Vallibonavenatrix
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''Vallibonavenatrix'' (meaning "
Vallibona Vallibona () is a small town and Municipalities of Spain, municipality located in the Ports (comarca), Ports comarca, province of Castelló (province), Castelló, part of the Valencia (autonomous community), autonomous community of Valencia, Spa ...
huntress" after the town near where its remains were found) is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
spinosaurid Spinosauridae (or spinosaurids) is a clade or Family (taxonomy), family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. Spinosaurid fossils have been recovered worldwide, including Africa, Europe, South America, and Asia. ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
from the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
(
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 125.77 Ma (million years ago) and 121.4 ± 1.0 Ma (Historically, this stage was placed at 129.4 million to approximately 125 million years ago) It is a ...
) Arcillas de Morella Formation of Castellón,
Spain Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...
. 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. * ...
and only
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
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, located in the
Province of Castellón Castellón (); co-officially in ) 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 Sea to the east. T ...
, Spain. In one of these—the Santa Águeda locality in the town of
Vallibona Vallibona () is a small town and Municipalities of Spain, municipality located in the Ports (comarca), Ports comarca, province of Castelló (province), Castelló, part of the Valencia (autonomous community), autonomous community of Valencia, Spa ...
—he excavated numerous
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
remains dating to the
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
era An era is a span of time. Era or ERA may also refer to: * Era (geology), a subdivision of geologic time * Calendar era Education * Academy of European Law (German: '), an international law school * ERA School, in Melbourne, Australia * E ...
, among which were
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
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 as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
Fernando Gómez-Fernández and colleagues published a provisional description on the
pelvis The pelvis (: pelves or pelvises) is the lower part of an Anatomy, anatomical Trunk (anatomy), trunk, between the human abdomen, abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also c ...
of a
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
from Forner's collection. Further fossils from the same specimen included a cervical (neck)
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
, 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 The fus ...
(back) vertebrae, an almost complete
sacrum The sacrum (: sacra or sacrums), in human anatomy, is a 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 of the pelvic cavity, ...
, 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 th ...
(tail) vertebrae, ten partial
ribs The rib cage or thoracic cage is an endoskeletal enclosure in the thorax of most vertebrates that comprises the ribs, vertebral column and sternum, which protect the vital organs of the thoracic cavity, such as the heart, lungs and great vessels ...
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 la ...
, an almost complete left ilium (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 the city of Naples. It is the largest of the Phlegrean Islands. Although inhabited since the Bronze Age, as a Ancient G ...
(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 any type of communication that aims to make vivid a place, object, person, group, or other physical entity. It is one of four rhetorical modes (also known as ''modes of discourse''), along with exposition, argumentation, and narr ...
of a new
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
and
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
spinosaurid Spinosauridae (or spinosaurids) is a clade or Family (taxonomy), family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. Spinosaurid fossils have been recovered worldwide, including Africa, Europe, South America, and Asia. ...
dinosaur, ''Vallibonavenatrix cani'', with the partial skeleton as the
holotype specimen A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was Species description, formally described. It is either the single such physical example (or illus ...
. 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 ( ), also known as Iberia, is a peninsula in south-western Europe. Mostly separated from the rest of the European landmass by the Pyrenees, it includes the territories of peninsular Spain and Continental Portugal, comprisin ...
.'''' Other Iberian spinosaurids include '' Iberospinus'', ''
Camarillasaurus ''Camarillasaurus'' (meaning " Camarillas lizard") is a genus of theropod dinosaur from the Early Cretaceous period (Barremian) of Camarillas, Teruel Province, in what is now northeastern Spain. Described in 2014, it was originally identified as ...
'', '' Protathlitis'', and '' Riojavenatrix''.


Description

''Vallibonavenatrix'' was a moderately-sized
bipedal Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) 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'' ...
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
. The upwards-projecting
neural spines Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
of its dorsal vertebrae were moderately tall, and one known spine is expanded from bottom to top in a
trapezoid In geometry, a trapezoid () in North American English, or trapezium () in British English, is a quadrilateral that has at least one pair of parallel sides. The parallel sides are called the ''bases'' of the trapezoid. The other two sides are ...
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 120 and 113 million years ago, during the Aptian Stage (geology), stage of the Early Cretaceous Period (geologic time), period. It is k ...
''. 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 may b ...
on ''Vallibonavenatrix'''s back, as is seen in some other spinosaurids. The sacrum of ''Vallibonavenatrix'' had deep pleurocoelous fossae (depressions) and
pneumatic Pneumatics (from Greek 'wind, breath') is the use of gas or pressurized air in mechanical systems. Pneumatic systems used in Industrial sector, industry are commonly powered by compressed air or compressed inert gases. A centrally located a ...
(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 ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of large-bodied
tetanuran Tetanurae (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, and coelurosaurs (which includes tyrannosauroids, ornithomimosaurs, compsognathids and maniraptoran ...
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 botanical subfamily names with "-oideae", and zool ...
: 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 125.77 Ma (million years ago) and 121.4 ± 1.0 Ma (Historically, this stage was placed at 129.4 million to approximately 125 million years ago) It is a ...
of Spain and England, ''Vallibonavenatrix'' was found to be more closely related to spinosaurines such as ''
Spinosaurus ''Spinosaurus'' (; ) is a genus of large spinosaurid theropod dinosaurs that lived in what now is North Africa during the Cenomanian faunal stage, stage of the Late Cretaceous Period (geology), period, about 100 to 94 annum, million year ...
'' and ''
Irritator ''Irritator'' is a genus of Spinosauridae, spinosaurid dinosaur that lived in what is now Brazil during the Albian Geological stage, stage of the Early Cretaceous Geological period, Period, about 113 to 110 million years ago. It is known from ...
'' from the southern
supercontinent In geology, a supercontinent is the assembly of most or all of Earth's continent, continental blocks or cratons to form a single large landmass. However, some geologists use a different definition, "a grouping of formerly dispersed continents", ...
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
, and the Asian genus ''
Ichthyovenator ''Ichthyovenator'' is a genus of spinosaurid dinosaur that lived in what is now Laos, sometime between 120 and 113 million years ago, during the Aptian Stage (geology), stage of the Early Cretaceous Period (geologic time), period. It is k ...
''. Thus, Malafaia and colleagues placed it within Spinosaurinae. The results of their analysis are shown in the following
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
:'''' 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 ( ; c. 1701 – 1761) was an English statistician, philosopher, and Presbyterian minister. Bayesian ( or ) may be either any of a range of concepts and approaches that relate to statistical methods based on Bayes' theorem Bayes ...
) that they performed: outside of both Spinosaurinae and Baryonychinae as the most basal (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 Portugal, Portuguese dinosaur paleontologist and biologist Professor of Paleontology at the Faculdade de Ciências e Tecnologia da NOVA University Lisbon, Universidade Nova de Lisboa. He graduated in University of ...
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, which has been dated to the
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 125.77 Ma (million years ago) and 121.4 ± 1.0 Ma (Historically, this stage was placed at 129.4 million to approximately 125 million years ago) It is a ...
stage Stage, stages, or staging may refer to: Arts and media 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 Brit ...
of the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
period Period may refer to: Common uses * Period (punctuation) * Era, a length or span of time *Menstruation, commonly referred to as a "period" Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (o ...
, between 129.4 and 125
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
. It coexisted in this environment with other dinosaurs including the
ornithischia Ornithischia () is an extinct clade 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 ''Iguanodon'' ( ; meaning 'iguana-tooth'), named in 1825, is a genus of iguanodontian dinosaur. While many species found worldwide have been classified in the genus ''Iguanodon'', dating from the Late Jurassic to Early Cretaceous, Taxonomy (bi ...
'' 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 b ...
, and fellow spinosaurid '' Protathlitis''.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 Spinosauridae Dinosaur genera Barremian dinosaurs Fossil taxa described in 2019 Dinosaurs of Spain