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''Brachauchenius'' (meaning 'short neck') 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
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 ...
s that lived during 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 ...
in what are now
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
and
North Africa North Africa (sometimes Northern Africa) is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region. However, it is sometimes defined as stretching from the Atlantic shores of t ...
. Only one
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), ...
is known, ''B. lucasi'', initially described by
Samuel Wendell Williston Samuel Wendell Williston (July 10, 1852 – August 30, 1918) was an American educator, entomologist, and Paleontology, paleontologist who was the first to propose that birds developed flight Origin of birds#Origin of bird flight, cursorially (by ...
in 1903 from a partial
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 preserve ...
skeleton A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...
discovered in a quarry in
Kansas Kansas ( ) is a landlocked U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to the west. Kansas is named a ...
, United States. Many other fossil specimens attributed to the species were subsequently discovered, including an individual from
Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
whose presence was made official in 2016. Many contemporary pliosaur specimens were formerly attributed to ''Brachauchenius'', but have since been reidentified as belonging to other genera or are recognized as indeterminate.


Research history


Holotype and naming

In 1884, a partial
skeleton A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...
of a
marine reptile Marine reptiles are reptiles which have become secondarily adapted for an aquatic or semiaquatic life in a marine environment. Only about 100 of the 12,000 extant reptile species and subspecies are classed as marine reptiles, including mari ...
was discovered by the owner of a quarry near Delphos, in
Ottawa County, Kansas Ottawa County is a county located in the U.S. state of Kansas. Its county seat and largest city is Minneapolis. As of the 2020 census, the county population was 5,735. The county was named after the Odawa tribe. History Early history F ...
. News of this find thus reached
Charles H. Sternberg Charles Hazelius Sternberg (June 15, 1850 – July 20, 1943) was an American fossil collector and paleontology, paleontologist. He was active in both fields from 1876 to 1928, and collected fossils for Edward Drinker Cope and Othniel C. Marsh, ...
, who was then collecting fossils for
Othniel Charles Marsh Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of paleontology. A prolific fossil collector, Marsh was one of the preeminent paleontologists of the nineteenth century. Among his legacies are the discovery or ...
. Sternberg then wrote several letters to Marsh seeking his permission and funding to exhume the specimen, which he allowed. After exhumation, Sternberg gave the specimen to the
Peabody Museum of Natural History The Peabody Museum of Natural History at Yale University (also known as the Yale Peabody Museum of Natural History or the Yale Peabody Museum) is one of the oldest, largest, and most prolific university natural history museums in the world. It ...
for Marsh, but it turned out that Marsh would do little with it. The fossil was eventually transferred to the
United States National Museum of Natural History The Smithsonian Institution ( ), or simply the Smithsonian, is a group of museums, Education center, education and Research institute, research centers, created by the Federal government of the United States, U.S. government "for the increase a ...
and subsequently given the code name USNM 4989. This same specimen consists of a nearly complete
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
measuring long, followed by thirty-seven
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e and associated
rib In vertebrate anatomy, ribs () are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the thoracic cavity, enabling the lungs to expand and thus facilitate breathing by expanding the ...
s. Historical documentation from the museum where the specimen has since resided notes that Sternberg was allegedly helped by a certain “John Potts” in its exhumation. In 1903,
Samuel Wendell Williston Samuel Wendell Williston (July 10, 1852 – August 30, 1918) was an American educator, entomologist, and Paleontology, paleontologist who was the first to propose that birds developed flight Origin of birds#Origin of bird flight, cursorially (by ...
designated the fossil as 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 ...
of a new
plesiosaur The Plesiosauria or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared in the latest Triassic Period (geology), Period, possibly in the Rhaetian st ...
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), ...
under the name ''Brachauchenius lucasi''. The generic name ''Brachauchenius'' derives from the
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
words (, "short") and (, "neck"), followed by 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 ...
suffix In linguistics, a suffix is an affix which is placed after the stem of a word. Common examples are case endings, which indicate the grammatical case of nouns and adjectives, and verb endings, which form the conjugation of verbs. Suffixes can ca ...
''ius'', all meaning "short neck". Williston chose this name because this specimen was considered the shortest- necked plesiosaur ever described. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
''lucasi'' is named in honor of
Frederic Augustus Lucas Frederic Augustus Lucas (March 25, 1852 – February 9, 1929) was a zoologist and taxidermist who served as a curator of the Brooklyn Museum and director of the American Museum of Natural History. He was an expert on the osteology and anatomy of bir ...
for his notable contributions to American paleontology. In his article, Williston only mentions the origin locality of the specimen, and does not detail any elements regarding the discovery of the latter. Thus, the circumstances surrounding the discovery of USNM 4989 remained unclear until 2007, when letters from Sternberg obtained by
Kenneth Carpenter Kenneth Carpenter (born 21 September 1949) is an American 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 are armore ...
were formalized in an article written by his colleague Michael J. Everhart. The
stratigraphic Stratigraphy is a branch of geology concerned with the study of rock layers (strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithost ...
provenance of the holotype specimen remained also uncertain for decades. Williston notes that it comes from the "Benton Cretaceous", but it turns out that three
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 ...
s outcrop the locality where the specimen was discovered. In 1996, Carpenter listed the specimen as coming from the
Greenhorn Limestone The Greenhorn Limestone or Greenhorn Formation is a geologic formation in the Great Plains Region of the United States, dating to the Cenomanian and Turonian ages of the Late Cretaceous period. The formation gives its name to the Greenhorn cycle ...
, but he does not, however, support any evidence for this assertion. A microscopic examination of the fossils published in 2005 revealed that the specimen came from either the upper level of the Greenhorn Limestone or the lower level of the Fairport Chalk Member of the
Carlile Shale The Carlile Shale is a Turonian age Upper Cretaceous, Upper/Late Cretaceous series shale geologic Formation (geology), formation in the central-western United States, including in the Great Plains region of Colorado, Kansas, Nebraska, New Mexico ...
, suggesting that the specimen dates to the early Middle
Turonian The Turonian is, in the International Commission on Stratigraphy, ICS' geologic timescale, the second age (geology), age in the Late Cretaceous epoch (geology), Epoch, or a stage (stratigraphy), stage in the Upper Cretaceous series (stratigraphy), ...
.


Further specimens

In 2016, a pliosaur
mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
from
Turonian The Turonian is, in the International Commission on Stratigraphy, ICS' geologic timescale, the second age (geology), age in the Late Cretaceous epoch (geology), Epoch, or a stage (stratigraphy), stage in the Upper Cretaceous series (stratigraphy), ...
deposits near Goulmima,
Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
were referred to the species ''Brachauchenius lucasi''.


Formerly attributed specimens

In 1860,
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...
documented a partial skull having been discovered around an unspecified date by George Cubitt in
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 ...
sediments from the town of
Dorking Dorking () is a market town in Surrey in South East England about south-west of London. It is in Mole Valley, Mole Valley District and the non-metropolitan district, council headquarters are to the east of the centre. The High Street runs ro ...
, in the
county A county () is a geographic region of a country used for administrative or other purposesL. Brookes (ed.) '' Chambers Dictionary''. Edinburgh: Chambers Harrap Publishers Ltd, 2005. in some nations. The term is derived from the Old French denoti ...
of
Surrey Surrey () is a Ceremonial counties of England, ceremonial county in South East England. It is bordered by Greater London to the northeast, Kent to the east, East Sussex, East and West Sussex to the south, and Hampshire and Berkshire to the wes ...
, England. Based on teeth characteristics, Owen referred this specimen to '' Polyptychodon interruptus'', a species of
pliosaurid Pliosauridae is a family of plesiosaurian marine reptiles from the Latest Triassic to the early Late Cretaceous (Rhaetian to Turonian stages). The family is more inclusive than the archetypal short-necked large headed species that are placed in ...
that he himself had already named in 1841 from fossils dating from the same period which were discovered in the counties of
Sussex Sussex (Help:IPA/English, /ˈsʌsɪks/; from the Old English ''Sūþseaxe''; lit. 'South Saxons'; 'Sussex') is an area within South East England that was historically a kingdom of Sussex, kingdom and, later, a Historic counties of England, ...
and
Kent Kent is a Ceremonial counties of England, ceremonial county in South East England. It is bordered by Essex across the Thames Estuary to the north, the Strait of Dover to the south-east, East Sussex to the south-west, Surrey to the west, and Gr ...
. The following year, Owen cited that the specimen was discovered during the construction of a
railway tunnel Rail transport (also known as train transport) is a means of transport using wheeled vehicles running in railway track, tracks, which usually consist of two parallel steel railway track, rails. Rail transport is one of the two primary means of ...
through the
Chalk Group The Chalk Group (often just called the Chalk) is the lithostratigraphic unit (a certain number of rock strata) which contains the Upper Cretaceous limestone succession in southern and eastern England. The same or similar rock sequences occur ac ...
, near
Frome Frome ( ) is a town and civil parish in Somerset, England, on uneven high ground at the eastern end of the Mendip Hills and on the River Frome, south of Bath. The population of the parish was 28,559 in 2021. Frome was one of the largest tow ...
,
Somerset Somerset ( , ), Archaism, archaically Somersetshire ( , , ) is a Ceremonial counties of England, ceremonial county in South West England. It is bordered by the Bristol Channel, Gloucestershire, and Bristol to the north, Wiltshire to the east ...
. However, this locality was most likely confused with those of other specimens assigned to this genus, and with multiple reported evidence, it seems more likely that the first locality originally mentioned is the most likely. This specimen has since resided in the local museum of Dorking, where it is cataloged as DOKDM G/1-2. In a
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 ...
study published in 2013, Roger B. J. Benson and colleagues recovered it as being close to several other specimens assigned to ''Brachauchenius'', and therefore suggested that it would be better to refer it to that genus as well, although without specific affiliation. In his revision of ''Polyptychodon'' published in 2016, Daniel Madzia designates this genus as a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
'' due to the lack of accessibility and diagnostics regarding the holotype tooth of the taxon, the latter having possibly even been lost. However, he considers it likely that DOKDM G/1-2 could be designated as the
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 associated. In other words, a type is an example that serves to anchor or centralizes ...
of this genus in order to maintain its validity, but he refrains from doing so at this time since the descriptions provided by Owen are obsolete. In a 2017 conference, Madzia and colleagues judge the Dorking specimen to be potentially diagnostic despite its incomplete preservation. All recognized fossils of the related and contemporary genus '' Megacephalosaurus'' were first interpreted as coming from ''Brachauchenius''. The holotype specimen of the genus, cataloged as FHSM VP-321, was discovered in 1950 by teenage brothers Frank and Robert Jennrich near
Fairport, Kansas Fairport is an unincorporated community in Paradise Township, Russell County, Kansas, United States. It is located about 12 miles north of Gorham near the west county line and next the Saline River. History In 1879, Knight & Bradshaw built ...
. The Jenrich brothers, aided by George F. Sternberg and a local cow rancher named Jim Rouse, exhumed the skull later that year. Although initially thinking the specimen would be a
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Ancient Greek, Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains wer ...
, G. F. Sternberg later identified it as an imposing specimen of ''B. lucasi'', placing the skull and mandible on a single support showing only their dorsal sides. However, it was in 1996 that the skull was first described by Carpenter, who still identified it as such. A new expedition carried out in November 2003 by R. Jennrich led scientists to discover that the specimen was exhumed from Fairport Chalk. In 2007, Everhart discovered photos of the skull's underside taken prior to its mounting, which revealed underside features that were different from those found in other ''B. lucasi'' skulls. The specimen is then removed from its mount, and is officially described in 2013 by Bruce A. Schumacher, Carpenter and Everhart as the holotype of a new genus and species which they name ''Megacephalosaurus eulerti''. Shortly before the description made by Schumacher ''et al.'', the phylogenetic study by Benson ''et al.'' classified the taxon as ''B. eulerti''. The history of the
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype (biology), isotype ...
skull, cataloged UNSM 50136, is much more obscure. Indeed, the latter presents very little contextual information on its discovery and its precise locality of origin, the only information known is that the specimen also comes from Kansas. It was first described in the scientific literature by Schumacher in 2008, who provisionally identified it under the name ''
aff. ''Species affinis'' (commonly abbreviated to: ''sp.'', ''aff.'', or ''affin.'') is taxonomic terminology in zoology and botany. In open nomenclature it indicates that available material or evidence suggests that the proposed species is related t ...
B. lucasi''. Although its locality of origin is unknown,
matrix Matrix (: matrices or matrixes) or MATRIX may refer to: Science and mathematics * Matrix (mathematics), a rectangular array of numbers, symbols or expressions * Matrix (logic), part of a formula in prenex normal form * Matrix (biology), the m ...
analysis extracted from the fossil identifies nannofossil assemblages that are associated with limestone deposits of the Greenhorn Formation. When FHSM VP-321 was found to be a distinct genus, USNM 50136 was identified to be conspecific with it and was assigned to be its paratype. Everhart blogged that other specimens currently attributed to ''Brachauchenius'' may actually belong to ''Megacephalosaurus''. The most notable specimen in his line of sight being UNSM 112437, a partial skull having been collected from the Graneros Shale. In 2000, a relatively complete pliosaurid skeleton, since cataloged as VL17052004-1, was discovered in the town of
Villa de Leyva Villa de Leyva, also called Villa de Leiva, is a touristic colonial town and municipality, in the Ricaurte Province, part of the Boyacá Department of Colombia. The town is a Colombian National Heritage Town and is on the tentative list for UNE ...
in Boyacá, Colombia, before being exhumed during 2004-2005. Coming from the
Paja Formation The Paja Formation (, K1p, Kip, Kimp, b3b6p) is an Early Cretaceous formation (geology), geologic formation of central Colombia. The formation extends across the northern part of the Altiplano Cundiboyacense, the Colombian emeralds#Western belt, W ...
, it was dated to the Upper
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 125.77 Ma (million years ago) and 121.4 ± 1.0 Ma (Historically, this stage was placed at 129.4 million to approximately 125 million years ago) It is a ...
, making the latter the oldest known example of the
brachauchenine Pliosauridae is a family of plesiosaurian marine reptiles from the Latest Triassic to the early Late Cretaceous (Rhaetian to Turonian stages). The family is more inclusive than the archetypal short-necked large headed species that are placed in ...
s. The skeleton was first described in 2005 by Oliver Hampe as belonging to ''Brachauchenius'', before being designated as the holotype of a new genus and species under the name '' Stenorhynchosaurus munozi'' by María Eurídice Páramo and colleagues in 2016.


Description

''Brachauchenius'' has a morphology typical of the pliosaurids of the
thalassophonea Pliosauridae is a family of plesiosaurian marine reptiles from the Latest Triassic to the early Late Cretaceous (Rhaetian to Turonian stages). The family is more inclusive than the archetypal short-necked large headed species that are placed in ...
n group, which has a large elongated skull connected to a short
neck The neck is the part of the body in many vertebrates that connects the head to the torso. It supports the weight of the head and protects the nerves that transmit sensory and motor information between the brain and the rest of the body. Addition ...
, unlike many other
plesiosaur The Plesiosauria or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared in the latest Triassic Period (geology), Period, possibly in the Rhaetian st ...
s, which have a long neck and a small head. Like all other plesiosaurs, ''Brachauchenius'' has a short
tail The tail is the elongated section at the rear end of a bilaterian animal's body; in general, the term refers to a distinct, flexible appendage extending backwards from the midline of the torso. In vertebrate animals that evolution, evolved to los ...
, a massive
trunk Trunk may refer to: Biology * Trunk (anatomy), synonym for torso * Trunk (botany), a tree's central superstructure, and the stem of woody plants * Trunk of corpus callosum, in neuroanatomy * Elephant trunk, the proboscis of an elephant Comput ...
and two pairs of large flippers. Like most Turonian pliosaurs, the measurement of ''Brachauchenius'' is quite modest, the holotype specimen having a maximum length estimated at for a body mass estimated at .


Classification

The cladogram below is modified from Madzia ''et al.'' (2018):


Notes


References


Bibliography

* * * * * *


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 inv ...
*
Timeline of plesiosaur research This timeline of plesiosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, Taxonomy (biology), taxonomic revisions, and cultural portrayals of plesiosaurs, an order of marine reptiles ...


External links


Oceans of Kansas Paleontology
{{Taxonbar, from=Q135636 Late Cretaceous plesiosaurs of North America Fossil taxa described in 1903 Pliosauridae Turonian genus extinctions Taxa named by Samuel Wendell Williston Sauropterygian genera Late Cretaceous reptiles of Africa