Istiodactylus Sinensis
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''Istiodactylus'' 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
pterosaur Pterosaurs are 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 million to 66 million years ago). Pterosaurs are the earli ...
that lived during the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
period, about 120 million years ago. The first fossil was discovered on the English
Isle of Wight The Isle of Wight (Help:IPA/English, /waɪt/ Help:Pronunciation respelling key, ''WYTE'') is an island off the south coast of England which, together with its surrounding uninhabited islets and Skerry, skerries, is also a ceremonial county. T ...
in 1887, and in 1901 became the
holotype specimen A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was Species description, formally described. It is either the single such physical example (or illus ...
of a new species, ''O. latidens'' (Latin for "wide tooth"), in the genus '' Ornithodesmus''. This species was moved to its own genus, ''Istiodactylus'', in 2001; this name is Greek for "sail finger". More specimens were described in 1913, and ''Istiodactylus'' was the only pterosaur known from three-dimensionally preserved fossils for much of the 20th century. In 2006, a species from China, ''I. sinensis'', was assigned to ''Istiodactylus'', but it has also been suggested to belong to a different genus. ''Istiodactylus'' was a large pterosaur; estimates of its wingspan range from . Its skull was about long, and was relatively short and broad for a pterosaur. The front of the snout was low and blunt, and bore a semicircle of 48 teeth. The triangular teeth were closely spaced, interlocked, and formed a "razor-edged" outline. The lower jaw also had a tooth-like projection that occluded with the teeth. The skull had a very large naso-antorbital opening (which combined the
antorbital fenestra An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with Archosauriformes, archosauriforms, first appearing during the Triassic Period. Among Extant ...
and the opening for the bony nostril) and a slender eye socket. Some of the
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 ...
e were fused into a
notarium Notarium or os dorsale is a bone consisting of the fused vertebra of the shoulder in birds and some pterosaurs. The structure helps brace the chest against the forces generated by the wings. In birds, the vertebrae are only in contact with adjac ...
, to which the shoulder blades connected. It had very large forelimbs, with a wing-membrane distended by a long wing-finger, but the
hindlimb A hindlimb or back limb is one of the paired articulated appendages ( limbs) attached on the caudal ( posterior) end of a terrestrial tetrapod vertebrate's torso.http://www.merriam-webster.com/medical/hind%20limb, Merriam Webster Dictionary-H ...
s were very short. Until the 21st century, ''Istiodactylus'' was the only known pterosaur of its kind, and was placed in its own
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
,
Istiodactylidae Istiodactylidae is a small Family (biology), 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 t ...
, within the group
Ornithocheiroidea Pterodactyloidea ( ; derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", and ''δάκτυλος'' (''dáktylos'') "finger") is one of the two traditional suborders of pterosaurs ("wing lizards"), and contai ...
. ''Istiodactylus'' differed from other istiodactylids in having a proportionally shorter skull. The distinctive teeth of ''Istiodactylus'' indicate that it was a
scavenger Scavengers are animals that consume Corpse decomposition, dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a he ...
that may have used its teeth to sever morsels from large carcasses in the manner of a
cookie cutter A cookie cutter in North American English, also known as a biscuit cutter outside North America, is a tool to cut out cookie/biscuit dough in a particular shape. They are often used for seasonal occasions when well-known decorative shapes are ...
. The wings of ''Istiodactylus'' may have been adapted for soaring, which would have helped it find carcasses before
terrestrial Terrestrial refers to things related to land or the planet Earth, as opposed to extraterrestrial. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on o ...
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...
s. ''Istiodactylus'' is known from the
Wessex Formation The Wessex Formation is a fossil-rich England, English geological formation that dates from the Berriasian to Barremian Stage (stratigraphy), stages of the Early Cretaceous. It forms part of the Wealden Group and underlies the younger Vectis Form ...
and the younger
Vectis Formation The Vectis Formation is a geological Formation (geology), formation on the Isle of Wight and Swanage, England whose strata were formed in the Barremian, approximately 125 million years ago."Magnetostratigraphy of the Lower Cretaceous Vectis Forma ...
, which represent river and coastal environments that were shared with various pterosaurs,
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 ...
s, and other animals.


History of discovery

In 1887, the British
palaeontologist 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 ...
Harry G. 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 fa ...
described a fossil
synsacrum The synsacrum is a skeletal structure of birds and other dinosaurs, pterosaurs, as well as xenarthran mammals, in which the sacrum is extended by incorporation of additional fused or partially fused caudal or lumbar In tetrapod anatomy, lum ...
(fused vertebrae attached to the pelvis) from the
Wessex Formation The Wessex Formation is a fossil-rich England, English geological formation that dates from the Berriasian to Barremian Stage (stratigraphy), stages of the Early Cretaceous. It forms part of the Wealden Group and underlies the younger Vectis Form ...
of the
Isle of Wight The Isle of Wight (Help:IPA/English, /waɪt/ Help:Pronunciation respelling key, ''WYTE'') is an island off the south coast of England which, together with its surrounding uninhabited islets and Skerry, skerries, is also a ceremonial county. T ...
, an island off the coast of southern England. Though he compared it with those of
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 ...
s and
pterosaurs Pterosaurs are an extinct clade of flying reptiles in the Order (biology), order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosau ...
, he concluded that it belonged to a bird (though more dinosaur-like than any known bird), which he named '' Ornithodesmus cluniculus''. The British
geologist A geologist is a scientist who studies the structure, composition, and History of Earth, history of Earth. Geologists incorporate techniques from physics, chemistry, biology, mathematics, and geography to perform research in the Field research, ...
John W. Hulke suggested later that year that ''Ornithodesmus'' was a pterosaur, finding it similar to fossils that he had seen before, but Seeley disagreed. In his 1901 ''Dragons of the Air'', the first popular book about pterosaurs, Seeley reported another specimen (NHMUK R176 at the
Natural History Museum A natural history museum or museum of natural history is a scientific institution with natural history scientific collection, collections that include current and historical records of animals, plants, Fungus, fungi, ecosystems, geology, paleo ...
in London, formerly BMNH), found by
the Reverend The Reverend (abbreviated as The Revd, The Rev'd or The Rev) is an honorific style (form of address), style given to certain (primarily Western Christian, Western) Christian clergy and Christian minister, ministers. There are sometimes differen ...
William Fox in Atherfield on the Isle of Wight, and acquired by the
British Museum The British Museum is a Museum, public museum dedicated to human history, art and culture located in the Bloomsbury area of London. Its permanent collection of eight million works is the largest in the world. It documents the story of human cu ...
in 1882. Specimen NHMUK R176 had been assigned to the species '' Ornithocheirus nobilis'' by the English
naturalist Natural history is a domain of inquiry involving organisms, including animals, fungi, and plants, in their natural environment, leaning more towards observational than experimental methods of study. A person who studies natural history is cal ...
Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was a British naturalist, geologist and writer of numerous books on natural history. He was known for his contributions to zoology, paleontology, and biogeography. He worked extensively in cata ...
in 1888, but Seeley considered it another species of the genus ''Ornithodesmus'', which he now considered pterosaurian. Seeley named the new species ''O. latidens''; ''latus'' is Latin for "wide" and ''dens'' means "tooth", a name originally used by Fox and his friends. Seeley presumably assigned the new species to the existing genus due to similarities between their sacra, but with little explanation. Specimen NHMUK R176 is a poorly preserved skeleton consisting of the back of the skull, a neck vertebra, the sternum, the sacrum, the right humerus, the notarium, the left humerus, part of the
radius In classical geometry, a radius (: radii or radiuses) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The radius of a regular polygon is th ...
and ulna, carpals, metacarpals, and wing
phalanx bone The phalanges (: phalanx ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. Structu ...
s. Though Seeley did not designate a
holotype specimen A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was Species description, formally described. It is either the single such physical example (or illus ...
for ''O. latidens'', he described and illustrated parts of NHMUK R176, which makes the 1901 naming valid according to the
International Commission on Zoological Nomenclature The International Commission on Zoological Nomenclature (ICZN) is an organization dedicated to "achieving stability and sense in the scientific naming of animals". Founded in 1895, it currently comprises 26 commissioners from 20 countries. Orga ...
, and the specimen is considered the holotype today. Later researchers have been mystified by the fact that Seeley described the jaws and teeth of ''O. latidens'' and named it after the latter, when the only specimen available in 1901, NHMUK R176, does not appear to have had these elements. Only the back of the skull was listed as present by Lydekker in 1888, but it was rumoured that a set of jaws had been lost from Fox's collection, so it is possible that Seeley had examined them prior to this. In 1913, the British amateur palaeontologist
Reginald W. 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 ...
described two more specimens of ''O. latidens'', collected from the sea after a
rockfall A rockfall or rock-fallWhittow, John (1984). ''Dictionary of Physical Geography''. London: Penguin, 1984. . is a quantity of Rock (geology), rock that has fallen freely from a cliff face. The term is also used for collapse of rock from roof or wa ...
near Atherfield Point on the Isle of Wight in 1904, originating from the
Vectis Formation The Vectis Formation is a geological Formation (geology), formation on the Isle of Wight and Swanage, England whose strata were formed in the Barremian, approximately 125 million years ago."Magnetostratigraphy of the Lower Cretaceous Vectis Forma ...
. The first of these, NHMUK R3877, was collected as three blocks and consists of a skull, neck and trunk vertebrae, a shoulder blade, an
ischium The ischium (; : is ...
, and parts of the forelimbs. The second specimen, NHMUK R3878, was collected in one block, and includes parts of the
pectoral girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans, it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists o ...
and forelimbs. These specimens represent the most complete remains of
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
pterosaurs found in England, and NHMUK R3877 was one of the only known three-dimensionally preserved pterosaur skeletons for much of the 20th century (pterosaur bones are often flattened
compression fossil A compression fossil is a fossil preserved in sedimentary rock that has undergone physical compression. While it is uncommon to find animals preserved as good compression fossils, it is very common to find plants preserved this way. The reason f ...
s). Hooley discussed ''O. latidens'' in detail, and placed the genus ''Ornithodesmus'' in its own family, Ornithodesmidae. His article ended with a discussion wherein it was noted that the palaeontologist
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 ...
had expressed doubts as to whether ''O. latidens'' belonged in the genus ''Ornithodesmus'', as the vertebrae of the specimen that genus was based on differed markedly from those of Hooley's specimen. The American palaeontologist
Samuel W. 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 ...
subsequently reviewed Hooley's article, disagreeing with some of his conclusions about the anatomy and classification of the animal. After Hooley's
monograph A monograph is generally a long-form work on one (usually scholarly) subject, or one aspect of a subject, typically created by a single author or artist (or, sometimes, by two or more authors). Traditionally it is in written form and published a ...
, little was written about the animal for the rest of the 20th century, and no similar pterosaurs were found for decades.


New genus and assigned species

In 1993, the British palaeontologists Stafford C. Howse and Andrew C. Milner concluded that the holotype sacrum and only specimen of ''O. cluniculus'' did not belong to a pterosaur, but 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, Dromaeosauridae, Troodontidae, ...
n dinosaur (this conclusion had also been reached independently by the British palaeontologist Christopher Bennett). They pointed out that no detailed attempts had been made to compare the sacrum of ''O. cluniculus'' with those of pterosaurs, and that ''O. latidens'' had in effect been treated 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 ...
of the genus ''Ornithodesmus'', with one writer even treating the original species as a synonym of the newer. As a definite species of pterosaurs, "''O.''" ''latidens'' therefore required a new genus name. In 2001, Howse, Milner, and