Riparovenator
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''Riparovenator'' ("riverbank hunter") 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 baryonychine
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 ...
) period of Britain. The genus contains a single species, ''Riparovenator milnerae''.


Discovery and naming

Between 2013 and 2017, spinosaurid fossils were uncovered at the beach near the Chilton Chine before being brought to
Dinosaur Isle Dinosaur Isle is a purpose-built dinosaur museum located in Sandown on the Isle of Wight in southern England. The museum was designed by Isle of Wight architect Rainey Petrie Johns in the shape of a giant pterosaur. It claims to be the first c ...
. Such remains had historically been referred to ''
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 ...
'' but were understood later to represent two new species. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
remains of this taxon consist of IWCMS 2014.95.6 (
premaxilla 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 mammals h ...
ry bodies), IWCMS 2014.96.1, 2; 2020.448.1, 2 (a disarticulated
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, brain-pan, or brainbox, 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 calv ...
), and IWCMS 2014.96.3 (a partial lacrimal and prefrontal). Referred remains include a posterior nasal fragment (IWCMS 2014.95.7) and a caudal axial series of twenty-two vertebrae (IWCMS 2020.447.1-39), representing around fifty individual bones in total. All of the material was recovered from rocks in the Chilton Chine of the
Wessex Formation The Wessex Formation is a fossil-rich England, English geological formation that dates from the Berriasian to Barremian Stage (stratigraphy), stages of the Early Cretaceous. It forms part of the Wealden Group and underlies the younger Vectis Form ...
. In 2021, the
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
''Riparovenator milnerae'' was named and described by a team of
palaeontologists 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 ...
including Chris T. Barker, David W. E. Hone,
Darren Naish Darren William Naish (born 26 September 1975) is a British vertebrate palaeontologist, author and science communicator. As a researcher, he is best known for his work describing and reevaluating dinosaurs and other Mesozoic reptiles, including ...
, and others. The generic name is derived from the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
''rīpārius'', "of the river bank", and ''vēnātor'', "hunter". 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 ...
honors Angela Milner, deceased in August 2021.


Classification

In their phylogenetic analysis, Barker ''et al''. (2021) recovered ''Ceratosuchops'' within the
Baryonychinae Baryonychinae is an extinct clade or subfamily of spinosaurids from the Early Cretaceous of Europe and West Africa. The clade was named by Charig & Milner in 1986 and defined by Sereno et al. in 1998 and Holtz et al. in 2004 as all taxa more close ...
, as the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to the coeval ''Riparovenator''. They are, in turn, in a clade containing ''
Suchomimus ''Suchomimus'', from Ancient Greek ''σούχος'' (''soúkhos''), meaning "crocodile", and Latin ''mimus'', meaning "actor", is a genus of spinosaurid dinosaur that lived between 125 and 112 million years ago in what is now Niger, North A ...
'', which they name Ceratosuchopsini. In 2022, Sereno and colleagues tentatively combined ''Ceratosuchops'' and ''Riparovenator'' into a single taxonomic unit for their phylogenetic analysis. They reason that the different features between the two taxa could be attributed to individual variation, citing the cranial variation present in specimens of ''
Allosaurus fragilis ''Allosaurus'' ( ) is an extinct genus of theropoda, theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic Geologic time scale, period (Kimmeridgian to late Tithonian Geologic time scale, ages). The first fossil ...
''. Some of their supposed distinguishing features are also seen in parts of the braincase of ''Suchomimus'', their closest relative. The results of their phylogenetic analysis (with ''Ceratosuchops'' and ''Riparovenator'' scored together) yielded similar results to those of Barker ''et al''. (2021), with the Wessex baryonychine fossils recovered as the sister taxon to ''Suchomimus''.


Palaeoecology

''Riparovenator'' lived in a dry Mediterranean habitat in the
Wessex Formation The Wessex Formation is a fossil-rich England, English geological formation that dates from the Berriasian to Barremian Stage (stratigraphy), stages of the Early Cretaceous. It forms part of the Wealden Group and underlies the younger Vectis Form ...
, where rivers were home to riparian galleries. Like most spinosaurids, it would have fed on aquatic prey as well as other terrestrial prey in these areas. Other dinosaurs from the Wessex Formation of the Isle of Wight the theropods ''Ceratosuchops'', ''
Neovenator ''Neovenator'' ( nˈiːə͡ʊvˌɛne͡ɪtə; "new hunter") is a genus of carcharodontosaurian theropod dinosaur. It is known primarily from several skeletons found in the Early Cretaceous (Hauterivian-Barremian) Wessex Formation on the south co ...
'', ''
Eotyrannus ''Eotyrannus'' (meaning "dawn tyrant") is a genus of tyrannosauroid theropod dinosaur hailing from the Early Cretaceous Wessex Formation beds, included in Wealden Group, located in the southwest coast of the Isle of Wight, United Kingdom. The re ...
'', '' Aristosuchus'', '' Thecocoelurus'', '' Calamospondylus'', and '' Ornithodesmus''; the ornithopods ''Iguanodon'', ''
Hypsilophodon ''Hypsilophodon'' (; meaning "high-crested tooth") is a neornithischian dinosaur genus from the Early Cretaceous period of England. It has traditionally been considered an early member of the group Ornithopoda, but recent research has put this ...
'', and ''
Valdosaurus ''Valdosaurus'' ("Weald Lizard") is a genus of bipedal herbivorous iguanodont ornithopod dinosaur found on the Isle of Wight and elsewhere in England, and possibly also Spain and Romania. It lived during the Early Cretaceous. Discovery and namin ...
''; the sauropods ''
Ornithopsis ''Ornithopsis'' (meaning "bird-likeness") is a genus of sauropod dinosaur, from the Early Cretaceous of England and possibly Germany. The type species, which is the only species seen as valid today, is ''O. hulkei,'' which is only known from frag ...
'', ''
Eucamerotus ''Eucamerotus'' (meaning "well-chambered", in reference to the hollows of the vertebrae) was a genus of sauropod dinosaur from the Barremian-age Lower Cretaceous Wessex Formation ( Wealden) of the Isle of Wight, England. History and taxonomy J ...
'', and '' Chondrosteosaurus''; and the
ankylosaur Ankylosauria is a group of herbivorous dinosaurs of the clade 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 l ...
''
Polacanthus ''Polacanthus'', deriving its name from the Ancient Greek polys-/πολύς- "many" and akantha/ἄκανθα "thorn" or "prickle", is an early armoured, spiked, plant-eating ankylosaurian dinosaur from the early Cretaceous period of England. In ...
''. 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 closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. An initially interbreeding population that splits into two or more distinct spe ...
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).


References

{{Portal bar, Paleontology, Britain Spinosauridae Dinosaur genera Barremian dinosaurs Wessex Formation Taxa named by Darren Naish Fossil taxa described in 2021 Dinosaurs of the United Kingdom