Gastornis

File:PSM V21 D483 Skeleton of a gastornis eduardsii.jpg

Gastornis was first described in 1855 from a fragmentary skeleton. It was named after Gaston Planté, described as a "studious young man full of zeal", who had discovered the first fossils in clay ({{ill|Argile Plastique|fr}}) formation deposits at Meudon, near Paris.{{cite journal|author=Prévost, Constant|year=1855|title= Annonce de la découverte d'un oiseau fossile de taille gigantesque, trouvé à la partie inférieure de l'argile plastique des terrains parisiens ["Announcement of the discovery of a fossil bird of gigantic size, found in the lower Argile Plastique formation of the Paris region"]|journal=C. R. Acad. Sci. Paris|volume=40|pages= 554–557|language=fr|url=http://visualiseur.bnf.fr/CadresFenetre?O=NUMM-2997}} The discovery was notable, due to the large size of the specimens, and because, at the time, Gastornis represented one of the oldest known birds.Buffetaut, E., and Burrrraur, E. (1997). "New remains of the giant bird Gastornis from the Upper Paleocene of the eastern Paris Basin and the relationships between Gastornis and Diatryma." N. Jb. Geol. Palâont. Mh., (3): 179–190. [https://www.academia.edu/download/31029134/Buffetaut_1997_Gastornis_Reims.pdf]{{dead link|date=July 2022|bot=medic}}{{cbignore|bot=medic}} Additional bones of the first known species, G. parisiensis, were found in the mid-1860s. Somewhat more-complete specimens, then referred to the new species G. edwardsii (now considered a synonym of G. parisiensis), were found a decade later. These specimens, found in the 1870s, formed the basis for a widely- circulated and reproduced skeletal restoration by Lemoine. The skulls of these original Gastornis fossils were unknown, other than nondescript fragments and several bones used in Lemoine's illustration, which turned out to be those of other animals.{{cite journal | author = Martin L.D. | year = 1992 | title = The status of the Late Paleocene birds Gastornis and Remiornis | journal = Papers in Avian Paleontology Honoring Pierce Brodkorb. Natural History Museum of Los Angeles County, Science Series | volume = 36 | pages = 97–108 |url=}} Thus, this European specimen was long reconstructed as a sort of gigantic "crane-like" bird.{{cite book|author=Lemoine, V.|year=1881a|title= Recherches sur les oiseaux fossiles des terrains tertiaires inférieurs des environs de Reims |volume=2|pages= 75–170|publisher= Matot-Braine, Reims}}{{cite journal|author=Lemoine, V.|year=1881b|title= Sur le Gastornis Edwardsii et le Remiornis Heberti de l'éocène inférieur des environs de Reims ["On G. edwardsii and R. heberti from the Lower Eocene of the Reims area"]|journal=C. R. Acad. Sci. Paris|volume=93|pages= 1157–1159|language=fr|url=http://visualiseur.bnf.fr/CadresFenetre?O=NUMM-3049}}

In 1874, the American paleontologist Edward Drinker Cope discovered another fragmentary set of fossils at the Wasatch Formation, New Mexico. Cope considered the fossils to be of a distinct genus and species of giant ground bird; in 1876, he named the remains Diatryma gigantea ({{IPAc-en|ˌ|d|aɪ|.|ə|ˈ|t|r|aɪ|m|ə}} {{Respell|DY|ə|TRY|mə}}),The biologist's handbook of pronunciations (1960) from the Ancient Greek διάτρημα (diatrema), meaning "through a hole", in reference to the large foramina (perforations) that penetrated some of the foot bones.{{cite journal|author=Cope, Edward Drinker|year=1876|title= On a gigantic bird from the Eocene of New Mexico|journal=Proceedings of the Academy of Natural Sciences of Philadelphia|volume=28|issue=2|pages= 10–11}}{{cite book|author=Feduccia, Alan|year=1999|title= The Origin and Evolution of Birds|edition=2nd|publisher=Yale University Press|place= New Haven|isbn=0300078617}} In 1894, a single gastornithid toe bone from New Jersey was described by Cope's "rival" Othniel Charles Marsh, and classified as a new genus and species: Barornis regens. In 1911, it was recognized that this, too, could be considered a junior synonym of Diatryma (and therefore, later, Gastornis).{{cite journal |author1=Matthew W.D. |author2=Granger W. |author3=Stein W. | year = 1917 | title = The skeleton of Diatryma, a gigantic bird from the Lower Eocene of Wyoming | journal = Bulletin of the American Museum of Natural History | volume = 37 | issue = 11| pages = 307–354 }} Additional, fragmentary specimens were found in Wyoming in 1911, and assigned (in 1913) to the new species Diatryma ajax (also now considered a synonym of G. giganteus). In 1916, an American Museum of Natural History expedition to the Bighorn Basin (Willwood Formation, Wyoming) found the first nearly-complete skull and skeleton, which was described in 1917 and gave scientists their first clear picture of the bird. Matthew, Granger, and Stein (1917) classified this specimen as yet another new species, Diatryma steini.

File:Gastornis 1917.jpg-like plumage, 1917]]

After the description of Diatryma, most new European specimens were referred to this genus, instead of Gastornis; however, after the initial discovery of Diatryma, researchers recognized the similarity between the two genera as early as 1884 when Elliott Coues placed Diatryma gigantea under the genus Gastornis as G. giganteus, a synonymy agreed upon by the American Ornithologists' Union in 1886.{{cite book|author=American Ornithologists' Union|year=1886|chapter=Fossil Birds|title=The code of nomenclature and check-list of North American birds : adopted by the American Ornithologists' Union : being the report of the Committee of the Union on Classification and Nomenclature|page=362|doi=10.5962/bhl.title.1538|url=https://dn720200.ca.archive.org/0/items/codeofnomenclatu00amer/codeofnomenclatu00amer.pdf}} Further meaningful comparisons between Gastornis and Diatryma were made more difficult by Lemoine's incorrect skeletal illustration, the composite nature of which was not discovered until the early 1980s. Following this, several authors began to recognize a greater degree of similarity between the European and North American birds, often placing both in the same order (†Gastornithiformes) or even family (†Gastornithidae). This newly-realized degree of similarity caused many scientists to, tentatively, accept the animals' synonymy pending a comprehensive review of the anatomy of both genera, in which Gastornis has the taxonomic priority. Some subsequent studies either continued to use the genus Diatryma or argued against the synonymy, since a detailed comparison of type specimens has not been done yet and notable differences can be found in the species originally assigned to Diatryma from the type species of Gastornis.

File:Anserimorphs size comparison.png and Genyornis]]

Gastornis is known from a large amount of fossil remains, but the clearest picture of the bird comes from a few nearly complete specimens of the species G. giganteus. These were generally very large birds, with huge beaks and massive skulls superficially similar to the carnivorous South American "terror birds" (phorusrhacids). The largest known species, G. giganteus would have reached about {{convert|2|m|ftin|abbr=on}} in maximum height, and up to {{convert|175|kg|lb|abbr=on}} in mass.

The skull of G. giganteus was huge compared to the body and powerfully built. The beak was extremely tall and compressed (flattened from side to side). Unlike other species of Gastornis, G. giganteus lacked characteristic grooves and pits on the underlying bone. The 'lip' of the beak was straight, without a raptorial hook as found in the predatory phorusrhacids. The nostrils were small and positioned close to the front of the eyes about midway up the skull. The vertebrae were short and massive, even in the neck. The neck was relatively short, consisting of at least 13 massive vertebrae. The torso was relatively short. The wings were vestigial, with the upper wing-bones small and highly reduced, similar in proportion to the wings of the cassowary. A largely complete skull specimen (GMH XVIII-1178-1958) of G. geiselensis was also described in 2024 after its discovery in 1958. The upper beaks of G. geiselensis show possible sexual dimorphism and are wider than those of G. giganteus and proportionally longer than those of G. laurenti.

File:Gastornis.png

Gastornis and its close relatives are classified together in the family Gastornithidae, and were long considered to be members of the order Gruiformes. However, the traditional concept of Gruiformes has since been shown to be an unnatural grouping. Beginning in the late 1980s with the first phylogenetic analysis of gastornithid relationships, consensus began to grow that they were close relatives of the lineage that includes waterfowl and screamers, the Anseriformes.{{cite journal |author1=Mustoe G.E. |author2=Tucker D.S. |author3=Kemplin K.L. | year = 2012 | title = Giant Eocene bird footprints from northwest Washington, USA | journal = Palaeontology | volume = 55 | issue = 6| pages = 1293–1305 | doi=10.1111/j.1475-4983.2012.01195.x|bibcode=2012Palgy..55.1293M | doi-access = free }} A 2007 study showed that gastornithids were a very early-branching group of anseriformes, and formed the sister group to all other members of that lineage.Agnolin, F. (2007). "Brontornis burmeisteri Moreno & Mercerat, un Anseriformes (Aves) gigante del Mioceno Medio de Patagonia, Argentina." Revista del Museo Argentino de Ciencias Naturales, n.s. 9, 15–25

Recognizing the apparent close relationship between gastornithids and waterfowl, some researchers classify gastornithids within the anseriform group itself. Others restrict the name Anseriformes only to the crown group formed by all modern species, and label the larger group including extinct relatives of anseriformes, like the gastornithids, with the name Anserimorphae.{{cite journal | author = Andors A.V. | year = 1992 | title = Reappraisal of the Eocene ground bird Diatryma (Aves: Anserimorphae) | journal = Science Series Natural History Museum of Los Angeles County | volume = 36 | pages = 109–125 }} Gastornithids are therefore sometimes placed in their own order, Gastornithiformes.Buffetaut, E. (2002). "Giant ground birds at the Cretaceous-Tertiary boundary: Extinction or survival?" Special papers – Geological Society of America, 303–306. A 2024 study, however, found little support for Gastornithiformes and instead places Gastornis as a member of the Galliformes crown group, as more closely related to Phasianoidea than to megapodes, being sister to the extinct Sylviornithidae, a recently extinct group of medium-sized flightless birds known from subfossil deposits in the Western Pacific.{{cite journal |last1=McInerney |first1=Phoebe L. |last2=Blokland |first2=Jacob C. |last3=Worthy |first3=Trevor H. |title=Skull morphology of the enigmatic Genyornis newtoni Stirling and Zeitz, 1896 (Aves, Dromornithidae), with implications for functional morphology, ecology, and evolution in the context of Galloanserae |journal=Historical Biology |date=2 June 2024 |volume=36 |issue=6 |pages=1093–1165 |doi=10.1080/08912963.2024.2308212 |language=en |issn=0891-2963|doi-access=free |bibcode=2024HBio...36.1093M }}

A simplified version of the family tree found by Agnolin et al. in 2007 is reproduced below.

{{clade| style=font-size:100%;line-height:80%

|label1=Anseriformes

|1={{clade

|1=†Gastornithidae

|2={{clade

|1=†Brontornis

|2={{clade

|1=†Dromornithidae

|label2=Anseres

|2={{clade

|1=Anatoidea (ducks & geese)

|2={{clade

|1=Anseranatidae (magpie geese)

|2=Anhimidae (screamers)

}} }} }} }} }} }}

Image:Gastornis geiselensis.jpg

As of 2024, at least three species are confidently placed within the genus Gastornis: G. parisiensis (type species), G. russelli and G. laurenti. The type species, Gastornis parisiensis, was named and described by Hébert in two 1855 papers.{{cite journal|author=Hébert, E.|year=1855a|title= Note sur le tibia du Gastornis pariensis [sic] ["Note on the tibia of G. parisiensis"]|journal=C. R. Acad. Sci. Paris|volume=40|pages= 579–582|language=fr|url=http://visualiseur.bnf.fr/CadresFenetre?O=NUMM-2997}}{{cite journal|author=Hébert, E.|year=1855b|title= Note sur le fémur du Gastornis parisiensis ["Note on the femur of G. parisiensis"]|journal=C. R. Acad. Sci. Paris|volume=40|pages= 1214–1217|language=fr|url=http://visualiseur.bnf.fr/CadresFenetre?O=NUMM-2997}} It is known from fossils found in western and central Europe, dating from the late Paleocene to the early Eocene. Other species previously considered distinct, but which are now considered synonymous with G. parisiensis, include G. edwardsii (Lemoine, 1878) and G. klaasseni (Newton, 1885). Additional European species of Gastornis are G. russelli (Martin, 1992) from the late Paleocene of Berru, France, and G. sarasini (Schaub, 1929) from the early-middle Eocene. The supposed small species G. minor is considered to be a nomen dubium.

Named in 2020, G. laurenti is the most recently described species of Gastornis from southwestern France. The holotype (MHNT.PAL.2018.20.1) is a nearly complete mandible which differs from other species within the genus, and the paratypes consist of the maxilla, right quadrate, femur shaft, tibiotarsus (two left and one right) and six cervical vertebrae.{{Cite journal|author1=Cécile Mourer-Chauviré |author2=Estelle Bourdon |year=2020 |title=Description of a new species of Gastornis (Aves, Gastornithiformes) from the early Eocene of La Borie, southwestern France |journal=Geobios |volume=63 |pages=39–46 |doi=10.1016/j.geobios.2020.10.002 |bibcode=2020Geobi..63...39M |s2cid=228975095 |url=https://hal.archives-ouvertes.fr/hal-03493566/file/S0016699520300863.pdf }} A 2024 study attributed more postcranial remains from the same locality to G. laurenti.{{Cite journal|last1=Mourer-Chauviré |first1=C. |last2=Bourdon |first2=E. |last3=Duffaud |first3=S. |last4=Le Roux |first4=G. |last5=Laurent |first5=Y. |year=2024 |title=New avian remains from the early Eocene of La Borie, southern France |journal=Geobios |volume=83 |pages=61–84 |doi=10.1016/j.geobios.2022.10.004 |s2cid=257251285 }}

File:Gastornis xichuanensis Holotype.jpg

Gastornis giganteus (Cope, 1876), formerly Diatryma gigantea, dates from the middle Eocene of western North America. Its junior synonyms include Barornis regens (Marsh, 1894) and possibly Omorhamphus storchii (Sinclair, 1928). O. storchii was described based on fossils from lower Eocene rocks of Wyoming.{{cite journal|author=Sinclair, W. J. |year=1928|title=Omorhamphus, a New Flightless Bird from the Lower Eocene of Wyoming|jstor=984256|journal=Proc. Am. Philos. Soc.|volume= LXVII |issue=1|pages= 51–65}} The species was named in honor of T. C. von Storch, who found the fossils remains in Princeton 1927 Expedition.{{cite journal|journal=The Auk|url=http://sora.unm.edu/sites/default/files/journals/auk/v045n04/p0522-p0523.pdf |title=Recent Literature|year=1928|volume=45|issue=4|pages=522–523|doi=10.2307/4075674|jstor=4075674 }} The fossil bones originally described as Omorhamphus storchii are considered to be the remains of a juvenile Gastornis giganteus by Brodkorb (1967),{{cite journal|author=Brodkorb, Pierce|year=1967|title=Catalogue of Fossil Birds: Part 3 (Ralliformes, Ichthyornithiformes, Charadriiformes)|journal=Bulletin of the Florida State Museum|volume=11|issue=3|url=http://fulltext10.fcla.edu/cgi/t/text/text-idx?cc=feol;subview=fullcitation;idno=UF00001511|access-date=21 April 2008|archive-date=23 February 2008|archive-url=https://web.archive.org/web/20080223203144/http://fulltext10.fcla.edu/cgi/t/text/text-idx?cc=feol;subview=fullcitation;idno=UF00001511|url-status=dead}} but Louchart et al. (2021) argued that no definitive juvenile specimens of G. giganteus are known and that the two taxa have no known association, so there is no unambiguous evidence to support this synonymy.{{cite journal|vauthors=Louchart A, Bhullar BA, Riamon S, Field DJ|year=2021|title=The True Identity of Putative Tooth Alveoli in a Cenozoic Crown Bird, the Gastornithid Omorhamphus|journal=Frontiers in Earth Science|volume=9|pages=Article 661699|bibcode=2021FrEaS...9..333L|doi=10.3389/feart.2021.661699|doi-access=free|s2cid=234487888}} Specimen YPM PU 13258 from lower Eocene Willwood Formation rocks of Park County, Wyoming also seems to be a juvenile – perhaps also of G. giganteus, in which case it would be an even younger individual.{{cite journal|author=Wetmore, Alexander|year=1933|title= Fossil Bird Remains from the Eocene of Wyoming|journal=Condor|volume=35|issue=3|pages= 115–118|url=http://sora.unm.edu/sites/default/files/journals/condor/v035n03/p0115-p0118.pdf|doi=10.2307/1363436|jstor=1363436}}

G. geiselensis, from the middle Eocene of Messel, Germany, has been considered a synonym of G. sarasini;{{cite book|author=Mlíkovský, Jirí|year=2002|title=Cenozoic Birds of the World, Part 1: Europe|publisher=Ninox Press|place=Prague|url=http://www.nm.cz/download/JML-18-2002-CBE.pdf|access-date=21 April 2008|archive-url=https://web.archive.org/web/20110520101755/http://www.nm.cz/download/JML-18-2002-CBE.pdf|archive-date=20 May 2011|url-status=dead}} however, other researchers have stated that there is currently insufficient evidence to synonymize the two, and that they should be kept separate at least pending a more detailed comparison of all gastornithids.{{cite journal | author = Hellmund M | year = 2013 | title = Reappraisal of the bone inventory of Gastornis geiselensis (Fischer, 1978) from the Eocene Geiseltal Fossillagerstatte (Saxony-Anhalt, Germany) | journal = Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen | volume = 269 | issue = 2| pages = 203–220 | doi=10.1127/0077-7749/2013/0345}}

In 2024, Gerald Mayr and colleagues argued against the synonymy of Diatryma with Gastornis based on the distinct features of the coracoid and tarsometatarsus of G. giganteus and G. geiselensis, referred to as D. gigantea and D. geiselensis in the paper, when compared to those of G. parisiensis. They further suggested that these two features support the placement of G. sarasini within Diatryma as D. sarasini, and that assigning all species of gastornithiforms to the genus Gastornis would not properly reflect the interrelationships of this taxonomic group. A simplified version of their phylogenetic analysis is reproduced below:{{cite journal|last1=Mayr |first1=G. |last2=Mourer-Chauviré |first2=C. |last3=Bourdon |first3=E. |last4=Stache |first4=M. |title=Resurrecting the taxon Diatryma: A review of the giant flightless Eocene Gastornithiformes (Aves), with a report of the first skull of Diatryma geiselensis |year=2024 |journal=Palaeontologia Electronica |volume=27 |issue=3 |at=27.3.a57 |doi=10.26879/1438 |doi-access=free }}

{{clade|style=font-size:100%;line-height:100%

|label1=Gastornithiformes

|1={{clade

|1=Walbeck gastornithiform

|2=Gastornis russelli

|3={{clade

|1=Gastornis parisiensis

|2=Gastornis laurenti

|3=Louvois gastornithiform

|4={{clade

|1=London Clay Diatryma

|2=Diatryma sarasini

|3=Diatryma geiselensis

|4=Diatryma gigantea }} }} }} }}

A tibiotarsus (upper foot bone) originally described in 1980 as Zhongyuanus xichuanensis from the early Eocene of Henan, China,{{cite journal | author = Hou L | year = 1980 | title = New form of the Gastornithidae from the Lower Eocene of the Xichuan, Honan | journal = Vertebrata PalAsiatica | volume = 18 | pages = 111–115 }} was suggested to be an Asian species of Gastornis in 2013.{{cite journal | author = Buffetaut E | year = 2013 | title = The giant bird Gastornis in Asia: A revision of Zhongyuanus xichuanensis Hou, 1980, from the Early Eocene of China | journal = Paleontological Journal | volume = 47 | issue = 11| pages = 1302–1307 | doi=10.1134/s0031030113110051| bibcode = 2013PalJ...47.1302B | s2cid = 84611178 }} However, the 2024 study which argued against the synonymy of Diatryma with Gastornis suggested that this fragmentary Chinese taxon cannot be confidently assigned to either Diatryma or Gastornis, and thus more evaluation is required to clarify its taxonomic affinities.

File:Gastornis skull and mandible.jpg

A long-standing debate surrounding Gastornis is the interpretation of its diet. It has often been depicted as a predator of contemporary small mammals, which famously included the early horse Eohippus. However, with the size of Gastornis legs, the bird would have had to have been more agile to catch fast-moving prey than the fossils suggest it to have been. Consequently, Gastornis has been suspected to have been an ambush hunter and/or used pack hunting techniques to pursue or ambush prey; if Gastornis was a predator, it would have certainly needed some other means of hunting prey through the dense forest. Alternatively, it could have used its strong beak for eating large or strong vegetation.

The skull of Gastornis is massive in comparison to those of living ratites of similar body size. Biomechanical analysis of the skull suggests that the jaw-closing musculature was enormous. The lower jaw is very deep, resulting in a lengthened moment arm of the jaw muscles. Both features strongly suggest that Gastornis could generate a powerful bite.{{cite journal|author1=Witmer, Lawrence |author2=Rose, Kenneth |year=1991|title= Biomechanics of the jaw apparatus of the gigantic Eocene bird Diatryma: Implications for diet and mode of life|journal= Paleobiology|volume=17|issue=2|pages= 95–120|doi=10.1017/S0094837300010435 |bibcode=1991Pbio...17...95W |s2cid=18212799 |url=http://www.oucom.ohiou.edu/dbms-witmer/Downloads/1991_Witmer_%26_Rose_Diatryma.pdf|url-status=dead|archive-url=https://web.archive.org/web/20100728040104/http://www.oucom.ohiou.edu/dbms-witmer/Downloads/1991_Witmer_%26_Rose_Diatryma.pdf|archive-date=28 July 2010}} Some scientists have proposed that the skull of Gastornis was 'overbuilt' for a herbivorous diet and support the traditional interpretation of Gastornis as a carnivore that used its powerfully constructed beak to subdue struggling prey and crack open bones to extract marrow. Others have noted the apparent lack of predatory features in the skull, such as a prominently hooked beak, as evidence that Gastornis was a specialized herbivore (or even an omnivore) of some sort, perhaps having used its large beak to crack hard foods like nuts and seeds.{{cite journal|author=Andors, Allison|year=1992|title= Reappraisal of the Eocene groundbird Diatryma (Aves: Anserimorphae)|journal=Papers in Avian Paleontology Honoring Pierce Brodkorb–Natural History Museum of Los Angeles County Science Series|volume=36|pages= 109–125}} Footprints attributed to gastornithids (possibly a species of Gastornis itself), described in 2012, showed that these birds lacked strongly hooked talons on the hind legs, another line of evidence suggesting that they did not have a predatory lifestyle.{{cite journal|author1=Mustoe, George E |author2=Tucker David S |author3=Kemplin, Keith L |year=2012|title=Giant Eocene Bird Footprints From Northwest Washington, USA|journal=Palaeontology|volume=55|issue=6|pages= 1293–1305|doi=10.1111/j.1475-4983.2012.01195.x|bibcode=2012Palgy..55.1293M |doi-access=free}}

Recent evidence suggests that Gastornis was likely a true herbivore.{{cite journal |author1=Angst D. |author2=Lécuyer C. |author3=Amiot R. |author4=Buffetaut E. |author5=Fourel F. |author6=Martineau F. |author7=Legendre S. |author8=Abourachid A. |author9=Herrel A. | year = 2014 | title = Isotopic and anatomical evidence of an herbivorous diet in the Early Tertiary giant bird Gastornis. Implications for the structure of Paleocene terrestrial ecosystems | journal = Naturwissenschaften | volume = 101|issue=4 | pages = 313–322| doi = 10.1007/s00114-014-1158-2 | pmid=24563098|bibcode=2014NW....101..313A |s2cid=18518649 }} Studies of the calcium isotopes in the bones of specimens of Gastornis by Thomas Tutken and colleagues showed no evidence that it had meat in its diet. The geochemical analysis further revealed that its dietary habits were similar to those of both herbivorous dinosaurs and mammals when it was compared to known fossil carnivores, such as Tyrannosaurus rex, leaving phorusrhacids and bathornithids as the only major carnivorous flightless birds.{{cite web|url=https://www.sciencedaily.com/releases/2013/08/130829214559.htm |title=Terror bird's beak was worse than its bite: 'Terror bird' was probably a herbivore |publisher=Sciencedaily.com |date=29 August 2013 |access-date=10 September 2013}} The first in situ preserved gastroliths in a specimen of G. geiselensis (or D. geiselensis) also conforms to its herbivorous diet.

In Late Paleocene deposits of Spain and early Eocene deposits of France, shell fragments of huge eggs have turned up, namely in Provence.{{cite journal|author1=Dughi, R. |author2=Sirugue, F.|year=1959|title= Sur des fragments de coquilles d'oeufs fossiles de l'Eocène de Basse-Provence ["On fossil eggshell fragments from the Eocene of Basse-Provence"]|journal=C. R. Acad. Sci. Paris|volume=249|pages= 959–961|language=fr|url=http://gallica.bnf.fr/ark:/12148/bpt6k3201c}}{{cite journal|author1=Fabre-Taxy, Suzanne |author2=Touraine, Fernand|year=1960|title= Gisements d'œufs d'Oiseaux de très grande taille dans l'Eocène de Provence ["Deposits of eggs from birds of very large size from the Eocene of Provence"]|journal=C. R. Acad. Sci. Paris|volume=250|issue=23|pages= 3870–3871|language=fr|url=http://gallica.bnf.fr/ark:/12148/bpt6k750m}} These were described as the ootaxon Ornitholithus and are presumably from Gastornis. While no direct association exists between Ornitholithus and Gastornis fossils, no other birds of sufficient size are known from that time and place; while the large Diogenornis and Eremopezus are known from the Eocene, the former lived in South America (still separated from North America by the Tethys Ocean then) and the latter is only known from the Late Eocene of North Africa, which also was separated by an (albeit less wide) stretch of the Tethys Ocean from Europe.{{cite journal|author=Buffetaut, Eric|year=2004|title= Footprints of Giant Birds from the Upper Eocene of the Paris Basin: An Ichnological Enigma|journal=Ichnos|volume=11|issue=3–4|pages= 357–362|doi=10.1080/10420940490442287|bibcode=2004Ichno..11..357B }}

Some of these fragments were complete enough to reconstruct a size of 24 by 10 cm (about 9.5 by 4 inches) with shells 2.3–2.5 mm (0.09–0.1 in) thick, roughly half again as large as an ostrich egg and very different in shape from the more rounded ratite eggs. If Remiornis is indeed correctly identified as a ratite (which is quite doubtful, however), Gastornis remains as the only known animal that could have laid these eggs. At least one species of Remiornis is known to have been smaller than Gastornis, and was initially described as Gastornis minor by Mlíkovský in 2002. This would nicely match the remains of eggs a bit smaller than those of the living ostrich, which have also been found in Paleogene deposits of Provence, were it not for the fact that these eggshell fossils also date from the Eocene, but no Remiornis bones are yet known from that time.

File:Gastornis restoration.jpg

Several sets of fossil footprints are suspected to belong to Gastornis. One set of footprints was reported from late Eocene gypsum at Montmorency and other locations of the Paris Basin in the 19th century, from 1859 onwards. Described initially by Jules Desnoyers, and later on by Alphonse Milne-Edwards, these trace fossils were celebrated among French geologists of the late 19th century. They were discussed by Charles Lyell in his Elements of Geology as an example of the incompleteness of the fossil record – no bones had been found associated with the footprints.{{cite book|author=Lyell, Charles|year=1865|title= Elements of Geology |edition=6th |publisher=J. Murray|url=https://archive.org/details/elementsgeology01lyelgoog}} Unfortunately, these fine specimens, which sometimes even preserved details of the skin structure, are now lost. They were brought to the Muséum national d'histoire naturelle when Desnoyers started to work there, and the last documented record of them deals with their presence in the geology exhibition of the MNHN in 1912. The largest of these footprints, although only consisting of a single toe's impression, was 40 cm (16 in) long. The large footprints from the Paris Basin could also be divided into huge and merely large examples, much like the eggshells from southern France, which are 20 million years older.

Another footprint record consists of a single imprint that still exists, though it has proven to be even more controversial. It was found in late Eocene Puget Group rocks in the Green River valley near Black Diamond, Washington. After its discovery, it raised considerable interest in the Seattle area in May–July 1992, being subject of at least two longer articles in the Seattle Times.{{cite web|author=Dietrich, B.|title= 'Big Bird' Footprint Has Scientists Aflutter – If Proven, Fossil Find Would Be A State First|work=Seattle Times|date=3 May 1992a|page= B1–2|url=https://archive.seattletimes.com/archive/19920503/1489721/big-bird-footprint-has-scientists-aflutter----if-proven-fossil-find-would-be-a-state-first }}{{cite web|author=Dietrich, B.|title= Track Is Hoax, Paleontologists Say – Expert On Prehistoric Bird Casts Doubt On Discovery In State Park|work=Seattle Times|date=17 July 1992b|page= B4 |url=https://archive.seattletimes.com/archive/19920717/1502608/track-is-hoax-paleontologists-say----expert-on-prehistoric-bird-casts-doubt-on-discovery-in-state-park }} Variously declared a hoax or genuine, this apparent impression of a single bird foot measures about {{cvt|27|cm|in}} wide by {{cvt|32|cm|in}} long and lacks a hallux (hind toe); it was described as the ichnotaxon Ornithoformipes controversus. Fourteen years after the initial discovery, the debate about the find's authenticity was still unresolved.{{cite web|author=Doughton, Sandi|title= Big birds on the Green River? The debate continues|work=Seattle Times|date=6 December 2004|url=https://archive.seattletimes.com/archive/20041206/bigbird06m/big-birds-on-the-green-river-the-debate-continues}} The specimen is now at Western Washington University.{{cite web|author=Bigelow, Phil|title=Controversial Patterson "Diatryma footprint" slab has been moved|work=Dinosaur Mailing List|date=2 April 2006|url=http://dml.cmnh.org/2006Apr/msg00010.html|access-date=21 April 2008|archive-date=3 March 2016|archive-url=https://web.archive.org/web/20160303204115/http://dml.cmnh.org/2006Apr/msg00010.html|url-status=dead}}{{cite journal|author1=Patterson, John |author2=Lockley, Martin|year=2004|title= A Probable Diatryma Track from the Eocene of Washington: An Intriguing Case of Controversy and Skepticism|journal=Ichnos|volume=11|issue=3–4|pages= 341–347|doi=10.1080/10420940490442278|bibcode=2004Ichno..11..341P }}Image:Cyperacites filiferus (syn Diatryma? filifera) type specimen.jpgThe problem with these early trace fossils is that no fossil of Gastornis has been found to be younger than about 45 million years. In North America, the fossil record of unequivocal gastornithids seems to end even earlier than in Europe. However, in 2009, a landslide near Bellingham, Washington exposed at least 18 tracks on 15 blocks in the Eocene Chuckanut Formation. The anatomy and age (about 53.7 Ma old{{cite thesis|last1=Breedlovestrout |first1=R. |year=2011 |title= Paleofloristic Studies in the Paleogene Chuckanut Basin, Western Washington, USA |work= Unpublished Phd Dissertation |publisher= University of Idaho, Moscow, Idaho}}) of the tracks suggest that the track maker was Gastornis. Although these birds have long been considered to be predators or scavengers, the absence of raptor-like claws supports earlier suggestions that they were herbivores. The Chuckanut tracks are named as the ichnotaxon Rivavipes giganteus, inferred to belong to the extinct family Gastornithidae. At least 10 of the tracks are on display at Western Washington University.{{cite journal |last1=Mustoe |first1=G. E. |last2=Tucker |first2=D. S. |last3=Kemplin |first3=K. L. |year=2012 |title=Giant Eocene bird footprints from northwest Washington, USA |journal=Palaeontology |volume=55 |issue=6 |pages=1293–1305 |doi=10.1111/j.1475-4983.2012.01195.x |bibcode=2012Palgy..55.1293M |doi-access=free}}

File:Green River Formation Gastornis Feather.jpg. By Darren Naish|alt=]]

The plumage of Gastornis has generally been depicted in art as a hair-like covering similar to some ratites. This has been based in part on some fibrous strands recovered from a Green River Formation deposit at Roan Creek, Colorado, which were initially believed to represent Gastornis feathers and named Diatryma? filifera.{{cite journal|last1=Cockerell |first1=Theodore Dru Alison|year=1923 |title= The Supposed Plumage of the Eocene Bird Diatryma |journal=American Museum Novitates |issue=62 |pages= 1–4 |url=http://digitallibrary.amnh.org/dspace/bitstream/2246/3259/1/N0062.pdf}} Subsequent examination has shown the fossil material to not actually be feathers,{{cite journal|author=Wetmore, Alexander|author-link=Alexander Wetmore|year=1930|title= The Supposed Plumage of the Eocene Diatryma|journal=Auk|volume=47|issue=4|pages= 579–580|url=http://sora.unm.edu/sites/default/files/journals/auk/v047n04/p0579-p0580.pdf|doi=10.2307/4075897|jstor=4075897}} but root fibers and the species renamed as Cyperacites filiferus.{{cite journal |last1=Becker |first1=H. F. |year=1962 |title=Reassignment of Eopuntia to Cyperacites |journal=Bulletin of the Torrey Botanical Club |volume=89 |issue=5 |pages=319–330|doi=10.2307/2482937 |jstor=2482937 }}

A second possible Gastornis feather has since been identified, also from the Green River Formation. Unlike the filamentous plant material, this single isolated feather resembles the body feathers of flighted birds, being broad and vaned. It was tentatively identified as a possible Gastornis feather based on its size; the feather measured {{convert|240|mm|in|abbr=on}} long and must have belonged to a gigantic bird.{{cite book|author=Grande, L.|year=2013|chapter=Birds|title=The Lost World of Fossil Lake: Snapshots from Deep Time|publisher=University of Chicago Press|pages=215–222|url=https://press.uchicago.edu/dam/ucp/books/pdf/9780226922966_sample_pages.pdf}}{{Cite web|url=https://birds.fieldmuseum.org/media-gallery/detail/316/1466|title=Fossilized Feathers {{!}} Zoology, Division of Birds|website=birds.fieldmuseum.org|access-date=22 February 2020}}

It has been argued that Gastornis has a Holarctic distribution with fossils found in western Europe, North America (including an indeterminate specimen identified as Gastornis sp. from Arctic Canada), and possibly central China.{{Cite journal|first1=Thomas A. |last1=Stidham |first2=Jaelyn J. |last2=Eberle |year=2016 |title=The palaeobiology of high latitude birds from the early Eocene greenhouse of Ellesmere Island, Arctic Canada |journal=Scientific Reports |volume=6 |pages=Article number 20912 |doi=10.1038/srep20912 |pmid=26867798 |pmc=4751439|bibcode=2016NatSR...620912S }} The earliest (Paleocene) fossils all come from Europe, and it is likely that the genus originated there. Europe in this epoch was an island continent, and Gastornis was the largest terrestrial tetrapod of the landmass. This offers parallels with the Malagasy elephant birds, herbivorous birds that were similarly the largest land animals in the isolated landmass of Madagascar, in spite of otherwise mammalian megafauna.{{cite journal | author = Buffetaut Eric, Angst Delphine | title = Stratigraphic distribution of large flightless birds in the Palaeogene of Europe and its palaeobiological and palaeogeographical implications | journal = Earth-Science Reviews | year = 2014 | volume = 138 | pages = 394–408| doi = 10.1016/j.earscirev.2014.07.001 | bibcode = 2014ESRv..138..394B }}

All other fossil remains are from the Eocene, though it is currently unknown how the genus Gastornis dispersed out of Europe and into other continents, and whether such assertion is even true given the potential validity of Diatryma. Given the possible presence of Gastornis fossils in the early Eocene of western China, these birds may have spread east from Europe and crossed into North America via the Bering land bridge. Gastornis also may have spread both east and west, arriving separately in eastern Asia and in North America across the Turgai Strait. Direct landbridges with North America are also known. European Gastornis survived somewhat longer than their North American counterparts, which seems to coincide with a period of increased isolation of the continent.

The reason for the extinction of Gastornis is currently unclear. Competition with mammals has often been cited as a possible factor, but Gastornis did occur in faunas dominated by mammals, and did co-exist with several megafaunal forms like pantodonts. Likewise, extreme climatic events like the Paleocene–Eocene Thermal Maximum (PETM) appear to have had little impact. Nonetheless, the extended survival in Europe is thought to coincide with increased isolation of the landmass.

{{Reflist|30em}}

{{Commons category|Gastornis}}

• {{cite web |url =http://2dgf.dk/publikationer/dgf_on_line/vol_1/gastdk.htm |title =The unfinished story of the Early Tertiary giant bird Gastornis |publisher =Geological Society of Denmark |access-date =11 April 2008 |url-status =dead |archive-url =https://web.archive.org/web/20070928142111/http://2dgf.dk/publikationer/dgf_on_line/vol_1/gastdk.htm |archive-date =28 September 2007 }}

{{Maniraptora|N.}}

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Category:Gastornithiformes

Category:Paleogene birds of Europe

Category:Paleogene birds of North America

Category:Fossil taxa described in 1855

Category:Paleocene birds

Category:Eocene birds

Category:Paleocene first appearances

Category:Eocene genus extinctions

Category:Green River Formation

Category:Puget Group