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''Saltriovenator'' (meaning "
Saltrio Saltrio is a ''comune'' (municipality) in the Province of Varese in the Italian region Lombardy, located about northwest of Milan and about northeast of Varese, on the border with Switzerland. As of 31 December 2004, it had a population of 2,92 ...
hunter") is a genus of
ceratosaurian Ceratosaurs are members of the clade Ceratosauria, a group of dinosaurs defined as all theropods sharing a more recent common ancestor with ''Ceratosaurus'' than with birds. The oldest known ceratosaur, ''Saltriovenator'', dates to the earliest ...
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 during the
Sinemurian In the geologic timescale, the Sinemurian is an age and stage in the Early or Lower Jurassic Epoch or Series. It spans the time between 199.3 ± 2 Ma and 190.8 ± 1.5 Ma (million years ago). The Sinemurian is preceded by the Hettangian and is ...
stage 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 * Sta ...
of the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
in what is now
Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical re ...
. The type and only species is ''Saltriovenator zanellai''; in the past, the species had been known under the informal name "saltriosaur". Although a full skeleton has not yet been discovered, ''Saltriovenator'' is thought to have been a large, bipedal carnivore similar to ''
Ceratosaurus ''Ceratosaurus'' (from Greek κέρας/κέρατος, ' meaning "horn" and σαῦρος ' meaning "lizard") was a carnivorous theropod dinosaur in the Late Jurassic period ( Kimmeridgian to Tithonian). The genus was first described in 1 ...
''.


Discovery and naming

On 4 August 1996, the first remains of ''Saltriovenator'' were discovered by amateur paleontologist Angelo Zanella, searching for
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 in the ''Salnova'' marble quarry in
Saltrio Saltrio is a ''comune'' (municipality) in the Province of Varese in the Italian region Lombardy, located about northwest of Milan and about northeast of Varese, on the border with Switzerland. As of 31 December 2004, it had a population of 2,92 ...
, northern Italy. Zanella had already been working for the ''
Museo Civico di Storia Naturale di Milano The Museo Civico di Storia Naturale di Milano (Milan Natural History Museum) is a museum in Milan, Italy. It was founded in 1838 when naturalist Giuseppe de Cristoforis donated his collections to the city. Its first director was Giorgio Jan. ...
'' and this institution after being informed sent out a team to investigate the find.
Cristiano Dal Sasso Cristiano Dal Sasso (born 12 September 1965) is an Italian paleontologist. Biography He was born in Monza, Italy and has been working since 1991 for the Milan Natural History Museum where he is the curator of fossil reptiles and birds. He was ...
and the volunteers of the Paleontological Group of Besano, under the direction of Giorgio Teruzzi managed to salvage a number of chalk blocks visibly containing bones. The skeleton had shortly before its discovery been blown to pieces by explosives used in the quarry to break the marble layers. Blocks that had been secured were inserted into a bath of
formic acid Formic acid (), systematically named methanoic acid, is the simplest carboxylic acid, and has the chemical formula HCOOH and structure . It is an important intermediate in chemical synthesis and occurs naturally, most notably in some ants. Es ...
for 1,800 hours to free the bones. Initially, 119 bone fragments were reported to have been collected in total; this was later increased to 132. However, most cannot be exactly identified. In 2000, the museum opened a special exhibition of the bones. On this occasion, Dal Sasso provisionally gave the dinosaur, now thought to be a species new to science, the Italian name ''Saltriosauro''. Although this has been occasionally Latinised to "Saltriosaurus", even in the scientific literature, in both the Italian and Latin form it remained an invalid '' nomen nudum''. In December 2018, Dal Sasso, Simone Maganuco and Andrea Cau named and described the specimen as 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 ...
''Saltriovenator zanellai''. The generic name combines a reference to Saltrio with
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
, ''venator'', "hunter", a common suffix in the names of theropods. The authors pointed out that a '' venator'' is also a type of Roman
gladiator A gladiator ( la, gladiator, "swordsman", from , "sword") was an armed combatant who entertained audiences in the Roman Republic and Roman Empire in violent confrontations with other gladiators, wild animals, and condemned criminals. Some gla ...
. The specific name honours Zanella. Because the article was published in an
electronic publication Electronic publishing (also referred to as publishing, digital publishing, or online publishing) includes the digital publication of e-books, digital magazines, and the development of digital libraries and catalogues. It also includes the editing ...
,
Life Science Identifiers Life Science Identifiers are a way to name and locate pieces of information on the web. Essentially, an LSID is a unique identifier for some data, and the LSID protocol specifies a standard way to locate the data (as well as a standard way of descr ...
were necessary to make the name valid. These are 8C9F3B56-F622-4C39-8E8B-C2E890811E74 for the genus and BDD366A7-6A9D-4A32-9841-F7273D8CA00B for the species. ''Saltriovenator'' is the third dinosaur named from Italy, the first from the Alps and the second theropod from Italy, after ''
Scipionyx ''Scipionyx'' ( ) was a genus of theropod dinosaur from the Early Cretaceous of Italy, around 113 million years ago. There is only one fossil known of ''Scipionyx'', discovered in 1981 by an amateur paleontologist and brought to the attention o ...
''. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
, MSNM V3664, was found in a layer of the
Saltrio Formation The Saltrio Formation is a geological formation in Italy. It dates back to the middle Sinemurian, and would have represented a pelagic or near-epicontinental environment, judging by the presence of marine fauna such as the nautiloid '' Cenoceras' ...
dating from the earliest early
Sinemurian In the geologic timescale, the Sinemurian is an age and stage in the Early or Lower Jurassic Epoch or Series. It spans the time between 199.3 ± 2 Ma and 190.8 ± 1.5 Ma (million years ago). The Sinemurian is preceded by the Hettangian and is ...
, 199 million years old. It consists of a fragmentary skeleton with a lower jaw. About 10% of the skeleton has been discovered, including a tooth, a right splenial, a right prearticular, a neck rib, fragments of the dorsal ribs and
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...
e, a well preserved but incomplete
furcula The (Latin for "little fork") or wishbone is a forked bone found in most birds and some species of non-avian dinosaurs, and is formed by the fusion of the two pink clavicles. In birds, its primary function is in the strengthening of the thoracic ...
,
humeri The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a round ...
,
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 ...
II,
phalanx The phalanx ( grc, φάλαγξ; plural phalanxes or phalanges, , ) was a rectangular mass military formation, usually composed entirely of heavy infantry armed with spears, pikes, sarissas, or similar pole weapons. The term is particularly use ...
II-1, phalanx III-1, phalanx III-2, manual
ungual An ungual (from Latin ''unguis'', i.e. ''nail'') is a highly modified distal toe bone which ends in a hoof, claw, or nail. Elephants and ungulates have ungual phalanges, as did the sauropod Sauropoda (), whose members are known as sauropods (; ...
III, a distal tarsal III, a distal tarsal IV and the
proximal 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 pro ...
second to fifth metatarsals. The holotype individual likely died on the shores of an ancient beach before being washed out to sea. After death, the skeletal remains suffered from prolonged transport, during which many bones were lost and the remaining ones highly fragmented. Although ''Saltriovenator'' was not aquatic, the environment in which the carcass was deposited was likely
pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...
, judging by the associated
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. The locality is also rich in
crinoid Crinoids are marine animals that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their adult form are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, which are ...
s,
gastropods 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. Ther ...
,
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,
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 and
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. Deposition occurred on a slope between a shallow carbonate platform and a deeper basin. Various scratches, grooves, and striations indicate that the carcass was subject to scavenging by marine invertebrates. The specimen represents a subadult individual, nearing its maximum size, of which the age has been estimated at twenty-four years.


Description

Because of the fragmentary nature of the remains, it was impossible to directly measure the size of the animal. The describing authors therefore compared the fossils with those of two theropods of a roughly similar volume. Comparing with the skeletal elements of MOR 693, an ''
Allosaurus fragilis ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch ( Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alludi ...
'' specimen, they conservatively concluded that the ''Saltriovenator'' holotype individual was at least seven to eight metres long. This would make ''Saltriovenator'' the largest known theropod living before the
Aalenian The Aalenian () is a subdivision of the Middle Jurassic Epoch/Series of the geologic timescale that extends from about 174.1 Ma to about 170.3 Ma (million years ago). It was preceded by the Toarcian and succeeded by the Bajocian. Stratigraphi ...
stage, 25% longer than ''
Ceratosaurus ''Ceratosaurus'' (from Greek κέρας/κέρατος, ' meaning "horn" and σαῦρος ' meaning "lizard") was a carnivorous theropod dinosaur in the Late Jurassic period ( Kimmeridgian to Tithonian). The genus was first described in 1 ...
'' from the late Jurassic. Comparing with ''Ceratosaurus'' itself, resulted in a body length of 730 centimetres, a hip height of 220 centimetres and a skull length of eighty centimetres. The thighbone length would then have been about eighty to eighty-seven centimetres, which indicates a body weight of 1160 to 1524 kilogrammes. Another method consisted in extrapolating from the known length of the forelimb. Applying the usual limb ratio indicated a hindlimb length of 198 centimetres. The thighbone would then have been 822 to 887 millimetres long, indicating a weight of 1269 to 1622 kilogrammes.


Classification

The precise systematic position of ''Saltriovenator'' has been traditionally uncertain, but it is known to be a
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 ...
.The Theropod DatabaseMatthew T. Carrano, Roger B. J. Benson, Scott D. Sampson
The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. Bd. 10, Nr. 2, 2012
Dal Sasso originally referred it to the
Tetanurae Tetanurae (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, tyrannosauroids, ornithomimosaurs, compsognathids and maniraptorans (including birds). Tetanurans ar ...
He later considered that it may represent an
allosauroid Allosauroidea is a superfamily or clade of theropod dinosaurs which contains four families — the Metriacanthosauridae, Allosauridae, Carcharodontosauridae, and Neovenatoridae. Allosauroids, alongside the family Megalosauroidea, were among ...
, although in either case it would predate other members of the clades by roughly 20-30 million years. Benson considered it a member of
Coelophysoidea Coelophysoidea were common dinosaurs of the Late Triassic and Early Jurassic periods. They were widespread geographically, probably living on all continents. Coelophysoids were all slender, carnivorous forms with a superficial similarity to the ...
in his review of ''
Magnosaurus ''Magnosaurus'' (meaning 'large lizard') was a genus of theropod dinosaur from the Middle Jurassic of England. It is based on fragmentary remains and has often been confused with or included in ''Megalosaurus''. History and taxonomy In 1923, Fr ...
''. The presence of a
wishbone Wishbone commonly refers to: * Furcula, a fork-shaped bone in birds and some dinosaurs Wishbone may also refer to: * Wish-Bone, an American salad dressing and condiment company * Wishbone formation, a type of offense in American football * Wish ...
may support its placement as a tetanuran, although wishbones have been reported from coelophysoids. The 2018 description paper ran a large phylogenetic analysis, and found it to be a basal
ceratosaur Ceratosaurs are members of the clade Ceratosauria, a group of dinosaurs defined as all theropods sharing a more recent common ancestor with ''Ceratosaurus'' than with birds. The oldest known ceratosaur, ''Saltriovenator'', dates to the earlies ...
, the sister-taxon of '' Berberosaurus''. The phylogenetic analysis is as shown:


Provenance and Paleoenvironment

''Saltriovenator'' was found on an open marine environment, where it was probably washed from the nearest mainland, being scavenged by invertebrates as proven by the presence of '' Sedilichnus'' sp. on the bones. This depositional environment, part of the
Saltrio Formation The Saltrio Formation is a geological formation in Italy. It dates back to the middle Sinemurian, and would have represented a pelagic or near-epicontinental environment, judging by the presence of marine fauna such as the nautiloid '' Cenoceras' ...
is considered as part of a proximal slope or ramp that was probably an open subtidal zone reached by the effects of storm waves and with constant bottom currents. Since the beginning of the Jurassic, from Hettangian to earliest
Sinemurian In the geologic timescale, the Sinemurian is an age and stage in the Early or Lower Jurassic Epoch or Series. It spans the time between 199.3 ± 2 Ma and 190.8 ± 1.5 Ma (million years ago). The Sinemurian is preceded by the Hettangian and is ...
on the western Lombardy Basin there was a notorious continental area that was found to be wider than previously thought, where a warm humid paleoclimate developed. The Dinosaur Fossils found on the Saltrio formation can have been translated from this area, or alternatively, the Arbostora swell (that was located at the north of the Saltrio formation, on
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 ...
). This was an emerged structural high close to the ''Saltrio Formation'', that caused a division between two near subsiding basins located at Mt. Nudo (East) and Mt. Generoso (West). It settled over a carbonate platform linked with other wider areas that appear along the west to the southeast, developing a large shallow water gulf to the north, where the strata deposited was controlled by a horst and tectonic gaben. Several outcrops of the so-called “terra rossa” paleosoils were also found, including at Castello Cabiaglio-Orino, a dozen of kilometers West of Saltrio. These outcrops show that the emerged areas that on the Hettangian-Sinemurian, the current location of the modern Maggiore Lake was covered with forests, what was proven by the presence of large plant fragments on the Moltrasio Formation. The plants have been recovered between the locations of Cellina and Arolo (eastern side of Lake Maggiore), from rocks that have been found to be coeval in age to the ''Saltrio Formation''. The Flora includes genera such as
Bennettitales Bennettitales (also known as cycadeoids) is an extinct order of seed plants that first appeared in the Permian period and became extinct in most areas toward the end of the Cretaceous. Bennettitales are among the most common Mesozoic seed plants ...
(''
Ptilophyllum ''Ptilophyllum'' is an extinct form genus of leaves belonging to the extinct seed plant order Bennettitales. The leaves, like other Bennettitales morphogenera are generally pinnate Pinnation (also called pennation) is the arrangement of feath ...
''), terrestrial
Araucariaceae Araucariaceae – also known as araucarians – is an extremely ancient family of coniferous trees. The family achieved its maximum diversity during the Jurassic and Cretaceous periods and the early Cenozoic, when it was distributed almost worldw ...
(''
Pagiophyllum ''Pagiophyllum'' is a form genus of fossil coniferous plant foliage. Plants of the genus have been variously assigned to several different conifer groups including Araucariaceae and Cheirolepidiaceae. They were found around the globe during the ...
''), and
Cheirolepidiaceae Cheirolepidiaceae is an extinct family of conifers. They first appeared in the Triassic, and were widespread during most of the Mesozoic era. They are united by the possession of a distinctive pollen type assigned to the form genus '' Classopolli ...
(''
Brachyphyllum ''Brachyphyllum'' (meaning "short leaf") is a form genus of fossil coniferous plant foliage. Plants of the genus have been variously assigned to several different conifer groups including Araucariaceae and Cheirolepidiaceae. They are known from ...
''), that developed on inland areas with dry-warm conditions. ''Saltriovenator'' probably come from this nearby landmass, as other emerged zones, such as the Trento Platform where it's far of the location of discovery. If so, this theropod was probably the largest predator on the region.


See also

*
Timeline of ceratosaur research This timeline of ceratosaur research is a chronological listing of events in the history of paleontology focused on the ceratosaurs, a group of relatively primitive, often horned, predatory theropod dinosaurs that became the apex predators of ...


References

{{Taxonbar, from1=Q60146997, from2=Q658283 Ceratosaurs Early Jurassic dinosaurs of Europe Jurassic Italy Sinemurian life Fossils of Italy Fossil taxa described in 2018