospondylus'' was found to be a more basal form. This has been supported by later analyses, including Müller (2019) which found it in a clade with ''Coloradisaurus'' and ''Lufengosaurus'', while ''Massospondylus'', ''Sarahsaurus'', '' Pradhania'', and '' Xingxiulong'' were more basal in the family. The paleontologist Oliver W. M. Rauhut and colleagues found ''Lufengosaurus'' to be most closely related to ''Glacialisaurus'' in 2020, and the following
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 ...
shows the placement they recovered for ''Glacialisaurus'' within the sauropodomorph group
Massopoda Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the Sauropoda, sauropods and their ancestral relatives. Sauropods generally grew to very large siz ...
: The discovery of ''Glacialisaurus'' is important to the study of the early distribution of sauropod dinosaurs. The presence of this primitive sauropodomorph in the Hanson Formation (which has also yielded remains attributed to true sauropods) shows that both primitive and advanced members of this lineage existed side by side in the early Jurassic Period.


Paleoenvironment

''Glacialisaurus'' is known from the Hanson Formation, which is one of only two major dinosaur-bearing rock formations found on Antarctica. The specimens were discovered in tuffaceous
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility. Although its permeabil ...
deposited in the Sinemurian to Pliensbachian stage of the Early Jurassic,Hammer, W.R.; Hickerson, W.J. (1999). Tomida, Y.; Rich, T.H.; Vickers-Rich, Y. (eds.). "Gondwana Dinosaurs from the Jurassic of Antarctica". Proceedings of the Second Gondwana Dinosaur Symposium National Science Museum Monographs. 15: 211–217. dating to about 194–188 million years ago. This
geological formation A geological formation, or simply formation, is a body of rock having a consistent set of physical characteristics (lithology) that distinguishes it from adjacent bodies of rock, and which occupies a particular position in the layers of rock expo ...
is part of the Victoria Group of the Transantarctic Mountains, which is approximately above sea level. The high altitude of this site supports the idea that early Jurassic Antarctica had
forests A forest is an ecosystem characterized by a dense community of trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological functio ...
populated by a diverse range of species, at least along the coast.Dodson, P. (1997). "Distribution and Diversity". In Currie, P.J.; Padian, K. (eds.). Encyclopedia of Dinosaurs. Academic Press. pp. 10–13. . The Hanson Formation was deposited in an active volcano−tectonic rift system formed during the breakup of the
supercontinent In geology, a supercontinent is the assembly of most or all of Earth's continent, continental blocks or cratons to form a single large landmass. However, some geologists use a different definition, "a grouping of formerly dispersed continents", ...
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 ...
. Local volcanism and evidence of wildfires is known from some paleobotanical sites in the Hanson Formation. Models of Jurassic air flow indicate that coastal areas probably never dropped much below freezing, although more extreme conditions existed inland. ''Glacialisaurus'' was found about from the
South Pole The South Pole, also known as the Geographic South Pole or Terrestrial South Pole, is the point in the Southern Hemisphere where the Earth's rotation, Earth's axis of rotation meets its surface. It is called the True South Pole to distinguish ...
, which was about or so farther north at the time. This formation has yielded the remains of the large theropod ''Cryolophosaurus'', a crow-sized dimorphodontid pterosaur, a rat-sized tritylodont
synapsid Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
, and two small unnamed sauropodomorphs. Many plant genera have also been recovered from the Shafer Peak section of the Hanson Formation that suggest forests similar to the open woodlands of
North Island The North Island ( , 'the fish of Māui', historically New Ulster) is one of the two main islands of New Zealand, islands of New Zealand, separated from the larger but less populous South Island by Cook Strait. With an area of , it is the List ...
, New Zealand. Known plants include Cheirolepidiaceaen conifers, '' Equisetites'' horsetails, and '' Cladophlebis'' ferns that have also been found or are similar to plants found in other Early Jurassic sites that represent warm climates. Basal sauropodomorphs like ''Glacialisaurus'' were the first very large dinosaurs and, due to their height, the first herbivores to high browse.


See also

* List of Australian and Antarctic dinosaurs * South Polar region of the Cretaceous


References

{{Taxonbar, from=Q431390 Massospondylidae Dinosaur genera Pliensbachian dinosaurs Fossil taxa described in 2007 Dinosaurs of Antarctica