Kunbarrasaurus
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''Kunbarrasaurus'' 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 small herbivorous
ankylosaurian 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. ...
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 ...
from the
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 ...
of Australia.


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 648 people. It is the administrative centre of the Shire of Richmond. Toponymy The origin of the name ...
, the skeleton was discovered of an ankylosaurian. In January 1990 it was secured by a team led by
Ralph Molnar Ralph E. Molnar is a paleontologist who had been Curator of Mammals at the Queensland Museum and more recently associated with the Museum of Northern Arizona. He is also a research associate at the Texas natural Science Centre. He co-authored descr ...
. 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 Kenneth Carpenter (born September 21, 1949, in Tokyo, Japan) is a paleontologist. He is the former director of the USU Eastern Prehistoric Museum and author or co-author of books on dinosaurs and Mesozoic life. His main research interests ...
, Lawrence M. Witmer and Steven W. Salisbury named and described the
type species In 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 species that contains the biological type specime ...
''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 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 sever ...
, QM F1801, was found in a layer of the
Allaru Formation The Allaru Formation, also known as the Allaru Mudstone, is a geological formation in Queensland, Australia, whose strata date back to the Early Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.Weisham ...
, marine sediments dating from the late
Albian The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/ Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0 ...
, or possibly the early Cenomanian. 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 (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 Boulia () is an outback town and locality in the Shire of Boulia, Queensland, Australia. In the , Boulia had a population of 301 people. Boulia is the administrative centre of the Boulia Shire, population approximately 600, which covers an area ...
, 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 of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/ Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0 ...
. 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
armoured 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 f ...
dinosaur, that was
quadrupedal 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 ...
and had a long tail. 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 bone is a small and fragile bone of the facial skeleton; it is roughly the size of the little fingernail. It is situated at the front part of the medial wall of the orbit. It has two surfaces and four borders. Several bony landmarks of ...
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. A core component of the cochlea is the Organ of Corti, the sensory or ...
not being made of bone and the vestibule being so large that the
semicircular canal In mathematics (and more specifically geometry), a semicircle is a one-dimensional locus of points that forms half of a circle. The full arc of a semicircle always measures 180° (equivalently, radians, or a half-turn). It has only one line o ...
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 and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amphi ...
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, 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 arrangement of armour is unclear on the tail, although the triangular plates may have run on the sides of the tail, with long scutes forming a row along the top of the tail.


Phylogeny

''Kunbarrasaurus'' was in 2015 placed in the
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. ...
. The same year
Victoria Arbour Victoria Megan Arbour is a Canadian evolutionary biologist and vertebrate palaeontologist at Royal BC Museum, where she is Curator of Palaeontology. An "expert on the armoured dinosaurs known as ankylosaurs", Arbour analyzes fossils and creates ...
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, w ...
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–Pa ...
, QM F1801 had a basal position in
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. ...
, 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–Pa ...
or Nodosauridae. 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 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 ...
.


Paleobiology

As with other ankylosaurians, ''Kunbarrasaurus'' was
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 ...
. 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 The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away f ...
plant tissue,
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
ing bodies, spherical
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
s, and vesicular tissue (possibly from
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
sporangia A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cyc ...
). The most common remains are the
fibrous Fiber or fibre (from la, fibra, links=no) is a #Natural fibers, natural or Fiber#Artificial fibers, artificial substance that is significantly longer than it is wide. Fibers are often used in the manufacture of other materials. The stronge ...
or vascular fragments, which are typically rather uniform in size at long and have clean cuts at their ends,
perpendicular In elementary geometry, two geometric objects are perpendicular if they intersect at a right angle (90 degrees or π/2 radians). The condition of perpendicularity may be represented graphically using the ''perpendicular symbol'', ⟂. It ca ...
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 oth ...
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, emus, and
geese A goose ( : geese) is a bird of any of several waterfowl species in the family Anatidae. This group comprises the genera '' Anser'' (the grey geese and white geese) and ''Branta'' (the black geese). Some other birds, mostly related to the she ...
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 focused on the ankylosaurs, quadrupedal herbivorous dinosaurs who were protected by a covering bony plates and spikes and sometimes by a club ...


References

{{Taxonbar, from=Q21685996 Ankylosaurs Early Cretaceous dinosaurs of Australia Paleontology in Queensland Fossil taxa described in 2015 Taxa named by Ralph Molnar Taxa named by Kenneth Carpenter Ornithischian genera