Ichthyornis

Ichthyornis Temporal range: Late Cretaceous, 95–85 Ma PreЄ Є O S D C P T J K Pg N
Artist's restoration
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Aves
clade:	Carinatae
Subclass:	†Ichthyornithes Marsh, 1873
Order:	†Ichthyornithiformes Fürbringer, 1888
Family:	†Ichthyornithidae Marsh, 1873
Genus:	†Ichthyornis Marsh, 1872b
Species:	†Ichthyornis dispar Marsh, 1872b
Synonyms
see text

Ichthyornis (meaning "fish bird", after its fish-like vertebrae) is a genus of toothed seabirds from the Late Cretaceous of North America. Its fossil remains are known from the chalks of Alberta, Alabama, Kansas, New Mexico, Saskatchewan, and Texas, in strata that were laid down in the Western Interior Seaway during the Turonian-Campanian ages, about 95-85 mya). Ichthyornis is a common component of the Niobrara Formation fauna, and numerous specimens of both adult and immature individuals have been found.^[1]

Ichthyornis has been historically important in shedding light on bird evolution. It was the first discovered prehistoric bird 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 birds known from more than a few specimens.^[2]

Description

It is thought that Ichthyornis was the Cretaceous ecological equivalent of modern seabirds such as gulls, petrels, and skimmers. It 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).^[3]

Ichthyornis is notable primarily for its combination of vertebrae which are concave both in front and back (similar to some fish, which is where the bird 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 and other primitive birds, 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.^[4] 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.^[2]

Although the wings and breastbone were very modern in appearance (suggesting strong flight ability and placing it with modern birds in the advanced group Carinatae), the jaws had numerous small, sharp teeth. Unlike earlier birds such as the enantiornithines, it appears to have matured to adulthood in a rather short, continuous process.^[1]

Timespan and evolution

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 85 million years ago).^[5] 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),^[3] and the Cenomanian of Saskatchewan, dating to about 95 million years ago.^[2]

History of study

Ichthyornis was first 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, USA. Mudge was a prolific fossil collector who shipped his discoveries to prominent scientists for study.^[6] 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.^[6]

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, 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."^[7] Later in 1872, Marsh described the toothed jaws as a new species of marine reptile, named Colonosaurus mudgei after their discoverer.^[8]

Skeleton restoration of I. victor (now I. dispar) by O.C. Marsh

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 the bird 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.^[9] 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.^[10]

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.^[2] (While Archaeopteryx was the first known Mesozoic bird and is now known to have also had teeth, the first specimen with a skull was not described until 1884).^[11] 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.^[2] 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.^[2]

Mounted specimens

Cast of the original composite panel mount of I. victor (now I. dispar), Peabody Museum of Natural History

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).^[2]

At some point before 1937, the specimen of the actual I. victor type specimen was re-assigned to the panel mount. Later reports of the specimen, even by the Peabody Museum's staff, therefore mistakenly stated that the original I. dispar specimen comprised most of the skeleton, when it was in fact only three bones.^[2] 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.^[2] A full re-description of these specimens was published by paleontologist Julia Clarke in 2004.^[2]

Classification

Ichthyornis is close to the ancestry of modern birds, the Neornithes, 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. New data on the radiation of the latter, which is now known to have started already in the Cretaceous (see Vegavis), could shed more light on the exact relationship of these taxa. In Clarke's 2004 review, the former order Ichthyornithiformes and the family Ichthyornithidae are now superseded by the subclass 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.^[2]

Of the several described species, only one, Ichthyornis dispar, is currently recognized, following the seminal review by Julia Clarke.^[2] 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. ancesps should have been considered the correct name for the only recognized Ichthyornis species.^[12]

There has been considerable confusion about the attribution of the fossil material.^[2] The similarity of the lower jaw and teeth to those of mosasaurs (fish-eating marine lizards) is so great that as late as 1952, it was argued that it really belonged to a diminutive species or young individual of or related to the genus Clidastes.^[13]

The presumed "Ichthyornis" lentus actually belongs into the early galliform genus Austinornis.^[2] "Ichthyornis" minusculus from the Bissekty Formation (Late Cretaceous) of Kyzyl Kum, Uzbekistan, is probably an enantiornithine.

Synonyms

Subclass-level

• Odontotormae Marsh, 1875

Genus-level

• Angelinornis Kashin 1972
• Colonosaurus Marsh, 1872c
• Plegadornis Wetmore, 1962 (non C.L. Brehm, 1855 - preoccupied)

Species-level

• Angelinornis antecessor (Wetmore, 1962)
• Colonosaurus mudgei Marsh, 1872c
• Graculavus agilis Marsh, 1873b
• Graculavus anceps Marsh, 1872a
• Ichthyornis antecessor (Wetmore, 1962) Olson, 1975
• Ichthyornis anceps (Marsh, 1872a) Marsh, 1880
• Ichthyornis agilis (Marsh, 1873b) Marsh, 1880
• Ichthyornis tener Marsh, 1880
• Ichthyornis validus Marsh, 1880
• Ichthyornis victor Marsh, 1876
• Plegadornis antecessor Wetmore, 1962

Footnotes

1. ^ ^{a b} Chinsamy, A., Martin, L.D. and Dobson, P. (1998). "Bone microstructure of the diving Hesperornis and the volant Ichthyornis from the Niobrara Chalk of western Kansas." Cretaceous Research, 19(2): 225-235. doi:10.1006/cres.1997.0102
2. ^ ^{a b c d e f g h i j k l m n} Clarke, J.A. (2004). "Morphology, phylogenetic taxonomy, and systematics of Ichthyornis and Apatornis (Avialae: Ornithurae)." Bulletin of the American Museum of Natural History, 286: 1-179. PDF fulltext
3. ^ ^{a b} Shimada, K. and Fernandes, M.V. (2006). "Ichthyornis sp. (Aves: Ichthyornithiformes) from the lower Turonian (Upper Cretaceous) of western Kansas." Transactions of the Kansas Academy of Science, 109(1/2): 21-26.
4. ^ Lamb, J.P. Jr. (1997). "Marsh was right: Ichthyornis had a beak." Journal of Vertebrate Paleontology, 17: 59A.
5. ^ Carpenter, K. (2003). "Vertebrate Biostratigraphy of the Smoky Hill Chalk (Niobrara Formation) and the Sharon Springs Member (Pierre Shale)." High-Resolution Approaches in Stratigraphic Paleontology, 21: 421-437. doi:10.1007/978-1-4020-9053-0
6. ^ ^{a b} Williston, S.W. (1898). "A brief history of fossil collecting in the Niobrara Chalk prior to 1900. Addenda to Part I." The University Geological Survey of Kansas, 4: 28-32.
7. ^ Marsh, O.C., (1872b). "Notice of a new and remarkable fossil bird." American Journal of Science, Series 3, 4(22): 344.
8. ^ Marsh, O.C. (1872). "Notice of a new reptile from the Cretaceous." American Journal of Science, Series 3, 4(23): 406.
9. ^ Marsh, O.C. (1873a). "On a new sub-class of fossil birds (Odontornithes)." American Journal of Science, Series 3, 5(25): 161-162.
10. ^ Mudge, B.F. (1877). "Annual Report of the committee on Geology, for the year ending November 1, 1876." Kansas Academy of Science, Transactions, Ninth Annual Meeting, pp. 4-5.
11. ^ Switek, B. (2010). "Thomas Henry Huxley and the reptile to bird transition." Pp. 251-264 in Moody, R.T.J., Buffetaut, E., Naish, D. and Martill, D.M. (eds.) Dinosaurs and Other Extinct Saurians: A Historical Perspective. Geological Society Special Publication 343.
12. ^ Mortimer, M. (2010). "Ichthyornis." The Theropod Database. Accessed online 29 December 2010, http://home.comcast.net/~eoraptor/Ornithuromorpha.htm#Ichthyornisanceps
13. ^ Gregory, J.T. (1952). "The jaws of the Cretaceous toothed birds, Ichthyornis and Hesperornis." Condor, 54(2): 73-88. PDF fulltext

References

• Chiappe, Luis M.; Lamb, James P. Jr. & Ericson, Per G.P. (2002): New enantiornithine bird from the marine Upper Cretaceous of Alabama. J. Vertebr. Paleontol. 22(1): 170-174. DOI:10.1671/0272-4634(2002)022[0170:NEBFTM]2.0.CO;2
• Marsh, Othniel Charles (1880): Odontornithes, a Monograph on the Extinct Birds of North America. Government Printing Office, Washington.

External links

Paleontology portal

• Ichthyornis dispar: A toothed, flying bird from the Late Cretaceous of Kansas by Mike Everhart, Oceans of Kansas website. Retrieved 2006-09-16.