''Bothriospondylus'' ("excavated

vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...

") is a

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 extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

neosauropod Neosauropoda is a clade within Dinosauria, coined in 1986 by Argentina, Argentine paleontologist José Bonaparte and currently described as ''Saltasaurus loricatus'', ''Diplodocus longus'', and all animals directly descended from their most recen ...

sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their bo ...

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 ...

. It lived during the Late

Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...

Discovery and naming

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 specimen ...

, ''Bothriospondylus suffossus'', was described by

Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Owe ...

in 1875. The

specific epithet In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...

''suffossus'' means "undermined" in

Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...

, a reference to the fact that

pleurocoels Skeletal pneumaticity is the presence of air spaces within bones. It is generally produced during development by excavation of bone by pneumatic diverticula (air sacs) from an air-filled space, such as the lungs or nasal cavity. Pneumatization is h ...

had hollowed out the sides of the vertebra. It is often incorrectly spelled as "suffosus". Owen based the species on

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 several ...

NHMUK R.44592-5, a set of four dorsal vertebrae found in

Wiltshire Wiltshire (; abbreviated Wilts) is a historic and ceremonial county in South West England with an area of . It is landlocked and borders the counties of Dorset to the southwest, Somerset to the west, Hampshire to the southeast, Gloucestershire ...

in stratum from the

Kimmeridgian In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...

, the

Kimmeridge Clay The Kimmeridge Clay is a sedimentary deposit of fossiliferous marine clay which is of Late Jurassic to lowermost Cretaceous age and occurs in southern and eastern England and in the North Sea. This rock formation is the major source rock for North ...

. Also three unfused sacral vertebrae were referred. Ornithopsis hulkei lectotype

At the same time Owen named three other species of ''Bothriospondylus''. ''B. robustus'' was based on NHMUK R.22428, a dorsal from the same location. ''B. elongatus'' was based on a vertebra from

Sussex Sussex (), from the Old English (), is a historic county in South East England that was formerly an independent medieval Anglo-Saxon kingdom. It is bounded to the west by Hampshire, north by Surrey, northeast by Kent, south by the English ...

, NHMUK R.2239, an original

syntype In biological nomenclature, a syntype is any one of two or more biological types that is listed in a description of a taxon where no holotype was designated. Precise definitions of this and related terms for types have been established as part of ...

of ''

Ornithopsis ''Ornithopsis'' (meaning "bird-likeness") was a medium-sized Early Cretaceous sauropod dinosaur, from England. The type species, which is the only species seen as valid today, is ''O. hulkei''. History of discovery Gideon Algernon Mantell descr ...

hulkei''. Finally, '' Bothriospondylus magnus'' was a new name for another syntype of ''

hulkei'' Seeley 1870, the present

lectotype 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 ...

NHMUK 28632. Owen himself in an addendum to the same publication renamed ''B. robustus'' to ''

Marmarospondylus ''Marmarospondylus'' ("marble eference to the Forest Marble Formation">Forest_Marble_Formation.html" ;"title="eference to the Forest Marble Formation">eference to the Forest Marble Formationvertebra") is a nomen dubium, dubious genus of sauropod ...

robustus''.

Friedrich von Huene Friedrich von Huene, born Friedrich Richard von Hoinigen, (March 22, 1875 – April 4, 1969) was a German paleontologist who renamed more dinosaurs in the early 20th century than anyone else in Europe. He also made key contributions about v ...

in 1908 referred the material to ''

Pelorosaurus ''Pelorosaurus'' ( ; meaning "monstrous lizard") is a genus of titanosauriform sauropod dinosaur. Remains referred to ''Pelorosaurus'' date from the Early Cretaceous period, about 140-125 million years ago, and have been found in England and Port ...

'' and in 1922 made ''B. suffossus'' into a ''Ornithopsis suffossa'' because the latter generic name has priority. More complete material from

Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...

was named by

Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was an English naturalist, geologist and writer of numerous books on natural history. Biography Richard Lydekker was born at Tavistock Square in London. His father was Gerard Wolfe Lydekker, ...

as a fifth species, ''Bothriospondylus madagascariensis''.

Franz Nopcsa Franz may refer to: People * Franz (given name) * Franz (surname) Places * Franz (crater), a lunar crater * Franz, Ontario, a railway junction and unorganized town in Canada * Franz Lake, in the state of Washington, United States – see ...

in 1902 assigned to ''Bothriospondylus'' a vertebra from

Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...

that later would be renamed ''

Nopcsaspondylus ''Nopcsaspondylus'' (meaning " Nopcsa's vertebra", in reference to the original describer) is a genus of rebbachisaurid sauropod dinosaur (a type of large, long-necked quadrupedal herbivorous dinosaur) from the Cenomanian-age (Upper Cretaceous) Ca ...

''. A skeleton from

France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...

was assigned to ''Bothriospondylus madagascariensis'', but has been described as a new genus and species of brachiosaurid, ''

Vouivria ''Vouivria'' is a genus of herbivorous sauropod dinosaur, belonging to the Brachiosauridae, that lived in the area of present France during the Late Jurassic. The type species is ''Vouivria damparisensis''. History In 1926, the Solvay company ...

''. In 1986 José Fernando Bonaparte moved material from "B." ''sp.'' into its own genus, ''

Lapparentosaurus ''Lapparentosaurus'' is a genus of sauropod dinosaur from the Middle Jurassic. Its fossils were found in Madagascar (Isalo III Formation). The type species is ''L. madagascariensis''. Discovery and naming In 1895 Richard Lydekker named a new s ...

''. The material, sometimes referred to ''B. madagascariensis'', was separated into a few dorsal vertebrae to be the new

of ''Lapparentosaurus''. A revision in 2010 by Philip Mannion concluded that ''Bothriospondylus'' is 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 s ...

''. However, ''"Bothriospondylus" madagascariensis'' was treated as valid and distinct from other Middle Jurassic sauropods from Madagascar.

Classification

''Bothriospondylus'' has over the years been assigned to many groups — even in a Bothriospondylidae of its own — with

Brachiosauridae The Brachiosauridae ("arm lizards", from Greek ''brachion'' (βραχίων) = "arm" and ''sauros'' = "lizard") are a family or clade of herbivorous, quadrupedal sauropod dinosaurs. Brachiosaurids had long necks that enabled them to access the ...

lately being the most popular designation. However, the sparse and eroded material shows no

synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...

of the Brachiosauridae and cannot be further determined than a very general

Neosauropoda Neosauropoda is a clade within Dinosauria, coined in 1986 by Argentine paleontologist José Bonaparte and currently described as ''Saltasaurus loricatus'', ''Diplodocus longus'', and all animals directly descended from their most recent common ...

Current species

Only the type species of ''Bothriospondylus'' can be referred, because of the dubious nature of the genus. * ''Bothriospondylus suffossus'' Owen, 1875 (

) = ''

suffosa'' (Owen, 1875) von Huene, 1922 * "Bothriospondylus" madagascariensis Lydekker, 1895 NHM R2598 NHM R2596–2615, R16588, R16589

Isalo III Formation The Isalo III Formation is a geological formation in Madagascar, off the eastern coast of Africa. It dates back to the Middle Jurassic.Weishampel et al., 2004, pp.542-543 The use of the term "Isalo III" is somewhat controversial as the two prior u ...

, Madagascar, Middle Jurassic, Non neosauropod-

eusauropod Eusauropoda (meaning "true sauropods") is a derived clade of sauropod dinosaurs. Eusauropods represent the node-based group that includes all descendant sauropods starting with the basal eusauropods of ''Shunosaurus'', and possibly ''Barapasaurus ...

distinct from ''Lapparentosaurus''.

Reassigned species

Several species have been referred to ''Bothriospondylus'' over its existence, with nearly all being reassigned to a new genus. * '' Bothriospondylus robustus'' Owen, 1875 renamed ''

'' (Owen, 1875) Owen, 1875 * ''Bothriospondylus elongatus'' Owen, 1875 non ''Bothriospondylus'' * '' Bothriospondylus magnus'' Owen, 1875 objective synonym of ''

Ornithopsis hulkei ''Ornithopsis'' (meaning "bird-likeness") was a medium-sized Early Cretaceous sauropod dinosaur, from England. The type species, which is the only species seen as valid today, is ''O. hulkei''. History of discovery Gideon Algernon Mantell descr ...

'' Seeley, 1870 * '' Bothriospondylus madagascariensis'' Lydekker, 1895 renamed ''

'' (Mannion, Allain & Moine, 2017) * "Bothriospondylus madagascariensis" Lydekker, 1895 removed from species and named ''

'' Bonaparte, 1986 * "Bothriospondylus" Lydekker, 1895

Discovery and naming

Classification

Current species

Reassigned species

References

Further reading