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''Kunbarrasaurus'' (meaning "shield lizard") is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
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 small ankylosaurian
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
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 ...
of
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
. The genus contains a single species, ''K. ieversi''.


Discovery

In November 1989, at Marathon Station near
Richmond, Queensland Richmond is a rural town and locality in the Shire of Richmond, Queensland, Australia. In the , the locality of Richmond had a population of 578 people. It is the administrative centre of the Shire of Richmond. Toponymy The origin of the ...
, the skeleton was discovered of an ankylosaurian. In January 1990 it was secured by a team led by Ralph Molnar. In 1996, in a provisional description, Molnar concluded that it could be referred to the genus '' Minmi'' as a ''Minmi'' sp. Subsequently, the specimen was further prepared by an acid bath and investigated by a CAT scan. The new information led to the conclusion that the species could be named in a separate genus of ankylosaur. In 2015, Lucy G. Leahey, Ralph E. Molnar, Kenneth Carpenter, Lawrence M. Witmer and Steven W. Salisbury named and described the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
''Kunbarrasaurus ieversi''. The genus name is derived from ''Kunbarra'' - the word for 'shield' in the Mayi language of the local Wunumara people. The specific name ''ieversi'' honours Mr Ian Ivers, the property manager who originally found the fossil. The description was limited to the skull. ''Kunbarrasaurus'' was one of eighteen dinosaur taxa from 2015 to be described in open access or free-to-read journals. 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 ...
, QM F1801, was found in a layer of the Allaru Formation, marine sediments dating from the late
Albian The Albian is both an age (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...
, or possibly the early
Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
. It consists of an almost complete skeleton with skull, containing the vertebral column up to the middle tail, the left shoulder girdle, the left arm minus the hand, the pelvis, both thighbones and most of the body armour. Both the bones and the armour are largely articulated. In the belly region remains have been found of the animal's last meal. The specimen represents the most complete dinosaur skeleton ever found in Eastern
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 ...
(Australia, Antarctica, Madagascar and India) and the most complete ankylosaurian skeleton from the entirety of the Gondwanan continents. Five specimens earlier referred to a ''Minmi'' sp. were not referred to ''Kunbarrasaurus'' by the 2015 description. In 2022, a specimen collected in 2005 by Benjamin P. Kear from Warra Station near Boulia, Queensland, SAMA P40536 was described and referred to ''Kunbarrasaurus.'' However, the describing authors refrained from referring it to ''K. leversi'' because it did not share any unique features with the type specimen. It was found in a layer belonging to the Toolebuc Formation, which directly underlies the Allaru Mudstone and dates to the middle-upper
Albian The Albian is both an age (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...
. The specimen consists of a partial skull associated with elements from the postcrania. Unfortunately, most of the postcranial skeleton remains in limestone and therefore not ready for description.


Description

''Kunbarrasaurus'' was a small quadrupedal
armoured Armour (Commonwealth 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 ...
dinosaur. Its length has been estimated at . In 2015, some distinguishing traits of the skull of ''Kunbarrasaurus'' were established. The roof of the skull is almost perfectly flat, apart from a limited convex profile of the postorbital bone and the nasal bone. The edges of the skull top, formed by the prefrontal, supraorbital and postorbital bones, make a right angle with the skull sides. The supraorbital is made up of one bone instead of two or three. The prefrontal is only exposed on the skull roof and does not reach the eye socket. The nasal bone does not reach the snout side and is limited to the snout top and the large, more centrally placed, opening around the nostril. This opening, which is completely located in the nasal bone, is large compared to the maxillary part of the snout and fully accessible from above and the side. The maxilla vertically attains the full height of the skull, reaching to the prefrontal on the skull roof. The hindmost tooth is positioned under the rear edge of the eye socket. The
lacrimal bone The lacrimal bones are two small and fragile bones of the facial skeleton; they are roughly the size of the little fingernail and situated at the front part of the medial wall of the orbit. They each have two surfaces and four borders. Several bon ...
is directed vertically. The pterygoid bones do not touch each other with their rear ends at the braincase, totally separated by the basisphenoid. The quadrate is vertically oriented. The coronoid process of the lower jaw is strongly protruding. The side of the braincase largely consists of cartilage instead of bone, so that many brain nerves must have had their exits in a single large opening, rather than separate small ones. The inner ear is very large compared with the skull as a whole and differs from that of all other known Dinosauria in the ear vestibule not being separated from the brain cavity, the floor for the
cochlea The cochlea is the part of the inner ear involved in hearing. It is a spiral-shaped cavity in the bony labyrinth, in humans making 2.75 turns around its axis, the modiolus (cochlea), modiolus. A core component of the cochlea is the organ of Cort ...
not being made of bone and the vestibule being so large that the
semicircular canal The semicircular canals are three semicircular interconnected tubes located in the innermost part of each ear, the inner ear. The three canals are the lateral, anterior and posterior semicircular canals. They are the part of the bony labyrinth, ...
s are shortened. The skull
osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of Extant taxon, extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, Temnospondyli, ...
are flat or at most have a low keel. There are no squamosal or quadratojugal horns or bosses on the upper skull corners or the cheeks.


Osteoderms

''Kunbarrasaurus'' had bony protrusions, also known as body
armour Armour (Commonwealth English) or armor (American English; see American and British English spelling differences#-our, -or, spelling differences) is a covering used to protect an object, individual, or vehicle from physical injury or damage, e ...
, in the skin on its head, back, abdomen, legs and along the tail. Several types of armour are known in place in ''Kunbarrasaurus'', including small ossicles, small keeled scutes on the body ordered in parallel longitudinal rows, large scutes without keels on the snout, large keeled scutes on the neck, shoulders, and possibly tail, spike-like scutes on the hips, and a combination of ridged and keeled scutes and triangular plates on the tail. There was one preserved ring of scutes around the neck. A sacral shield is absent. The exact arrangement of armour on the tail is unclear, although it appears that there were triangular plates that ran on the sides of the tail, with long scutes forming a row along the top of the tail. While most of the tail is not preserved, some of the relatives of ''Kunbarrasaurus'', including ''
Stegouros ''Stegouros'' (, meaning "roofed tail") is an extinct genus of ankylosaurian dinosaurs from the Late Cretaceous Dorotea Formation of southern Chile. The genus contains a single species, ''Stegouros elengassen'', known from a semi-articulated, ne ...
'' and ''
Antarctopelta ''Antarctopelta'' (; meaning 'Antarctic shield') is a genus of ankylosaurian dinosaur, a group of large, quadrupedal herbivores, that lived during the Maastrichtian stage of the Late Cretaceous period on what is now James Ross Island, Antarctica ...
'', had a large macuahuitl-like structure at the end of the tail, formed by expanded triangular flattened osteoderms.


Phylogeny

''Kunbarrasaurus'' was in 2015 placed in the
Ankylosauria Ankylosauria is a group of herbivorous dinosaurs of the clade 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 same year Victoria Arbour e.a. had entered QM F1801 and the ''Minmi'' holotype as separate
operational taxonomic unit An operational taxonomic unit (OTU) is an operational definition used to classify groups of closely related individuals. The term was originally introduced in 1963 by Robert R. Sokal and Peter H. A. Sneath in the context of numerical taxonomy, wh ...
s in their analysis. Whereas ''Minmi'' was recovered as a basal member of the
Ankylosauridae 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 ...
, QM F1801 had a basal position in
Ankylosauria Ankylosauria is a group of herbivorous dinosaurs of the clade 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 ...
, i.e. was too "primitive" to be included in either the
Ankylosauridae 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 ...
or
Nodosauridae Nodosauridae is a family of ankylosaurian dinosaurs known from the Late Jurassic to the Late Cretaceous periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as a monophyletic clade as the ...
.Arbour VM, Currie PJ. 2015. Systematics, phylogeny and palaeobiogeography of the ankylosauriddinosaurs. Journal of Systematic Palaeontology In the 2015 description of ''Kunbarrasaurus'' on qualitative considerations such a position was indeed deemed likely. In 2021, Sergio Soto-Acuña et al. found ''Kunbarrasaurus'' to belong to a distinctive basal lineage of Southern Hemisphere ankylosaurs, the
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 ...
. Their
phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...
placed it as the
sister taxon 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 ...
to the clade formed by the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
''Stegouros'' and ''Antarctopelta''.


Paleobiology

As with other ankylosaurians, ''Kunbarrasaurus'' was
herbivorous A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
. Unlike most herbivorous dinosaurs, there is direct evidence of the diet of ''Kunbarrasaurus'': gut contents are known from the well-preserved nearly complete holotype specimen, found in the abdominal cavity in front of the left ilium.


Diet

The gut contents consist of fragments of fibrous or
vascular Vascular can refer to: * blood vessels, the vascular system in animals * vascular tissue Vascular tissue is a complex transporting tissue, formed of more than one cell type, found in vascular plants. The primary components of vascular tissue ...
plant tissue,
fruit In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering. Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...
ing bodies, spherical
seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s, and vesicular tissue (possibly from
fern The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
sporangia A sporangium (from Late Latin, ; : sporangia) is an enclosure in which spores are formed. It can be composed of a unicellular organism, single cell or can be multicellular organism, multicellular. Virtually all plants, fungus, fungi, and many ot ...
). The most common remains are the
fibrous Fiber (spelled fibre in British English; from ) is a natural or artificial substance that is significantly longer than it is wide. Fibers are often used in the manufacture of other materials. The strongest engineering materials often inco ...
or vascular fragments, which are typically rather uniform in size at long and have clean cuts at their ends,
perpendicular In geometry, two geometric objects are perpendicular if they intersect at right angles, i.e. at an angle of 90 degrees or π/2 radians. The condition of perpendicularity may be represented graphically using the '' perpendicular symbol'', � ...
to a given fragment's long axis. Because of the small size of the fragments, they have been interpreted as having been nibbled from plants or chopped in the mouth, evidence of some method of retaining food in the mouth. These small fragments may have come from twigs or stems, but their size is more suggestive of vascular bundles in leaves. The clean cuts and lack of
gastrolith A gastrolith, also called a stomach stone or gizzard stone, is a rock held inside a gastrointestinal tract. Gastroliths in some species are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. In ...
s suggest that the animal relied on oral processing instead of gastroliths or grit to grind food. The seeds (0.3 mm .01 inacross) and fruiting bodies (4.5 mm .18 inacross) were apparently swallowed whole. Comparisons with gut contents and scat from other modern herbivores like
lizards Lizard is the common name used for all squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The ...
,
emu The emu (; ''Dromaius novaehollandiae'') is a species of flightless bird endemism, endemic to Australia, where it is the Tallest extant birds, tallest native bird. It is the only extant taxon, extant member of the genus ''Dromaius'' and the ...
s, and
geese A goose (: geese) is a bird of any of several waterfowl species in the family Anatidae. This group comprises the genera '' Anser'' (grey geese and white geese) and ''Branta'' (black geese). Some members of the Tadorninae subfamily (e.g., Egyp ...
indicate that this ''Kunbarrasaurus'' individual had a more sophisticated process for cutting up plant material.


See also

*
Timeline of ankylosaur research This timeline of ankylosaur research is a chronological listing of events in the History of paleontology, history of paleontology focused on the ankylosaurs, quadrupedal herbivorous dinosaurs who were protected by a covering bony plates and spik ...


References

{{Taxonbar, from=Q21685996 Parankylosauria Dinosaur genera Albian dinosaurs Fossil taxa described in 2015 Taxa named by Ralph Molnar Taxa named by Kenneth Carpenter Dinosaurs of Australia