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''Elopteryx'' 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 paravian theropod
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 ...
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 preserve ...
s found in
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
rocks of
Romania Romania is a country located at the crossroads of Central Europe, Central, Eastern Europe, Eastern and Southeast Europe. It borders Ukraine to the north and east, Hungary to the west, Serbia to the southwest, Bulgaria to the south, Moldova to ...
. The single
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), ...
, ''Elopteryx nopcsai'', was known only from very incomplete material until new specimens were reported in the 21st century. '' Balaur bondoc'' might represent a
junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...
of this taxon.


History of discovery and naming


Initial finds

In the late nineteenth or early twentieth century, the famous Hungarian
paleontologist 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 ...
Franz Nopcsa von Felső-Szilvás found near Sînpetru, in what is now the Romanian region of
Transylvania Transylvania ( or ; ; or ; Transylvanian Saxon dialect, Transylvanian Saxon: ''Siweberjen'') is a List of historical regions of Central Europe, historical and cultural region in Central Europe, encompassing central Romania. To the east and ...
, some bone fragments of a small theropod. These were acquired by the British Museum of Natural History. In 1913, curator Charles William Andrews named these as 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 ...
''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 classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
''helos'' (ἕλος), "marsh" + ''pteryx'' (πτέρυξ), "wing". The specific name honors Nopcsa. Initially, ''Elopteryx'' was described from its
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 ...
, a proximal left
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
, 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 ...
was also tentatively assigned to this
taxon In biology, a taxon (back-formation from ''taxonomy''; : 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 name and ...
; 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 International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
( 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 conven ...
, circa 70-66
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 ...
, originating from the Sânpetru Formation of the Hațeg Island. The animal was by Andrews believed to be a pelecaniform
seabird Seabirds (also known as marine birds) are birds that are adaptation, adapted to life within the marine ecosystem, marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent ...
.Harrison & Walker (1975) In 1929 the Hungarian paleontologist Kálmán Lambrecht 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 that includes organisms with mixed evolutionary origin but does not include their most recent common ancestor. The term is often applied to groups that share similar features known as Homoplasy, homoplasies ...
" Pelecaniformes" – in 1963 by Pierce Brodkorb in his fossil bird catalogue, and the
Cenozoic The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...
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 '' Dasornis'' 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 gannets 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 removed from ''Elopteryx'', redescribed as '' Bradycneme draculae'' and '' Heptasteornis andrewsi'' respectively, and used to establish a supposed family of gigantic two metre tall owls, 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 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 ...
bones were not of birds but of non-avian dinosaurs. In 1981, Dan Grigorescu and Eugen Kessler stated that ''Elopteryx'' was a non-avian coelurosaurian dinosaur. They also referred a supposed distal femur (FGGUB R.351) to ''Elopteryx'', but both researchers (with Zoltan Csiki) later identified this specimen as a hadrosauroid distal metatarsal. In 1992, it was proposed by Jean Le Loeuff e.a. that ''Bradycneme'' and ''Heptasteornis'' should be synonymized with ''E. nopcsai'' again, and a femur (MDE-D203), an
anterior Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
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 ...
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 ...
(MDE-D01), a posterior sacral vertebra (MDE collection, unnumbered) and some dorsal rib fragments from the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
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 coelurosaurian theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from ...
or a family or
subfamily 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 zo ...
(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, another (distal) femur piece, FGGUB R.1957, has been described as a new specimen of ''Elopteryx'' on the basis of the bone texture. In 2019, a potential pelvis specimen identified as cf. ''E. nopcsai'' was reported. In 2024, a new femur specimen from
Romania Romania is a country located at the crossroads of Central Europe, Central, Eastern Europe, Eastern and Southeast Europe. It borders Ukraine to the north and east, Hungary to the west, Serbia to the southwest, Bulgaria to the south, Moldova to ...
was attributed to ''Elopteryx'', and the specimen shows that it was secondarily flightless.


Phylogeny

Modern interpretations have differed on the question whether the '' Bradycneme'' 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. Previously, some researchers proposed ''Elopteryx'' was a member of the
Dromaeosauridae Dromaeosauridae () is a family of feathered coelurosaurian theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from ...
or Troodontidae, without being able to support this with much empirical evidence. In 1998 Csiki & Grigorescu suggested that ''Elopteryx'' belonged to the Maniraptora, while ''Bradycneme'' had a more basal position in the Tetanurae. Since the 21st century, ''Elopteryx'' has been supported widely as a member of the Maniraptora, with most studies recovering it as a member of the Paraves. In 2004, Darren Naish and Gareth Dyke considered ''Elopteryx'' as a Eumaniraptora ''
incertae sedis or is a term used for a taxonomy (biology), 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, uncertainty ...
'', possibly either a non- ornithuromorphan pygostylian bird or a troodontid, while ''Bradycneme'' would be a maniraptoran, and the dubious ''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, Grigorescu and Csiki reunited all the material in ''Elopteryx'' but considered it an alvarezsaurid. In a 2011 classification, Tom Holtz assigned ''Elopteryx'' to the Troodontidae.Holtz, Thomas R. Jr. (2011) ''Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages,'
Winter 2010 Appendix.
/ref> In 2013, Stephen L. Brusatte and colleagues mentioned a possibility of ''Elopteryx'' and the paravian ''Balaur bondoc'' being the same taxon, though the authors did not consider it the most likely case. Later, in 2019, two studies have found it to be an avialan once again, but a basal one; Hartman and colleagues recover it as a confuciusornithiform while Mayr and colleagues note similarities with '' Gargantuavis'' and '' Balaur'', suggesting they form a clade native to the Late Cretaceous European archipelago. Mayr and colleagues also claimed that the synonymy of ''Elopteryx'' and ''Balaur'' remains possible and that more work is needed for confirmation. In 2024, Stoicescu and colleagues suggested that ''Elopteryx'' is indeed a paravian and a member of the
Avialae Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds, and their closest relatives. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to Deinonychosauria, deinonychosaurs, though ...
based on its new specimen, and that ''Balaur bondoc'' is probably a
junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...
of ''Elopteryx''.


See also

* Timeline of troodontid research


References


Bibliography

* (1913): On some bird remains from the Upper Cretaceous of Transylvania. '' Geological Magazine'' 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 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 geo ...
'' 18(3): 563–570. * (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 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 geo ...
'' 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 Paraves 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