Baryonyx
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''Baryonyx'' () is a genus of theropod dinosaur which lived in the Barremian stage of 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 ...
period, about 130–125 million years ago. The first skeleton was discovered in 1983 in the
Smokejack Clay Pit Smokejack Clay Pit is a geological Site of Special Scientific Interest east of Cranleigh in Surrey. It is a Geological Conservation Review site. This site exposes Lower Cretaceous rocks of the Weald Clay Group. Fossils of six orders of insects h ...
, of
Surrey Surrey () is a ceremonial and non-metropolitan county in South East England, bordering Greater London to the south west. Surrey has a large rural area, and several significant urban areas which form part of the Greater London Built-up Area. ...
, England, in sediments of the
Weald Clay Formation Weald Clay or the Weald Clay Formation is a Lower Cretaceous sedimentary rock unit underlying areas of South East England, between the North and South Downs, in an area called the Weald Basin. It is the uppermost unit of the Wealden Group of ro ...
, and became the holotype specimen of ''Baryonyx walkeri'', named by
palaeontologists 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 ...
Alan J. Charig Alan Jack Charig (1 July 1927 – 15 July 1997) was an English palaeontologist and writer who popularised his subject on television and in books at the start of the wave of interest in dinosaurs in the 1970s. Charig was, though, first and fo ...
and
Angela C. Milner Angela Cheryl Milner (3 October 1947 – 13 August 2021) was a British paleontologist who, in 1986 alongside Alan Charig, described the dinosaur ''Baryonyx''. Early life Milner was born Angela Girven in Gosforth, daughter of Cyril and Luc ...
in
1986 The year 1986 was designated as the International Year of Peace by the United Nations. Events January * January 1 ** Aruba gains increased autonomy from the Netherlands by separating from the Netherlands Antilles. **Spain and Portugal ente ...
. The generic name, ''Baryonyx'', means "heavy claw" and alludes to the animal's very large claw on the first finger; 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 ...
, ''walkeri'', refers to its discoverer, amateur
fossil collector Fossil collecting (sometimes, in a non-scientific sense, fossil hunting) is the collection of fossils for scientific study, hobby, or profit. Fossil collecting, as practiced by amateurs, is the predecessor of modern paleontology and many stil ...
William J. Walker. The holotype specimen is one of the most complete theropod skeletons from the UK (and remains the most complete
Spinosaurid The Spinosauridae (or spinosaurids) are a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. They came into prominence during the Cretaceous period. Spinosaurid fossils have been recovered worldwide, includi ...
), and its discovery attracted media attention. Specimens later discovered in other parts of the United Kingdom and Iberia have also been assigned to the genus, though many have since been moved to new genera. The holotype specimen, which may not have been fully grown, was estimated to have been between long and to have weighed between . ''Baryonyx'' had a long, low, and narrow snout, which has been compared to that of a
gharial The gharial (''Gavialis gangeticus''), also known as gavial or fish-eating crocodile, is a crocodilian in the family Gavialidae and among the longest of all living crocodilians. Mature females are long, and males . Adult males have a distinct b ...
. The tip of the snout expanded to the sides in the shape of a rosette. Behind this, the upper jaw had a notch which fitted into the lower jaw (which curved upwards in the same area). It had a triangular
crest Crest or CREST may refer to: Buildings *The Crest (Huntington, New York), a historic house in Suffolk County, New York *"The Crest", an alternate name for 63 Wall Street, in Manhattan, New York *Crest Castle (Château Du Crest), Jussy, Switzerla ...
on the top of its
nasal bones The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...
. ''Baryonyx'' had a large number of finely serrated, conical teeth, with the largest teeth in front. The neck formed an S-shape, and the
neural spines 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 ...
of its dorsal vertebrae increased in height from front to back. One elongated neural spine indicates it may have had a hump or ridge along the centre of its back. It had robust forelimbs, with the eponymous first-finger claw measuring about long. Now recognised as a member of the family Spinosauridae, ''Baryonyx'' affinities were obscure when it was discovered. Some researchers have suggested that ''
Suchosaurus cultridens ''Suchosaurus'' (meaning "crocodile lizard") is a spinosaurid dinosaur from Cretaceous England and Portugal, originally believed to be a genus of crocodile. The type material, consisting of teeth, was used by British palaeontologist Richard Owen ...
'' is a senior synonym (being an older name), and that ''
Suchomimus tenerensis ''Suchomimus'' (meaning "crocodile mimic") is a genus of spinosaurid dinosaur that lived between 125 and 112 million years ago in what is now Niger, during the Aptian to early Albian stages of the Early Cretaceous period. It was named a ...
'' belongs in the same genus; subsequent authors have kept them separate. ''Baryonyx'' was the first theropod dinosaur demonstrated to have been piscivorous (fish-eating), as evidenced by fish scales in the stomach region of the holotype specimen. It may also have been an active predator of larger prey and a
scavenger Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding b ...
, since it also contained bones of a juvenile
iguanodontid Iguanodontidae is a family of iguanodontians belonging to Styracosterna, a derived clade within Ankylopollexia. Characterized by their elongated maxillae, they were herbivorous and typically large in size. This family exhibited locomotive dynam ...
. The creature would have caught and processed its prey primarily with its forelimbs and large claws. ''Baryonyx'' may have had
semi-aquatic In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below. Semiaquatic animals Semiaquatic animals include: * Verte ...
habits, and coexisted with other theropod,
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 ...
, and
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 ...
dinosaurs, as well as
pterosaurs 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 ...
, crocodiles, turtles and fishes, in a
fluvial In geography and geology, fluvial processes are associated with rivers and streams and the deposits and landforms created by them. When the stream or rivers are associated with glaciers, ice sheets, or ice caps, the term glaciofluvial or fluviog ...
environment.


History of discovery

In January 1983, the plumber and amateur
fossil collector Fossil collecting (sometimes, in a non-scientific sense, fossil hunting) is the collection of fossils for scientific study, hobby, or profit. Fossil collecting, as practiced by amateurs, is the predecessor of modern paleontology and many stil ...
William J. Walker explored the
Smokejack Clay Pit Smokejack Clay Pit is a geological Site of Special Scientific Interest east of Cranleigh in Surrey. It is a Geological Conservation Review site. This site exposes Lower Cretaceous rocks of the Weald Clay Group. Fossils of six orders of insects h ...
, a
clay pit A clay pit is a quarry or mine for the extraction of clay, which is generally used for manufacturing pottery, bricks or Portland cement. Quarries where clay is mined to make bricks are sometimes called brick pits. A brickyard or brickworks is of ...
in the
Weald Clay Formation Weald Clay or the Weald Clay Formation is a Lower Cretaceous sedimentary rock unit underlying areas of South East England, between the North and South Downs, in an area called the Weald Basin. It is the uppermost unit of the Wealden Group of ro ...
near
Ockley Ockley is a rural village in Surrey. It lies astride the A29, the modern road using the alignment of Stane Street (Chichester). The A29 diverges from the A24 from London about 2.5 miles northeast and takes the alignment of Stane Street a mile ...
in Surrey, England. He found a rock wherein he discovered a large claw, but after piecing it together at home, he realised the tip of the claw was missing. Walker returned to the same spot in the pit some weeks later, and found the missing part after searching for an hour. He also found a and part of a . Walker's son-in-law later brought the claw to the
Natural History Museum of London 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 ...
, where it was examined by the
palaeontologists 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 ...
Alan J. Charig Alan Jack Charig (1 July 1927 – 15 July 1997) was an English palaeontologist and writer who popularised his subject on television and in books at the start of the wave of interest in dinosaurs in the 1970s. Charig was, though, first and fo ...
and
Angela C. Milner Angela Cheryl Milner (3 October 1947 – 13 August 2021) was a British paleontologist who, in 1986 alongside Alan Charig, described the dinosaur ''Baryonyx''. Early life Milner was born Angela Girven in Gosforth, daughter of Cyril and Luc ...
, who identified it as belonging to a theropod dinosaur. The palaeontologists found more bone fragments at the site in February, but the entire skeleton could not be collected until May and June due to weather conditions at the pit. A team of eight museum staff members and several volunteers excavated of rock
matrix Matrix most commonly refers to: * ''The Matrix'' (franchise), an American media franchise ** ''The Matrix'', a 1999 science-fiction action film ** "The Matrix", a fictional setting, a virtual reality environment, within ''The Matrix'' (franchis ...
in 54 blocks over a three-week period. Walker donated the claw to the museum, and the Ockley Brick Company (owners of the pit) donated the rest of the skeleton and provided equipment. The area had been explored for 200 years, but no similar remains had been found before. Most of the bones collected were encased in
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility.Blatt ''et al.'' 1980, p ...
nodules Nodule may refer to: *Nodule (geology), a small rock or mineral cluster *Manganese nodule, a metallic concretion found on the seafloor *Nodule (medicine), a small aggregation of cells *Root nodule Root nodules are found on the roots of plants, p ...
surrounded by fine sand and silt, with the rest lying in clay. The bones were and scattered over a area, but most were not far from their natural positions. The position of some bones was disturbed by a
bulldozer A bulldozer or dozer (also called a crawler) is a large, motorized machine equipped with a metal blade to the front for pushing material: soil, sand, snow, rubble, or rock during construction work. It travels most commonly on continuous track ...
, and some were broken by mechanical equipment before they were collected. Preparing the specimen was difficult, due to the hardness of the siltstone matrix and the presence of siderite; acid preparation was attempted, but most of the matrix was removed mechanically. It took six years of almost constant preparation to get all the bones out of the rock, and by the end, dental tools and air mallets had to be used under a microscope. The specimen represents about 65 per cent of the skeleton, and consists of partial skull bones, including
premaxillae The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
(first bones of the upper jaw); the left
maxillae The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
(second bone of the upper jaw); both
nasal Nasal is an adjective referring to the nose, part of human or animal anatomy. It may also be shorthand for the following uses in combination: * With reference to the human nose: ** Nasal administration, a method of pharmaceutical drug delivery ** ...
bones; the left lacrimal; the left prefrontal; the left postorbital; the
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
including the occiput; both
dentaries In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
(the front bones of the lower jaw); various bones from the back of the lower jaw; teeth;
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 **cervic ...
(neck), dorsal (back), and
caudal Caudal may refer to: Anatomy * Caudal (anatomical term) (from Latin ''cauda''; tail), used to describe how close something is to the trailing end of an organism * Caudal artery, the portion of the dorsal aorta of a vertebrate that passes into the ...
(tail) ; ribs; a ; both (shoulder blades); both ; both (upper arm bones); the left and (lower arm bones); finger bones and (claw bones); hip bones; the upper end of the left (thigh bone) and lower end of the right; right (of the lower leg); and foot bones including an ungual. The original specimen number was BMNH R9951, but it was later re-catalogued as NHMUK VP R9951. In
1986 The year 1986 was designated as the International Year of Peace by the United Nations. Events January * January 1 ** Aruba gains increased autonomy from the Netherlands by separating from the Netherlands Antilles. **Spain and Portugal ente ...
, Charig and Milner named a new genus and species with the skeleton as holotype specimen: ''Baryonyx walkeri''. The generic name derives from ancient Greek; βαρύς (''barys'') means "heavy" or "strong", and ὄνυξ (''onyx'') means "claw" or "talon". 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 ...
honours Walker, for discovering the specimen. At that time, the authors did not know if the large claw belonged to the hand or the foot (as in dromaeosaurs, which it was then assumed to be). The dinosaur had been presented earlier the same year during a lecture at a conference about dinosaur systematics in Drumheller, Canada. Due to ongoing work on the bones (70 per cent had been prepared at the time), they called their article preliminary and promised a more detailed description at a later date. ''Baryonyx'' was the first large
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 ...
theropod found anywhere in the world by that time. Before the discovery of ''Baryonyx'' the last significant theropod find in the United Kingdom was '' Eustreptospondylus'' in 1871, and in a 1986 interview Charig called ''Baryonyx'' "the best find of the century" in Europe. ''Baryonyx'' was widely featured in international media, and was nicknamed "Claws" by journalists punning on the title of the film ''
Jaws Jaws or Jaw may refer to: Anatomy * Jaw, an opposable articulated structure at the entrance of the mouth ** Mandible, the lower jaw Arts, entertainment, and media * Jaws (James Bond), a character in ''The Spy Who Loved Me'' and ''Moonraker'' * ...
''. Its discovery was the subject of a 1987 BBC documentary, and a cast of the skeleton is mounted at the Natural History Museum in London. In 1997, Charig and Milner published a
monograph A monograph is a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monograph ...
describing the holotype skeleton in detail. The holotype specimen remains the most completely known
spinosaurid The Spinosauridae (or spinosaurids) are a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. They came into prominence during the Cretaceous period. Spinosaurid fossils have been recovered worldwide, includi ...
skeleton.


Assigned specimens

Fossils from other parts of the UK and Iberia, mostly isolated teeth, have subsequently been attributed to ''Baryonyx'' or similar animals. Isolated teeth and bones from the Isle of Wight, including hand bones reported in 1998 and a vertebra reported by the palaeontologists Steve Hutt and Penny Newbery in 2004, have been attributed to this genus. A maxilla fragment from La Rioja, Spain, was attributed to ''Baryonyx'' by the palaeontologists Luis I. Viera and José Angel Torres in 1995 (although the palaeontologist
Thomas R. Holtz Thomas Richard Holtz Jr. (born September 13, 1965) is an American vertebrate palaeontologist, author, and principal lecturer at the University of Maryland, College Park, University of Maryland's Department of Geology. He has published extensively ...
and colleagues raised the possibility that it could have belonged to ''Suchomimus'' in 2004). In 1999, a postorbital, , tooth, vertebral remains, (hand bones), and a phalanx from the Salas de los Infantes deposit in Burgos Province, Spain, were attributed to an immature ''Baryonyx'' (though some of these elements are unknown in the holotype) by the palaeontologist Carolina Fuentes Vidarte and colleagues. Dinosaur tracks near Burgos have also been suggested to belong to ''Baryonyx'' or a similar theropod. In 2011, a specimen (Catalogued as ML1190 in
Museu da Lourinhã Museu da Lourinhã is a museum in the town of Lourinhã, west Portugal. It was founded in 1984 by GEAL - Grupo de Etnologia e Arqueologia da Lourinhã (Lourinhã's Group of Ethnology and Archeology). The president of the Direction Board is Lubélia ...
) from the Papo Seco Formation in
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, Portugal, with a fragmentary dentary, teeth, vertebrae, ribs, hip bones, a scapula, and a phalanx bone, was attributed to ''Baryonyx'' by the palaeontologist Octávio Mateus and colleagues, the most complete Iberian remains of the animal. The skeletal elements of this specimen are also represented in the more complete holotype (which was of similar size), except for the mid-neck vertebrae. In 2018, the palaeontologist Thomas M. S. Arden and colleagues found that the Portuguese skeleton did not belong to ''Baryonyx'', since the front of its dentary bone was not strongly upturned. This specimen was made the basis of the new genus ''
Iberospinus ''Iberospinus'' or (meaning " Iberian spine") is an extinct genus of spinosaurid dinosaur from the Early Cretaceous (Barremian) Papo Seco Formation of Portugal. The genus contains a single species, ''I. natarioi'', known from several assorted bo ...
'' by Mateus and Darío Estraviz-López in 2022. Additional spinosaurid remains from Iberia may belong to taxa other than ''Baryonyx'', such as ''
Vallibonavenatrix ''Vallibonavenatrix'' (meaning "Vallibona huntress" after the town near where its remains were found) is a genus of spinosaurid dinosaur from the Early Cretaceous (Barremian) Arcillas de Morella Formation of Castellón, Spain. The type and onl ...
'' and '' Protathlitis'', or may be indeterminate. In 2021, the palaeontologist Chris T. Barker and colleagues described two new spinosaurid genera from the Wessex Formation of the Isle of Wight, ''
Ceratosuchops ''Ceratosuchops'' (meaning "horned crocodile face") is a genus of spinosaurid from the Early Cretaceous (Barremian) of Britain. Discovery and naming In 2021, the type species ''C. inferodios'' was named and described by a team of paleonto ...
'' and ''
Riparovenator ''Riparovenator'' ("riverbank hunter") is a genus of baryonychine spinosaurid dinosaur from the Early Cretaceous (Barremian) period of Britain, the type species is ''Riparovenator milnerae''. Discovery and naming Between 2013 and 2017, spin ...
'' (the latter named ''R. milnerae'' honouring Milner for her contributions to spinosaurid research), and stated that spinosaurid material from there that had previously been attributed to the contemporary ''Baryonyx'' could have belonged to other taxa instead. These specimens had previously been assigned to ''Baryonyx'' in a 2017 conference abstract. Barker and colleagues stated that the recognition of the Wessex Formation specimens as new genera renders the presence of ''Baryonyx'' there ambiguous, and most of the previously assigned isolated material from the
Wealden Supergroup The Wealden Group, occasionally also referred to as the Wealden Supergroup, is a group (a sequence of rock strata) in the lithostratigraphy of southern England. The Wealden group consists of paralic to continental (freshwater) facies sedimentary ...
is therefore indeterminate. A 2023 study of an isolated tooh by Barker and colleagues found that it and other teeth from the Wealden Supergroup that have previously been assigned to ''Baryonyx'' probably do not belong to the genus, based on their morphology and age.


Possible synonyms

In 2003, Milner noted that some teeth at the Natural History Museum previously identified as belonging to the genera ''
Suchosaurus ''Suchosaurus'' (meaning "crocodile lizard") is a spinosaurid dinosaur from Cretaceous England and Portugal, originally believed to be a genus of crocodile. The type material, consisting of teeth, was used by British palaeontologist Richard Owen ...
'' (the first named spinosaurid) and '' Megalosaurus'' probably belonged to ''Baryonyx''. The type species of ''Suchosaurus'', ''S. cultridens'', was named by the biologist
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 1841, based on teeth discovered by the geologist Gideon A. Mantell in
Tilgate Forest Tilgate Forest is a Local Nature Reserve in Crawley in West Sussex. It is owned and managed by Crawley Borough Council and is part of Tilgate Park. This site has woods, tall herb and fern, and heathland. The most common trees in areas of natural ...
,
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 ...
. Owen originally thought the teeth to have belonged to a
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to inclu ...
; he was yet to name the group Dinosauria, which happened the following year. A second species, ''S. girardi'', was named by the palaeontologist
Henri Émile Sauvage Henri Émile Sauvage (22 September 1842 in Boulogne-sur-Mer – 3 January 1917 in Boulogne-sur-Mer) was a French paleontologist, ichthyologist, and herpetologist. He was a leading expert on Mesozoic fish and reptiles.Éric Buffetaut considered the teeth of ''S. girardi'' very similar to those of ''Baryonyx'' (and ''S. cultridens'') except for the stronger development of the flutes (or "ribs"; lengthwise ridges), suggesting that the remains belonged to the same genus. Buffetaut agreed with Milner that the teeth of ''S. cultridens'' were almost identical to those of ''B. walkeri'', but with a ribbier surface. The former taxon might be a senior synonym of the latter (since it was published first), depending on whether the differences were within a taxon or between different ones. According to Buffetaut, since the holotype specimen of ''S. cultridens'' is a single tooth and that of ''B. walkeri'' is a skeleton, it would be more practical to retain the newer name. In 2011, Mateus and colleagues agreed that ''Suchosaurus'' was closely related to ''Baryonyx'', but considered both species in the former genus ''
nomina dubia 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 ...
'' (dubious names) since their holotype specimens were not considered diagnostic (lacking distinguishing features) and could not be definitely equated with other taxa. Barker and colleagues agreed with this in 2023. In 1997, Charig and Milner noted that two fragmentary spinosaurid snouts from the Elrhaz Formation of Niger (reported by the palaeontologist Philippe Taquet in 1984) were similar enough to ''Baryonyx'' that they considered them to belong to an indeterminate species of the genus (despite their much younger
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...
geological age). In 1998, these fossils became the basis of the genus and species ''
Cristatusaurus lapparenti ''Cristatusaurus'' is a genus of theropod dinosaur that lived during the Early Cretaceous Period of what is now Niger, 112 million years ago. It was a baryonychine member of the Spinosauridae, a group of large bipedal carnivores with wel ...
'', named by Taquet and the palaeontologist
Dale Russell Dale Alan Russell (27 December 1937 – 21 December 2019) was an American-Canadian geologist and palaeontologist. Throughout his career Russell worked as the Curator of Fossil Vertebrates at the Canadian Museum of Nature, Research Professor a ...
. The palaeontologist Paul Sereno and colleagues named the new genus and species ''
Suchomimus tenerensis ''Suchomimus'' (meaning "crocodile mimic") is a genus of spinosaurid dinosaur that lived between 125 and 112 million years ago in what is now Niger, during the Aptian to early Albian stages of the Early Cretaceous period. It was named a ...
'' later in 1998, based on more complete fossils from the Elrhaz Formation. In 2002, the German palaeontologist
Hans-Dieter Sues Hans-Dieter Sues (born January 13, 1956) is a German-born American paleontologist who is Senior Scientist and Curator of Vertebrate Paleontology at the National Museum of Natural History of the Smithsonian Institution in Washington, DC. He receiv ...
and colleagues proposed that ''Suchomimus tenerensis'' was similar enough to ''Baryonyx walkeri'' to be considered a species within the same genus (as ''B. tenerensis''), and that ''Suchomimus'' was identical to ''Cristatusaurus''. Milner concurred that the material from Niger was indistinguishable from ''Baryonyx'' in 2003. In a 2004
conference abstract An abstract is a brief summary of a research article, thesis, review, academic conference, conference proceedings, proceeding, or any in-depth analysis of a particular subject and is often used to help the reader quickly ascertain the paper's purpo ...
, Hutt and Newberry supported the synonymy based on a large theropod vertebra from the Isle of Wight which they attributed to an animal closely related to ''Baryonyx'' and ''Suchomimus''. Later studies have kept ''Baryonyx'' and ''Suchomimus'' separate, whereas ''Cristatusaurus'' has been proposed to be either a ''nomen dubium'' or possibly distinct from both. A 2017
review paper A review article is an article that summarizes the current state of understanding on a topic within a certain discipline. A review article is generally considered a secondary source since it may analyze and discuss the method and conclusions i ...
by the palaeontologist Carlos Roberto A. Candeiro and colleagues stated that this debate was more in the realm of semantics than science, as it is generally agreed that ''B. walkeri'' and ''S. tenerensis'' are distinct, related species. Barker and colleagues found ''Suchomimus'' to be closer related to the British genera ''Riparovenator'' and ''Ceratosuchops'' than to ''Baryonyx'' in 2021.


Description

''Baryonyx'' is estimated to have been between long, in hip height, and to have weighed between . The fact that elements of the skull and vertebral column of the ''B. walkeri'' holotype specimen (NHM R9951) do not appear to have
co-ossified Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in t ...
(fused) suggests that the individual was not fully grown, and the mature animal may have been much larger (as is the case for some other spinosaurids). On the other hand, the specimen's fused sternum indicates that it may have been mature.


Skull

The skull of ''Baryonyx'' is incompletely known, and much of the middle and hind portions are not preserved. The full length of the skull is estimated to have been long, based on comparison with that of the related genus ''Suchomimus'' (which was 20% larger). It was elongated, and the front of the premaxillae formed a long, narrow, and low snout ( rostrum) with a smoothly rounded upper surface. The (bony nostrils) were long, low, and placed far back from the snout tip. The front of the snout expanded into a spatulate (spoon-like), "terminal rosette", a shape similar to the rostrum of the modern
gharial The gharial (''Gavialis gangeticus''), also known as gavial or fish-eating crocodile, is a crocodilian in the family Gavialidae and among the longest of all living crocodilians. Mature females are long, and males . Adult males have a distinct b ...
. The front of the lower margin of the premaxillae was downturned (or hooked), whereas that of the front portion of the maxillae was upturned. This morphology resulted in a sigmoid or S shaped margin of the lower upper tooth row, in which the teeth from the front of the maxilla were projecting forward. The snout was particularly narrow directly behind the rosette; this area received the large teeth of the mandible. The maxilla and premaxilla of ''Baryonyx'' fit together in a complex articulation, and the resulting gap between the upper and lower jaw is known as the . A downturned premaxilla and a sigmoid lower margin of the upper tooth row was also present in distantly related theropods such as '' Dilophosaurus''. The snout had extensive (openings), which would have been exits for blood vessels and nerves, and the maxilla appears to have housed sinuses. ''Baryonyx'' had a rudimentary , similar to crocodiles but unlike most theropod dinosaurs. A rugose (roughly wrinkled) surface suggests the presence of a horny pad in the roof of the mouth. The nasal bones were fused, which distinguished ''Baryonyx'' from other spinosaurids, and a
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
was present above the eyes, on the upper mid-line of the nasals. This crest was triangular, narrow, and sharp in its front part, and was distinct from those of other spinosaurids in ending hind wards in a cross-shaped process. The lacrimal bone in front of the eye appears to have formed a horn core similar to those seen, for example, in '' Allosaurus'', and was distinct from other spinosaurids in being solid and almost triangular. The occiput was narrow, with the paroccipital processes pointing outwards horizontally, and the were lengthened, descending far below the (the lowermost bone of the occiput). Sereno and colleagues suggested that some of ''Baryonyx'' cranial bones had been misidentified by Charig and Milner, resulting in the occiput being reconstructed as too deep, and that the skull was instead probably as low, long and narrow as that of ''Suchomimus''. The front of the dentary in the mandible sloped upwards towards the curve of the snout. The dentary was very long and shallow, with a prominent
Meckelian groove The Meckelian groove (or Meckel's groove, Meckelian fossa, or Meckelian foramen, or Meckelian canal) is an opening in the medial (inner) surface of the mandible (lower jaw) which exposes the Meckelian cartilage. Most of the teeth found with the holotype specimen were not in articulation with the skull; a few remained in the upper jaw, and only small replacement teeth were still borne by the lower jaw. The teeth had the shape of recurved cones, where slightly flattened from sideways, and their curvature was almost uniform. The roots were very long, and tapered towards their extremity. The carinae (sharp front and back edges) of the teeth were finely serrated with denticles on the front and back, and extended all along the crown. There were around six to eight denticles per mm (0.039 in), a much larger number than in large-bodied theropods like '' Torvosaurus'' and '' Tyrannosaurus''. Some of the teeth were fluted, with six to eight ridges along the length of their inner sides and fine-grained (outermost layer of teeth), while others bore no flutes; their presence is probably related to position or ontogeny (development during growth). The inner side of each tooth row had a bony wall. The number of teeth was large compared to most other theropods, with six to seven teeth in each premaxilla and thirty-two in each dentary. Based on the closer packing and smaller size of the dentary teeth compared to those in the corresponding length of the premaxilla, the difference between the number of teeth in the upper and lower jaws appears to have been more pronounced than in other theropods. The terminal rosette in the upper jaw of the holotype had thirteen (tooth sockets), six on the left and seven on the right side, showing tooth count asymmetry. The first four upper teeth were large (with the second and third the largest), while the fourth and fifth progressively decreased in size. The diameter of the largest was twice that of the smallest. The first four alveoli of the dentary (corresponding to the tip of the upper jaw) were the largest, with the rest more regular in size. Small subtriangular were present between the alveoli.


Postcranial skeleton

Initially thought to have lacked the sigmoid curve typical of theropods, the neck of ''Baryonyx'' does appear to have formed an S shape, though straighter than in other theropods. The cervical vertebrae of the neck tapered towards the head and became progressively longer front to back. Their (the processes that connected the vertebrae) were flat, and their (processes to which neck muscles attached) were well developed. The (the second neck vertebra) was small relative to the size of the skull and had a well-developed . The of the cervical vertebrae were not always sutured to the (the bodies of the vertebrae), and the there were low and thin. The were short, similar to those of crocodiles, and possibly overlapped each other somewhat. The centra of the dorsal vertebrae of the back were similar in size. Like in other theropods, the skeleton of ''Baryonyx'' showed
skeletal pneumaticity 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 ...
, reducing its weight through (openings) in the neural arches and (hollow depressions) in the centra (primarily near the ). From front to back, the neural spines of the dorsal vertebrae changed from short and stout to tall and broad. One isolated dorsal neural spine was moderately elongated and slender, indicating that ''Baryonyx'' may have had a hump or ridge along the centre of its back (though incipiently developed compared to those of other spinosaurids). ''Baryonyx'' was unique among spinosaurids in having a marked constriction from side to side in a vertebra that either belonged to the or front of the tail. The coracoid tapered hind-wards when viewed in profile, and, uniquely among spinosaurids, connected with the scapula in a peg-and-notch articulation. The scapulae were robust and the bones of the forelimb were short in relation to the animal's size, but broad and sturdy. The humerus was short and stout, with its ends broadly expanded and flattened—the upper side for the and muscle attachment and the lower for articulation with the radius and ulna. The radius was short, stout and straight, and less than half the length of the humerus, while the ulna was a little longer. The ulna had a powerful and an expanded lower end. The hands had three fingers; the first finger bore a large claw measuring about along its curve in the holotype specimen. The claw would have been lengthened by a keratin (horny) sheath in life. Apart from its size, the claw's proportions were fairly typical of a theropod, i.e. it was
bilaterally symmetric Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pla ...
, slightly compressed, smoothly rounded, and sharply pointed. A groove for the sheath ran along the length of the claw. The other claws of the hand were much smaller. The (main hip bone) of the pelvis had a prominent , an anterior process that was slender and vertically expanded, and a posterior process that was long and straight. The ilium also had a prominent and a deep grove that faced downwards. The (the socket for the femur) was long from front to back. The (lower and rearmost hip bone) had a well developed at the upper part. The margin of the blade at the lower end was turned outward, and the pubic foot was not expanded. The femur lacked a groove on the fibular condyle, and, uniquely among spinosaurids, the fibula had a very shallow fibular (depression).


Classification

In their original description, Charig and Milner found ''Baryonyx'' unique enough to warrant a new family of theropod dinosaurs: Baryonychidae. They found ''Baryonyx'' to be unlike any other theropod group, and considered the possibility that it was a thecodont (a grouping of early archosaurs now considered unnatural), due to having apparently primitive features, but noted that the articulation of the maxilla and premaxilla was similar to that in ''Dilophosaurus''. They also noted that the two snouts from Niger (which later became the basis of ''Cristatusaurus''), assigned to the family Spinosauridae by Taquet in 1984, appeared almost identical to that of ''Baryonyx'' and they referred them to Baryonychidae instead. In 1988, the palaeontologist Gregory S. Paul agreed with Taquet that ''Spinosaurus'', described in 1915 based on fragmentary remains from Egypt that were destroyed in World War II, and ''Baryonyx'' were similar and (due to their kinked snouts) possibly late-surviving dilophosaurs. Buffetaut also supported this relationship in 1989. In 1990, Charig and Milner dismissed the spinosaurid affinities of ''Baryonyx'', since they did not find their remains similar enough. In 1997, they agreed that Baryonychidae and Spinosauridae were related, but disagreed that the former name should become a synonym of the latter, because the completeness of ''Baryonyx'' compared to ''Spinosaurus'' made it a better type genus for a family, and because they did not find the similarities between the two significant enough. Holtz and colleagues listed Baryonychidae as a synonym of Spinosauridae in 2004. Discoveries in the 1990s shed more light on the relationships of ''Baryonyx'' and its relatives. In 1996, a snout from Morocco was referred to ''Spinosaurus'', and '' Irritator'' and ''Angaturama'' from Brazil (the two are possible synonyms) were named. ''Cristatusaurus'' and ''Suchomimus'' were named based on fossils from Niger in 1998. In their description of ''Suchomimus'', Sereno and colleagues placed it and ''Baryonyx'' in the new
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 ...
Baryonychinae Baryonychinae is an extinct clade or subfamily of spinosaurids from the Early Cretaceous (Valanginian-Albian) of Britain, Portugal, and Niger. In 2021, it consisted of six genera: ''Ceratosuchops'', ''Cristatusaurus'', ''Riparovenator'', ''Suchomi ...
within Spinosauridae; ''Spinosaurus'' and ''Irritator'' were placed in the subfamily
Spinosaurinae The Spinosauridae (or spinosaurids) are a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. They came into prominence during the Cretaceous period. Spinosaurid fossils have been recovered worldwide, includi ...
. Baryonychinae was distinguished by the small size and larger number of teeth in the dentary behind the terminal rosette, the deeply keeled front dorsal vertebrae, and by having serrated teeth. Spinosaurinae was distinguished by their straight tooth crowns without serrations, small first tooth in the premaxilla, increased spacing of teeth in the jaws, and possibly by having their nostrils placed further back and the presence of a deep
neural spine sail A neural spine sail is a large, flattish protrusion from the back of an animal formed of a sequence of extended vertebral spinous processes and associated tissues. Such structures are comparatively rare in modern animals, but have been identified ...
. They also united the spinosaurids and their closest relatives in the superfamily Spinosauroidea, but in 2010, the palaeontologist Roger Benson considered this a junior synonym of Megalosauroidea (an older name). In a 2007 conference abstract, the palaeontologist Denver W. Fowler suggested that since ''Suchosaurus'' is the first named genus in its group, the clade names Spinosauroidea, Spinosauridae, and Baryonychinae should be replaced by Suchosauroidea, Suchosauridae, and Suchosaurinae, regardless of whether or not the name ''Baryonyx'' is retained. A 2017 study by the palaeontologists Marcos A. F. Sales and Cesar L. Schultz found that the clade Baryonychinae was not well supported, since serrated teeth may be an
ancestral trait In phylogenetics, a primitive (or ancestral) character, trait, or feature of a lineage or taxon is one that is inherited from the common ancestor of a clade (or clade group) and has undergone little change since. Conversely, a trait that appears ...
among spinosaurids. Barker and colleagues found support for a Baryonychinae-Spinosaurinae split in 2021, and the following cladogram shows the position of ''Baryonyx'' within Spinosauridae according to their study:


Evolution

Spinosaurids appear to have been widespread from the Barremian to the
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
stages Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly British theatre newspaper * S ...
of the Cretaceous period, about 130 to 95 million years ago, while the oldest known spinosaurid remains date to 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 ...
. They shared features such as long, narrow, crocodile-like skulls; sub-circular teeth, with fine to no serrations; the terminal rosette of the snout; and a secondary palate that made them more resistant to torsion. In contrast, the primitive and typical condition for theropods was a tall, narrow snout with blade-like (ziphodont) teeth with serrated carinae. The skull adaptations of spinosaurids converged with those of
crocodilian 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 ...
s; early members of the latter group had skulls similar to typical theropods, later developing elongated snouts, conical teeth, and secondary palates. These adaptations may have been the result of a dietary change from terrestrial prey to fish. Unlike crocodiles, the post-cranial skeletons of baryonychine spinosaurids do not appear to have aquatic adaptations.Supplementary Information
/ref> Sereno and colleagues proposed in 1998 that the large thumb-claw and robust forelimbs of spinosaurids evolved in the Middle Jurassic, before the elongation of the skull and other adaptations related to fish-eating, since the former features are shared with their
megalosaurid Megalosauridae is a monophyletic family of carnivorous theropod dinosaurs within the group Megalosauroidea. Appearing in the Middle Jurassic, megalosaurids were among the first major radiation of large theropod dinosaurs. They were a relative ...
relatives. They also suggested that the spinosaurines and baryonychines diverged before the Barremian age of the Early Cretaceous. Several theories have been proposed about the
biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
of the spinosaurids. Since ''Suchomimus'' was more closely related to ''Baryonyx'' (from Europe) than to ''Spinosaurus''—although that genus also lived in Africa—the distribution of spinosaurids cannot be explained as
vicariance Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...
resulting from
continental rifting In geology, a rift is a linear zone where the lithosphere is being pulled apart and is an example of extensional tectonics. Typical rift features are a central linear downfaulted depression, called a graben, or more commonly a half-graben wi ...
. Sereno and colleagues proposed that spinosaurids were initially distributed across the supercontinent Pangea, but split with the opening of the Tethys Sea. Spinosaurines would then have evolved in the south (Africa and South America: in
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
) and baryonychines in the north (Europe: in
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pan ...
), with ''Suchomimus'' the result of a single north-to-south
dispersal event Biological dispersal refers to both the movement of individuals ( animals, plants, fungi, bacteria, etc.) from their birth site to their breeding site ('natal dispersal'), as well as the movement from one breeding site to another ('breeding ...
. Buffetaut and the Tunisian palaeontologist Mohamed Ouaja also suggested in 2002 that baryonychines could be the ancestors of spinosaurines, which appear to have replaced the former in Africa. Milner suggested in 2003 that spinosaurids originated in Laurasia during the Jurassic, and dispersed via the Iberian land bridge into Gondwana, where they radiated. In 2007, Buffetaut pointed out that
palaeogeographical Palaeogeography (or paleogeography) is the study of historical geography, generally physical landscapes. Palaeogeography can also include the study of human or cultural environments. When the focus is specifically on landforms, the term paleo ...
studies had demonstrated that Iberia was near northern Africa during the Early Cretaceous, which he found to confirm Milner's idea that the Iberian region was a
stepping stone Stepping stone(s) may refer to: * Stepping stones, stones placed to allow pedestrians to cross a watercourse Places * Stepping Stone, Virginia, US, an unincorporated community * Stepping Stones (islands), Antarctic and sub-Antarctic Buildings ...
between Europe and Africa, which is supported by the presence of baryonychines in Iberia. The direction of the dispersal between Europe and Africa is still unknown, and subsequent discoveries of spinosaurid remains in Asia and possibly Australia indicate that it may have been complex. Candeiro and colleagues suggested in 2017 that spinosaurids of northern Gondwana were replaced by other predators, such as
abelisauroids Abelisauroidea is typically regarded as a Cretaceous group, though the earliest abelisauridae remains are known from the Middle Jurassic of Argentina (classified as the species Eoabelisaurus mefi) and possibly Madagascar (fragmentary remains o ...
, since no definite spinosaurid fossils are known from after the Cenomanian anywhere in the world. They attributed the disappearance of spinosaurids and other shifts in the fauna of Gondwana to changes in the environment, perhaps caused by transgressions in sea level. Malafaia and colleagues stated in 2020 that ''Baryonyx'' remains the oldest unquestionable spinosaurid, while acknowledging that older remains had also been tentatively assigned to the group. Barker and colleagues found support for a European origin for spinosaurids in 2021, with an expansion to Asia and Gondwana during the first half of the Early Cretaceous. In contrast to Sereno, these authors suggested there had been at least two dispersal events from Europe to Africa, leading to ''Suchomimus'' and the African part of Spinosaurinae.


Palaeobiology


Diet and feeding

In 1986, Charig and Milner suggested that its elongated snout with many finely serrated teeth indicated that ''Baryonyx'' was piscivorous (fish-eating), speculating that it crouched on a riverbank and used its claw to gaff fish out of the water (similar to the modern
grizzly bear The grizzly bear (''Ursus arctos horribilis''), also known as the North American brown bear or simply grizzly, is a population or subspecies of the brown bear inhabiting North America. In addition to the mainland grizzly (''Ursus arctos horri ...
). Two years earlier, Taquet pointed out that the spinosaurid snouts from Niger were similar to those of the modern gharial and suggested a behaviour similar to herons or storks. In 1987, the biologist Andrew Kitchener disputed the piscivorous behaviour of ''Baryonyx'' and suggested that it would have been a
scavenger Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding b ...
, using its long neck to feed on the ground, its claws to break into a carcass, and its long snout (with nostrils far back for breathing) for investigating the body cavity. Kitchener argued that ''Baryonyx'' jaws and teeth were too weak to kill other dinosaurs and too heavy to catch fish, with too many adaptations for piscivory. According to the palaeontologist Robin E. H. Reid, a scavenged carcass would have been broken up by its predator and large animals capable of doing so—such as grizzly bears—are also capable of catching fish (at least in shallow water). In 1997, Charig and Milner demonstrated direct dietary evidence in the stomach region of the ''B. walkeri'' holotype. It contained the first evidence of piscivory in a theropod dinosaur, acid-etched scales and teeth of the common fish '' Scheenstia mantelli'' (then classified in the genus '' Lepidotes''), and abraded or etched bones of a young
iguanodontid Iguanodontidae is a family of iguanodontians belonging to Styracosterna, a derived clade within Ankylopollexia. Characterized by their elongated maxillae, they were herbivorous and typically large in size. This family exhibited locomotive dynam ...
. They also presented circumstantial evidence for piscivory, such as crocodile-like adaptations for catching and swallowing prey: long, narrow jaws with their "terminal rosette", similar to those of a gharial, and the downturned tip and notch of the snout. In their view, these adaptations suggested that ''Baryonyx'' would have caught small to medium-sized fish in the manner of a crocodilian: gripping them with the notch of the snout (giving the teeth a "stabbing function"), tilting the head backwards, and swallowing them headfirst. Larger fish would be broken up with the claws. That the teeth in the lower jaw were smaller, more crowded and numerous than those in the upper jaw may have helped the animal grip food. Charig and Milner maintained that ''Baryonyx'' would primarily have eaten fish (although it would also have been an active predator and opportunistic scavenger), but it was not equipped to be a
macro-predator An apex predator, also known as a top predator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the highest trophic lev ...
like ''Allosaurus''. They suggested that ''Baryonyx'' mainly used its forelimbs and large claws to catch, kill and tear apart larger prey. An apparent gastrolith ( gizzard stone) was also found with the specimen. The German palaeontologist Oliver Wings suggested in 2007 that the low number of stones found in theropods like ''Baryonyx'' and ''Allosaurus'' could have been ingested by accident. In 2004, a pterosaur neck vertebra from Brazil with a spinosaurid tooth embedded in it reported by Buffetaut and colleagues confirmed that the latter were not exclusively piscivorous. A 2005 beam-theory study by the palaeontologist François Therrien and colleagues was unable to reconstruct force profiles of ''Baryonyx'', but found that the related ''Suchomimus'' would have used the front part of its jaws to capture prey, and suggested that the jaws of spinosaurids were adapted for hunting smaller terrestrial prey in addition to fish. They envisaged that spinosaurids could have captured smaller prey with the rosette of teeth at the front of the jaws, and finished it by shaking it. Larger prey would instead have been captured and killed with their forelimbs instead of their bite, since their skulls would not be able to resist the bending stress. They also agreed that the conical teeth of spinosaurids were well-developed for impaling and holding prey, with their shape enabling them to withstand bending loads from all directions. A 2007
finite element analysis The finite element method (FEM) is a popular method for numerically solving differential equations arising in engineering and mathematical modeling. Typical problem areas of interest include the traditional fields of structural analysis, heat ...
of
CT scanned A computed tomography scan (CT scan; formerly called computed axial tomography scan or CAT scan) is a medical imaging technique used to obtain detailed internal images of the body. The personnel that perform CT scans are called radiographers ...
snouts by the palaeontologist Emily J. Rayfield and colleagues indicated that the biomechanics of ''Baryonyx'' were most similar to those of the gharial and unlike those of the American alligator and more-conventional theropods, supporting a piscivorous diet for spinosaurids. Their secondary palate helped them resist bending and torsion of their tubular snouts. A 2013 beam-theory study by the palaeontologists Andrew R. Cuff and Rayfield compared the biomechanics of CT-scanned spinosaurid snouts with those of extant crocodilians, and found the snouts of ''Baryonyx'' and ''Spinosaurus'' similar in their resistance to bending and torsion. ''Baryonyx'' was found to have relatively high resistance in the snout to dorsoventral bending compared with ''Spinosaurus'' and the gharial. The authors concluded (in contrast to the 2007 study) that ''Baryonyx'' performed differently than the gharial; spinosaurids were not exclusive piscivores, and their diet was determined by their individual size. In a 2014 conference abstract, the palaeontologist Danny Anduza and Fowler pointed out that grizzly bears do not gaff fish out of the water as was suggested for ''Baryonyx'', and also ruled out that the dinosaur would not have darted its head like herons, since the necks of spinosaurids were not strongly S-curved, and their eyes were not well-positioned for binocular vision. Instead, they suggested the jaws would have made sideways sweeps to catch fish, like the gharial, with the hand claws probably used to stamp down and impale large fish, whereafter they manipulated them with their jaws, in a manner similar to grizzly bears and fishing cats. They did not find the teeth of spinosaurids suitable for dismembering prey, due to their lack of serrations, and suggested they would have swallowed prey whole (while noting they could also have used their claws for dismemberment). A 2016 study by the palaeontologist Christophe Hendrickx and colleagues found that adult spinosaurs could displace their mandibular rami (halves of the lower jaw) sideways when the jaw was depressed, which allowed the pharynx (opening that connects the mouth to the oesophagus) to be widened. This jaw-articulation is similar to that seen in pterosaurs and living pelicans, and would likewise have allowed spinosaurids to swallow large prey such as fish and other animals. They also reported that some possible Portuguese ''Baryonyx'' fossils were found associated with isolated '' Iguanodon'' teeth, and listed it along with other such associations as support for opportunistic feeding behaviour in spinosaurs. Another 2016 study by the palaeontologist Romain Vullo and colleagues found that the jaws of spinosaurids were convergent with those of
pike conger eel The Muraenesocidae, or pike congers, are a small family of marine eels found worldwide in tropical and subtropical seas. Some species are known to enter brackish water. Pike congers have cylindrical bodies, scaleless skin, narrow heads with large ...
s; these fish also have jaws that are compressed side to side (whereas the jaws of crocodilians are compressed from top to bottom), an elongated snout with a "terminal rosette" that bears enlarged teeth, and a notch behind the rosette with smaller teeth. Such jaws likely evolved for grabbing prey in aquatic environments with low light, and may have helped in prey detection. A 2023 study by Barker and colleagues based on CT scans of the braincases of ''Baryonyx'' and ''Ceratosuchops'' found that the brain anatomy of these baryonychines was similar to that of other non-
maniraptoriform Maniraptoriformes is a clade of dinosaurs with pennaceous feathers and wings that contains ornithomimosaurs and maniraptorans. This group was named by Thomas Holtz, who defined it as "the most recent common ancestor of ''Ornithomimus'' and bir ...
theropods. Their neurosensory capabilities such as hearing and
olfaction The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
(sense of smell) were unexceptional, and their gaze stabilisation less developed than those of spinosaurines, so their behavioural adaptations were probably comparable to those of other large-bodied terrestrial theropods. This suggests that their transition from terrestrial hypercarnivores to
semi-aquatic In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below. Semiaquatic animals Semiaquatic animals include: * Verte ...
“generalists” during their evolution did not require substantial modification of their brain and sensory systems. This could mean that spinosaurids were either pre-adapted for detection and capture of aquatic prey, or that their transition to semi-aquatic lifestyles only required modifications to the bones associated with the mouth. Their reptile encephalization quotient values imply that the cognitive capacity and behavioural sophistication of baryonychines did not deviate much from that of other
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...
theropods.


Motion and semi-aquatic habits

In their original description, Charig and Milner did not consider ''Baryonyx'' to be aquatic (due to its nostrils being on the sides of its snout—far from the tip—and the form of the post-cranial skeleton), but thought it was capable of swimming, like most land vertebrates. They speculated that the elongated skull, long neck, and strong humerus of ''Baryonyx'' indicated that the animal was a facultative quadruped, unique among theropods. In their 1997 article they found no skeletal support for this, but maintained that the forelimbs would have been strong enough for a quadrupedal posture and it would probably have caught aquatic prey while crouching—or on all fours—near (or in) water. A 2014 re-description of ''Spinosaurus'' by the palaeontologist
Nizar Ibrahim Nizar Ibrahim (born in 1982) is a German-Moroccan vertebrate paleontologist and comparative anatomist. He is currently a senior lecturer at the University of Portsmouth. Ibrahim has led several expeditions to Africa's Sahara and is notable for h ...
and colleagues based on new remains suggested that it was a quadruped, based on its anterior
centre of body mass In physics, the center of mass of a distribution of mass in space (sometimes referred to as the balance point) is the unique point where the weighted relative position of the distributed mass sums to zero. This is the point to which a force may ...
. The authors found quadrupedality unlikely for ''Baryonyx'', since the better-known legs of the closely related ''Suchomimus'' did not support this posture. In 2017, the palaeontologists David E. Hone and Holtz hypothesized that the head crests of spinosaurids were probably used for sexual or threat display. The authors also pointed out that (like other theropods) there was no reason to believe that the forelimbs of ''Baryonyx'' were able to pronate (crossing the radius and ulna bones of the lower arm to turn the hand), and thereby make it able to rest or walk on its palms. Resting on or using the forelimbs for locomotion may have been possible (as indicated by tracks of a resting theropod), but if this was the norm, the forelimbs would probably have showed adaptations for this. Hone and Holtz furthermore suggested that the forelimbs of spinosaurids do not seem optimal for trapping prey, but instead appear similar to the forelimbs of digging animals. They suggested that the ability to dig would have been useful when excavating nests, digging for water, or to reach some kinds of prey. Hone and Holtz also believed that spinosaurids would have waded and dipped in water rather than submerging themselves, due to their sparsity of aquatic adaptations. A 2010 study by the palaeontologist Romain Amiot and colleagues proposed that spinosaurids were semi-aquatic, based on the
oxygen isotope There are three known stable isotopes of oxygen (8O): , , and . Radioactive isotopes ranging from to have also been characterized, all short-lived. The longest-lived radioisotope is with a half-life of , while the shortest-lived isotope is ...
composition of spinosaurid teeth from around the world compared with that of other theropods and extant animals. Spinosaurids probably spent much of the day in water, like crocodiles and hippopotamuses, and had a diet similar to the former; both were opportunistic predators. Since most spinosaurids do not appear to have anatomical adaptations for an aquatic lifestyle, the authors proposed that submersion in water was a means of thermoregulation similar to that of crocodiles and hippopotamuses. Spinosaurids may also have turned to aquatic habitats and piscivory to avoid competition with large, more-terrestrial theropods. In 2016, Sales and colleagues statistically examined the fossil distribution of spinosaurids, abelisaurids, and
carcharodontosaurids Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
, and concluded that spinosaurids had the strongest support for association with coastal palaeoenvironments. Spinosaurids also appear to have inhabited inland environments (with their distribution there being comparable to carcharodontosaurids), which indicates they may have been more generalist than usually thought. Sales and Schultz agreed in 2017 that spinosaurids were semi-aquatic and partially piscivorous, based on skull features such as conical teeth, snouts that were compressed from side to side, and retracted nostrils. They interpreted the fact that histological data indicates some spinosaurids were more terrestrial than others as reflecting ecological niche partitioning among them. As some spinosaurids have smaller nostrils than others, their
olfactory The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
abilities were presumably lesser, as in modern piscivorous animals, and they may instead have used other senses (such as vision and
mechanoreception A mechanoreceptor, also called mechanoceptor, is a sensory receptor that responds to mechanical pressure or distortion. Mechanoreceptors are innervated by sensory neurons that convert mechanical pressure into electrical signals that, in animals, a ...
) when hunting fish. Olfaction may have been more useful for spinosaurids that also fed on terrestrial prey, such as baryonychines. A 2022 study by the palaeontologist Matteo Fabbri and colleagues revealed that ''Baryonyx'' possessed dense bones that would have allowed it to dive underwater. This same adaptation was revealed in the related ''Spinosaurus'', and they are believed to have been subaqueous foragers that dived after aquatic prey, while ''Suchomimus'' was better adapted to a non-diving lifestyle by comparison according to the provided analysis. This discovery also showcases the unique and ecologically disparate lifestyles spinosaurids had, with more hollow-boned genera preferring to hunt in shallower water.


Palaeoenvironment

The Weald Clay Formation consists of sediments of
Hauterivian The Hauterivian is, in the geologic timescale, an age in the Early Cretaceous Epoch or a stage in the Lower Cretaceous Series. It spans the time between 132.9 ± 2 Ma and 129.4 ± 1.5 Ma (million years ago). The Hauterivian is preceded by the Va ...
(Lower Weald Clay) to Barremian (Upper Weald Clay)
age Age or AGE may refer to: Time and its effects * Age, the amount of time someone or something has been alive or has existed ** East Asian age reckoning, an Asian system of marking age starting at 1 * Ageing or aging, the process of becoming older ...
, about 130–125 million years old. The original ''Baryonyx'' specimen was found in the latter, in clay representing non-marine still water, which has been interpreted as a
fluvial In geography and geology, fluvial processes are associated with rivers and streams and the deposits and landforms created by them. When the stream or rivers are associated with glaciers, ice sheets, or ice caps, the term glaciofluvial or fluviog ...
or
mudflat Mudflats or mud flats, also known as tidal flats or, in Ireland, slob or slobs, are coastal wetlands that form in intertidal areas where sediments have been deposited by tides or rivers. A global analysis published in 2019 suggested that tidal fl ...
environment with shallow water, lagoons, and marshes. During the Early Cretaceous, the Weald area of Surrey, Sussex, and Kent was partly covered by the large, fresh-to- brackish water
Wealden Lake Wealden Lake was a shallow freshwater to brackish lake which existed over the lands of what is now northern France and southern England during the Berriasian to Barremian ages of the Early Cretaceous epoch (approximately 145 to 125 mya). The sedime ...
. Two large rivers drained the northern area (where London now stands), flowing into the lake through a river delta; the Anglo-Paris Basin was in the south. Its climate was sub-tropical, similar to the present Mediterranean region. Since the Smokejack Clay Pit consists of different stratigraphic levels, fossil taxa found there are not necessarily contemporaneous. Dinosaurs from the locality include the ornithopods '' Mantellisaurus'', ''Iguanodon'', and small sauropods. Other vertebrates from the Weald Clay include crocodiles, pterosaurs, lizards (such as ''
Dorsetisaurus ''Dorsetisaurus'' is a genus of extinct lizard, known from the Late Jurassic of North America, and the Late Jurassic-earliest Cretaceous of Europe. The genus was first reported from the Early Cretaceous (Berriasian) Lulworth Formation of the Pur ...
''), amphibians, sharks (such as ''
Hybodus ''Hybodus'' (from el, ύβος , 'crooked' and el, ὀδούς 'tooth') is an extinct genus of hybodont, a group of shark-like elasmobranchs that lived from the Late Devonian to the end of the Cretaceous. Species closely related to the type sp ...
''), and bony fishes (including ''Scheenstia''). Members of ten orders of insects have been identified, including '' Valditermes'', '' Archisphex'', and '' Pterinoblattina''. Other invertebrates include
ostracods 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, typical ...
, isopods,
conchostracans Clam shrimp are a group of bivalved branchiopod crustaceans that resemble the unrelated Bivalvia, bivalved molluscs. They are extant and also known from the fossil record, from at least the Devonian period and perhaps before. They were originally ...
, and
bivalves 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 ...
. The plants ''
Weichselia ''Weichselia'' is an extinct genus of fern. They were abundant from the Middle Jurassic to Early Cretaceous.herbaceous Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition of t ...
'' Bevhalstia'' were common. Other plants found include ferns, horsetails,
club mosses Lycopodiopsida is a class of vascular plants known as lycopods, lycophytes or other terms including the component lyco-. Members of the class are also called clubmosses, firmosses, spikemosses and quillworts. They have dichotomously branching s ...
, and
conifers Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All extan ...
. Other dinosaurs from the Wessex Formation of the Isle of Wight where ''Baryonyx'' may have occurred include the theropods ''Riparovenator'', ''Ceratosuchops'', ''
Neovenator ''Neovenator'' (nee-o-ven-a-tor meaning "new hunter") is a genus of carcharodontosaurian theropod dinosaur. It is known from several skeletons found in the Early Cretaceous (Barremian~130-125 million years ago) Wessex Formation on the south coa ...
'', '' Eotyrannus'', '' Aristosuchus'', ''
Thecocoelurus ''Thecocoelurus'' is a nomen dubium, dubious genus of theropod dinosaur from the early Cretaceous period of England. The phylogenetic placement of this genus is uncertain, and it has been referred to an oviraptosaur, an ornithomimosaur, or a ther ...
'', ''
Calamospondylus ''Calamospondylus'' (meaning "quill vertebrae") is a genus of theropod dinosaur. It lived during the Early Cretaceous and its fossils were found on the Isle of Wight in southern England. The type species is ''C. oweni''. The holotype was collec ...
'', and ''
Ornithodesmus ''Ornithodesmus'' (meaning "bird link") is a genus of small, dromaeosaurid dinosaur from the Isle of Wight in England, dating to about 125 million years ago. The name was originally assigned to a bird-like sacrum (a series of vertebrae fused to ...
''; the ornithopods ''Iguanodon'', '' Hypsilophodon'', and ''
Valdosaurus ''Valdosaurus'' ("Weald Lizard") is a genus of bipedal herbivorous iguanodont ornithopod dinosaur found on the Isle of Wight and elsewhere in England, Spain and possibly also Romania. It lived during the Early Cretaceous. Discovery and naming I ...
''; the sauropods '' Ornithopsis'', '' Eucamerotus'', and ''
Chondrosteosaurus ''Chondrosteosaurus'' (meaning "cartilage and bone lizard") was a sauropod dinosaur from Early Cretaceous England. The type species, ''Chondrosteosaurus gigas'', was described and named by Richard Owen in 1876. The fossils of ''Chondrosteosau ...
''; and 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. ...
'' Polacanthus''. Barker and colleagues stated in 2021 that the identification of the two additional spinosaurids from the Wealden Supergroup, ''Riparovenator'' and ''Ceratosuchops'', has implications for potential ecological separation within Spinosauridae if these and ''Baryonyx'' were contemporary and interacted. They cautioned that it is possible the Upper Weald Clay and Wessex Formations and the spinosaurids known from them were separated in time and distance. It is generally thought that large predators occur with small taxonomic diversity in any area due to ecological demands, yet many Mesozoic assemblages include two or more
sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species sh ...
theropods that were comparable in size and morphology, and this also appears to have been the case for spinosaurids. Barker and colleagues suggested that high diversity within Spinosauridae in a given area may have been the result of environmental circumstances benefiting their niche. While it has been generally assumed that only identifiable anatomical traits related to resource partitioning allowed for coexistence of large theropods, Barker and colleagues noted that this does not preclude that similar and closely related taxa could coexist and overlap in ecological requirements. Possible niche partitioning could be in time (seasonal or daily), in space (between habitats in the same ecosystems), or depending on conditions, and they could also have been separated by their choice of habitat within their regions (which may have ranged in climate).


Taphonomy

Charig and Milner presented a possible scenario explaining the taphonomy (changes during decay and fossilisation) of the ''B. walkeri'' holotype specimen. The fine-grained sediments around the skeleton, and the fact that the bones were found close together (skull and forelimb elements at one end of the excavation area and the pelvis and hind-limb elements at the other), indicates that the environment was quiet at the time of deposition, and water currents did not carry the carcass far—possibly because the water was shallow. The area where the specimen died seems to have been suitable for a piscivorous animal. It may have caught fish and scavenged on the mud plain, becoming mired before it died and was buried. Since the bones are well-preserved and had no gnaw marks, the carcass appears to have been undisturbed by scavengers (suggesting that it was quickly covered by sediment). The disarticulation of the bones may have been the result of soft-tissue decomposition. Parts of the skeleton seem to have weathered to different degrees, perhaps because water levels changed or the sediments shifted (exposing parts of the skeleton). The girdle and limb bones, the dentary, and a rib were broken before fossilisation, perhaps from trampling by large animals while buried. Most of the tail appears to have been lost before fossilisation, perhaps due to scavenging, or having rotted and floated off. The orientation of the bones indicates that the carcass lay on its back (perhaps tilted slightly to the left, with the right side upwards), which may explain why all the lower teeth had fallen out of their sockets and some upper teeth were still in place.


References


External links


Natural History Museum – "''Baryonyx'': the discovery of an amazing fish-eating dinosaur"
– four minute video presented by Angela C. Milner {{Academic peer reviewed, Q63252951 Spinosaurids Monotypic dinosaur genera Barremian life Early Cretaceous dinosaurs of Europe Cretaceous England Fossils of England Fossil taxa described in 1986 Taxa named by Alan J. Charig Taxa named by Angela Milner Articles containing video clips