''Pyroraptor'' (meaning "fire thief") is an extinct

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

paravian Paraves are a widespread group of theropod dinosaurs that originated in the Middle Jurassic period. In addition to the extinct dromaeosaurids, troodontids, anchiornithids, and possibly the scansoriopterygids, the group also contains the aviala ...

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 ...

, probably a

dromaeosaurid Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...

or unenlagiid (considering that unenlagiids are a separate family), from the

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...

Ibero-Armorican island, of what is now southern

France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...

and northern Spain. It lived during the late

Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campanian s ...

and early

Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...

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 ...

, approximately 70.6 million years ago. It is known from a single partial specimen that was found in

Provence Provence (, , , , ; oc, Provença or ''Prouvènço'' , ) is a geographical region and historical province of southeastern France, which extends from the left bank of the lower Rhône to the west to the Italian border to the east; it is bor ...

in 1992, after a forest fire. The animal was named ''Pyroraptor olympius'' by Allain and Taquet in

2000 File:2000 Events Collage.png, From left, clockwise: Protests against Bush v. Gore after the 2000 United States presidential election; Heads of state meet for the Millennium Summit; The International Space Station in its infant form as seen from ...

Discovery and naming

Naming and Material

The first remains of ''Pyroraptor olympius'', or P. olympius, were discovered in southeastern France, at the La Boucharde locality of the Arc Basin in

. It was described and named by French

paleontologists 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 ...

Ronan Allain and

Philippe Taquet Philippe Taquet (b. April 25, 1940 Saint-Quentin, Aisne) is a French paleontologist who specializes in dinosaur systematics of finds primarily in northern Africa. He is a member of the French Academy of Sciences since November 30, 2004, president ...

, 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 ...

and so far the only species is ''Pyroraptor olympius''. The

name is

Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...

for "Fire thief", due to its remains being discovered after a forest fire that occurred in 1992. The

species In biology, a species is the basic unit of 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 the appropriate s ...

name is derived from Mont Olympe, a mountain in Provence at the foot of which the animal's remains were unearthed.Allain, R., and Taquet, P. (2000). "A new genus of Dromaeosauridae (Dinosauria, Theropoda) from the Upper Cretaceous of France." ''Journal of Vertebrate Paleontology'', 20: 404-407. une 27, 2000/ref> 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 ...

, MNHN BO001, consists of the second toe claw of the left foot. The assigned

paratypes In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). Of ...

include the equivalent claw of the right foot; the left second

metatarsal The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the med ...

; another, more complete second toe claw; a right

ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...

(long forearm bone); and two teeth. Additional material from the Vitória Formation and the

Tremp Group The Tremp Formation ( es, Formación de Tremp, ca, Formació de Tremp), alternatively described as Tremp Group ( es, Grupo Tremp), is a geological formation in the comarca Pallars Jussà, Lleida, Spain. The formation is restricted to the Trem ...

, both in Spain, was referred to ''Pyroraptor'', including five pedal digits, one manual digit, a piece of a

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 ...

, a right

radius In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...

, a dorsal vertebra, and a tail vertebra. Some teeth 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 ...

of North Eastern Spain have been compared to those referred to ''Pyroraptor'', suggesting that ''Pyroraptor'' may have also inhabited Spain; however, a 2022 reevaluation of these teeth states that they cannot be confidently assigned to ''Pyroraptor'' and may belong to a whole different variety of European

dromaeosaurids Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...

Relation to ''Variraptor''

Finds of dromaeosaurid dinosaur remains are rare in Europe and typically provide little taxonomic information. The first dromaeosaurid fossils found in France were those of ''

Variraptor mechinorum ''Variraptor'' ( ; "Var thief") is a genus of dromaeosaurid theropod dinosaur from the Late Cretaceous of France. Discovery Between 1992 and 1995 amateur paleontologists Patrick Méchin and Annie Méchin-Salessy uncovered the remains of a small ...

'', described by Jean Le Lœuff and Eric Buffetaut. Allain and Taquet considered ''Variraptor mechinorum'' to be a

nomen dubium 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 ...

on the basis that fossil material of the species was collected from localities different from that of 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 ...

, and that the material did not have enough diagnostic traits to warrant the naming of a new species. However, one of the authors that named ''Variraptor'', Jean Le Lœuff, points out in a conversation that the same reasons given for considering ''Variraptor mechinorum'' to be a nomen dubium would apply to ''Pyroraptor olympius'' as well, since ''Pyroraptor'''s remains present the exact same problems. In a 2009 description of new dromaeosaurid remains assigned to ''Variraptor mechinorum'', Phornphen Chanthasit and Eric Buffetaut addressed the claims of Allain & Taquet by mentioning that the original description of ''Variraptor mechinorum'' very clearly distinguished the holotype from any referred material and argue that some of the material very clearly belongs to the same individual while also elucidating that the situation with ''Pyroraptor'' is quite similar. However, due to the disarticulated nature of ''Pyroraptor'' remains, it's unknown if they belong to the same individual. This suspicion is affirmed by a 2012 study that states the known ''Pyroraptor'' material to belong to at least two different individuals. The other criticism of Allain and Taquet, that there are not enough diagnostic features to support the validity of ''Variraptor'', is also addressed by the description of new and overlapping ''Variraptor'' remains from the same locality as the holotype ''Variraptor'' specimen, some of which are stated to undeniably belong to the holotype individual, thus adding new diagnostic traits and establishing the validity of ''Variraptor mechinorum''. The authors also bring light to the fact that Allain and Taquet did not provide enough diagnosable traits to establish the holotype of ''Pyroraptor'' as a valid species, noting that most features stated for diagnosis are not unique to ''Pyroraptor'' but are simply widespread traits of dromaeosaurs. A 2012 study also agrees that ''Pyroraptor's'' "unique" traits are widespread features of dromaeosauridae while also ruling out some of the other "unique traits" as results of preservation deformation. Since the two existed at the same time and place, the possibility of a synonymy between ''Pyroraptor'' and ''Variraptor'' is raised but it is concluded that due to the current lack of overlap in material, this possible synonymy cannot yet be tested and more remains are needed.

Description

''Pyroraptor olympius'' was a dromaeosaurid, a small, bird-like predatory

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 ...

that possessed enlarged curved claws on the second toe of each foot for predation; these claws were 6.5 centimeters (2.5 in) long for Pyroraptor. As in other dromaeosaurids, these claws might have been used as weapons or as climbing aids.Manning, Phil L., Payne, David., Pennicott, John., Barrett, Paul M., Ennos, Roland A. (2005) "Dinosaur killer claws or climbing crampons?" Biology Letters (2006) 2; pg. 110-112 Its two known teeth are flattened and curved backwards, with their rear margins having finer

serrations Serration is a saw-like appearance or a row of sharp or tooth-like projections. A serrated cutting edge has many small points of contact with the material being cut. By having less contact area than a smooth blade or other edge, the applied pr ...

than at the front. As a dromaeosaurid, ''Pyroraptor'' likely had well-developed forelimbs with curved claws, and probably balanced the body with a long, thin tail. ''Pyroraptor'' was also covered in feathers, as many of its relatives, like ''

Microraptor ''Microraptor'' (Greek, μικρός, ''mīkros'': "small"; Latin, ''raptor'': "one who seizes") is a genus of small, four-winged dromaeosaurid dinosaurs. Numerous well-preserved fossil specimens have been recovered from Liaoning, China. They dat ...

'' and ''

Sinornithosaurus ''Sinornithosaurus'' (derived from a combination of Latin and Greek, meaning 'Chinese bird-lizard') is a genus of feathered dromaeosaurid dinosaur from the early Cretaceous Period (late Barremian) of the Yixian Formation in what is now China. It ...

'', also had plumage.

Scott Sampson Scott Donald Sampson (born April 22, 1961) is a Canadian paleontologist and science communicator. Sampson is currently the Executive Director of California Academy of Sciences in San Francisco, California. He was previously Vice President of Res ...

Discovery Channel Discovery Channel (known as The Discovery Channel from 1985 to 1995, and often referred to as simply Discovery) is an American cable channel owned by Warner Bros. Discovery, a publicly traded company run by CEO David Zaslav. , Discovery Channe ...

's 2003 documentary series '' Dinosaur Planet'', ep. 2: "Pod's Travels".

Classification

Although ''Pyroraptor olympius'' has been shown to be a

, in its initial description it was noted that due to the paucity of remains, not enough diagnostic features can be gathered to create a good comparison with other dromaeosaurids. This concern has been echoed by others, most notably in a 2012 phylogeny of paravia where ''Pyroraptor'' was found to wildly fluctuate in its position within Dromaeosauridae. Due to its status as a "wildcard taxa", ''Pyroraptor'' is known to create issues in phylogenies which are only fixed when it is pruned from the analysis. Hence, this taxon has rarely been used since in phylogenies, which causes difficulty in pinning its precise classification. ''Pyroraptor'' was included in a 2014 phylogeny of ''

'', as with most trees including ''Pyroraptor'', this caused a

polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...

as seen in the cladogram below. This

was also repeated in a phylogeny of ''

Archaeopteryx ''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird''), is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaīos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...

'' from the same year. In the 2019 description of ''

Hesperornithoides ''Hesperornithoides'' (meaning "western bird form"; nicknamed "Lori") is a genus of troodontid theropod dinosaur that lived in North America during the Late Jurassic period. Discovery Although several Troodontid teeth were found, with the trood ...

'', a phylogeny tree found ''Pyroraptor'' to be an

unenlagiine Unenlagiinae is a subfamily of long-snouted paravian theropods. They are traditionally considered to be members of Dromaeosauridae, though some authors place them into their own family, Unenlagiidae, alongside the subfamily Halszkaraptorinae. ...

. The study noted that this is biostratigraphically and geographically consistent with Cretaceous Europe showing a trend of hosting typically

Gondwanan 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 ...

species. This is also consistent with some previous studies, such as a 2005 study where it was suggested that some

laurasian 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 ...

dromaeoaurids like ''Pyroraptor'' could be considered part of

lineages, but scant material from European forms prevented this idea from being tested at the time. A 2012 phylogeny study noted that despite being an unstable taxa, ''Pyroraptor'' never fell within any of the

dromaeosaur groups. The 2022 description of ''Vectiraptor greeni'' noted that the foot morphology of ''Pyroraptor olympius'' had more similarities with unenlagiines than with

eudromaeosaurs Eudromaeosauria ("true dromaeosaurs") is a subgroup of terrestrial Dromaeosauridae, dromaeosaurid theropod dinosaurs. They were relatively small to medium-sized, feathered hypercarnivores (with diets consisting almost entirely of other terrestr ...

, lending support to the idea that European raptors were closely related to Southern forms from Africa and South America. Another subsequent 2022 study that describes a newly discovered dromaeosaurid sickle claw from the same

geologic formation A geological formation, or simply formation, is a body of rock having a consistent set of physical characteristics ( lithology) that distinguishes it from adjacent bodies of rock, and which occupies a particular position in the layers of rock exp ...

as ''Pyroraptor'', compared the sickle claws with those of ''Pyroraptor'' and other dromaeosaurids, and more similarities between ''Pyroraptor'' and other unenlagiines were found, this time in the anatomy of the claws. Two separate studies from 2011 regarding unenlagiines, where the teeth of ''Pyroraptor'' were compared to those of South American Unenlagiines found that although not significantly, some similarities in cross section were noticeable. But Unenlagiine teeth do not have any serrations on their edges and are usually fluted, a trait that is in direct contradiction with what is seen in ''Pyroraptor'' teeth which are serrated, as is typical of dromaeosaurs, the 2021 description of '' Ypupiara lopai'' rules out ''Pyroraptor'' as being an unenlagiid due to this dissimilarity. However, a 2022 study points out that if ''Pyroraptor'' is an unenlagiid, then this would mean that not all unenlagiids would have fluted, unserrated teeth. During the initial description of ''Pyroraptor olympius'', the suggested model of

evolution proposed that the

dromaeosaurs Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...

originated from North America or Euro-America, and then dispersed into Asia. On the basis of newly discovered Early Cretaceous material from China, this dispersal model has since lost support, so Allain and Taquet postulated that perhaps European dromaeosaurids such as ''Pyroraptor olympius'' were the remnants of Late Jurassic or Early Cretaceous fauna that evolved separately during the late Cretaceous isolation of Southern Europe. However, recent studies show that ''Pyroraptor'' along with other European raptors, likely originated from the Southern continent of

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 ...

. as its anatomy is more consistent with the

Unenlagiines Unenlagiinae is a subfamily of long-snouted paravian theropods. They are traditionally considered to be members of Dromaeosauridae, though some authors place them into their own family, Unenlagiidae, alongside the subfamily Halszkaraptorinae. Un ...

than with

. This is congruent with other

European fauna, which are typically

lineages.

Paleoecology

Paleogeography of Europe in the Maastrichtian

''Pyroraptor olympius'' is known from the

Argiles et Grès à Reptiles Formation Argiles et Grès à Reptiles Formation (meaning Reptile Clay and Sandstone) also known as the Argiles Rutilantes FormationTabuce et al., 2013 is an early MaastrichtianWeishampel et al., 2004, pp.588-593 France, French geologic formation in the Depa ...

of what is now Southern France. During the Late Cretaceous, it was one part of the island landmass known as Ibero-Armorica, formed from what is today Southern France and Northern Spain in the

Tethys Ocean The Tethys Ocean ( el, Τηθύς ''Tēthús''), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean that covered most of the Earth during much of the Mesozoic Era and early Cenozoic Era, located between the ancient continents ...

. At that time, ''Pyroraptor'' would have coexisted alongside a variety of species such as ''

Rhabdodon priscus ''Rhabdodon'' (meaning "fluted tooth") is a genus of ornithopod dinosaur that lived in Europe approximately 70-66 million years ago in the Late Cretaceous. It is similar in build to a very robust "hypsilophodont" (non-iguanodont ornithopod), thou ...

'', ''

Zalmoxes ''Zalmoxes'' is an extinct genus of rhabdodontid ornithopod dinosaur from the Maastrichtian of Romania. The genus is known from specimens first named as the species '' Mochlodon robustum'' in 1899 by Franz Nopcsa before being reclassified as ...

'', ''

Ampelosaurus atacis ''Ampelosaurus'' ( ; meaning "vine lizard") is a titanosaurian sauropod dinosaur from the Late Cretaceous Period of what is now France. Its type species is ''A. atacis'', named by Le Loeuff in 1995. A possible unnamed species has given ''Ampel ...

'', '' Lirainosaurus astibiae'', '' Atsinganosaurus velauciensis'', some undescribed titanosaurids, '' Arcovenator escotae'', '' Tarascosaurus salluvicus'', ''Struthiosaurus sp''., '' Variraptor mechanorum'', '' Gargantuavis philoinos,'' '' Martinaves cruzyensis'', and non-dinosaurs such as the pterosaur ''

Azhdarcho lancicollis ''Azhdarcho'' is a genus of azhdarchid pterosaur from the late Cretaceous Period of the Bissekty Formation (middle Turonian stage, about 92 million years ago) of Uzbekistan, as well as the Zhirkindek Formation of Kazakhstan. It is known from ...

'', the crocodylomorphs '' Musturzabalsuchus buffetauti,

Massaliasuchus ''Massaliasuchus'' is an extinct monospecific genus of allodaposuchid eusuchian crocodyliform that is known from fossils found in Santonian–Campanian-age Upper Cretaceous rocks of southeastern France. Systematics ''Massaliasuchus'' was first ...

,'' ''

Allodaposuchus ''Allodaposuchus'' is an extinct genus of crocodyliforms that lived in what is now southern Europe during the Campanian and Maastrichtian stages of the Late Cretaceous. Although generally classified as a non-crocodylian eusuchian crocodylomorph, ...

'' the turtles '' Dortoka'', and Solemydid turtles,

zhelestid Zhelestidae is a lineage of extinct eutherian mammals. Occurring in the Late Cretaceous from the Turonian to the Maastrichtian, they were an extremely successful group, with representatives present in Europe, Asia, India (and subsequently in Mad ...

eutherians, palaeobatrachid anurans, batrachosauroidid urodeles,

amphisbaenian Amphisbaenia (called amphisbaenians or worm lizards) is a group of usually legless squamates, comprising over 200 extant species. Amphisbaenians are characterized by their long bodies, the reduction or loss of the limbs, and rudimentary eyes. As ...

and/or anguid squamates, and derived alethinophidian snakes.

References

External links

''Pyroraptor'' official site
in

French French (french: français(e), link=no) may refer to: * Something of, from, or related to France ** French language, which originated in France, and its various dialects and accents ** French people, a nation and ethnic group identified with Franc ...

''Pyroraptor'' computer animation
by Meteor Studios for

''Pyroraptor'' computer animation stills
by Meteor Studios for Discovery Channel {{Taxonbar, from=Q132542 Dromaeosaurs Late Cretaceous dinosaurs of Europe Campanian life Maastrichtian life Cretaceous France Fossils of France Cretaceous Spain Fossils of Spain Tremp Formation Fossil taxa described in 2000 Taxa named by Philippe Taquet