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''Vallibonavenatrix'' (meaning " Vallibona huntress" after the town near where its remains were found) is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
of
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 ...
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 ...
from 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 Pr ...
(
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma ( million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is pre ...
)
Arcillas de Morella Formation The Arcillas de Morella Formation is a geological formation in Spain whose strata date back to the Barremian stage of the Early Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.Weishampel, David B; et a ...
of Castellón,
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = '' Plus ultra'' ( Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , ...
. The
type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * Ty ...
and only
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
is ''Vallibonavenatrix cani'', known from a partial skeleton.


Discovery and naming

During the late 1980s and early 1990s, amateur fossil collector Juan Cano Forner was recovering bones from various localities in
Els Ports Natural Park Ports de Tortosa-Beseit (), also known as Ports de Beseit, or simply as Els Ports or Lo Port by locals, is a limestone mountain massif located at the north-eastern end of the Sistema Ibérico, a complex system of mountain ranges and massifs in ...
, located in the
Province of Castellón Castellón (officially in ca-valencia, Castelló) is a province in the northern part of the Valencian Community. It is bordered by the provinces of Valencia to the south, Teruel to the west, Tarragona to the north, and by the Mediterranean S ...
, Spain. In one of these—the Santa Águeda locality in the town of Vallibona—he excavated numerous
vertebrate Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
remains dating to the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Creta ...
era An era is a span of time defined for the purposes of chronology or historiography, as in the regnal eras in the history of a given monarchy, a calendar era used for a given calendar, or the geological eras defined for the history of Earth. Com ...
, among which were
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 ...
fossils A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
. Forner housed these fossils in a private collection at Sant Mateu, which the
Generalitat Valenciana The Generalitat Valenciana is the generic name covering the different self-government institutions under which the Spanish autonomous community of Valencia is politically organized. It consists of seven institutions including the ''Corts Valen ...
acknowledged as a museographic collection in 1994. In 2007, the Spanish
palaeontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Fernando Gómez-Fernández and colleagues published a provisional description on the pelvis of 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 ...
from Forner's collection. Further fossils from the same specimen included a
cervical In anatomy, cervical is an adjective that has two meanings: # of or pertaining to any neck. # of or pertaining to the female cervix: i.e., the ''neck'' of the uterus. *Commonly used medical phrases involving the neck are **cervical collar **cervic ...
(neck)
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characterist ...
, six
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal co ...
(back) vertebrae, an almost complete
sacrum The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part ...
, fragments of neurapophyses, four
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) vertebrae, ten partial
ribs The rib cage, as an enclosure that comprises the ribs, vertebral column and sternum in the thorax of most vertebrates, protects vital organs such as the heart, lungs and great vessels. The sternum, together known as the thoracic cage, is a semi ...
and rib fragments, three incomplete
chevrons Chevron (often relating to V-shaped patterns) may refer to: Science and technology * Chevron (aerospace), sawtooth patterns on some jet engines * Chevron (anatomy), a bone * '' Eulithis testata'', a moth * Chevron (geology), a fold in rock l ...
, an almost complete left
ilium Ilium or Ileum may refer to: Places and jurisdictions * Ilion (Asia Minor), former name of Troy * Ilium (Epirus), an ancient city in Epirus, Greece * Ilium, ancient name of Cestria (Epirus), an ancient city in Epirus, Greece * Ilium Building, a ...
(main hip bone), fragments of the ventral part of a right ilium, incomplete right and left
ischia Ischia ( , , ) is a volcanic island in the Tyrrhenian Sea. It lies at the northern end of the Gulf of Naples, about from Naples. It is the largest of the Phlegrean Islands. Roughly trapezoidal in shape, it measures approximately east to wes ...
(lower and rearmost hip bone), and a fragment interpreted as belonging to the proximal part of a pubis (pubic bone).'''' In 2019, Elisabete Malafaia and colleagues published their
description Description is the pattern of narrative development that aims to make vivid a place, object, character, or group. Description is one of four rhetorical modes (also known as ''modes of discourse''), along with exposition, argumentation, and na ...
of a new
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
and
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
of
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 ...
dinosaur, ''Vallibonavenatrix cani'', with the partial skeleton as the
holotype specimen 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 ...
. The generic name refers to the town of Vallibona, with the Latin suffix "''-venatrix''", meaning "huntress". The specific name honours Cano Forner, the fossil's discoverer. ''Vallibonavenatrix'' represents the most complete spinosaurid specimen recovered from the
Iberian Peninsula The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, defi ...
.'''' Other Iberian spinosaurids include ''
Baryonyx ''Baryonyx'' () is a genus of theropod dinosaur which lived in the Barremian stage of the Early Cretaceous period, about 130–125 million years ago. The first skeleton was discovered in 1983 in the Smokejack Clay Pit, of Surrey, England, in ...
'', '' Camarillasaurus'', and '' Iberospinus''.


Description

''Vallibonavenatrix'' was a moderately-sized
bipedal Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' ...
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
. The upwards-projecting
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 i ...
of its dorsal vertebrae were moderately tall, and one known spine is expanded from bottom to top in a
trapezoid A quadrilateral with at least one pair of parallel sides is called a trapezoid () in American and Canadian English. In British and other forms of English, it is called a trapezium (). A trapezoid is necessarily a convex quadrilateral in Eucli ...
al, fan-like shape, similar to those of the spinosaurid ''
Ichthyovenator ''Ichthyovenator'' is a genus of spinosaurid dinosaur that lived in what is now Laos, sometime between 125 and 113 million years ago, during the Aptian stage of the Early Cretaceous period. It is known from fossils collected from the ...
''. Similarities between the
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
of the neural spines of these two taxa may indicate the presence of elongated spines forming a
sail A sail is a tensile structure—which is made from fabric or other membrane materials—that uses wind power to propel sailing craft, including sailing ships, sailboats, windsurfers, ice boats, and even sail-powered land vehicles. Sails ma ...
on ''Vallibonavenatrix'''s back, as is seen in some other spinosaurids. The sacrum of ''Vallibonavenatrix'' had deep pleurocoelous fossae (depressions) and pneumatic (air-filled) openings. The ilium of the pelvis was also highly pneumatic, and had large internal chambers.''''


Classification

In 2019, Malafaia and her team classified ''Vallibonavenatrix'' within the Spinosauridae, a
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
of large-bodied
tetanuran Tetanurae (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, tyrannosauroids, ornithomimosaurs, compsognathids and maniraptorans (including birds). Tetanurans are ...
dinosaurs traditionally separated into two
subfamilies 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 ...
: Spinosaurinae and Baryonychinae. Though closer in proximity to European baryonychines such as ''Baryonyx'' from the
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma ( million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is pre ...
of Spain and England, ''Vallibonavenatrix'' was found to be more closely related to spinosaurines such as ''
Spinosaurus ''Spinosaurus'' (; ) is a genus of spinosaurid dinosaur that lived in what now is North Africa during the Cenomanian to upper Turonian stages of the Late Cretaceous period, about 99 to 93.5 million years ago. The genus was known first ...
'' and ''
Irritator ''Irritator'' is a genus of spinosaurid dinosaur that lived in what is now Brazil during the Albian stage of the Early Cretaceous Period, about 113 to 110 million years ago. It is known from a nearly complete skull found in the Romualdo Fo ...
'' from the southern
supercontinent In geology, a supercontinent is the assembly of most or all of Earth's continental blocks or cratons to form a single large landmass. However, some geologists use a different definition, "a grouping of formerly dispersed continents", which lea ...
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 st ...
, and the Asian genus ''
Ichthyovenator ''Ichthyovenator'' is a genus of spinosaurid dinosaur that lived in what is now Laos, sometime between 125 and 113 million years ago, during the Aptian stage of the Early Cretaceous period. It is known from fossils collected from the ...
''. Thus, Malafaia and colleagues placed it within Spinosaurinae. The results of their analysis are shown in the following
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
:'''' However, in 2021, Chris Barker and colleagues instead found ''Vallibonavenatrix'' in one of two positions, depending on the type of analysis ( parsimony-based or
Bayesian Thomas Bayes (/beɪz/; c. 1701 – 1761) was an English statistician, philosopher, and Presbyterian minister. Bayesian () refers either to a range of concepts and approaches that relate to statistical methods based on Bayes' theorem, or a follower ...
) that they performed: outside of both Spinosaurinae and Baryonychinae as the most
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 ...
(least specialized) spinosaurid, or as a member of the Baryonychinae. They concluded that its phylogenetic position was unstable. Like the results of Barker ''et al.'', a 2022 study by
Octávio Mateus Octávio Mateus (born 1975) is a Portuguese dinosaur paleontologist and biologist Professor of Paleontology at the Faculdade de Ciências e Tecnologia da Universidade Nova de Lisboa. He graduated in Universidade de Évora and received his PhD at ...
and Darío Estraviz-López did not find ''Vallibonavenatrix'' to be a spinosaurine. However, certain anatomical characteristics may indicate an affinity with that group, rather than with the baryonychines.


Palaeoenvironment

''Vallibonavenatrix'' hails from the
Arcillas de Morella Formation The Arcillas de Morella Formation is a geological formation in Spain whose strata date back to the Barremian stage of the Early Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.Weishampel, David B; et a ...
, which has been dated to the
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma ( million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is pre ...
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 Pr ...
period, between 129.4 and 125
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
. It coexisted in this environment with other dinosaurs including the
ornithischia Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek ...
ns '' Iguanodon bernissartensis'' and '' Morelladon beltrani'', an indeterminate
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 ...
, and an indeterminate baryonychine.Weishampel, David B; et al. (2004). "Dinosaur distribution (Early Cretaceous, Europe)." In: Weishampel, David B.; Dodson, Peter; and Osmólska, Halszka (eds.): The Dinosauria, 2nd, Berkeley: University of California Press. Pp. 562. .


References

{{Taxonbar, from=Q66614821 Spinosaurids Barremian life Early Cretaceous dinosaurs of Europe Cretaceous Spain Fossil taxa described in 2019