Ferrodraco
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''Ferrodraco'' ("Iron Dragon" after the
ironstone Ironstone is a sedimentary rock, either deposited directly as a ferruginous sediment or created by chemical replacement, that contains a substantial proportion of an iron ore compound from which iron (Fe) can be smelted commercially. Not to be con ...
the fossil was found in) is an extinct
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
anhanguerid Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a late ...
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...
known 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'', ...
Winton Formation The Winton Formation is a Cretaceous geological formation in central-western Queensland, Australia. It is late Albian to early Turonian in age. The formation blankets large areas of central-western Queensland. It consists of sedimentary rocks suc ...
of
Queensland ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , establishe ...
, Australia, containing the single species ''F. lentoni''. The species was named after the former mayor of Winton, Graham Thomas ‘Butch’ Lenton. It is the most complete pterosaur fossil from Australia, being known from the holotype specimen AODF 876, consisting primarily of the anterior portion of the skull and dentary, cervical vertebral centra and a partial wing. Its wingspan was estimated to be about . ''Ferrodraco'' was found to have been within the subfamily Ornithocheirinae, as sister taxon to '' Mythunga''. A recent study also recovered ''Ferrodraco'' as sister taxon to ''Mythunga'', but both placed within the family
Anhangueridae Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a later ...
, more specifically within the subfamily Tropeognathinae. ''Ferrodraco'' is also the latest surviving member of
Anhangueria Anhangueria (or anhanguerians) is a group of pterosaurs belonging to the clade Pteranodontoidea. Fossil remains of this group date back from the Early to Late Cretaceous periods (Valanginian to Turonian stages), around 140 to 92.5 million years ...
.


Discovery and naming

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 sever ...
specimen was initially discovered in April 2017 when cattle farmer Robert A. Elliott was spraying herbicide near Belmont Station. It was excavated by a team led by Adele H. Pentland. Nicknamed 'Butch', it was further prepared by volunteer Ali Calvey. In 2019, 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 ...
''Ferrodraco lentoni'' was named and described by Adele H. Pentland, Stephen Francis Poropat, Travis R. Tischler, Trish Sloan, Robert A. Elliott, Harry A. Elliott, Judy A. Elliott and David A. Elliott. It was subsequently X-ray CT scanned at high-resolution using the Imaging and Medical Beamline at the Australian Synchrotron and osteology described in detail. The generic name is derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
''ferrum'', "iron", and ''draco'', "dragon", in reference to the fact that the skeleton was found in
ironstone Ironstone is a sedimentary rock, either deposited directly as a ferruginous sediment or created by chemical replacement, that contains a substantial proportion of an iron ore compound from which iron (Fe) can be smelted commercially. Not to be con ...
. The specific name honors the late mayor of Winton Shire, Graham Thomas ‘Butch’ Lenton, for his work for the local community and his support for the
Australian Age of Dinosaurs Australian Age of Dinosaurs Ltd (AAOD) is a not for profit organisation located in Winton, Queensland and founded by David Elliott and Judy Elliott in 2002. The organisation's activities include operation of the Australian Age of Dinosaurs Museu ...
. He died in 2017. The holotype, AODF 876, was found in a layer of the
Winton Formation The Winton Formation is a Cretaceous geological formation in central-western Queensland, Australia. It is late Albian to early Turonian in age. The formation blankets large areas of central-western Queensland. It consists of sedimentary rocks suc ...
dating from the Cenomanian - lower
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded b ...
, about ninety-six million years old. It consists of a partial skeleton with skull and lower jaws. It contains the front part of the head with the premaxillae, the maxillae and the dentaries; the left
frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, pa ...
, the rear part of the left lower jaw; forty single teeth; five neck vertebrae; the right shoulder joint; the left ulna; the left radius; the proximal and distal left wrist bones; two fourth metacarpals; phalanges from the first to third fingers of the left hand; and the first phalanx of the fourth finger. It represents a fully-grown but not yet mature animal. The skeleton has largely been preserved three-dimensionally due to the ironstone, but some bones however, have been crushed. It was probably fossilized in articulation but got some time prior to the discovery dispersed by erosion and cattle. ''Ferrodraco'' is the only pterosaur fossil known from the Winton Formation, and is the most complete pterosaur ever found in Australia, a continent where such finds are rare.


Description


Size and distinguishing traits

The
wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the other wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingspan o ...
of ''Ferrodraco'' was estimated at . The describing authors indicated two
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
(distinguishing traits) that ''Ferrodraco'' had. The first tooth pair in both the premaxillae of the snout and in the front lower jaws is smaller than the other front teeth. The fourth to seventh tooth pairs are smaller than the third and eight pair. Additionally, a unique combination is present of traits that in themselves are not unique. The front edge of the premaxilla is flattened and triangular. The first tooth pair in the premaxillae is directed vertically and is slightly set-off to above from the jawline. The front parts of the upper and lower jaws are not expanded sideways. The rear teeth are directed vertically, gradually declining in size. The tooth sockets are swollen relative to the outer wall of the jaw bones. The snout bears a premaxillary crest, the front edge of which continues the line of the snout tip, steeply rising under an angle of 60 degrees, and ending in a rounded crest top.


Skeleton

The length of the skull has been estimated to be about . The snout bears a relatively high crest, and it probably had a triangular profile in side view; the rear edge has not been preserved. The crest is very thin transversely however, only up to thick. It is hollow inside, the smooth bone walls being connected by small struts. The crest has a base length of and a height of . The symphysis of the lower jaws, their front fused area, probably extended to below in a second crest. There is an estimated total of twelve teeth in the upper jaw and thirteen teeth in the lower jaw for a total of fifty in the head as a whole. The teeth are formed as conical spikes with an oval cross-section, transversely flattened. The swollen tooth sockets cause an undulating profile of the jawlines in top view.


Classification

''Ferrodraco'' was the youngest known member of the clade
Anhangueria Anhangueria (or anhanguerians) is a group of pterosaurs belonging to the clade Pteranodontoidea. Fossil remains of this group date back from the Early to Late Cretaceous periods (Valanginian to Turonian stages), around 140 to 92.5 million years ...
, and proves that the clade did not become extinct during 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'', ...
. In 2010 however, an ornithocheirid jaw fragment that includes two teeth (specimen WAM 68.5.11) had been reported from Australia, the same place where ''Ferrodraco'' was found, and this specimen was unearthed in the Molecap Greensand, layers that possibly have an even younger age than ''Ferrodraco''.Kear, B.P., Deacon, G.L. & Siverson, M. 2010. "Remains of a Late Cretaceous pterosaur from the Molecap Greensand of Western Australia". ''Alcheringa'' 34: 273–279 The 2019 study of ''Ferrodraco'' made by Pentland ''et al.'' contained the results of two
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analyses, trying to determine the position of ''Ferrodraco'' in the evolutionary tree. The first analysis found ''Ferrodraco'' within the clade
Ornithocheirae Ornithocheirae is an extinct clade of pteranodontoid pterosaurs from the Early Cretaceous to the Late Cretaceous (Valanginian to Turonian stages) of Asia, Europe, North America and South America. It was named by Harry Seeley in 1870 as a family ...
, more precisely within the subfamily Ornithocheirinae as the
sister species 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 ...
of '' Mythunga'', another Australian pterosaur from somewhat older layers. The second analysis placed ''Ferrodraco'' as a basal member of the Anhangueria, and sister taxon to the
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 ...
that comprises ''
Anhanguera Anhanguera may refer to: People * Bartolomeu Bueno da Silva (1672–1740), a bandeirante Places in Brazil * Anhanguera, Goiás, a municipality in the state of Goiás * Anhanguera (district of São Paulo), a district in São Paulo * Parque Anhan ...
'', ''
Coloborhynchus ''Coloborhynchus'' is a genus of pterodactyloid pterosaur belonging to the family Anhangueridae, though it has also been recovered as a member of the Ornithocheiridae in some studies. ''Coloborhynchus'' is known from the Lower Cretaceous of Engl ...
'' and ''
Ornithocheirus ''Ornithocheirus'' (from Ancient Greek "ὄρνις", meaning ''bird'', and "χεῖρ", meaning ''hand'') is a pterosaur genus known from fragmentary fossil remains uncovered from sediments in the UK and possibly Morocco. Several species have ...
''. A more recent study made in 2020 by Borja Holgado and Rodrigo Pêgas placed ''Ferrodraco'' within the family
Anhangueridae Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a later ...
, more specifically within the subfamily Tropeognathinae, although still the sister taxon to ''Mythunga''. In 2022, Pentland et al. published a detailed description of ''Ferrodraco'' (particularly the post-cranial skeleton) and a renewed phylogenetic appraisal that unequivocally demonstrated that it belongs in the family Anhangueridae as proposed by Holgado and Pêgas. This study suggested that the precise position of ''Ferrodraco'' and ''Mythunga'' within Anhangueridae still remains uncertain, and that the diversity of Australian pterosaur fauna has been greatly underestimated. Topology 1: First analysis by Pentland ''et al.'' (2019). Topology 2: Holgado & Pêgas (2020).


See also

*
Timeline of pterosaur research This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic era. Although pterosaurs w ...


References

{{Portal bar, Paleontology, Cretaceous, Australia Fossil taxa described in 2019 Pterosaurs of Australia Pteranodontoids