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''Kronosaurus'' ( ; meaning "lizard of Kronos") is a potentially
dubious Doubt is a mental state in which the mind remains suspended between two or more contradictory propositions, unable to be certain of any of them. Doubt on an emotional level is indecision between belief and disbelief. It may involve uncertainty ...
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 extinct short-necked
pliosaur Pliosauroidea is an extinct clade of plesiosaurs, known from the earliest Jurassic to early Late Cretaceous. They are best known for the subclade Thalassophonea, which contained crocodile-like short-necked forms with large heads and massive toot ...
. With an estimated length of , it was among the largest pliosaurs, and is named after the leader of the Greek Titans, Kronos. It lived in the Early Cretaceous period ( Aptian to Late Albian).Hampe O. 1992. Ein großwüchsiger Pliosauride (Reptilia: Plesiosauria) aus der Unterkreide (oberes Aptium) von Kolumbien. ''Courier Forschungsinstitut Senckenberg'' 145: 1–32.Kear BP. 2003
Cretaceous marine reptiles of Australia: a review of taxonomy and distribution
''Cretaceous Research'' 24: 277–303.
Fossil material has been recovered from the
Toolebuc Formation The Toolebuc Formation is a geological formation that extends from Queensland across South Australia and the Northern Territory in Australia, whose strata date back to the Albian stage of the Early Cretaceous. Dinosaurs,Weishampel ''et al.'', 20 ...
(middle to late Albian) and
Wallumbilla Formation The Wallumbilla Formation is an Aptian geologic formation found in Australia. Plesiosaur and theropod remains are among the fossils that have been recovered from its strata. Description The formation is present in the Northern Territory, Que ...
s (Aptian) of Queensland and New South Wales in Australia.


Discovery

In 1899, Andrew Crombie of
Hughenden Hughenden may refer to: *Hughenden, Queensland, a town in Australia *Hughenden, Alberta, a village in central Alberta, Canada *Hughenden Valley Hughenden Valley (formerly called Hughenden or Hitchendon) is an extensive village and civil parish in ...
discovered a "scrap of bone" containing six conical teeth, and gave this fragmentary
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 ...
to the
Queensland Museum The Queensland Museum is the state museum of Queensland, dedicated to natural history, cultural heritage, science and human achievement. The museum currently operates from its headquarters and general museum in South Brisbane with specialist m ...
. Twenty-five years later, then-director Heber Longman formally described the specimen as 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 ...
of a new
species In biology, a species is the basic unit of 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 the appropriate s ...
: ''Kronosaurus queenslandicus''.Mather, Patricia, with Agnew, N.H. et al. ''The History of the Queensland Museum, 1862–1986'' Retrieved fro
archive.org
/ref> More ''Kronosaurus'' material, including a partial skull, was discovered in 1929, in the same location as Crombie's original find.


Taxonomic validity

The validity of ''Kronosaurus'' was a longstanding issue due to the fragmentary nature of the holotype. Welles (1962) declared the taxon as a '' nomen vanum'' and recommended the designation of a
neotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes the ...
specimen that would preserve the taxon's validity. McHenry (2009) subsequently proposed that two partial skeletons (QM F18827 and QM F10113) that matched the holotype's features and were from its same geological horizon (
Toolebuc Formation The Toolebuc Formation is a geological formation that extends from Queensland across South Australia and the Northern Territory in Australia, whose strata date back to the Albian stage of the Early Cretaceous. Dinosaurs,Weishampel ''et al.'', 20 ...
) could be candidate neotypes. However, no formal petition to the ICZN to designate a neotype was ever submitted. In 2021, Noè and Gómez-Pérez formally restricted ''Kronosaurus'' to the holotype. They subsequently erected a new taxon named ''Eiectus longmani'' with the Harvard skeleton (MCZ 1285) as its holotype and reassigned all specimens previously referred as ''K. queenslandicus'' to it. This move, which was initially interpreted as renaming the entire species to ''E. longmani'', faced criticism by paleontologists.
Darren Naish Darren William Naish is a British vertebrate palaeontologist, author and science communicator. As a researcher, he is best known for his work describing and reevaluating dinosaurs and other Mesozoic reptiles, including '' Eotyrannus'', '' Xenop ...
commented that the apparent renaming was "not a good idea" that discourages stability while paleontologist Daniel Madzia dubbed the situation "Kronogate" and criticized the decision to not petition for a neotype. However, the reassignment of all ''Kronosaurus'' material to the new taxon is tentative. Noè & Gómez-Pérez (2021) noted the likelihood that the fossils actually represent multiple pliosaur species, meaning that the Toolebuc Formation specimens could be distinct from ''E. longmani'', and previous studies already suggested that the Harvard skeleton might be distinct from ''K. queenslandicus''.


Harvard's completed skeleton

In 1931 the
Harvard Museum of Comparative Zoology Harvard University is a Private university, private Ivy League research university in Cambridge, Massachusetts. Founded in 1636 as Harvard College and named for its first benefactor, the History of the Puritans in North America, Puritan cler ...
(MCZ) sent an expedition to Australia for the dual purpose of procuring specimens – the museum being "weak in Australian animals and...desires ngto complete its series" – and to engage in "the study of the animals of the region when alive." The
Harvard Australian Expedition (1931–1932) The Harvard Australian Expedition of 1931–1932 was a six-man venture sent by then Harvard Museum of Comparative Zoology (MCZ) director Thomas Barbour to Australia for the dual purpose of procuring specimens and studying native (living) wildlife i ...
, as it became known, was a six-man venture led by Harvard Professor
William Morton Wheeler William Morton Wheeler (March 19, 1865 – April 19, 1937) was an American entomologist, myrmecologist and Harvard professor. Biography Early life and education William Morton Wheeler was born on March 19, 1865, to parents Julius Morton Wheel ...
, with the others being Dr. P. Jackson Darlington Jr. (a renowned coleopterist), Dr.
Glover Morrill Allen Glover Morrill Allen (February 8, 1879 – February 14, 1942) was an American zoologist. He was born at Walpole, New Hampshire, the son of Reverend Nathaniel Glover Allen and Harriet Ann (Schouler) Allen, and studied at Harvard University. While s ...
and his student Ralph Nicholson Ellis, medical officer Dr. Ira M. Dixon, and William E. Schevill (a graduate-student in his twenties and Associate Curator of Invertebrate Palaeontology).About the Exhibits by Elizabeth Hall and Max Hall (Museum of Comparative Zoology "Agazziz Museum" Harvard University. Third Edition, Copyright 1964, 1975, 1985, by the President and Fellows of Harvard College MCZ director
Thomas Barbour Thomas Barbour (August 19, 1884 – January 8, 1946) was an American herpetologist. From 1927 until 1946, he was director of the Harvard Museum of Comparative Zoology (MCZ) founded in 1859 by Louis Agassiz at Harvard University in Cambridge, ...
said at the time "We shall hope for specimens' of the kangaroo, the wombat, the Tasmanian devil and Tasmanian wolf," and the mission was a success with over 300 mammal and thousands of insect specimens returning to the United States. Yet Mr. Schevill, the team's fossil enthusiast, remained in Australia after the others had departed and, in the winter of 1932, was told by the rancher R.W.H. Thomas of rocks with something "odd" poking out of them on his property near Hughenden. The rocks were
limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
nodules containing the most complete skeleton of ''Kronosaurus'' ever discovered.Meyers, Troy. ''Kronosaurus Chronicles''. Australian Age of Dinosaurs, Issue 3, 2005. Retrieved fro
australianageofdinosaurs.com
/ref> After dynamiting the nodules out of the ground (and into smaller pieces weighing approximately four tons) with the aid of a British migrant trained in the use of explosives,The Rarest of the Rare: Stories Behind the Treasures at the Harvard Museum of Natural History (Hardcover) – 26 October 2004 William Schevill had the fossils shipped back to Harvard for examination and preparation. The skull—which matched the holotype jaw fragment of ''K. queenslandicus''—was prepared right away, but time and budget constraints put off restoration of the nearly complete skeleton – most of the bones of which remained unexcavated within the limestone blocks – for 20 years. This interim ended when they came to the attention of
Godfrey Lowell Cabot Godfrey Lowell Cabot (February 26, 1861 – November 2, 1962) was an American industrialist who founded the Cabot Corporation. Early life Godfrey Lowell Cabot was born in Boston, Massachusetts and attended Boston Latin School. His father was S ...
– Boston industrialist, philanthropist, and founder of the
Cabot Corporation Cabot Corporation is an American specialty chemicals and performance materials company headquartered in Boston, Massachusetts. The company operates in over 20 countries with 36 manufacturing plants, eight research and development facilities and ...
– "who was then in his nineties" and "had been interested in sea serpents since childhood." Having formerly questioned MCZ director
Alfred Romer Alfred Sherwood Romer (December 28, 1894 – November 5, 1973) was an American paleontologist and biologist and a specialist in vertebrate evolution. Biography Alfred Romer was born in White Plains, New York, the son of Harry Houston Romer an ...
about the existence and reports of sea serpents, it thus occurred to Dr. Romer to tell Mr. Cabot about the skeleton in the museum closet. Godfrey Cabot thus asked how much a restoration would cost and "Romer, pulling a figure out of the musty air, replied, 'Oh, about $10,000.'" Romer may not have been serious but the philanthropist clearly was because the check for said sum came shortly thereafter. Two years – and more than $10,000 – later, following the careful labor of the museum preparators, the restored and mounted skeleton was displayed at Harvard in 1959. However, Dr. Romer and MCZ preparator Arnold Lewis confirmed that same year in the institution's journal ''Breviora'' that "erosion had destroyed a fair fraction of this once complete and articulated skeleton...so that approximately a third of the specimen as exhibited is plaster restoration." Furthermore, the original (real) bones are also layered in plaster; a fact that, while keeping the fossils safe, makes it difficult for paleontologists to study it – an issue which factors into the controversial question of the true size of the ''Kronosaurus queenslandicus''. In 2021, a revision of ''K. boyacensis'' also transferred most of the remains of ''K. queenslandicus'', including the Harvard remains, to a new genus and species, ''
Eiectus ''Eiectus'' is a potentially dubious genus of extinct short-necked pliosaur that lived in the Early Cretaceous period.Hampe O. (1992). Ein großwüchsiger Pliosauride (Reptilia: Plesiosauria) aus der Unterkreide (oberes Aptium) von Kolumbien. ''C ...
longmani''. The revision limits the genus ''Kronosaurus'' to the holotype mandible, and treats it as a ''nomen dubium''.


Size issues

Body-length estimates, largely based on the 1959 Harvard reconstruction, had previously put the total length of ''Kronosaurus'' at .Romer AS, Lewis AD. 1959. A mounted skeleton of the giant plesiosaur ''Kronosaurus''. ''Breviora'' 112: 1–15. However, more recent studies, comparing fossil specimens of ''Kronosaurus'' to other pliosaurs suggests that the Harvard reconstruction may have included too many vertebrae, exaggerating the previous estimate, with the true length probably only . In 2022, the specimens referred to ''Eiectus longmani'' were estimated at in length and in body mass.


Description

Like other pliosaurs, ''Kronosaurus'' was a marine reptile. It had an elongated head, a short neck, a stiff body propelled by four flippers, and a relatively short tail. The posterior flippers were larger than the anterior. ''Kronosaurus'' was carnivorous, and had many long, sharp, conical teeth. A feature of the genus ''Kronosaurus'' is that the first three maxillary teeth are enlarged to fangs. Current estimates put ''Kronosaurus'' at around in length. In 2009, ''K. queenslandicus'' was estimated to weigh about .McHenry, Colin R. 2009. Devourer of Gods: The palaeoecology of the Cretaceous pliosaur ''Kronosaurus queenslandicus''. The University of Newcastle, N.S.W. Australia, Web

/ref> The skull length of ''Kronosaurus'' was estimated to be . All Sauropterygians had a modified
pectoral girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists of ...
that supported a powerful swimming stroke.Palaios – Sauropterygia
/ref> ''Kronosaurus'' and other plesiosaurs/pliosaurs had a similarly adapted pelvic girdle,
/ref> allowing them to push hard against the water with all four flippers. Between its two limb girdles was a massive mesh of
gastralia Gastralia (singular gastralium) are dermal bones found in the ventral body wall of modern crocodilians and tuatara, and many prehistoric tetrapods. They are found between the sternum and pelvis, and do not articulate with the vertebrae. In thes ...
(belly ribs) that provided additional strength and support. The strength of the limb girdles, combined with evidence of large, powerful swimming muscles, indicates that ''Kronosaurus'' was likely a fast, active swimmer. ''Kronosaurus queenslandicus'' has four pairs of
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
ry teeth. The first three pairs of its maxillary teeth are large caniniform teeth, and in its lower jaw there are also three pairs of large caniniform teeth which are located immediately in front of the three maxillary caniniforms when the mouth is closed. Two pairs of the lower jaw caniniforms occlude between the last pair of premaxillary teeth and first pair of maxillary teeth in a
diastema A diastema (plural diastemata, from Greek διάστημα, space) is a space or gap between two teeth. Many species of mammals have diastemata as a normal feature, most commonly between the incisors and molars. More colloquially, the condition ...
(gap in the tooth row). The enlargement of these two pairs of lower jaw teeth in ''K. queenslandicus'' may be related to the absence of a fifth pair of premaxillary teeth, which are present in a number of other pliosaurs.


Teeth

''Kronosaurus'' teeth exceed in length (the largest up to long with crowns). However, they lack carinae (cutting edges) and the distinct trihedral (three facets) of ''
Pliosaurus ''Pliosaurus'' (meaning 'more lizard') is an extinct genus of thalassophonean pliosaurid known from the Kimmeridgian and Tithonian stages (Late Jurassic) of Europe and South America. Their diet would have included fish, cephalopods, and marin ...
'' and ''
Liopleurodon ''Liopleurodon'' (; meaning 'smooth-sided teeth') is an extinct genus of large, carnivorous marine reptile belonging to the Thalassophonea, a clade of short-necked pliosaurid plesiosaurs. ''Liopleurodon'' lived from the Callovian Stage of the Mi ...
'' teeth. The combination of large size, conical shape and lack of cutting edges allows for easy identification of ''Kronosaurus'' teeth in Cretaceous formations from Australia.Massare JA. 1997. Introduction – faunas, behaviour and evolution. In: Callaway JM, Nicholls EL. (Eds.), ''Ancient Marine Reptiles''. Academic Press, San Diego, pp. 401–421.


Classification

The
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 ...
below follows a 2011 analysis by paleontologists Hilary F. Ketchum and Roger B. J. Benson, and reduced to genera only.


Palaeobiology

Fossil stomach contents from Northern Queensland show that ''Kronosaurus'' preyed on turtles and plesiosaurs. Fossil remains of large octobrachians have been found in the same area as ''Kronosaurus''. While no direct evidence of the animal predating on these octobrachians exists they fall within the projected size range of prey ''Kronosaurus'' would potentially have pursued. Large, round bite-marks have been found on the skull of an Albian-age Australian
elasmosaurid Elasmosauridae is an extinct family of plesiosaurs, often called elasmosaurs. They had the longest necks of the plesiosaurs and existed from the Hauterivian to the Maastrichtian stages of the Cretaceous, and represented one of the two groups of ...
(''
Eromangasaurus ''Eromangasaurus'' is an extinct genus of elasmosaurid known from northern Queensland of Australia. Description ''Eromangasaurus'' is a medium-sized elasmosaurid, measuring in length and in body mass. It is known from the holotype QM F11 ...
'') that could be from a ''Kronosaurus'' attack.Sachs S. 2005. ''Tuarangisaurus australis'' sp. nov. (Plesiosauria: Elasmosauridae) from the Lower Cretaceous of northeastern Queensland, with additional notes on the phylogeny of the Elasmosauridae. ''Memoirs of the Queensland Museum''. 50 (2): 425–440.Kear, B. P. 2007. Taxonomic Clarification of the Australian Elasmosaurid Genus ''Eromangasaurus'', with Reference to Other Austral Elasmosaur Taxa. ''Memoirs of the Queensland Museum''. 27 (1): 241–246.


Paleoecology

''Kronosaurus'' is known from remains in Australia. The area was covered by a shallow inland sea called the Eromanga Sea which ''Kronosaurus'' inhabited.Queensland Museum – ''Kronosaurus''
/ref> This environment was notably cold, experiencing near freezing temperatures and seasonal ice in certain regions.


See also

*
List of plesiosaur genera This list of plesiosaurs is a comprehensive listing of all genera that have ever been included in the order Plesiosauria, excluding purely vernacular terms. The list includes all commonly accepted genera, but also genera that are now considered in ...
*
Timeline of plesiosaur research This timeline of plesiosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, taxonomic revisions, and cultural portrayals of plesiosaurs, an order of marine reptiles that flourished dur ...
* ''
Platypterygius ''Platypterygius'' is a historically paraphyletic genus of platypterygiine ichthyosaur from the Cretaceous period. It was historically used as a wastebasket taxon, and most species within ''Platypterygius'' likely are undiagnostic at the genus ...
''


References


External links


Genus ''Kronosaurus''
The Plesiosaur Directory. * {{Authority control Pliosaurids Early Cretaceous plesiosaurs Aptian life Albian life Early Cretaceous reptiles of Australia Fossils of Australia Early Cretaceous reptiles of South America Cretaceous Colombia Fossils of Colombia Paja Formation Altiplano Cundiboyacense Fossil taxa described in 1924 Fossil taxa described in 1992 Taxa named by Albert Heber Longman Sauropterygian genera