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''Elopteryx'' 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 nom ...
of, perhaps troodontid,
maniraptora Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to ''Ornithomimus velox''. It contains the major subgroups Avialae, Deinonychosauria, Oviraptoros ...
n
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 ...
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 ...
based on fragmentary
fossil 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 ...
s found in late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
Period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
rocks of
Romania Romania ( ; ro, România ) is a country located at the crossroads of Central, Eastern, and Southeastern Europe. It borders Bulgaria to the south, Ukraine to the north, Hungary to the west, Serbia to the southwest, Moldova to the east, and ...
. The single
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 ...
, ''Elopteryx nopcsai'', is known only from very incomplete material, and therefore is considered a '' nomen dubium'' ("dubious name") by most paleontologists.


History of discovery and naming


Initial finds

In the late nineteenth or early twentieth century, the famous Hungarian
paleontologist 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 ...
Franz Nopcsa von Felső-Szilvás Baron Franz Nopcsa von Felső-Szilvás (also Baron Nopcsa von Felső-Szilvás, Baron Nopcsa, Ferenc Nopcsa, báró felsőszilvási Nopcsa Ferenc, Baron Franz Nopcsa, and Franz Baron Nopcsa; May 3, 1877 – April 25, 1933) was a Hungarian aristoc ...
found near Sînpetru, in what is now the Romanian region of
Transylvania Transylvania ( ro, Ardeal or ; hu, Erdély; german: Siebenbürgen) is a historical and cultural region in Central Europe, encompassing central Romania. To the east and south its natural border is the Carpathian Mountains, and to the west the Ap ...
, some bone fragments of a small theropod. These were acquired by the
British Museum of Natural History The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum ...
. In 1913, curator
Charles William Andrews Charles William Andrews (30 October 1866 – 25 May 1924) F.R.S., was a British palaeontologist whose career as a vertebrate paleontologist, both as a curator and in the field, was spent in the services of the British Museum, Department of Ge ...
named these 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 specime ...
''Elopteryx nopcsai''. The genus name ''Elopteryx'' is from
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic p ...
''helos'' (ἕλος), "marsh" + ''pteryx'' (πτέρυξ), "wing". The specific name honors Nopcsa. Initially, ''Elopteryx'' was described from its
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 sever ...
, a
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 ...
left
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
, specimen BMNH A1234. A second upper left thighbone fragment, BMNH A1235, was referred. A distal left
tibiotarsus The tibiotarsus is the large bone between the femur and the tarsometatarsus in the leg of a bird. It is the fusion of the proximal part of the tarsus with the tibia. A similar structure also occurred in the Mesozoic Heterodontosauridae. These s ...
was also tentatively assigned to this
taxon In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
; it was initially classified with the same specimen number as the holotype and was found in close proximity, but may not be from the same individual (see below). This has since been relabeled and is now specimen BMNH A4359. The exact location and time of the discoveries are today unknown. The fossils date from the early-late
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 ...
( Begudian)
faunal stage In chronostratigraphy, a stage is a succession of rock strata laid down in a single age on the geologic timescale, which usually represents millions of years of deposition. A given stage of rock and the corresponding age of time will by convent ...
, circa 71-66
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) ...
, originating from the
Sânpetru Formation The Sânpetru Formation is an early Maastrichtian geologic formation. Dinosaur remains are among the fossils that have been recovered from the formation.Weishampel, et al. (2004). "Dinosaur distribution." Pp. 517-607. It is located in Romania, ne ...
of the
Hațeg Island Hațeg Island was a large offshore island in the Tethys Sea which existed during the Late Cretaceous period, probably from the Cenomanian to the Maastrichtian ages. It was situated in an area corresponding to the region around modern-day Hațeg, H ...
. The animal was by Andrews believed to be a
pelecaniform The Pelecaniformes are an order of medium-sized and large waterbirds found worldwide. As traditionally—but erroneously—defined, they encompass all birds that have feet with all four toes webbed. Hence, they were formerly also known by such n ...
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
.Harrison & Walker (1975) In 1929 the Hungarian paleontologist
Kálmán Lambrecht Kálmán Lambrecht (1889–1936) was a Hungarian palaeontologist, best known for his work on fossil birds. He authored the “Handbuch der Palaeornithologie”, an exhaustive review of fossil birds published in 1933. Positions held include librar ...
referred two more specimens: BMNH A PAL.1528 and BMNH A PAL.1588, respectively a left and a right tibiotarsus. In 1933 Lambrecht named a separate family Elopterygidae. The supposed family Elopterygidae was initially placed in the suborder Sulae – then still in the
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...
"
Pelecaniformes The Pelecaniformes are an order of medium-sized and large waterbirds found worldwide. As traditionally—but erroneously—defined, they encompass all birds that have feet with all four toes webbed. Hence, they were formerly also known by such n ...
" – in 1963 by
Pierce Brodkorb William Pierce Brodkorb (September 29, 1908, Chicago – July 18, 1992, Gainesville, Florida) was an American ornithologist and paleontologist. Interested in birds since childhood, he was taught to prepare birds at the age of 16. Later, he rec ...
in his fossil bird catalogue, and the Cenozoic genera '' Argillornis'' and '' Eostega'' were moved to it. These two are unequivocal derived neornith birds and the latter indeed seems to be an ancient sulid, whereas ''Argillornis'' has turned out to be referrable to the giant
pseudotooth bird The Pelagornithidae, commonly called pelagornithids, pseudodontorns, bony-toothed birds, false-toothed birds or pseudotooth birds, are a prehistoric family of large seabirds. Their fossil remains have been found all over the world in rocks dating ...
''
Dasornis ''Dasornis'' is a genus of the prehistoric pseudotooth birds. These were probably rather close relatives of either pelicans and storks, or of waterfowl, and are here placed in the order Odontopterygiformes to account for this uncertainty. ...
'' which was almost certainly not very closely related to the Sulae. Reconstruction attempts of ''E. nopcsai'' lik
this
are based on this presumed affiliation with
gannet Gannets are seabirds comprising the genus ''Morus'' in the family Sulidae, closely related to boobies. Gannets are large white birds with yellowish heads; black-tipped wings; and long bills. Northern gannets are the largest seabirds in the ...
s and cormorants. But more recent studies would result in radically different interpretations.


Later finds

In 1975, the distal tibiotarsi BMNH A1588 and BMNH A1528, together with BMNH A4359, were by Colin James Oliver Harrison and
Cyril Alexander Walker Cyril Alexander Walker (8 February 1939 – 6 May 2009) was a British palaeontologist, curator of fossil birds in the Natural History Museum. He was also interested in fossil turtles. book, '' Smithsonian Handbook of Fossils''. He has also cont ...
removed from ''Elopteryx'', redescribed as '' Bradycneme draculae'' and ''
Heptasteornis andrewsi ''Heptasteornis'' is the name given to a potentially dubious genus of alvarezsaurid dinosaur from the Late Cretaceous. The type (and only known) species is ''Heptasteornis andrewsi'', described as a presumed gigantic prehistoric owl in 1975. It w ...
'' respectively, and used to establish a supposed family of gigantic two metre tall
owl Owls are birds from the order Strigiformes (), which includes over 200 species of mostly solitary and nocturnal birds of prey typified by an upright stance, a large, broad head, binocular vision, binaural hearing, sharp talons, and feathers a ...
s, the Bradycnemidae. In 1978 Brodkorb had changed his opinion after the supposed ''Elopteryx'' material was divided among three species in total, and was actually the first scholar in modern times to suggest that these
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 Cretace ...
bones were not of birds but of non-avian dinosaurs. In 1981, Dan Grigorescu and Eugen Kessler stated that ''Elopteryx'' was a non-avian
coelurosauria Coelurosauria (; from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, t ...
n dinosaur. They also referred a supposed distal femur (FGGUB R.351) to ''Elopteryx'', but this was eventually identified as a
hadrosauroid Hadrosauroidea is a clade or superfamily of ornithischian dinosaurs that includes the "duck-billed" dinosaurs, or hadrosaurids, and all dinosaurs more closely related to them than to ''Iguanodon''. Their remains have been recovered in Asia, Eu ...
distal
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 me ...
.Kessler, E., Grigorescu D. and Csiki, Z., 2005, "''Elopteryx'' revisited - a new bird-like specimen from the Maastrichtian of the Hateg Basin", ''Acta Palaeontologica Romaniae''. 5: 249-258 In 1992, it was proposed by
Jean Le Loeuff Jean may refer to: People * Jean (female given name) * Jean (male given name) * Jean (surname) Fictional characters * Jean Grey, a Marvel Comics character * Jean Valjean, fictional character in novel ''Les Misérables'' and its adaptations * Jean ...
e.a. that ''Bradycneme'' and ''Heptasteornis'' should be synonymized with ''E. nopcsai'' again, and a femur (MDE-D203), an anterior
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 c ...
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 characteristi ...
(MDE-D01), a posterior
sacral vertebra 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 ...
(MDE collection, unnumbered) and some dorsal rib fragments from the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
Grès à Reptiles formation of France were described as an indeterminate species of ''Elopteryx''; that study placed all this material in the
Dromaeosauridae 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 a family or subfamily (Elopteryginae) very close to these. The vertebrae were in 1998 separated again and assigned to a new dromaeosaurid, '' Variraptor mechinorum''. The French femur is similar in general appearance to the ''Elopteryx'' type but it differs in diagnostic traits, e.g. lacking a fourth trochanter. Also, neither the ribs nor the tibiotarsi can be compared to the type specimen of ''Elopteryx'', there being no overlapping material. In 2005, by Kessler yet another (distal) femur piece, FGGUB R.1957, has been placed with ''Elopteryx'' on the basis of the bone texture.


Phylogeny

Modern interpretations have differed on the question whether the ''
Bradycneme ''Bradycneme'' (meaning "ponderous leg") is a genus of theropod dinosaur from the Maastrichtian-age Upper Cretaceous Sânpetru Formation of the Hațeg Basin, Transylvania, Romania. The genus contains a single species, ''Bradycneme draculae'', k ...
'' and '' Heptasteornis'' material should be included — they have meanwhile been synonymized and split from each other and ''Elopteryx'' many times — and what the exact affiliations of the material would be. Various solutions were proposed for this problem. During the eighties some researchers proposed ''Elopteryx'' were a member of the
Troodontidae Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil discov ...
, without being able to support this with much empirical evidence. In 1998 Csiki & Grigorescu suggested that ''Elopteryx'' belonged to the
Maniraptora Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to ''Ornithomimus velox''. It contains the major subgroups Avialae, Deinonychosauria, Oviraptoros ...
, while ''Bradycneme'' had a more basal position in 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 ...
. In 2004 ''Elopteryx'' was by
Darren Naish Darren William Naish 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 '' Eotyrannus'', '' Xenop ...
and
Gareth Dyke Gareth John Dyke is a paleontologist whose work is concerned with the evolutionary history of birds and their dinosaurian relatives. His specific research interests include the phylogenetics of birds, the functional morphology of aves and non-avia ...
considered a
eumaniraptora 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 avial ...
n ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
'', possibly either a non-
ornithuromorpha Euornithes (from Greek ' meaning "true birds") is a natural group which includes the most recent common ancestor of all avialans closer to modern birds than to ''Sinornis''. Description Clarke ''et al''. (2006) found that the most primitive know ...
n
pygostylia Pygostylia is a group of avialans which includes the Confuciusornithidae and all of the more advanced species, the Ornithothoraces. Definition The group Pygostylia was intended to encompass all avialans with a short, stubby tail, as opposed to ...
n bird or a
troodontid Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil disco ...
, while ''Bradycneme'' would be a maniraptor, and ''Heptasteornis'' (at least its holotype BMNH A4359) a member of the Alvarezsauridae. Thus ''E. nopcsai'' seems to be some sort of birdlike eumaniraptoran, but not related to modern birds. In 2005 Kessler however, reunited all the material in ''Elopteryx'' but considered it an alvarezsaurid. Later, in 2019, two studies have found it to be a bird once again, but a basal one; Hartman ''et al.'' recover it as a confuciusornithiform while Mayr ''et al.'' note similarities with ''
Gargantuavis ''Gargantuavis'' is an extinct genus of large, primitive bird containing the single species ''Gargantuavis philoinos''. It is the only member of the monotypic family Gargantuaviidae. Its fossils were discovered in several formations dating to 73 ...
'' and ''
Balaur A balaur ( pl. ''balauri'') in Romanian folklore is a type of many-headed dragon or monstrous serpent, sometimes said to be equipped with wings. The number of heads is usually around three, but they can also have seven heads or even twelve hea ...
'', suggesting they form a clade native to the Late Cretaceous European archipelago.


See also

*
Timeline of troodontid research This timeline of troodontid research is a chronological listing of events in the History of paleontology, history of paleontology focused on the troodontids, a group of bird-like theropod dinosaurs including animals like ''Troodon''. Troodontid re ...


References


Bibliography

* (1913): On some bird remains from the Upper Cretaceous of Transylvania. ''
Geological Magazine The ''Geological Magazine'' is a peer-reviewed scientific journal established in 1864, covering the earth sciences. It publishes original scientific research papers on geological topics. The journal is published bimonthly by Cambridge University ...
'' 5: 193–196. * (1963): Catalogue of fossil birds. Part 1 (Archaeopterygiformes through Ardeiformes). ''Bulletin of the Florida State Museum, Biological Sciences'' 7(4): 179–293
PDF fulltext
* (1978): Catalogue of fossil birds, Part 5 (Passeriformes). ''Bulletin of the Florida State Museum, Biological Sciences'' 23(3): 139–228. * (1998): Small theropods from the Late Cretaceous of the Hateg Basin (western Romania) - an unexpected diversity at the top of the food chain. '' Oryctos'' 1: 87-104. * (1975): The Bradycnemidae, a new family of owls from the Upper Cretaceous of Romania. '' Palaeontology'' 18(3): 563–570
PDF fulltext
* (1998): A new dromaeosaurid theropod from the Upper Cretaceous of Southern France. ''Oryctos'' 1: 105–112. * (1992): The first record of dromaeosaurid dinosaurs (Saurischia, Theropoda) in the Maastrichtian of southern Europe: palaeobiogeographical implications. ''Bulletin de la Société géologique de la France'' 163(3): 337–343. * (2008): A skull of the giant bony-toothed bird ''Dasornis'' (Aves: Pelagornithidae) from the Lower Eocene of the Isle of Sheppey. '' Palaeontology'' 51(5): 1107–1116. (HTML abstract) * (2004): ''Heptasteornis'' was no ornithomimid, troodontid, dromaeosaurid or owl: the first alvarezsaurid (Dinosauria: Theropoda) from Europe. ''Neues Jahrbuch für Geologie und Paläontologie - Monatshefte'' 7: 385–401. * (1988): ''Predatory Dinosaurs of the World.'' New York, Simon & Schuster. * (1991): The dinosaurs of Transylvania. ''National Geographic Research and Exploration'' 7(2): 196–215
PDF fulltext
{{Taxonbar, from=Q2344524 Prehistoric maniraptorans Maastrichtian life Late Cretaceous dinosaurs of Europe Cretaceous Romania Fossils of Romania Hațeg fauna Fossils of Serbia Fossil taxa described in 1913 Taxa named by Charles William Andrews Nomina dubia