Ichthyornis Temporal range: Late Cretaceous, [1] | |
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Cast skeleton, Rocky Mountain Dinosaur Resource Center | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Clade: | Avialae |
Clade: | Ornithurae |
Clade: | † Ichthyornithes |
Genus: | † Ichthyornis Marsh, 1873 |
Species: | †I. dispar |
Binomial name | |
†Ichthyornis dispar (Marsh, 1872) | |
Synonyms | |
AngelinornisKashin 1972 Contents
Species synonymy
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Ichthyornis (meaning "fish bird", after its fish-like vertebrae) is an extinct genus of toothy seabird-like ornithuran from the late Cretaceous period of North America. Its fossil remains are known from the chalks of Alberta, Alabama, Kansas (Greenhorn Limestone), New Mexico, Saskatchewan, and Texas, in strata that were laid down in the Western Interior Seaway during the Turonian through Campanian ages, about 95–83.5 million years ago. Ichthyornis is a common component of the Niobrara Formation fauna, and numerous specimens have been found.
Ichthyornis has been historically important in shedding light on bird evolution. It was the first known prehistoric bird relative preserved with teeth, and Charles Darwin noted its significance during the early years of the theory of evolution. Ichthyornis remains important today as it is one of the few Mesozoic era ornithurans known from more than a few specimens.
It is thought that Ichthyornis was the Cretaceous ecological equivalent of modern seabirds such as gulls, petrels, and skimmers. An average specimen was the size of a pigeon, 24 centimetres (9.4 in) long, with a skeletal wingspan (not taking feathers into account) of around 43 centimetres (17 in), [2] though there is considerable size variation among known specimens, with some smaller and some much larger than the type specimen of I. dispar. [1]
Ichthyornis is notable primarily for its combination of vertebrae which are concave both in front and back (similar to some fish, which is where it gets its name) and several more subtle features of its skeleton which set it apart from its close relatives. Ichthyornis is perhaps most well known for its teeth. The teeth were present only in the middle portion of the upper and lower jaws. The jaw tips had no teeth and were covered in a beak. The beak of Ichthyornis, like the hesperornithids, was compound and made up of several distinct plates, similar to the beak of an albatross, rather than a single sheet of keratin as in most modern birds. [3] The teeth were more flattened than the rounded teeth found in crocodilians, though they became wider towards the base of the crown. The tips of the teeth were curved backward and lacked any serrations. [1] They were arranged in a groove, much like those of marine reptiles. [4]
The wings and breastbone were very modern in appearance, suggesting strong flight ability and placing it with modern birds in the advanced group Carinatae. Unlike earlier avialans such as the enantiornithines, the species appears to have matured to adulthood in a rather short, continuous process. [5]
A study on an Ichthyornis endocast reveals that it had a relatively basal brain compared to modern birds, similar to that of Archaeopteryx and other non-avian theropods. Conversely, it had a palate remarkably convergent with that of modern neognaths. [6]
Ichthyornis fossils have been found in almost all levels of the Niobrara Chalk, from beds dating to the late Coniacian age (about 89 million years ago) to the Campanian age (about 83.5 million years ago). [1] [7] Even earlier remains attributed to Ichthyornis have been found in the Greenhorn Formation of Kansas, dating to the early Turonian age (about 93 million years ago). [2] Specimens of Ichthyornis from earlier eras were, on average, smaller than later ones. The holotype specimen of Ichthyornis dispar, YPM 1450, had a humerus about 58 millimetres (2.3 in) long. In many geologically younger specimens like YPM 1742, the same wing bone was 71.5 millimetres (2.81 in) long. Both the older, smaller specimens, and the more recent, larger specimens show signs that they had reached skeletal maturity and were adults, and came from the same geographic area. It is likely that Ichthyornis dispar as a species increased in size over the several million years it inhabited the Western Interior Seaway ecosystem. [1]
Ichthyornis was one of the first Mesozoic avialans ever found and the first one known to have had teeth, making it an important discovery in the early history of paleontology. It remains important today, as it represents one of the closest non-avian relatives of modern birds, and one of a handful of Mesozoic bird relatives represented by numerous specimens. [1] Ichthyornis was discovered in 1870 by Benjamin Franklin Mudge, a professor from Kansas State Agricultural College who recovered the initial fossils from the North Fork of the Solomon River in Kansas, United States. Mudge was a prolific fossil collector who shipped his discoveries to prominent scientists for study. [8] Mudge had previously had a close partnership with paleontologist Edward Drinker Cope of the Academy of Natural Sciences in Philadelphia. However, as described by S.W. Williston in 1898, Mudge was soon contacted by Othniel Charles Marsh, Cope's rival in the so-called Bone Wars, a rush to collect and identify fossils in the American West. Marsh wrote to Mudge in 1872 and offered to identify any important fossils free of charge, and to give Mudge sole credit for their discovery. Marsh had been a friend of Mudge when they were younger, so when Mudge learned of Marsh's request, he changed the address on the shipping crate containing the Ichthyornis specimen (which had already been addressed to Cope and was ready to be sent), and shipped it to Marsh instead. Marsh had narrowly won the prestige of studying and naming the important fossil at the expense of his rival. [8]
However, Marsh did not initially recognize the true importance of the fossil. Soon after receiving it, he reported back to Mudge his opinion that the chalk slab contained the bones of two distinct animals: a small bird animal, and the toothed jaws of some unknown reptile. Marsh considered the unusual vertebrae of the bird to resemble those of a fish, so he named it Ichthyornis, or "fish bird." [9] Later in 1872, Marsh described the toothed jaws as a new species of marine reptile, named Colonosaurus mudgei after their discoverer. [10] The similarity of the lower jaw and teeth to those of mosasaurs is so great that as late as 1952, J.T. Gregory argued that it really belonged to a diminutive species or young individual related to the genus Clidastes . [11]
By early in 1873, Marsh had recognized his error. Through further preparation and exposure of skull bones from the rock, he found that the toothed jaws must have come from the bird itself and not a marine reptile. Due to the previously unknown features of Ichthyornis (vertebrae concave on either side and teeth), Marsh chose to classify it in an entirely new sub-class of birds he called the Odontornithes (or "toothed birds"), and in the new order Ichthyornithes (later Ichthyornithiformes). The only other bird Marsh included in these groups was the newly named Apatornis , which he had previously named as a species of Ichthyornis, I. celer. [12] Mudge later noted the rare and unique quality of these toothed birds (including Hesperornis, which was found to also have teeth by 1877), and the irony of their association with the remains of toothless pterosaurs, flying reptiles which were only known to have had teeth in other regions of the world at that time. [13]
Soon after these discoveries, Ichthyornis was recognized for its significance to the theory of evolution recently published by Charles Darwin. Darwin himself told Marsh in an 1880 letter that Ichthyornis and Hesperornis offered "the best support for the theory of evolution" since he had first published On the Origin of Species in 1859. [1] (While Archaeopteryx was the first known Mesozoic avialan and is now known to have also had teeth, the first specimen with a skull was not described until 1884). [14] Others at the time also recognized the implications of a nearly modern bird with reptilian teeth, and feared the controversy it caused. One Yale student described various men and women urging Marsh to conceal Ichthyornis from the public because it lent too much support to evolutionary theory. [1] Many accused Marsh of having tampered with the fossils or intentionally created a hoax by associating reptilian jaws with the body of a bird, accusations that continued to surface even as late as 1967. However, an overwhelming majority of researchers have demonstrated that Marsh's interpretation of the fossils was correct, and he was fully vindicated by later finds. [1]
At the turn of the 20th century, the Peabody Museum of Natural History at Yale University, where most Ichthyornis specimens were housed, began placing many of its most interesting or important specimens on display in the museum's Great Hall. Two panel mounts (that is, pieces where the skeleton is arranged and set into a plaster slab) were created for Ichthyornis; one for I. dispar, and one for "I. victor". Both were created by Hugh Gibb, who prepared many of Marsh's fossils for study and display. The I. dispar mount contained only the holotype fossils, while the "I. victor" mount was a composite incorporating a variety of different specimens to make the piece appear more complete (it did not, however, contain any part of the actual "I. victor" holotype specimen). [1]
At some point before 1937, the catalogue number of the actual "I. victor" type specimen was mistakenly reassigned to the panel mount. Later reports of the specimen, even by the Peabody Museum's staff, therefore mistakenly stated that the original "I. victor" specimen comprised most of the skeleton, when it was in fact only three bones. [1] By 1997, the situation had become so confused that Jacques Gauthier, the current curator of the museum's vertebrate paleontology collection, authorized the dismantling of both panel mounts. This allowed the bones to be properly sorted out and studied in three dimensions, which had been impossible previously when they were embedded in plaster. [1] A full re-description of these specimens was published by paleontologist Julia Clarke in 2004. [1]
Ichthyornis is close to the ancestry of modern birds, the Aves, but represents an independent lineage. It was long believed that it was closely related to some other Cretaceous taxa known from very fragmentary remains – Ambiortus , Apatornis , Iaceornis and Guildavis – but these seem to be closer to the ancestors of modern birds than to Ichthyornis dispar. In Clarke's 2004 review, the former order Ichthyornithiformes and the family Ichthyornithidae are now superseded by the clade Ichthyornithes, which in the paper was also defined according to phylogenetic taxonomy as all descendants of the most recent common ancestor of Ichthyornis dispar and modern birds. [1]
Of the several described species, only one, Ichthyornis dispar, is currently recognized, following the seminal review by Julia Clarke. [1] Marsh had previously named a specimen now attributed to I. dispar as Graculavus anceps. Clarke argued that because the rules for naming animals laid out by the ICZN state that a type species for a genus must have originally been included in that genus, Ichthyornis anceps is ineligible to replace I. dispar as the type species and so must be considered a junior synonym even though it was named first. However, Michael Mortimer pointed out that this is incorrect; while I. anceps cannot become the type species of Ichthyornis, the ICZN does not preclude it from becoming the senior synonym of the type species I. dispar. Therefore, I. anceps should have been considered the correct name for the only recognized Ichthyornis species. [15] All other supposed species of Ichthyornis have not been supported as valid. The presumed "Ichthyornis" lentos, for example, actually belongs into the early galliform genus Austinornis . [1] "Ichthyornis" minusculus from the Bissekty Formation (Late Cretaceous) of Kyzyl Kum, Uzbekistan, is probably an enantiornithine. All other Ichthyornis species are synonymous with I. dispar. [1]
The cladogram below is the result of a 2014 analysis by Michael Lee and colleagues that expanded on data from an earlier study by O’Connor & Zhou in 2012. The clade names are positioned based on their definitions. [16]
Ornithurae |
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Pteranodon ; from Ancient Greek πτερόν and ἀνόδων is a genus of pterosaur that included some of the largest known flying reptiles, with P. longiceps having a wingspan of over 6 m (20 ft). They lived during the late Cretaceous geological period of North America in present-day Kansas, Nebraska, Wyoming, South Dakota and Alabama. More fossil specimens of Pteranodon have been found than any other pterosaur, with about 1,200 specimens known to science, many of them well preserved with nearly complete skulls and articulated skeletons. It was an important part of the animal community in the Western Interior Seaway.
Cretoxyrhina is an extinct genus of large mackerel shark that lived about 107 to 73 million years ago during the late Albian to late Campanian of the Late Cretaceous. The type species, C. mantelli, is more commonly referred to as the Ginsu shark, first popularized in reference to the Ginsu knife, as its theoretical feeding mechanism is often compared with the "slicing and dicing" when one uses the knife. Cretoxyrhina is traditionally classified as the likely sole member of the family Cretoxyrhinidae but other taxonomic placements have been proposed, such as within the Alopiidae and Lamnidae.
Hesperornithes is an extinct and highly specialized group of aquatic avialans closely related to the ancestors of modern birds. They inhabited both marine and freshwater habitats in the Northern Hemisphere, and include genera such as Hesperornis, Parahesperornis, Baptornis, Enaliornis, and Potamornis, all strong-swimming, predatory divers. Many of the species most specialized for swimming were completely flightless. The largest known hesperornithean, Canadaga arctica, may have reached a maximum adult length of 2.2 metres (7.2 ft).
Hesperornis is a genus of cormorant-like Ornithuran that spanned throughout the Campanian age, and possibly even up to the early Maastrichtian age, of the Late Cretaceous period. One of the lesser-known discoveries of the paleontologist O. C. Marsh in the late 19th century Bone Wars, it was an early find in the history of avian paleontology. Locations for Hesperornis fossils include the Late Cretaceous marine limestones from Kansas and the marine shales from Canada. Nine species are recognised, eight of which have been recovered from rocks in North America and one from Russia.
Tylosaurus is a genus of russellosaurine mosasaur that lived about 92 to 66 million years ago during the Turonian to Maastrichtian stages of the Late Cretaceous. Its fossils have been found primarily around North Atlantic Ocean including in North America, Europe, and Africa. The earliest discoveries were possibly made by Native American tribes in the Great Plains, whose creation myths spoke of giant serpentine water monsters turned to stone in ancient times. Paleontologist Edward Drinker Cope first scientifically described fossils of the genus from Kansas in 1869, but the name Tylosaurus was coined by Cope's rival Othniel Charles Marsh.
Odontornithes is an obsolete and disused taxonomic term proposed by Othniel Charles Marsh for birds possessing teeth, notably the genera Hesperornis and Ichthyornis from the Cretaceous deposits of Kansas.
The UW–Madison Geology Museum (UWGM) is a geology and paleontology museum housed in Weeks Hall, in the southwest part of the University of Wisconsin–Madison campus. The museum's main undertakings are exhibits, outreach to the public, and research. It has the second highest attendance of any museum at the University of Wisconsin–Madison, exceeded only by the Chazen Museum of Art. The museum charges no admission.
Protosphyraena is a fossil genus of swordfish-like marine fish, that thrived worldwide during the Upper Cretaceous Period (Coniacian-Maastrichtian). Though fossil remains of this taxon have been found in both Europe and Asia, it is perhaps best known from the Smoky Hill Member of the Niobrara Chalk Formation of Kansas. Protosphyraena was a large fish, averaging 2–3 metres in length. Protosphyraena shared the Cretaceous oceans with aquatic reptiles, such as mosasaurs and plesiosaurs, as well as with many other species of extinct predatory fish. The name Protosphyraena is a combination of the Greek word protos ("early") plus Sphyraena, the genus name for barracuda, as paleontologists initially mistook Protosphyraena for an ancestral barracuda. Recent research shows that the genus Protosphyraena is not at all related to the true swordfish-family Xiphiidae, but belongs to the extinct family Pachycormidae.
Ichthyornithes is an extinct group of toothed avialan dinosaurs very closely related to the common ancestor of all modern birds. They are known from fossil remains found throughout the late Cretaceous period of North America, though only two genera, Ichthyornis and Janavis, are represented by complete enough fossils to have been named. Ichthyornitheans became extinct at the Cretaceous–Paleogene boundary, along with enantiornitheans, all other non-avian dinosaurs, and many other animal and plant groups.
Baptornis is a genus of flightless, aquatic birds from the Late Cretaceous, some 87-80 million years ago. The fossils of Baptornis advenus, the type species, were discovered in Kansas, which at its time was mostly covered by the Western Interior Seaway, a shallow shelf sea. It is now known to have also occurred in today's Sweden, where the Turgai Strait joined the ancient North Sea; possibly, it occurred in the entire Holarctic.
Polarornis is a genus of prehistoric bird, possibly an anserimorph. It contains a single species Polarornis gregorii, known from incomplete remains of one individual found on Seymour Island, Antarctica, in rocks which are dated to the Late Cretaceous.
Benjamin Franklin Mudge was an American lawyer, geologist and teacher. Briefly the mayor of Lynn, Massachusetts, he later moved to Kansas where he was appointed the first State Geologist. He led the first geological survey of the state in 1864, and published the first book on the geology of Kansas. He lectured extensively, and was department chair at the Kansas State Agricultural College.
Apatornis is a genus of ornithuran dinosaurs endemic to North America during the late Cretaceous. It currently contains a single species, Apatornis celer, which lived around the Santonian-Campanian boundary, dated to about 83.5 million years ago. The remains of this species were found in the Smoky Hill Chalk of the Niobrara Formation in Kansas, United States. It is known from a single fossil specimen: a synsacrum, the fused series of vertebrae over the hips.
Parahesperornis is a genus of prehistoric flightless birds from the Late Cretaceous. Its range in space and time may have been extensive, but its remains are rather few and far between, at least compared with its contemporary relatives in Hesperornis. Remains are known from central North America, namely the former shallows of the Western Interior Seaway in Kansas. Found only in the upper Niobrara Chalk, these are from around the Coniacian-Santonian boundary, 85–82 million years ago (mya).
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 Smoky Hill Chalk Member is the uppermost of the two structural units of the Niobrara Chalk. It is underlain by the Fort Hays Limestone Member and overlain by the Pierre Shale. The Smoky Hill Chalk outcrops in parts of northwest Kansas, its most famous localities for fossils, and in southeastern Nebraska. Large well-known fossils excavated from the Smoky Hill Chalk include marine reptiles such as plesiosaurs, large bony fish such as Xiphactinus, mosasaurs, flying reptiles or pterosaurs, flightless marine birds such as Hesperornis, and turtles. Many of the most well-known specimens of the marine reptiles were collected by dinosaur hunter Charles H. Sternberg and his son George. The son collected a unique fossil of the giant bony fish Xiphactinus audax with the skeleton of another bony fish, Gillicus arcuatus inside the larger one. Another excellent skeleton of Xiphactinus audax was collected by Edward Drinker Cope during the late nineteenth century heyday of American paleontology and its Bone Wars.
During the time of the deposition of the Niobrara Chalk, much life inhabited the seas of the Western Interior Seaway. By this time in the Late Cretaceous many new lifeforms appeared such as mosasaurs, which were to be some of the last of the aquatic lifeforms to evolve before the end of the Mesozoic. Life of the Niobrara Chalk is comparable to that of the Dakota Formation, although the Dakota Formation, which was deposited during the Cenomanian, predates the chalk by about 10 million years.
Iaceornis is a genus of marine ornithuran dinosaurs closely related to modern birds. It was endemic to North America during the Late Cretaceous, living about 83.5 million years ago. It is known from a single fossil specimen found in Gove County, Kansas (USA), and consisting of a partial skeleton lacking a skull.
Pteranodontoidea is an extinct clade of ornithocheiroid pterosaurs from the Early to Late Cretaceous of Asia, Africa, Europe, North America and South America. It was named by Alexander Wilhelm Armin Kellner in 1996. In 2003, Kellner defined the clade as a node-based taxon consisting of the last common ancestor of Anhanguera, Pteranodon and all its descendants. The clade Ornithocheiroidea is sometimes considered to be the senior synonym of Pteranodontoidea, however it depends on its definition. Brian Andres in his analyses, converts Ornithocheiroidea using the definition of Kellner (2003) to avoid this synonymy.
This timeline of mosasaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of mosasaurs, a group of giant marine lizards that lived during the Late Cretaceous Epoch. Although mosasaurs went extinct millions of years before humans evolved, humans have coexisted with mosasaur fossils for millennia. Before the development of paleontology as a formal science, these remains would have been interpreted through a mythological lens. Myths about warfare between serpentine water monsters and aerial thunderbirds told by the Native Americans of the modern western United States may have been influenced by observations of mosasaur fossils and their co-occurrence with creatures like Pteranodon and Hesperornis.
Tingmiatornis is a genus of flighted and possibly diving ornithurine dinosaur from the High Arctic of Canada. The genus contains a single species, T. arctica, described in 2016, which lived during the Turonian epoch of the Cretaceous.
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