Lithornis Promiscuus
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''Lithornis'' 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 extinct paleognathous birds. Although ''Lithornis'' was able to fly well, their closest relatives are the extant
tinamou Tinamous () are members of the order Tinamiformes (), and family Tinamidae (), divided into two distinct subfamily, subfamilies, containing 46 species found in Mexico, Central America, and South America. The word "tinamou" comes from the Carib la ...
s (which are poor flyers) and
ratite Ratites () are a polyphyletic group consisting of all birds within the infraclass Palaeognathae that lack keels and cannot fly. They are mostly large, long-necked, and long-legged, the exception being the kiwi, which is also the only nocturnal ...
s (which are
flightless bird Flightless birds are birds that cannot Bird flight, fly, as they have, through evolution, lost the ability to. There are over 60 extant species, including the well-known ratites (ostriches, emus, cassowary, cassowaries, Rhea (bird), rheas, an ...
s). Fossils of ''Lithornis'' are known with certainty from the Upper
Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
through the
Middle Eocene The Eocene ( ) is a geological epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''Ēṓs'', ' Dawn') a ...
, but their fossil record may extend to the late Cretaceous. ''Lithornis'' is from ancient Greek for 'stone bird', as it is one of the first fossil birds to become widely discussed. Presumably closely related genera are '' Paracathartes'' and '' Pseudocrypturus''. Some researchers consider '' Calciavis'' as a species of ''Lithornis'' (''L. grandei'').


Species

Six
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
have been recognized in modern times; undescribed ones are also likely to exist. The supposed
tarsometatarsus The tarsometatarsus is a bone that is only found in the lower leg of birds and some non-avian dinosaurs. It is formed from the fusion of several bird bones found in other types of animals, and homologous to the mammalian tarsus (ankle bones) a ...
piece from which ''"Lithornis" emuinus'' was described is actually a
humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
fragment of the giant
pseudotooth bird The Pelagornithidae, commonly called pelagornithids, pseudodontorns, bony-toothed birds, false-toothed birds or pseudotooth birds, are a prehistoric family of large seabirds. Their fossil remains have been found all over the world in rocks dating ...
''
Dasornis ''Dasornis'' is a genus of prehistoric pseudotooth birds. These were probably close relatives of either pelicans and storks or waterfowl; they are placed in the order Odontopterygiformes to account for this uncertainty. Almost all known mater ...
''.Houde, P. (1988)


''Lithornis celetius''

''L. celetius'' is from the Bangtail Quarry, Sedan Quadrangle,
Park County, Montana Park County is a county in the U.S. state of Montana. At the 2020 census, the population was 17,191. Its county seat is Livingston. A small part of Yellowstone National Park is in the southern part of the county. History The Territorial Leg ...
, and was described by Peter Houde (1988). It is from the
Fort Union Formation The Fort Union Formation is a geologic unit containing sandstones, shales, and coal beds in Wyoming, Montana, and parts of adjacent states. In the Powder River Basin, it contains important economic deposits of coal, uranium, and coalbed methane. ...
, which is earliest
Tiffanian The Tiffanian North American Stage on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 60,200,000 to 56,800,000 years BP lasting . It is usually co ...
, Late
Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
. The type fossil is USNM 290601. ''L. celetius'' was of average size for the genus, and the name ''Celetius'' is derived from the
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
word ''keletion'' a race horse for which the type locality is also named.


''Lithornis hookeri''

Harrison and Walker originally labeled the fossil as belonging to the species ''Pediorallus barbarae'' in 1977. In 1984, Harrison redescribed the fossil as coming from a new species ''Pediorallus hookeri'', and later that year it was moved to ''Lithornis hookeri''.Mayr, G. (2008) ''L. hookeri'' is the smallest of the Lithornithidae.


''Lithornis nasi''

W. George collected the original
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
in the Division A
London Clay The London Clay Formation is a Sediment#Shores and shallow seas, marine formation (geology), geological formation of Ypresian (early Eocene Epoch, c. 54-50 million years ago) age which outcrop, crops out in the southeast of England. The London C ...
,
North Sea Basin The North Sea lies between Great Britain, Denmark, Norway, Germany, the Netherlands, Belgium, and France. A sea on the European continental shelf, it connects to the Atlantic Ocean through the English Channel in the south and the Norwegian Se ...
at
Walton on the Naze Walton-on-the-Naze is a seaside town on the North Sea coast. It is part of the parish of Frinton and Walton, in the Tendring district in Essex, England. The town is located north of Clacton and south of the port of Harwich; Frinton-on-Sea lie ...
,
Essex Essex ( ) is a Ceremonial counties of England, ceremonial county in the East of England, and one of the home counties. It is bordered by Cambridgeshire and Suffolk to the north, the North Sea to the east, Kent across the Thames Estuary to the ...
,
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
. The fossil was described by Harrison in 1984 as a rail, ''Pediorallus nasi'', but was later moved to ''Lithornis nasi''. ''L. nasi'' is larger than ''L. hookeri'' and ''L. plebius''. In 2015, ''L. nasi'' was proposed to be a
junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...
of ''L. vulturinus'' because its differences in shape and size from the latter species were concluded to more likely represent intraspecific variation. Mayr and Kitchner (2025) disagreed with the proposed synonymy based on their notable size difference.


''Lithornis plebius''

''L. plebius'' is from the same locality as ''L. promiscuus'' and was also described by Peter Houde (1988). The type specimen is USNM 336534.


''Lithornis promiscuus''

''L. promiscuus'' has
type specimen In biology, a type is a particular wikt:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to ancho ...
USNM 336535 and was described by Peter Houde (1988). It is from the Clark Quadrangle, Park County, Wyoming, USA. It is from the Willwood Formation, which is earliest
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
in age. An egg, USNM 336570, is known for ''L. celetius'' as well.


''Lithornis vulturinus''

''L. vulturinus'' was described as a vulture by Owen (1840) from the holotype fossil 955 738 - TM 024 717. The fossil was collected from
Early Eocene In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
London Clay The London Clay Formation is a Sediment#Shores and shallow seas, marine formation (geology), geological formation of Ypresian (early Eocene Epoch, c. 54-50 million years ago) age which outcrop, crops out in the southeast of England. The London C ...
deposits on the Isle of Sheppey, Kent, England by J. Hunter before 1793. This fossil was destroyed by bombing in
World War II World War II or the Second World War (1 September 1939 – 2 September 1945) was a World war, global conflict between two coalitions: the Allies of World War II, Allies and the Axis powers. World War II by country, Nearly all of the wo ...
. Numerous isolated fossil bones of ''Lithornis vulturinus'' were incorrectly described anew, such as ''Parvigyps praecox'' and ''Promusophaga magnifica'' - the supposed earliest vulture and turaco, while others were referred to existing families of neognathous birds. A
neotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to anchor or centralizes ...
(BMNH A 5204) was erected to replace the holotype in 1988 by Houde, who for the first time diagnosed it as a paleognath based on complete three-dimensional skulls and skeletons of congeners from North America. An exceptionally preserved specimen was collected from Denmark and cataloged as MGUH 26770.Leonard, L. & Van Tuinen, M. (2005)


Palaeobiology

''Lithornis wing bones are similar to those of storks and vultures, meaning that unlike modern tinamous it was capable of soaring flight.Houde, Peter W. (1988). "Paleognathous Birds from the Early Tertiary of the Northern Hemisphere". Publications of the Nuttall Ornithological Club (Cambridge, MA)Paleogene Fossil Birds
/ref> In a study about
ratite Ratites () are a polyphyletic group consisting of all birds within the infraclass Palaeognathae that lack keels and cannot fly. They are mostly large, long-necked, and long-legged, the exception being the kiwi, which is also the only nocturnal ...
endocasts, ''Lithornis'' ranks among the taxa with well developed olfactory lobes. This is consistent with a nocturnal, forest-dwelling lifestyle, though as much all volant birds it retains large optical lobes. Unlike modern tinamous, ''Lithornis'' has toe claws and reversed halluxes that allow for efficient perching. Several egg fossils have been attributed to ''Lithornis''. Their eggshells are, perhaps unsurprisingly, noted as being "ratite-like".Gerald Grellet-Tinner and Gareth J. Dyke, The eggshell of the Eocene bird Lithornis, Acta Palaeontologica Polonica 50 (4), 2005: 831-835


References

Footnotes Sources * * * * {{Taxonbar, from=Q6648154 Fossil taxa described in 1840 Lithornithidae Paleocene birds Eocene birds of Europe Paleogene birds of Europe Bartonian extinctions Paleogene birds of North America