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 ...
of
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 ...
that lived between 166 and 164 million years ago during the
Callovian
In the geologic timescale, the Callovian is an age and stage in the Middle Jurassic, lasting between 166.1 ± 4.0 Ma (million years ago) and 163.5 ± 4.0 Ma. It is the last stage of the Middle Jurassic, following the Bathonian and preceding the ...
(
Middle Jurassic
The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
Period
Period may refer to:
Common uses
* Era, a length or span of time
* Full stop (or period), a punctuation mark
Arts, entertainment, and media
* Period (music), a concept in musical composition
* Periodic sentence (or rhetorical period), a concept ...
) in what is now
England
England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe b ...
. A
herbivore
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 mouthpart ...
, ''Cetiosauriscus'' had — by sauropod standards — a moderately long
tail
The tail is the section at the rear end of certain kinds of animals’ bodies; in general, the term refers to a distinct, flexible appendage to the torso. It is the part of the body that corresponds roughly to the sacrum and coccyx in mammals, r ...
, and longer
forelimb
A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial ( anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used instead. ...
s, making them as long as its
hindlimb
A hindlimb or back limb is one of the paired articulated appendages (limbs) attached on the caudal ( posterior) end of a terrestrial tetrapod vertebrate's torso.http://www.merriam-webster.com/medical/hind%20limb, Merriam Webster Dictionary-Hindl ...
s. It has been estimated as about long and between in weight.
The only known
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 ...
includes most of the rear half of a skeleton as well as a hindlimb (NHMUK R3078). Found in
Cambridgeshire
Cambridgeshire (abbreviated Cambs.) is a Counties of England, county in the East of England, bordering Lincolnshire to the north, Norfolk to the north-east, Suffolk to the east, Essex and Hertfordshire to the south, and Bedfordshire and North ...
in the 1890s, it was described by
Arthur Smith Woodward
Sir Arthur Smith Woodward, FRS (23 May 1864 – 2 September 1944) was an English palaeontologist, known as a world expert in fossil fish. He also described the Piltdown Man fossils, which were later determined to be fraudulent. He is not relate ...
in 1905 as a new specimen of the
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 ...
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 ...
found NHMUK R3078 and the ''C. leedsi''
type specimen
In biology, a type is a particular wiktionary:en:specimen, 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 a ...
to be too different from ''
Cetiosaurus
''Cetiosaurus'' () meaning 'whale lizard', from the Greek '/ meaning 'sea monster' (later, 'whale') and '/ meaning 'lizard', is a genus of herbivorous sauropod dinosaur from the Middle Jurassic Period, living about 168 million years ago in what ...
'', warranting its own genus, which he named ''Cetiosauriscus'', meaning "''Cetiosaurus''-like". ''Cetiosauriscus leedsi'' was referred to the sauropod
family
Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Diplodocidae
Diplodocids, or members of the family Diplodocidae ("double beams"), are a group of sauropod dinosaurs. The family includes some of the longest creatures ever to walk the Earth, including ''Diplodocus'' and '' Supersaurus'', some of which may hav ...
because of similarities in the tail and foot, and had the dubious or indeterminate species "Cetiosauriscus" ''greppini'', "C." ''longus'', and "C." ''glymptonensis'' assigned to it. In 1980, Alan Charig named a new species of ''Cetiosauriscus'' for NHMUK R3078 because of the lack of comparable material to the type of ''C. leedsi''; this species was named ''Cetiosauriscus stewarti''. Because of the poor state of preservation of the ''Cetiosauriscus leedsi'' fossil, Charig sent a petition to the
International Commission on Zoological Nomenclature
The International Commission on Zoological Nomenclature (ICZN) is an organization dedicated to "achieving stability and sense in the scientific naming of animals". Founded in 1895, it currently comprises 26 commissioners from 20 countries.
Orga ...
to instead make ''C. stewarti'' the type species. ''Cetiosauriscus stewarti'' became the oldest confirmed diplodocid until a
phylogenetic analysis
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
published in 2003 instead found the species to belong to
Mamenchisauridae
Mamenchisauridae is a family of sauropod dinosaurs belonging to Eusauropoda known from the Jurassic and Early Cretaceous of Asia and Africa. Some members of the group reached gigantic sizes, amongst the largest of all sauropods.
Classification
T ...
, and followed by studies in 2005 and 2015 that found it outside
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 ...
, while not a mamenchisaurid proper.
''Cetiosauriscus'' was found in the marine deposits of the
Oxford Clay Formation
The Oxford Clay (or Oxford Clay Formation) is a Jurassic marine sedimentary rock formation underlying much of southeast England, from as far west as Dorset and as far north as Yorkshire. The Oxford Clay Formation dates to the Jurassic, specifica ...
alongside many different
invertebrate
Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
groups, marine
ichthyosaurs
Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, alt ...
,
plesiosaurs
The Plesiosauria (; Greek: πλησίος, ''plesios'', meaning "near to" and ''sauros'', meaning "lizard") or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia.
Plesiosaurs first appeared ...
and
crocodylians
Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest living ...
, a single
pterosaur
Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
, and various dinosaurs: the
ankylosaur
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. ...
''
Sarcolestes
''Sarcolestes'' (meaning "flesh robber") is an extinct genus of ankylosaurian ornithischian dinosaur from the Oxford Clay of England. The current type and only species is ''S. leedsi'', and the holotype is a single partial left mandible. The ...
'', the
stegosaur
Stegosauria is a group of herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere, predominantly in what is now North America, Europe, ...
s ''
Lexovisaurus
''Lexovisaurus'' is a genus of stegosaur from mid-to-Late Jurassic Europe, 165.7-164.7 mya. Fossils of limb bones and armor fragments have been found in middle to late Jurassic-aged strata of England.
Discovery and taxonomy
In the early ...
'' and ''
Loricatosaurus
''Loricatosaurus'' (meaning "armored lizard") is a Stegosaurid genus from Callovian-age ( Middle Jurassic) rocks of England and France.
Discovery and naming
''Loricatosaurus'' is known from remains previously assigned to '' Lexovisaurus'' tha ...
'', the
ornithopod
Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (), that started out as small, bipedal running grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous world ...
''
Callovosaurus
''Callovosaurus'' (meaning "Callovian lizard") is a genus of iguanodontian dinosaur known from most of a left thigh bone discovered in Middle Jurassic-age rocks of England. At times, it has been considered dubious or a valid genus of basal igu ...
'', as well as some unnamed
taxa
In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
. The
theropod
Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
s ''
Eustreptospondylus
''Eustreptospondylus'' ( ; meaning "true ''Streptospondylus''") is a genus of megalosaurid Theropoda, theropod dinosaur, from the Oxfordian stage of the Late Jurassic period (some time between 163 and 154 million years ago) in southern Eng ...
'' and ''
Metriacanthosaurus
''Metriacanthosaurus'' (meaning "moderately-spined lizard") is a genus of metriacanthosaurid dinosaur from the upper Oxford Clay of England, dating to the Late Jurassic period, about 160 million years ago (lower Oxfordian).
History of discovery ...
'' are known from the formation, although probably not from the same level as ''Cetiosauriscus''.
History of discovery
Background
The fossil later known as ''Cetiosauriscus'' was originally ascribed to the genus ''
Cetiosaurus
''Cetiosaurus'' () meaning 'whale lizard', from the Greek '/ meaning 'sea monster' (later, 'whale') and '/ meaning 'lizard', is a genus of herbivorous sauropod dinosaur from the Middle Jurassic Period, living about 168 million years ago in what ...
''—one of the first sauropods to be named, in
1842
Events
January–March
* January
** Michael Alexander takes office, as the first appointee to the Anglican-German Bishopric in Jerusalem.
** American medical student William E. Clarke of Berkshire Medical College becomes the first pe ...
by
palaeontologist
Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
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 ...
, and one with a complicated history due to many unfounded referrals of species and specimens, involving almost all English sauropod specimens. 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 ...
of ''Cetiosaurus'' has changed throughout history because of incomplete remains and the taxon's significance, and many aspects of its anatomy and relationships are still uncertain. ''Cetiosaurus'' was originally named to include ''C. medius'', ''C. brevis'', ''C. brachyurus'' and ''C. longus'', which span from the
Middle Jurassic
The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
to the
Early Cretaceous
The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma.
Geology
Pro ...
of various localities across England. As none of these species are truly diagnostic, and ''Cetiosaurus'' is a historically and taxonomically important taxon, the more complete Middle Jurassic species ''C. oxoniensis'' named by
geologist
A geologist is a scientist who studies the solid, liquid, and gaseous matter that constitutes Earth and other terrestrial planets, as well as the processes that shape them. Geologists usually study geology, earth science, or geophysics, althou ...
1871
Events January–March
* January 3 – Franco-Prussian War – Battle of Bapaume: Prussians win a strategic victory.
* January 18 – Proclamation of the German Empire: The member states of the North German Confederation and the sout ...
became the type species. ''C. glymptonensis'' was also named in the same publication by Phillips, but is less complete and of questionable validity.
Another English taxon, ''
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 ...
'', was named in
1870
Events
January–March
* January 1
** The first edition of ''The Northern Echo'' newspaper is published in Priestgate, Darlington, England.
** Plans for the Brooklyn Bridge are completed.
* January 3 – Construction of the Broo ...
by palaeontologist
Harry Govier Seeley
Harry Govier Seeley (18 February 1839 – 8 January 1909) was a British paleontologist.
Early life
Seeley was born in London on 18 February 1839, the second son of Richard Hovill Seeley, a goldsmith, and his second wife Mary Govier. When his fat ...
for vertebrae from the Early Cretaceous
Wessex Formation
The Wessex Formation is a fossil-rich English geological formation that dates from the Berriasian to Barremian stages (about 145–125 million years ago) of the Early Cretaceous. It forms part of the Wealden Group and underlies the younger Vecti ...
, younger than the existing species of ''Cetiosaurus''. Seeley considered ''Ornithopsis'' to be closely related to ''Cetiosaurus'', but different due to the internal bone structure. An additional species, '' Ornithopsis leedsii'' was named in
1887
Events
January–March
* January 11 – Louis Pasteur's anti-rabies treatment is defended in the Académie Nationale de Médecine, by Dr. Joseph Grancher.
* January 20
** The United States Senate allows the Navy to lease Pearl Har ...
by
John Hulke
John Whitaker Hulke FRCS FRS FGS (6 November 1830 – 19 February 1895) was a British surgeon, geologist and fossil collector. He was the son of a physician in Deal, who became a Huxleyite despite being deeply religious.
Hulke became Huxley's ...
for a
pelvis
The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton).
The ...
, vertebrae and
rib
In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...
s collected by
Alfred Nicholson Leeds
Alfred Nicholson Leeds (9 March 184725 August 1917) was an English amateur palaeontologist.
Biography
Leeds was born at Eyebury, Peterborough, the youngest of the eight children of Edward Thurlow Leeds (180251) and Eliza Mary Leeds (née Nichol ...
, an English farmer and amateur fossil collector who throughout his life compiled numerous collections of fossils from the Oxford Clay. ''O. leedsii'', from the
Late Jurassic
The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987.
In European lithostratigraphy, the name ...
, showed similarities to older ''
Cetiosaurus oxoniensis
''Cetiosaurus'' () meaning 'whale lizard', from the Greek '/ meaning 'sea monster' (later, 'whale') and '/ meaning 'lizard', is a genus of herbivorous sauropod dinosaur from the Middle Jurassic Period, living about 168 million years ago in what ...
'' as well as younger ''O. hulkei''. It was described in more detail by Seeley in
1889
Events
January–March
* January 1
** The total solar eclipse of January 1, 1889 is seen over parts of California and Nevada.
** Paiute spiritual leader Wovoka experiences a vision, leading to the start of the Ghost Dance movement in the ...
, where he considered ''O. hulkei'', ''C. oxoniensis'' and ''O. leedsii'' to all be in the same genus, bearing the name ''Cetiosaurus''. But naturalist
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, ...
discussed with Seeley, before the publication of Seeley's 1889 paper, that ''Cetiosaurus'' and ''Ornithopsis'' were not the same taxon. Lydekker suggested that Wealden fossils (including ''O. hulkei'') belonged to ''Ornithopsis'' and the Jurassic remains (including ''O. leedsii'' and ''C. oxoniensis'') to ''Cetiosaurus''. Lydekker in
1895
Events
January–March
* January 5 – Dreyfus affair: French officer Alfred Dreyfus is stripped of his army rank, and sentenced to life imprisonment on Devil's Island.
* January 12 – The National Trust for Places of Histor ...
changed his mind and referred the species ''O. leedsii'' 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 ...
'' (known already from the species ''P. brevis'', once named ''Cetiosaurus brevis'')—as ''P. leedsi''—and referred the genus to Atlantosauridae. Lydekker's classification of the species was not supported by later authors like palaeontologist
Arthur Smith Woodward
Sir Arthur Smith Woodward, FRS (23 May 1864 – 2 September 1944) was an English palaeontologist, known as a world expert in fossil fish. He also described the Piltdown Man fossils, which were later determined to be fraudulent. He is not relate ...
in
1905
As the second year of the massive Russo-Japanese War begins, more than 100,000 die in the largest world battles of that era, and the war chaos leads to the 1905 Russian Revolution against Nicholas II of Russia (Shostakovich's 11th Symphony i ...
, who followed Seeley's classification scheme.
Discovery and naming
The sauropod fossil today known as ''Cetiosauriscus stewarti'' was discovered in May
1898
Events
January–March
* January 1 – New York City annexes land from surrounding counties, creating the City of Greater New York as the world's second largest. The city is geographically divided into five boroughs: Manhattan, B ...
by clay workers in the area around
Fletton
Fletton is an area of the city of Peterborough, in the Peterborough district, in the ceremonial county of Cambridgeshire, England, south of the River Nene.
Notable for its large brickworks, the area has given its name to "Fletton bricks",
Admin ...
to the south of
Peterborough
Peterborough () is a cathedral city in Cambridgeshire, east of England. It is the largest part of the City of Peterborough unitary authority district (which covers a larger area than Peterborough itself). It was part of Northamptonshire until ...
sedimentary rock
Sedimentary rocks are types of rock that are formed by the accumulation or deposition of mineral or organic particles at Earth's surface, followed by cementation. Sedimentation is the collective name for processes that cause these particles ...
s of the
marine
Marine is an adjective meaning of or pertaining to the sea or ocean.
Marine or marines may refer to:
Ocean
* Maritime (disambiguation)
* Marine art
* Marine biology
* Marine debris
* Marine habitats
* Marine life
* Marine pollution
Military
* ...
Oxford Clay
The Oxford Clay (or Oxford Clay Formation) is a Jurassic marine sedimentary rock formation underlying much of southeast England, from as far west as Dorset and as far north as Yorkshire. The Oxford Clay Formation dates to the Jurassic, specifical ...
, which is of middle
Callovian
In the geologic timescale, the Callovian is an age and stage in the Middle Jurassic, lasting between 166.1 ± 4.0 Ma (million years ago) and 163.5 ± 4.0 Ma. It is the last stage of the Middle Jurassic, following the Bathonian and preceding the ...
age and today regarded as one of the classic
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 exp ...
s of British palaeontology. The sauropod fossil possibly stems from NPBCL pit No.1, which was the northernmost pit operated by the New Peterborough Brick Company Limited, and which produced the most vertebrate fossils. The discovery was brought to the attention of Leeds, who, after excavation, took the sauropod specimen to Eyebury, the Leeds' family home. In mid-August, after some cleaning and repairing of the specimen, geologist Henry Woodward visited Eyebury and produced a life-sized drawing of the remains for presentation at the British Association for the Advancement of Science Meeting. Following this presentation, on 17 August 1898, Henry Woodward returned with American palaeontologist
Othniel Charles Marsh
Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of Paleontology in Yale College and President of the National Academy of Sciences. He was one of the preeminent scientists in the field of paleontology. Among h ...
, who considered the sauropod to be closely related to the North American taxon ''
Diplodocus
''Diplodocus'' (, , or ) was a genus of diplodocid sauropod dinosaurs, whose fossils were first discovered in 1877 by S. W. Williston. The generic name, coined by Othniel Charles Marsh in 1878, is a neo-Latin term derived from Greek διπ ...
''. Alfred Leeds offered the sauropod to the
British Museum of Natural History
The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum an ...
(BMNH, now abbreviated as NHMUK) for £250, which would equate to about £30,529 in 2017. The NHMUK had earlier in 1890 and 1892 bought the First and Second Collections of Alfred Leeds, respectively. Woodward, Keeper of Geology at the NHMUK, had "great pleasure" to recommend to the Trustees of the NHMUK the fossil be purchased. The purchase was sanctioned on 25 February 1899, along with the purchase of assorted other remains for just over £357 (~£43,596 now), where the Leeds sauropod gained the accession number BMNH R3078 (now NHMUK R3078).
The amount of material made NHMUK R3078 the most complete sauropod specimen from the
United Kingdom
The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and North ...
, comparable only later to the " Rutland Dinosaur" (referred to ''Cetiosaurus'') discovered in 1967. Known regions of the specimen include the
forelimb
A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial ( anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used instead. ...
,
hindlimb
A hindlimb or back limb is one of the paired articulated appendages (limbs) attached on the caudal ( posterior) end of a terrestrial tetrapod vertebrate's torso.http://www.merriam-webster.com/medical/hind%20limb, Merriam Webster Dictionary-Hindl ...
and
vertebral column
The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordata, ...
. The forelimb lacks the manus (hand) and part of the
radius
In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...
and
ulna
The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
, although the hindlimb lacks only a few bones in the pes (foot) and fragments of the
tibia
The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
,
fibula
The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity is ...
and ilium. The vertebrae known are four parts of dorsal vertebrae, the neural spines of the
sacrum
The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30.
The sacrum situates at the upper, back part ...
, multiple
anterior
Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
caudal vertebrae
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 ...
(tail bones), and a series of 27 nearly complete vertebrae from the middle of the tail with associated or articulated
chevrons
Chevron (often relating to V-shaped patterns) may refer to:
Science and technology
* Chevron (aerospace), sawtooth patterns on some jet engines
* Chevron (anatomy), a bone
* '' Eulithis testata'', a moth
* Chevron (geology), a fold in rock l ...
(ribs along the underside of the tail), although the vertebral series is not continuous. A tail tip (NHMUK R1967) from the same locality, but a different individual was thought by palaeontologist Alan Charig in 1980 to belong to ''Cetiosauriscus''. The assignment of NHMUK R1967 to ''Cetiosauriscus'' was considered unlikely in alternate studies by palaeontologists
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 ...
, Paul Upchurch and
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'', ''Xenopos ...
because of the lack of overlap and uncertain phylogenetic positions. In
1903
Events January
* January 1 – Edward VII is proclaimed Emperor of India.
* January 19 – The first west–east transatlantic radio broadcast is made from the United States to England (the first east–west broadcast having been ...
, the skeleton was mounted as preserved in the British Museum, so it could be more easily compared with other mounted sauropods from
North America
North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
. The mount of ''Cetiosauriscus'' was put on display just prior to the cast skeleton of ''Diplodocus'', and was displayed with the dorsal vertebrae NHMUK R1984 and some isolated teeth from a
camarasaurid
Camarasauridae is a family of sauropod dinosaurs. Among sauropods, camarasaurids are small to medium-sized, with relatively short necks. They are visually identifiable by a short skull with large nares, and broad, spatulate teeth filling a thic ...
(possibly referable to ''Cetiosauriscus''), making it the first sauropod skeleton mounted in the United Kingdom.
NHMUK R3078 was referred in 1905 by Arthur Woodward to the species ''Cetiosaurus leedsii'', as it was from the same geologic formation as other specimens that were assigned to ''C. leedsii''. Woodward also referred the dorsal vertebrae NHMUK R1984 and the tail tip NHMUK R1967 to the species. In 1927, Huene briefly described the anatomy of the species ''C. leedsii'', where he noted that it shared many similarities with ''
Haplocanthosaurus
''Haplocanthosaurus'' (meaning "simple spined lizard") is a genus of intermediate sauropod dinosaur. Two species, ''H. delfsi'' and ''H. priscus'', are known from incomplete fossil skeletons. It lived during the late Jurassic period (Kimmeridgia ...
'' and was most likely between ''Cetiosaurus'' proper and the former genus. For this reason, Huene proposed the new genus name ''Cetiosauriscus'' for the species. To the genus he referred the specimens NHMUK R1984–R1988 and NHMUK R3078.
Misassigned species
Huene (1927) assigned ''"Ornithopsis" greppini'', which he had named in
1922
Events
January
* January 7 – Dáil Éireann (Irish Republic), Dáil Éireann, the parliament of the Irish Republic, ratifies the Anglo-Irish Treaty by 64–57 votes.
* January 10 – Arthur Griffith is elected President of Dáil Éirean ...
, to the genus ''Cetiosauriscus''. The known material, discovered in Late Jurassic (
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 ...
) deposits in the
Reuchenette Formation
The Reuchenette Formation is a Jurassic geologic formation in Switzerland. It is Kimmeridgian in age and predominantly consists of well stratified limestone, with lithology variable both laterally and stratigraphically including wackestones, pack ...
of
Switzerland
). Swiss law does not designate a ''capital'' as such, but the federal parliament and government are installed in Bern, while other federal institutions, such as the federal courts, are in other cities (Bellinzona, Lausanne, Luzern, Neuchâtel ...
, includes dorsal and caudal vertebrae, forelimb bones, and a hindlimb and partial pes, from at least two individuals. The long humerus was built like ''Cetiosauriscus stewarti'' (''C. leedsi'' of von Huene's 1927 usage), and the two species were originally distinguished from ''Cetiosaurus'' by having shorter dorsal vertebrae, a shorter forelimb, and a longer lower leg. Similarities such as the anatomy of the caudal vertebrae were suggested by Christian Meyer and Basil Thüring in 2003 to support the referral of ''greppini'' to ''Cetiosauriscus''. However, Weishampel et al. (2004) and Whitlock (2011) considered ''"Cetiosauriscus" greppini'' to be
Eusauropoda
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 ''Barapasaur ...
''
incertae sedis
' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
'', while Hofer (2005) and Schwarz et al. (2007) concluded that ''"Cetiosauriscus" greppini'' represents an unnamed genus of basal eusauropod. ''"Ornithopsis" greppini'' was finally named as the new genus ''
Amanzia
''Amanzia'' (after Swiss geologist Amanz Gressly) is a genus of turiasaurian sauropod dinosaur from the Reuchenette Formation in Moutier, Switzerland. The type and only species is ''Amanzia greppini'', originally named as a species of ''Orni ...
'' in 2020.
The species ''Cetiosaurus longus'', named in 1842 by Owen, was referred to the genus ''Cetiosauriscus'' without comment by sauropod palaeontologist John Stanton McIntosh in 1990. The species was named for a dorsal and caudal vertebrae from the
Portland Stone
Portland stone is a limestone from the Tithonian stage of the Jurassic period quarried on the Isle of Portland, Dorset. The quarries are cut in beds of white-grey limestone separated by chert beds. It has been used extensively as a building sto ...
of
Garsington
Garsington is a village and civil parish about southeast of Oxford in Oxfordshire. "A History of the County of Oxfordshire" provides a detailed history of the parish from 1082. The 2011 Census recorded the parish's population as 1,689. The vi ...
,
Oxfordshire
Oxfordshire is a ceremonial and non-metropolitan county in the north west of South East England. It is a mainly rural county, with its largest settlement being the city of Oxford. The county is a centre of research and development, primarily ...
(both now missing) and two other caudal vertebrae from the same deposit of nearby
Thame
Thame is a market town and civil parish in Oxfordshire, about east of the city of Oxford and southwest of Aylesbury. It derives its name from the River Thame which flows along the north side of the town and forms part of the county border wi ...
. Owen also referred a single vertebra and some
metatarsals
The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the medi ...
nomen nudum
In taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published with an adequate descr ...
'') to the species. One of the vertebrae (OUMNH J13871) may instead be a
cervical
In anatomy, cervical is an adjective that has two meanings:
# of or pertaining to any neck.
# of or pertaining to the female cervix: i.e., the ''neck'' of the uterus.
*Commonly used medical phrases involving the neck are
**cervical collar
**cerv ...
, as it has one mildly convex and one concave articular face. Characterised as having uniquely elongate vertebral
centra
Centra is a convenience shop chain that operates throughout Ireland. The chain operates as a symbol group owned by Musgrave Group, the food wholesaler, meaning the stores are all owned by individual franchisees.
The chain has three different f ...
(body of the vertebra), ''C. longus'' is not a
diagnostic
Diagnosis is the identification of the nature and cause of a certain phenomenon. Diagnosis is used in many different disciplines, with variations in the use of logic, analytics, and experience, to determine " cause and effect". In systems engine ...
taxon. As it lacks any diagnostic features of ''Cetiosauriscus'', the species should be referred to by its original designation, ''Cetiosaurus longus''.
''Cetiosaurus glymptonensis'', named on the basis of nine middle-distal caudal centra from the
Forest Marble Formation
The Forest Marble is a geological formation in England. Part of the Great Oolite Group, it dates to the late Bathonian stage of the Middle Jurassic.Weishampel, David B; et al. (2004). "Dinosaur distribution (Middle Jurassic, Europe)." In: Weisham ...
of Oxfordshire, England, was referred to ''Cetiosauriscus'' by McIntosh in 1990. These caudal vertebrae were considered to be more elongate than those of ''Cetiosaurus oxoniensis'', but caudal length proportions vary significantly throughout the tail and in different taxa, ''
Apatosaurus
''Apatosaurus'' (; meaning "deceptive lizard") is a genus of herbivorous sauropod dinosaur that lived in North America during the Late Jurassic period. Othniel Charles Marsh described and named the first-known species, ''A. ajax'', in 1877, an ...
'', ''Diplodocus'' and ''Cetiosauriscus'' having similarly elongate caudal vertebrae. The more anterior caudals have a large ridge two-thirds up the centrum and a smaller ridge one-third up. These ridges are similar to the middle caudals of ''Cetiosauriscus''. Still, they are absent in caudals of the same size and proportions, and because of this difference the species was concluded to be separate from ''Cetiosauriscus'' by Upchurch and Martin in 2003. "Cetiosaurus" ''glymptonensis'' is considered to be Eusauropoda ''incertae sedis'' by Upchurch and Martin (2003), Weishampel and colleagues (2004) and Whitlock (2011), and is in need of a new genus name because it has a single diagnostic feature, the lateral ridges.
In
1980
Events January
* January 4 – U.S. President Jimmy Carter proclaims a grain embargo against the USSR with the support of the European Commission.
* January 6 – Global Positioning System time epoch begins at 00:00 UTC.
* January 9 – ...
, Charig described a specimen of indeterminate diplodocid from the Early Cretaceous of England, and re-examined the holotype of ''Cetiosauriscus leedsii'' to compare its characteristics. In this publication he confirmed that the ilium of the holotype of ''C. leedsii'', NHMUK R1988, was too incomplete to be compared to the also incomplete ilium of the referred specimen NHMUK R3078. Because of the lack of overlap the referral of NHMUK R3078 to ''Cetiosauriscus leedsii'' was no longer verifiable, so Charig named the new species ''Cetiosauriscus stewarti'' for NHMUK R3078. The specific name was chosen to honour Sir Ronald Stewart, the chairman of the London Brick Company that owned the clay pit the fossils had been found in. Furthermore, Charig considered ''Cetiosauriscus leedsii'' and ''Cetiosauriscus greppini'' to be
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 ...
taxa, making ''C. stewarti'' the only valid species within ''Cetiosauriscus''. Because of the invalidity of the type species ''C. leedsii'', Charig made a petition to the
International Code of Zoological Nomenclature
The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its publisher, the ...
(ICZN) in
1993
File:1993 Events Collage.png, From left, clockwise: The Oslo I Accord is signed in an attempt to resolve the Israeli–Palestinian conflict; The Russian White House is shelled during the 1993 Russian constitutional crisis; Czechoslovakia is peace ...
to designate ''Cetiosauriscus stewarti'' as the type species of its genus, being the taxon containing the specimen Huene had originally named the genus for and distinguished from ''Cetiosaurus'', and also a taxonomically more stable name. This was accepted by the ICZN in
1995
File:1995 Events Collage V2.png, From left, clockwise: O.J. Simpson is O. J. Simpson murder case, acquitted of the murders of Nicole Brown Simpson and Ronald Goldman from the 1994, year prior in "The Trial of the Century" in the United States; The ...
, making ''Cetiosauriscus stewarti'' the type species of ''Cetiosauriscus''. The only specimen that can confidently be assigned to ''C. stewarti'' is the holotype NHMUK R3078, although it is possible that isolated teeth from the Oxford Clay could be from the taxon.
Description
''Cetiosauriscus'' was a moderately sized, quadrupedal eusauropod. It had a moderately long tail, and relatively long arms making the shoulders level with the hips. ''Cetiosauriscus'' was approximately long based on the known skeleton, comparable to possible relatives like long ''Cetiosaurus'', and long ''
Patagosaurus
''Patagosaurus'' (meaning "Patagonia lizard") is an extinct genus of eusauropod dinosaur from the Middle-Late Toarcian of Patagonia, Argentina. It was first found in deposits of the Cañadón Asfalto Formation, which date to around 179 to 177&n ...
''. The weight of ''Cetiosauriscus'' is less certain, depending on its phylogenetic placement. Restored as a diplodocid, ''Cetiosauriscus'' was estimated by Paul (2010) as , but restored as a cetiosaur it was estimated by Paul (2016) as .
Vertebrae
The dorsal vertebrae of NHMUK R3078 are incomplete or fragmentary. A partial anterior dorsal is known from a single centrum, which is about as long as wide, with a strong anterior articular ball (an opisthocoelous condition). On the
lateral
Lateral is a geometric term of location which may refer to:
Healthcare
*Lateral (anatomy), an anatomical direction
*Lateral cricoarytenoid muscle
*Lateral release (surgery), a surgical procedure on the side of a kneecap
Phonetics
*Lateral cons ...
surfaces (sides) of the centrum there are deep but small
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 ...
(depressions in the sides of vertebrae for
air sac
Air sacs are spaces within an organism where there is the constant presence of air. Among modern animals, birds possess the most air sacs (9–11), with their extinct dinosaurian relatives showing a great increase in the pneumatization (presence ...
s). A single middle dorsal centrum is preserved, slightly smaller than the anterior dorsal. The pleurocoel is more elongate, but like the anterior dorsal there is no
ventral
Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Th ...
(underside) concavity. A posterior dorsal is also known and is probably the last dorsal before the sacrum (vertebrae between the pelves). It preserves the entire centrum and most of the neural arch, and is significantly shortened in length compared to the other dorsals, although it is about as wide across as tall. A shallow pleurocoel is also present, but is placed higher on the side of the centrum and disappears into the
neural arch
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 i ...
. Unlike the anterior dorsal, the posterior dorsal is only very slightly opisthocoelous. A tall and narrow
hyposphene The hyposphene-hypantrum articulation is an accessory joint found in the vertebrae of several fossil reptiles of the group Archosauromorpha. It consists of a process on the backside of the vertebrae, the hyposphene, that fits in a depression in the ...
(thin vertical ridge below the anterior processes of the arch, providing additional vertebral articulation) is present and well expanded off the arch. A single dorsal
neural spine
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 i ...
is also preserved. It is flattened and not tall, with a narrowed tip, and the only noticeable laminae present are the spinopostzygapophyseal laminae running down the rear corners of the spine to the
postzygapophyses
The articular processes or zygapophyses (Greek ζυγον = "yoke" (because it links two vertebrae) + απο = "away" + φυσις = "process") of a vertebra are projections of the vertebra that serve the purpose of fitting with an adjacent vertebr ...
. This is unlike most diplodocoids where there are many laminar running along the length of the spines. Four neural spines of the sacrum are preserved, three of which form a single plate and the fourth of which is separate, like in ''Diplodocus''.
Of the anterior four caudal vertebrae, the anteriormost two are highly incomplete. Both the short, but wider-than-tall, centra preserve traces of the sideways projections (
transverse process
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 i ...
es) found in other vertebrae, which are very low on the sides compared to following caudals. The neural spines are very thin, thinning to a single ridge in front (the prespinal lamina), but having two spinopostzygapophyseal laminae like the dorsals. The fourth caudal is the most complete anterior caudal. The centrum is concave in front, but flat behind ( amphiplatyan). There are no pleurocoels unlike the dorsals, and the transverse processes begin in the top half of the centrum. The centrum is long, tall and wide, with the total vertebra being tall. Anterior caudals of the ''Cetiosaurus leedsi'' specimen NHMUK R1984 are very similar to those of ''Cetiosauriscus'', but the neural arches are not as tall in ''C. leedsi'', and the transverse processes lack a prominent ridge along the top of them. Middle and posterior caudals from a nearly continuous series of 27 bones are well preserved in ''Cetiosauriscus''. The later vertebrae are slightly more elongate than those in front, and slightly less concave in their anterior face. Moving towards the end of the tail the centra reduce in size and the transverse processes shrink until they are completely absent, with the neural spines becoming shorter, thinner, and more slanted. The seventh vertebrae of the series, at tall, is long, only one cm longer than the 21st of the same series that is tall. A distinguishing characteristic of ''Cetiosauriscus'' is the presence of a front-to-back concavity on the top of the anterior and middle caudal neural spines.
Appendicular skeleton
''Cetiosauriscus'' preserves a single right scapula (shoulder blade), which is elongate and slender. The scapula is long and wide at the middle, making it very narrow. The inner face is flat across, while the outer face is gently convex. There is no expansion of the far end of the blade. The bone thickens close to the humerus joint, where it also articulates with the coracoid. The coracoid is incomplete, but enough is preserved to show it is rectangular, and longer, at , than it is wide—. A long humerus is known, and complete with minimal crushing. The bone is short and stout, with a robust crest for the deltoid muscle along the upper half of the bone. The shape of the humerus is similar to the shortness of ''
Neuquensaurus
''Neuquensaurus'' (meaning "Neuquén lizard") is a genus of saltasaurid sauropod dinosaur that lived in the Late Cretaceous, about 80 million years ago in Argentina and Uruguay in South America. Its fossils were recovered from outcrops of the Ana ...
'', although overall the forelimb is long, as in ''Diplodocus'' and ''Cetiosaurus'', being 69% of the femur length. The distal end is roughened for a large
cartilage
Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck an ...
cap as found in some other eusauropods like "Cetiosauriscus" ''greppini''. The radius and ulna are broken, but complete they would have been long.
The hindlimb of ''Cetiosauriscus'' is about the length of the forelimb. Both ilia are very fragmentary, but the two sides supplement each other to give a reasonable idea of the proportions of the complete bone. The ilium is long, and has a long and slender pubic peduncle. It is proportionally lower than in ''Cetiosaurus'', being similar in proportions to ''Haplocanthosaurus'' and the later "
Titanosauridae
Lithostrotia is a clade of derived titanosaur sauropods that lived during the Early Cretaceous and Late Cretaceous. The group was defined by Upchurch ''et al.'' in 2004 as the most recent common ancestor of ''Malawisaurus'' and ''Saltasaurus'' an ...
". The left femur is complete, but part of the shaft is eroded away. It is very slender, being tall but only wide at the middle. This very gracile femoral morphology is shared with ''
Amphicoelias
''Amphicoelias'' (, meaning "biconcave", from the Greek ἀμφί, ''amphi'': "on both sides", and κοῖλος, ''koilos'': "hollow, concave") is a genus of herbivorous sauropod dinosaur that lived approximately 150 million years ago during t ...
'', ''
Shunosaurus
''Shunosaurus'', meaning "shu lizard", is a genus of sauropod dinosaur from Late Jurassic ( Oxfordian) beds in Sichuan Province in China, approximately 159±2 million years ago. The name derives from "Shu", an ancient name for the Sichuan p ...
'', ''
Ligabuesaurus
''Ligabuesaurus'' is a genus of somphospondylan sauropod dinosaur from the Early Cretaceous that lived in what is now Argentina. The type species, ''Ligabuesaurus leanzai'', was described by Jose Bonaparte, Gonzalez Riga, and Sebastián Apeste ...
'' and a specimen of ''Diplodocus'', being more gracile than ''Cetiosaurus'' and most other eusauropods. A prominent fourth trochanter is present, but the remaining shaft is very compressed. The tibia, fibula and pes are also preserved, but are fragmentary and disarticulated making comparisons difficult, the lower hindlimb being about upright. The foot is similar to ''Diplodocus'' and ''
Brontosaurus
''Brontosaurus'' (; meaning "thunder lizard" from Greek , "thunder" and , "lizard") is a genus of gigantic quadruped sauropod dinosaurs. Although the type species, ''B. excelsus'', had long been considered a species of the closely related ''A ...
'', where the first toes are large and clawed, and the outer ones are small and clawless. Metatarsal III is the longest, followed by metatarsal IV, II, V and I. Metatarsal I is the widest, and the width of the bones decreases numerically.
Classification
''Cetiosauriscus'' was originally classified by Huene as a genus in the
family
Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Cetiosauridae
Cetiosauridae is a family of sauropod dinosaurs which was first proposed by Richard Lydekker in 1888. While traditionally a wastebasket taxon containing various unrelated species, some recent studies have found that it may represent a natural c ...
, within the
subfamily
In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
Rhoetosaurus
''Rhoetosaurus'' (meaning "Rhoetos lizard"), named after Rhoetus, a titan in Greek Mythology, is a genus of sauropod dinosaur from the Jurassic ( Oxfordian) Walloon Coal Measures of what is now eastern Australia. ''Rhoetosaurus'' is estimated t ...
'', was founded upon the general basal features of elongate cervicals and shortened dorsals—both opisthocoelous, amphicoelous caudals that are rod-shaped distally, paired sternal plates, an ilium lacking the postacetabular process (region of the ilium behind the ischium joint and
acetabulum
The acetabulum (), also called the cotyloid cavity, is a concave surface of the pelvis. The head of the femur meets with the pelvis at the acetabulum, forming the hip joint.
Structure
There are three bones of the ''os coxae'' (hip bone) that c ...
), a very wide pubis, wide distal ischium, significantly shorter forelimb than hindlimb, fibula lacking the middle muscle attachment, and long
metacarpal
In human anatomy, the metacarpal bones or metacarpus form the intermediate part of the skeletal hand located between the phalanges of the fingers and the carpal bones of the wrist, which forms the connection to the forearm. The metacarpal bones ...
s and short
metatarsal
The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the med ...
s. This classification was emmended in 1932 when Huene concluded ''Cetiosauriscus'' was closer to ''Haplocanthosaurus'' than ''Cetiosaurus'' in the family, because of forelimb and hindlimb proportions. Conversely, in
1956
Events
January
* January 1 – The Anglo-Egyptian Sudan, Anglo-Egyptian Condominium ends in Sudan.
* January 8 – Operation Auca: Five U.S. evangelical Christian Missionary, missionaries, Nate Saint, Roger Youderian, Ed McCully, Jim ...
,
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 ...
synonymised ''Cetiosauriscus'' and ''Cetiosaurus'', a position that has not been followed by subsequent studies on the taxon.
David S. Berman and McIntosh in
1978
Events January
* January 1 – Air India Flight 855, a Boeing 747 passenger jet, crashes off the coast of Bombay, killing 213.
* January 5 – Bülent Ecevit, of Republican People's Party, CHP, forms the new government of Turkey (42nd go ...
referred ''Cetiosauriscus'' to the family
Diplodocidae
Diplodocids, or members of the family Diplodocidae ("double beams"), are a group of sauropod dinosaurs. The family includes some of the longest creatures ever to walk the Earth, including ''Diplodocus'' and '' Supersaurus'', some of which may hav ...
along with multiple other genera; ''Diplodocus'', ''Apatosaurus'', ''Barosaurus'', ''Mamenchisaurus'' ''
Dicraeosaurus
''Dicraeosaurus'' (Gr. , ' "bifurcated, double-headed" + Gr. , ' "lizard") is a genus of diplodocoid sauropod dinosaur that lived in what is now Lindi Region, Tanzania during the late Jurassic period. The genus was named for the neural spines on ...
'' and ''
Nemegtosaurus
''Nemegtosaurus'' (meaning 'Reptile from the Nemegt') was a sauropod dinosaur from the Late Cretaceous Period of what is now Mongolia. It was named after the Nemegt Basin in the Gobi Desert, where the remains — a single skull — were found. Th ...
''. Like other members of the family, ''Cetiosauriscus'' possesses wing-like transverse processes, divided chevrons with forward and backward projections, the tail is "whiplash"-like, the humerus is 2/3 the length of the femur, the
calcaneum
In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is the point of the hock.
St ...
is absent, metatarsal III and IV are the longest, and metatarsal I has a process on the bottom back corner. This referral would make ''Cetiosauriscus'', known from the Callovian, the oldest diplodocid, millions of years older than ''Diplodocus'', ''Barosaurus'' or ''Apatosaurus''. In the paper naming ''Cetiosauriscus stewarti'', Charig also described the chevrons of a new specimen and created the term "diplodociform" to describe them. This meant they were robust and double-beamed, as in ''Diplodocus'' and its relatives like ''Mamenchisaurus''. Because of the similarly "diplodociform" chevrons, Charig referred ''Cetiosauriscus'' to the Diplodocidae along with the new specimen. Elaborating upon his earlier paper, McIntosh (1990) weakly referred ''Cetiosauriscus'' to the subfamily
Diplodocinae
Diplodocinae is an extinct subfamily of diplodocid sauropods that existed from the Late Jurassic to Early Cretaceous of North America, Europe, Africa and South America, about 161.2 to 136.4 million years ago. Genera within the subfamily include ...
, characterised by more cervicals and fewer dorsals, tall sacrum neural spines, short forelimbs, no calcaneum, metatarsals III and IV being the longest, and a small process on the distal end of metatarsal I. The subfamily also included ''Diplodocus'', ''
Barosaurus
''Barosaurus'' ( ) was a giant, long-tailed, long-necked, plant-eating sauropod dinosaur closely related to the more familiar ''Diplodocus''. Remains have been found in the Morrison Formation from the Upper Jurassic Period of Utah and South Da ...
'' and ''Apatosaurus''. In
2004
2004 was designated as an International Year of Rice by the United Nations, and the International Year to Commemorate the Struggle Against Slavery and its Abolition (by UNESCO).
Events January
* January 3 – Flash Airlines Flight 6 ...
this placement was followed by Weishampel ''et al.'' without comment.
A
phylogenetic analysis
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
of ''Cetiosauriscus'' was conducted in 2003 by Julia Heathcote and Upchurch, based upon the two most inclusive matrices of the time, those of Jeffrey A. Wilson (2002) and Upchurch (1995), neither of which had included the taxon in the past. Added to the analysis of Upchurch, ''Cetiosauriscus'' placed as the sister taxon of '' Tehuelchesaurus'', in a group including ''
Mamenchisaurus
''Mamenchisaurus'' (or spelling pronunciation ) is a genus of sauropod dinosaur known for their remarkably long necks which made up nearly half the total body length. Numerous species have been assigned to the genus; however, many of these might ...
'', ''
Omeisaurus
''Omeisaurus'' (meaning "Omei lizard") is a genus of sauropod dinosaur from the Middle Jurassic Period (Bathonian-Callovian stage) of what is now China. Its name comes from Mount Emei, where it was discovered in the lower Shaximiao Formation of ...
'' and ''
Euhelopus
''Euhelopus'' is a genus of sauropod dinosaur that lived between 145 and 133 million years ago during the Berriasian and Valanginian stages of the Early Cretaceous in what is now Shandong Province in China. It was a large quadrupedal herbivore. U ...
'', and a placement within a group of ''Omeisaurus'' and ''Mamenchisaurus'' was also found by using the Wilson matrix. Based on these two results, Heathcote and Upchurch concluded ''Cetiosauriscus'' was not a diplodocid or even within
Diplodocoidea
Diplodocoidea is a superfamily of sauropod dinosaurs, which included some of the longest animals of all time, including slender giants like ''Supersaurus'', ''Diplodocus'', ''Apatosaurus'', and ''Amphicoelias''. Most had very long necks and long, ...
, instead being a more basal sauropod outside
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 ...
. The phylogenetic analysis of Rauhut ''et al.'' (2005) resolved ''Cetiosauriscus'' in a clade with ''Omeisaurus'', itself in a group with ''
Losillasaurus
''Losillasaurus'' (meaning " Losilla lizard") is a genus of sauropod dinosaur from the Late Jurassic and possibly Early Cretaceous (Kimmeridgian-?Berriasian) in the southeast of Spain. The type species of the turiasaurian ''Losillasaurus giganteu ...
'' and ''Mamenchisaurus'', outside of Neosauropoda. The phylogenetic relationships of ''Cetiosauriscus'' were also tested in
2015
File:2015 Events Collage new.png, From top left, clockwise: Civil service in remembrance of November 2015 Paris attacks; Germanwings Flight 9525 was purposely crashed into the French Alps; the rubble of residences in Kathmandu following the Apri ...
by Tschopp ''et al.'', as a potential diplodocid. Although the genus was found to be within
Diplodocimorpha
Diplodocoidea is a superfamily of sauropod dinosaurs, which included some of the longest animals of all time, including slender giants like ''Supersaurus'', ''Diplodocus'', ''Apatosaurus'', and ''Amphicoelias''. Most had very long necks and long, ...
with one analysis method, it was also found outside Neosauropoda. In both, ''Cetiosauriscus stewarti'' was found to be in a clade alone with '' Barosaurus affinis'', a dubious species known only from foot bones. Tschopp ''et al.'' concluded that ''Cetiosauriscus'' was not a diplodocid or a diplodocoid, as forcing it to be outside Neosauropoda was more
parsimonious
Occam's razor, Ockham's razor, or Ocham's razor ( la, novacula Occami), also known as the principle of parsimony or the law of parsimony ( la, lex parsimoniae), is the problem-solving principle that "entities should not be multiplied beyond neces ...
than forcing it to be within Diplodocoidea in all analyses. As the paper was only to test relationships within Diplodocidae, more solid conclusions regarding the position of ''Cetiosauriscus'' could not be made. The results of the favoured
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 d ...
of Tschopp ''et al.'' is shown below:
Palaeobiology
Palaeopathology
The series of distal caudal vertebrae NHMUK R1967, once referred to ''Cetiosauriscus'', is similar to the caudals of ''Diplodocus'', with two convex ends ( biconvex) and a long and thin centrum. These caudals display signs of injury at two points along the series of ten vertebrae, where there are signs of breakage that was later healed. These lesions were identified as the same form of pathologies as found on the tail of ''Diplodocus''. It has been suggested that the biconvex distal caudal vertebrae in sauropods were used for making whip-like cracking noise, being thin and delicate and not intended for impact, as the joints would be very vulnerable to damage rendering them useless.
Palaeoecology
''Cetiosauriscus'' lived during the Callovian, an
epoch
In chronology and periodization, an epoch or reference epoch is an instant in time chosen as the origin of a particular calendar era. The "epoch" serves as a reference point from which time is measured.
The moment of epoch is usually decided by ...
in the Middle Jurassic, about 166 to 164 mya.
The single specimen is known from the Lower Member of the Oxford Clay Formation, along with multiple other dinosaur genera and many other groups of animals, in the
biozone
In biostratigraphy, biostratigraphic units or biozones are intervals of geological strata that are defined on the basis of their characteristic fossil taxa, as opposed to a lithostratigraphic unit which is defined by the lithological properties of ...
of the
index fossil
Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock Stratum, strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictiona ...
'' Kosmoceras jason''. The Oxford Clay Formation is a marine deposit of southern and middle England, known for the high-quality preservation of some fossils and the large diversity of taxa. Sediments are generally brownish-grey
mudstone
Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from '' shale'' by its lack of fissility (parallel layering).Blatt, H., and R.J. Tracy, 1996, ''Petrology. ...
, organic-rich with plentiful crushed
ammonite
Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttlefish) ...
s and
bivalve
Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
s, at most thick. A large diversity of
flora
Flora is all the plant life present in a particular region or time, generally the naturally occurring (indigenous) native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' gut flora'' or '' skin flora''.
E ...
can be seen, preserved in the form of pollen and spores.
Gymnosperms
The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό ...
are present, along with
pteridophyte
A pteridophyte is a vascular plant (with xylem and phloem) that disperses spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as "cryptogams", meaning that their means of reproduction is hidden. Ferns, ...
s, unidentifiable wood fragments, other intermediate pollen, and miscellaneous organic plant material.
The intermediate sauropod '' Ornithopsis leedsi'' is known from the same section of the formation as ''Cetiosauriscus'', along with the stegosaurids '' Lexovisaurus durobrivensis'' and '' Loricatosaurus priscus'' (which are possibly synonyms), the basal
ankylosaur
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. ...
ornithopod
Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (), that started out as small, bipedal running grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous world ...
'' Callovosaurus leedsi'', and a second unnamed ornithopod taxon. Dinosaur eggs that have not yet been assigned to a taxon are also known from the Lower Oxford Clay. The
theropod
Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
s ''
Eustreptospondylus
''Eustreptospondylus'' ( ; meaning "true ''Streptospondylus''") is a genus of megalosaurid Theropoda, theropod dinosaur, from the Oxfordian stage of the Late Jurassic period (some time between 163 and 154 million years ago) in southern Eng ...
'' and possibly ''
Megalosaurus
''Megalosaurus'' (meaning "great lizard", from Greek , ', meaning 'big', 'tall' or 'great' and , ', meaning 'lizard') is an extinct genus of large carnivorous theropod dinosaurs of the Middle Jurassic period (Bathonian stage, 166 million years a ...
'' are also known from the Oxford Clay Formation, but slightly younger deposits (the Middle Member). In addition, the theropod ''
Metriacanthosaurus
''Metriacanthosaurus'' (meaning "moderately-spined lizard") is a genus of metriacanthosaurid dinosaur from the upper Oxford Clay of England, dating to the Late Jurassic period, about 160 million years ago (lower Oxfordian).
History of discovery ...
'' is from an unknown level and age in the formation.
Hundreds of
invertebrate
Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s are known from the marine deposits, including bivalves,
gastropod
The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda ().
This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
nautiloid
Nautiloids are a group of marine cephalopods ( Mollusca) which originated in the Late Cambrian and are represented today by the living ''Nautilus'' and ''Allonautilus''. Fossil nautiloids are diverse and speciose, with over 2,500 recorded species ...
,
foraminifera
Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
,
coelenterate
Radiata or Radiates is a historical taxonomic rank that was used to classify animals with radially symmetric body plans. The term Radiata is no longer accepted, as it united several different groupings of animals that do not form a monophyletic ...
s,
bryozoa
Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...
ns,
brachiopod
Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, w ...
s,
annelid
The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
s,
crustacean
Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s,
ostracod
Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typic ...
s,
cirripede
A barnacle is a type of arthropod constituting the subclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in eros ...
s and
echinoderm
An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...
s. Fish are known from the clades
Elasmobranchii
Elasmobranchii () is a subclass of Chondrichthyes or cartilaginous fish, including sharks (superorder Selachii), rays, skates, and sawfish (superorder Batoidea). Members of this subclass are characterised by having five to seven pairs of gil ...
,
Chimaera
Chimaeras are cartilaginous fish in the order Chimaeriformes , known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively.
At ...
, and
Actinopterygii
Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species.
The ray-finned fishes are so called because their fins are webs of skin supported by bony or h ...
, and the
ichthyosaur
Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, altho ...
''
Ophthalmosaurus
''Ophthalmosaurus'' (meaning "eye lizard" in Greek) is an ichthyosaur of the Jurassic period (165–150 million years ago). Possible remains from the Cretaceous, around 145 million years ago, are also known. It was a relatively medium-sized ichth ...
'', the
plesiosaur
The Plesiosauria (; Greek: πλησίος, ''plesios'', meaning "near to" and ''sauros'', meaning "lizard") or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia.
Plesiosaurs first appeared ...
s ''
Cryptoclidus
''Cryptoclidus'' ( ) is a genus of plesiosaur reptile from the Middle Jurassic Period (geology), period of England, France, and Cuba.
Discovery
''Cryptoclidus'' was a plesiosaur whose specimens include adult and juvenile skeletons, and remai ...
'', ''
Muraenosaurus
''Muraenosaurus'' (from the Latin "'' Muraena''" meaning "eel" and "''Sauros''" meaning lizard) is an extinct genus of cryptoclidid plesiosaur reptile from the Oxford Clay of Southern England. The genus was given its name due to the eel-like ...
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 ...
'', ''
Peloneustes
''Peloneustes'' (meaning "mud swimmer") is a genus of pliosaurid plesiosaur from the Middle Jurassic of England. Its remains are known from the Peterborough Member of the Oxford Clay Formation, which is Callovian in age. It was originally descri ...
'', ''
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 marine re ...
crocodilia
Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
ns ''
Metriorhynchus
''Metriorhynchus'' is an extinct genus of marine crocodyliform that lived in the oceans during the Late Jurassic. The type species, ''M. brevirostris'' was named in 1829 as a species of ''Steneosaurus'' before being named as a separate genus by ...
'' and ''
Steneosaurus
''Steneosaurus'' (from el, στενός , 'narrow' and el, σαῦρος , 'lizard') is a dubious genus of teleosaurid crocodyliform from the Middle or Late Jurassic ( Callovian or early Oxfordian) of France. The genus has been used as a w ...
'', and the
pterosaur
Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
''
Rhamphorhynchus
''Rhamphorhynchus'' (, from Ancient Greek ''rhamphos'' meaning "beak" and ''rhynchus'' meaning "snout") is a genus of long-tailed pterosaurs in the Jurassic period. Less specialized than contemporary, short-tailed pterodactyloid pterosaurs such ...