Pteranodontoidea (or pteranodontoids, from
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 ...
meaning "toothless wings") is an
extinct
Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
of
ornithocheiroid
Ornithocheiroidea (or ornithocheiroids) is a group of pterosaurs within the extinct suborder Pterodactyloidea. They were typically large pterosaurs that lived from the Early to Late Cretaceous periods (Valanginian to Maastrichtian stages), with fo ...
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 to ...
s from the
Early
Early may refer to:
History
* The beginning or oldest part of a defined historical period, as opposed to middle or late periods, e.g.:
** Early Christianity
** Early modern Europe
Places in the United States
* Early, Iowa
* Early, Texas
* Early ...
to
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 ...
(early
Valanginian
In the geologic timescale, the Valanginian is an age or stage of the Early or Lower Cretaceous. It spans between 139.8 ± 3.0 Ma and 132.9 ± 2.0 Ma (million years ago). The Valanginian Stage succeeds the Berriasian Stage of the Lower Cretaceou ...
to 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 from ...
stages) of
Asia
Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
,
Africa
Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
,
Europe
Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
,
North America
North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
and
South America
South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
.
It was named by
Alexander Wilhelm Armin Kellner in
1996
File:1996 Events Collage.png, From left, clockwise: A Centennial Olympic Park bombing, bomb explodes at Centennial Olympic Park in Atlanta, set off by a radical Anti-abortion violence, anti-abortionist; The center fuel tank explodes on TWA Flight 8 ...
. In 2003, Kellner defined the clade as a
node-based taxon
Phylogenetic nomenclature is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional approach, in which taxon names are defined by a '' type'', which ...
consisting of the last common ancestor of ''
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 Anhangue ...
'', ''
Pteranodon
''Pteranodon'' (); from Ancient Greek (''pteron'', "wing") and (''anodon'', "toothless") is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of . They lived during the late Cr ...
'' and all its descendants. The clade Ornithocheiroidea is sometimes considered to be the
senior synonym
The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently.
* In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linnae ...
of Pteranodontoidea, however it depends on its definition.
[Unwin, D. M., (2003): On the phylogeny and evolutionary history of pterosaurs. pp. 139-190. — ''in'' Buffetaut, E. & Mazin, J.-M., (eds.): ''Evolution and Palaeobiology of Pterosaurs''. Geological Society of London, Special Publications 217, London, 1-347] Brian Andres (2008, 2010, 2014) in his analyses, converts Ornithocheiroidea using the definition of Kellner (2003) to avoid this synonymy.
[A preview that shows the cladogram without clade names]
/ref>
Despite the meaning of the group's name, not all pteranodontoids were toothless, the earlier genera such as the istiodactylid
Istiodactylidae is a small family of pterosaurs. This family was named in 2001 after the type genus ''Istiodactylus'' was discovered not to be a member of the genus ''Ornithodesmus''.
Systematics and distribution
Remains of taxa that can be co ...
s and 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 later ...
s had teeth that they used to catch prey, and several conclusions had led to the idea that they later evolved into the toothless pteranodontid
The Pteranodontidae are a Family (biology), family of large pterosaurs of the Cretaceous Period of North America and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a distinctive, elongated crest jutting from the ...
s. Pteranodontoids were also the most diverse group of pterosaurs throughout the 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 th ...
period, and therefore the most successful flyers during their time. They were mostly replaced by the larger azhdarchid
Azhdarchidae (from the Persian word , , a dragon-like creature in Persian mythology) is a family of pterosaurs known primarily from the Late Cretaceous Period, though an isolated vertebra apparently from an azhdarchid is known from the Early Cret ...
s at the end of the Late Cretaceous however, which resulted in resource and terrain competitions, but even so, pteranodontoids still managed to thrive with them, and genera such as ''Alcione
Alcione may refer to:
* ''Alcione'', a 1638 literary work by Pierre du Ryer
* ''Alcione'', a 1787 musical drama by João de Sousa Carvalho
* '' CANT Z.1007 Alcione'', World War II Italian bomber aircraft
* ''Alcione'' (opera), a 1706 opera by ...
'', ''Barbaridactylus
''Barbaridactylus'' is a Nyctosauridae, nyctosaurid pterosaur from the Ouled Abdoun Basin of Morocco, a basin that dates back to the Maastrichtian stage of the Late Cretaceous. It was published in 2018 in paleontology, 2018 by paleontologists Nic ...
'', and ''Simurghia
''Simurghia'' is a genus Nyctosauridae, nyctosaurid pterosaur from the Ouled Abdoun Basin of Morocco, a basin that dates to the Maastrichtian stage of the Late Cretaceous period, about 66 million years ago. It was published in 2018 in paleontolog ...
'' lived until the Cretaceous-Paleogene extinction event. Pteranodontoids had similar a similar diet to modern-day soaring bird
This is a list of Lift (soaring), soaring birds, which are birds that can maintain flight without wing flapping, using rising air currents. Many gliding birds are able to "lock" their extended wings by means of a specialized tendon.
;Bird of prey ...
s such the albatross
Albatrosses, of the biological family Diomedeidae, are large seabirds related to the procellariids, storm petrels, and diving petrels in the order Procellariiformes (the tubenoses). They range widely in the Southern Ocean and the North Pacifi ...
, which mainly consisted of fish. Some genera of this group however, fed mainly on carrion
Carrion () is the decaying flesh of dead animals, including human flesh.
Overview
Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures, c ...
, this can be seen on the earlier genera such as ''Istiodactylus
''Istiodactylus'' is a genus of pterosaur that lived during the Early Cretaceous period, about 120 million years ago. The first fossil was discovered on the English Isle of Wight in 1887, and in 1901 became the holotype specimen of a new species ...
''. They were also known for their proportionally large 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 of ...
s, the largest recorded of any pteranodontoid reached , and it belongs to the genus ''Tropeognathus
''Tropeognathus'' (meaning "keel jaw") is a genus of large pterosaurs from the late Early Cretaceous of South America. This genus is considered to be a member of the family Anhangueridae, however, several studies have also recovered it within a ...
''.
History of early discovered genera
Several fossil remains were found in the Chalk Formation
The Chalk Group (often just called the Chalk) is the lithostratigraphic unit (a certain number of rock strata) which contains the Upper Cretaceous limestone succession in southern and eastern England. The same or similar rock sequences occur acro ...
of England
England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe b ...
, a chalk pit known for its wide variety of fossils belonging to different animals. British
British may refer to:
Peoples, culture, and language
* British people, nationals or natives of the United Kingdom, British Overseas Territories, and Crown Dependencies.
** Britishness, the British identity and common culture
* British English, ...
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 ...
James Scott Bowerbank
James Scott Bowerbank (14 July 1797 – 8 March 1877) was a British naturalist and palaeontologist.
Biography
Bowerbank was born in Bishopsgate, London, and succeeded in conjunction with his brother to his father's distillery, in which he was ...
named and described the specimens found as a new species of ''Pterodactylus'', ''P. giganteus'' due to its much larger size. ''P. giganteus'' is nowadays considered to belong to the genus called ''Lonchodraco
''Lonchodraco'' is a genus of lonchodraconid Pterodactyloidea, pterodactyloid pterosaur from the Late Cretaceous of southern England. The genus includes species that were previously assigned to other genera.
Discovery and naming
In 1846, Jame ...
''. In 1851, Bowerbank unearthed more specimens in the same chalk pit as ''P. giganteus'', and assigned these specimens to yet another newly named species of ''Pterodactylus'', known as ''P. cuvieri'', in honor of the German naturalist and zoologist
Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the Animal, animal kingdom, including the anatomy, structure, embryology, evolution, Biological clas ...
Georges Cuvier
Jean Léopold Nicolas Frédéric, Baron Cuvier (; 23 August 1769 – 13 May 1832), known as Georges Cuvier, was a French natural history, naturalist and zoology, zoologist, sometimes referred to as the "founding father of paleontology". Cuvier ...
. Later that year, British paleontologist Sir Richard Owen
Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils.
Owe ...
had also unearthed several fossil specimens in the Chalk Formation, in which he assigned them as ''Pterodactylus compressirostris'' during its description.[Owen, R. (1851). Monograph on the fossil Reptilia of the Cretaceous Formations. ''The Palaeontographical Society'' 5(11):1-118.] Owen later unearthed many fossil specimens from a different fossil site called the Cambridge Greensand
The Cambridge Greensand is a geological unit in England whose strata are earliest Cenomanian in age. It lies above the erosive contact between the Gault Formation and the Chalk Group in the vicinity of Cambridgeshire, and technically forms the low ...
, he then assigned these specimens into a new species called ''Pterodactylus simus''. With the naming of this new species, British paleontologist Harry Govier Seeley
Harry Govier Seeley (18 February 1839 – 8 January 1909) was a British paleontologist.
Early life
Seeley was born in London on 18 February 1839, the second son of Richard Hovill Seeley, a goldsmith, and his second wife Mary Govier. When his fat ...
created a new separate genus called ''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 ...
'' (from Ancient Greek "ὄρνις", meaning "bird", and "χεῖρ", meaning "hand"), due to the notion of the time, in which pterosaurs were the direct ancestors of birds. In 1870, Seeley then also reassigned ''Pterodactylus cuvieri'' into a new species called ''Ornithocheirus cuvieri''.[Seeley, H.G. (1870). ''The Ornithosauria: an Elementary Study of the Bones of Pterodactyles''. Cambridge, 130 pp.] This species is now considered to belong to the genus ''Cimoliopterus
''Cimoliopterus'' is a genus of pterosaur that lived during the Late Cretaceous in what is now England and the United States. The first known specimen, consisting of the front part of a snout including part of a crest, was discovered in the G ...
''. Back in 1874 however, controversies between the naming of species and genera of these pterosaurs started, with the paleontologists Harry Seeley and Richard Owen fighting over reassignings of different species as well as genera. Owen therefore created two new genera: ''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 Engla ...
'' (meaning "maimed beak") and ''Criorhynchus
''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 ...
'' (meaning "ram beak"), in reference to their unique convex "keeled" crests on top, and underside of their snout. Owen then created a type species for ''Coloborhynchus'', ''C. clavirostris'', and sunk three species which were assigned previously to ''Ornithocheirus'', and ''Criorhynchus'' consisted entirely of former ''Ornithocheirus'' species.[Owen, R. 1874, ''Monograph on the fossil Reptilia of the Mesozoic Formations''. Palaeontographical Society, London, 14 pp] Later in 1881 however, Seeley disagreed with Owen's conclusions, and he therefore assigned the species he named ''Ornithocheirus simus'' as the type species of ''Ornithocheirus''.
The naming of the different pterosaur species, as well as the genus ''Ornithocheirus'' in England resulted in further research in other places. In the United States
The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
, American paleontologist Othniel Charles Marsh
Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of Paleontology in Yale College and President of the National Academy of Sciences. He was one of the preeminent scientists in the field of paleontology. Among h ...
led an expedition to the Smoky Hill Chalk
The Smoky Hill Chalk Member of the Niobrara Chalk formation is a Cretaceous conservation Lagerstätte, or fossil rich geological formation, known primarily for its exceptionally well-preserved marine reptiles. Named for the Smoky Hill River, the S ...
deposits in western Kansas
Kansas () is a state in the Midwestern United States. Its capital is Topeka, and its largest city is Wichita. Kansas is a landlocked state bordered by Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to the ...
in 1870. Marsh unearthed the first ever specimens of pterosaur from North America
North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
(specimens YPM 1160 and YPM 1161), which now belong to the genus ''Pteranodon
''Pteranodon'' (); from Ancient Greek (''pteron'', "wing") and (''anodon'', "toothless") is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of . They lived during the late Cr ...
'', and consisted of partial wing remains and a tooth of the fish ''Xiphactinus
''Xiphactinus'' (from Latin and Greek for " sword-ray") is an extinct genus of large (Shimada, Kenshu, and Michael J. Everhart. "Shark-bitten Xiphactinus audax (Teleostei: Ichthyodectiformes) from the Niobrara Chalk (Upper Cretaceous) of Kansas. ...
'', which Marsh had once believed that it belonged to ''Pteranodon''. In 1871, he assigned these specimens to a new species called ''"Pterodactylus oweni"'' (meaning "Owen's wing finger"),[Marsh, O. C., 1871.
"Note on a new and gigantic species of Pterodactyle."
American Journal of Science, ser. 3, 1(6):472 ] in honor of Sir Richard Owen, but he realized that the name he had chosen had already been used for a different European pterosaur species described by Harry Seeley, so he renamed his discovery as ''Pterodactylus occidentalis'' (meaning "Western wing finger"), in reference to the location where it was found.[Marsh, O. C. 1872. Discovery of additional remains of Pterosauria, with descriptions of two new species. ''American Journal of Science'', Series 3, 3(16):241–248.] American paleontologist Edward Drinker Cope
Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontologist, comparative anatomist, herpetologist, and ichthyologist. Born to a wealthy Quaker family, Cope distinguished himself as a child prodigy interested ...
had also unearthed several remains of the large North American pterosaur, and based on these remains, Cope named two new species, ''Ornithochirus umbrosus'' and ''Ornithochirus harpyia'', in an attempt to assign them to the large European genus ''Ornithocheirus'', but he had missed the 'e' when describing them. The paper naming these species by Cope's paper naming had been published in 1872, just five days after the publication of Marsh's paper. This led to a dispute between the two paleontologists, which both fought over whose names had priority to the published literature, in what was obviously the same species of pterosaur.
In 1876, Marsh had described the first skull specimen of ''Pteranodon'', which as unearthed by American paleontologist Samuel Wendell Williston
Samuel Wendell Williston (July 10, 1852 – August 30, 1918) was an American educator, entomologist, and paleontologist who was the first to propose that birds developed flight cursorially (by running), rather than arboreally (by leaping from tr ...
in the Smoky Hill River
The Smoky Hill River is a river in the central Great Plains of North America, running through Colorado and Kansas.Smoky Hill River. (2009). In Encyclopædia Britannica. Retrieved July 22, 2009, from Encyclopædia Britannica Online: http://www.br ...
, located in the Wallace County (now Logan County) of Kansas, USA. Later that year, another skull specimen was found, though this time a bit smaller in size.[ Soon enough, Marsh recognized a major difference when he described these specimens, they lacked teeth and had bony crests on their skulls. He then described the specimens found as "distinguished from all previously known genera of the order Pterosauria by the entire absence of teeth." Marsh then coined the name ''Pteranodon'' (meaning "wing without tooth"), and pointed that the unique characteristics identified warranted a new generic name. Marsh had also reclassified all the previously named North American species from ''Pterodactylus'' to ''Pteranodon'', and considered the smaller skull to belong to the reassigned ''Pteranodon occidentalis'' based on its size.][Bennett, S.C. (2000). "Inferring stratigraphic position of fossil vertebrates from the Niobrara Chalk of western Kansas." ''Current Research in Earth Sciences: Kansas Geological Survey Bulletin'', 244(Part 1): 26 pp.] Specimens of smaller individuals were later uncovered, and Marsh had also named several additional species: ''Pteranodon comptus'' and ''Pteranodon nanus
''Nyctosaurus'' (meaning "night lizard" or "bat lizard") is a genus of nyctosaurid pterosaur from the Late Cretaceous period of what is now the Niobrara Formation of the mid-western United States, which, during the time ''Nyctosaurus'' was alive ...
'', based on these distinct and smaller fragmentary skeletons. He then created another separate species called ''Pteranodon gracilis'', which is now based on a wing bone that he previously mistook for a pelvic bone
The hip bone (os coxae, innominate bone, pelvic bone or coxal bone) is a large flat bone, constricted in the center and expanded above and below. In some vertebrates (including humans before puberty) it is composed of three parts: the ilium, ischi ...
. Marsh soon realized his mistake, and therefore reassigned that specimen he found into a separate genus which he named ''Nyctosaurus
''Nyctosaurus'' (meaning "night lizard" or "bat lizard") is a genus of nyctosaurid pterosaur from the Late Cretaceous period of what is now the Niobrara Formation of the mid-western United States, which, during the time ''Nyctosaurus'' was alive ...
'', and the species ''P. nanus'' was soon recognized as a ''Nyctosaurus'' specimen as well, this is due to the synapomorphies
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
seen in both.
[Witton, M. (2010). ''Pteranodon'' and beyond: The history of giant pterosaurs from 1870 onwards. Geological Society of London, Special Publications. 343. 313-323. ] Later in 1892, Samuel Williston had become the first scientist to critically evaluate all of the ''Pteranodon'' species classified by Cope and Marsh. He agreed with most of Marsh's reassignings and classifications, but several exceptions were still made. Williston revised his earlier conclusion in 1903, in which he stated that there were only three species of ''Pteranodon'', instead of seven. He also considered both ''P. comptus'' and ''P. nanus'' to be specimens of ''Nyctosaurus'', and divided the other species into small (''P. velox''), medium (''P. occidentalis''), and large species (''P. ingens''), based primarily on the shape of their upper arm bones. He also followed his first conclusion of ''P. longiceps'' being a synonym of either ''P. velox'' or ''P. occidentalis'' based on its size.[Williston, S. W. 1903. On the osteology of ''Nyctosaurus'' (''Nyctodactylus''), with notes on American pterosaurs. Field Mus. Publ. (Geological Ser.) 2(3):125–163, 2 figs., pls. XL-XLIV.] However, in 1994, Christopher Bennett concluded that ''P. longiceps'' and another species called '' P. sternbergi'' would be the only valid ''Pteranodon'' species, while the other ones were either considered ''nomina dubia
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 ...
'' or synonyms.[
A peculiar discovery was made in 1887 by Seeley in the ]Isle of Wight
The Isle of Wight ( ) is a county in the English Channel, off the coast of Hampshire, from which it is separated by the Solent. It is the largest and second-most populous island of England. Referred to as 'The Island' by residents, the Isle of ...
, an island off the coast of southern England. Seeley thought that these fossil remains belonged to bird-like creature, which he then named ''Ornithodesmus cluniculus
''Ornithodesmus'' (meaning "bird link") is a genus of small, dromaeosaurid dinosaur from the Isle of Wight in England, dating to about 125 million years ago. The name was originally assigned to a bird-like sacrum (a series of vertebrae fused to ...
''. Seeley also reported another specimen found on the same site. Seeley named the new species ''O. latidens'' in 1901 based on this discovery, the specific name translates to "wide tooth". He presumably assigned this new species to ''Ornithodesmus'' due to synapomorphies between their sacra, but his conclusion was not entirely clear. In 1913, English paleontologist Reginald Walter Hooley
Reginald Walter Hooley (5 September 1865 – 5 May 1923) was a businessman and amateur paleontologist, collecting on the Isle of Wight. He is probably best remembered for describing the dinosaur ''Iguanodon atherfieldensis'', now ''Mantellisaurus ...
reviewed ''O. latidens'' in detail, based on specimens he had found, and placed the genus ''Ornithodesmus'' within a newly created family, Ornithodesmidae. Later in 1993, the British paleontologists Stafford C. Howse and Andrew C. Milner concluded that the holotype sacrum and only specimen of the species ''O. cluniculus'' didn't belong to a pterosaur, but instead to a 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. They pointed out that no detailed attempts had been made to compare the sacrum of ''O. cluniculus'' with those of pterosaurs, and that the species ''O. latidens'' had in effect been treated as the type species of the genus ''Ornithodesmus'' in their conclusion. Now as a definite species of pterosaur, "''O.''" ''latidens'' thus required a new generic name. In 2001, Howse, Milner, and David Martill, reassigned "''O.''" ''latidens'' to a new genus called ''Istiodactylus
''Istiodactylus'' is a genus of pterosaur that lived during the Early Cretaceous period, about 120 million years ago. The first fossil was discovered on the English Isle of Wight in 1887, and in 1901 became the holotype specimen of a new species ...
'', resulting in a new combination known as ''Istiodactylus latidens''. The generic name ''Istiodactylus'' means "sail finger", in reference to its proportionally large wings.
Classification
Originally defined by Alexander Kellner
Alexander Wilhelm Armin Kellner (born September 26, 1961) is a Brazilian geologist and paleontologist who is a leading expert in the field of studying pterosaurs. His research has focused mainly on fossil reptiles from the Cretaceous Period, i ...
in 1996, Pteranodontoidea is sometimes considered a synonym of the clade Ornithocheiroidea according to several studies. The name Ornithocheiroidea was originally defined as an apomorphy
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
-based taxon by Christopher Bennett in 1994. In 2003 however, Kellner redefined Ornithocheiroidea to represent the node of ''Anhanguera'', ''Pteranodon'', ''Quetzalcoatlus
''Quetzalcoatlus'' is a genus of pterosaur known from the Late Cretaceous period of North America (Maastrichtian stage); its members were among the largest known flying animals of all time. ''Quetzalcoatlus'' is a member of the Azhdarchidae, ...
'' and ''Dsungaripterus
''Dsungaripterus'' is a genus of dsungaripterid pterosaur with an average wingspan of . ''Dsungaripterus'' lived during the Early Cretaceous in what is now China, and its first fossil was found in the Junggar Basin.
Description
''Dsungaripter ...
'', thus making Ornithocheiroidea a more inclusive group.[Kellner, A. W. A., (2003): Pterosaur phylogeny and comments on the evolutionary history of the group. pp. 105-137. — ''in'' Buffetaut, E. & Mazin, J.-M., (eds.): ''Evolution and Palaeobiology of Pterosaurs''. Geological Society of London, Special Publications 217, London, 1-347] Later that year, David Unwin suggested a different definition, the node that contains ''Pteranodon longiceps
''Pteranodon'' (); from Ancient Greek (''pteron'', "wing") and (''anodon'', "toothless") is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of . They lived during the late Cr ...
'' and ''Istiodactylus latidens
''Istiodactylus'' is a genus of pterosaur that lived during the Early Cretaceous period, about 120 million years ago. The first fossil was discovered on the English Isle of Wight in 1887, and in 1901 became the holotype specimen of a new specie ...
'', as a resulting, he considered Pteranodontoidea a junior synonym
The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently.
* In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...
of Ornithocheiroidea.[ Brian Andres (2008, 2010, 2014) in his analyses, converts Ornithocheiroidea using the definition of Kellner (2003) to avoid this synonymy.][ In 2013, Andres & Timothy Myers presented a ]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 ...
analysis that placed Pteranodontoidea within the group Pteranodontia
Pteranodontia is an extinct group of ornithocheiroid pterodactyloid pterosaurs that lived during the Late Cretaceous period (Coniacian to Maastrichtian stages) of North America and Africa. They were some of the most advanced pterosaurs, and poss ...
, as the sister taxon of the family Nyctosauridae
Nyctosauridae (meaning "night lizards" or "bat lizards") is a family of specialized soaring pterosaurs of the late Cretaceous Period of North America, Africa, and possibly Europe. It was named in 1889 by Henry Alleyne Nicholson and Richard Lydek ...
.[ In 2019 however, Kellner and colleagues revisited the classification, and concluded that Pteranodontoidea, which was sister taxon to ]Tapejaroidea
Tapejaroidea (or tapejaroids) is a group of pterosaurs belonging to the clade Ornithocheiroidea. Tapejaroids lived from the Early to Late Cretaceous periods, with one possible member, ''Tendaguripterus'', extending the fossil range to the Late Ju ...
, would be the more inclusive group containing both the Lanceodontia
Ornithocheiromorpha (from Ancient Greek, meaning "bird hand form") is a group of pterosaurs within the suborder Pterodactyloidea. Fossil remains of this group date back from the Early to Late Cretaceous periods (Valanginian to Turonian stag ...
and the Pteranodontia.
Topology 1: Andres & Myers (2013).
Topology 2: Kellner and colleagues (2019).
Paleobiology
Locomotion
Pteranodontoids, like other pterosaurs, are considered to have been skilled fliers as well as adept at moving on the ground. Evidence from footprints shows that most pterosaurs did not sprawl their limbs to a large degree, as in modern reptiles, but rather held the limbs relatively erect when walking, like 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 ...
s. While no pteranodontoid footprints are known, it is likely that they also walked erect. Among pterosaurs, pteranodontoids had unusually uneven limb proportions, with the forelimbs much longer than the hind limbs. This would likely have required them to use unique modes of locomotion when on the ground compared to other pterosaurs. It is possible that pteranodontoids ran (but not walked) bipedally, or that they used a hopping gait.[ Pterosaur researcher Mike Habib has noted that the limbs proportions of pteranodontoids like '']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 Anhangue ...
'' are consistent with hopping.[Habib, M. (2011).]
Dinosaur Revolution: ''Anhanguera''.
''H2VP: Paleobiomechanics''. Weblog entry, 20-SEP-2011. Accessed 28-SEP-2011: http://h2vp.blogspot.com/2011/09/dinosaur-revolution-anhanguera.html
References
{{Taxonbar, from=Q2597187
Fossil taxa described in 1996
Barremian first appearances
Campanian extinctions