Tympanonesiotes
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''Tympanonesiotes'' is a somewhat doubtfully valid
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 the
prehistoric Prehistory, also called pre-literary history, is the period of human history between the first known use of stone tools by hominins  million years ago and the beginning of recorded history with the invention of writing systems. The use o ...
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 ...
s. These were probably rather close relatives of either
pelican Pelicans (genus ''Pelecanus'') are a genus of large water birds that make up the family Pelecanidae. They are characterized by a long beak and a large throat pouch used for catching prey and draining water from the scooped-up contents before ...
s and
stork Storks are large, long-legged, long-necked wading birds with long, stout bills. They belong to the family Ciconiidae, and make up the order Ciconiiformes . Ciconiiformes previously included a number of other families, such as herons and ibise ...
s, or of
waterfowl Anseriformes is an order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (three species of screamers), Anseranatidae (the magpie goose), and Anatidae, the largest family, which i ...
, and are here placed in the
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
Odontopterygiformes to account for this uncertainty.


Species and taxonomy

Only a single
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), ...
, ''Tympanonesiotes wetmorei'', is known to date. The only known specimen (
USNM The National Museum of Natural History (NMNH) is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. With 4.4 ...
16809), a
distal Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position provi ...
right
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 ...
end, was found in the Cooper River near Drum Island (Charleston) at
Charleston, South Carolina Charleston is the List of municipalities in South Carolina, most populous city in the U.S. state of South Carolina. The city lies just south of the geographical midpoint of South Carolina's coastline on Charleston Harbor, an inlet of the Atla ...
(United States). At first it was believed to be from the
Early Miocene The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages: the Aquitanian age, Aquitanian and Burdigalian stages. The sub-epoch lasted from 23.03 ± 0.05 annum, Ma to ...
Hawthorne Formation, but as it seems its actual age cannot be precisely determined: For one thing, no Hawthorne Formation deposits were known in the region where the fossil was found. However, close to its type locality,
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 ...
s of
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
animals have been found reworked from a now-
eroded Erosion is the action of surface processes (such as water flow or wind) that removes soil, rock, or dissolved material from one location on the Earth's crust and then transports it to another location where it is deposited. Erosion is disti ...
layer of rock into older deposits, such as the
Chattian The Chattian is, in the geologic timescale, the younger of two ages or upper of two stages of the Oligocene Epoch/Series. It spans the time between . The Chattian is preceded by the Rupelian and is followed by the Aquitanian (the lowest stage ...
(Late Oligocene) sediments of the
Cooper Cooper, Cooper's, Coopers and similar may refer to: * Cooper (profession), a maker of wooden casks and other staved vessels Arts and entertainment * Cooper (producers), an alias of Dutch producers Klubbheads * "Cooper", a song by Roxette from ...
or
Chandler Bridge Formation The Chandler Bridge Formation is a Formation (geology), geologic formation in South Carolina. It preserves fossils dating back to the Chattian (Late Oligocene) of the Paleogene Period (geology), period, corresponding to the Arikareean in the Nort ...
where the specimen of ''T. wetmorei'' was presumably found. The much-worn pseudotooth bird bone may also be such a reworked specimen. The genus'
scientific name In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
references the type locality: it is derived from
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
''tympanon'' (drum) + ''nesiotes'' (islander). The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
honors the famous
ornithologist Ornithology, from Ancient Greek ὄρνις (''órnis''), meaning "bird", and -logy from λόγος (''lógos''), meaning "study", is a branch of zoology dedicated to the study of birds. Several aspects of ornithology differ from related discip ...
Alexander Wetmore Frank Alexander Wetmore (June 18, 1886 – December 7, 1978) was an American ornithologist and avian paleontologist. He was the sixth Secretary of the Smithsonian Institution. He was also an elected member of both the American Philosophical Soc ...
; thus the scientific name means roughly "Alexander Wetmore's Drum Island bird".


Description

The bone is not very well preserved; for most of its length only the
anterior Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
surface remains. What remains of the trochleae is still preserved in good detail however. Altogether, the bone is very similar to that of the
sympatric In biology, two closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. An initially interbreeding population that splits into two or more distinct spe ...
and probably contemporary ''
Palaeochenoides mioceanus ''Palaeochenoides'' is a genus of the prehistoric pseudotooth birds of somewhat doubtful validity. These were probably rather close relatives of either pelicans and storks, or of waterfowl, and are here placed in the order Odontopterygiformes to ...
'', only appearing a bit more
albatross Albatrosses, of the biological family Diomedeidae, are large seabirds related to the procellariids, storm petrels, and diving petrels in the order Procellariiformes (the tubenoses). They range widely in the Southern Ocean and the North Paci ...
-like. The spread of the toes must have resembled that found in a
fulmar The fulmars are tube-nosed seabirds in the family Procellariidae. The family includes two extant species, and two extinct fossil species from the Miocene. Fulmars superficially resemble gulls, but are readily distinguished by their flight on s ...
quite a lot, by contrast. The thin-walled bone has a second
toe Toes are the digits of the foot of a tetrapod. Animal species such as cats that walk on their toes are described as being ''digitigrade''. Humans, and other animals that walk on the soles of their feet, are described as being ''plantigrade''; ...
trochlea that attaches notably kneewards from the others and is angled slightly outwards while the
hallux Toes are the digits of the foot of a tetrapod. Animal species such as cats that walk on their toes are described as being ''digitigrade''. Humans, and other animals that walk on the soles of their feet, are described as being ''plantigrade''; ...
was vestigial or missing, as is typical for the pseudotooth birds. The fossil is about one-quarter smaller than that of ''Palaeochenoides'', with a maximum end width of (as far as the trochleae are preserved), and a shaft that is wide near the point where it flares into the trochleae. It was thus about half again as large as " Odontoptila inexpectata" from the
Late Paleocene The Thanetian is, in the International Commission on Stratigraphy, ICS Geologic timescale, the latest age (geology), age or uppermost stage (stratigraphy), stratigraphic stage of the Paleocene epoch (geology), Epoch or series (stratigraphy), Serie ...
/
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 ...
of the Ouled Abdoul Basin (
Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
), i.e. around the size of the Early Eocene '' Odontopteryx toliapica'' from
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 ...
or slightly bigger, or about the size of a
great albatross The great albatrosses are seabirds in the genus ''Diomedea'' in the albatross family (biology), family. The genus ''Diomedea'' formerly included all albatrosses except the sooty albatrosses, but in 1996 the genus was split, with the mollymawks a ...
(''Diomedea'') of our time. If the ''Tympanonesiotes'' fossil is indeed of Miocene age, it would be among the very few pseudotooth birds known from the
Neogene The Neogene ( ,) is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period million years ago. It is the second period of th ...
that was not of immense size.


Systematics

''T. wetmorei'' was initially placed in the presumed pelecaniform
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 ...
Cyphornithidae, which had been placed in the " pelecaniform"
suborder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and recognized ...
Cladornithes, together with two other misidentified pseudotooth birds – ''Palaeochenoides'' and the family's
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
'' Cyphornis''. The
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
of that supposed suborder, the enigmatic Late Oligocene '' Cladornis'' from the
Argentinian Argentines, Argentinians or Argentineans are people from Argentina. This connection may be residential, legal, historical, or cultural. For most Argentines, several (or all) of these connections exist and are collectively the source of their ...
part of
Patagonia Patagonia () is a geographical region that includes parts of Argentina and Chile at the southern end of South America. The region includes the southern section of the Andes mountain chain with lakes, fjords, temperate rainforests, and glaciers ...
, like the present species is only known from a distal right tarsometatarsus end. This was believed to be reminiscent of the (then still undescribed) ''Tympanonesiotes'', and thus it was argued that all four genera were closely related. But ''Cladornis'' is more generally held to be a
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 ...
bird of unclear affiliations rather than a
seabird Seabirds (also known as marine birds) are birds that are adaptation, adapted to life within the marine ecosystem, marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent ...
nowadays, and the Cladornithes are not used anymore by recent authors. As regards the supposed Cyphornithidae, most if not all pseudotooth birds placed there are probably closely related to the better-known ''
Pelagornis ''Pelagornis'' is an extinct genus of prehistoric pseudotooth birds, a group of extinct seabirds. Species span from the Oligocene to the Early Pleistocene. Members of ''Pelagornis'' represent among the largest pseudotooth birds, with one species ...
'',
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
of the family Pelagornithidae. And even if ''Cyphornis'' is the
senior 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 ''Palaeochenoides'' and ''Tympanoneisiotes'' (which is not overly likely), according to the rules of
zoological nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its formal author, t ...
the family name Pelagornithidae would not be affected. Thus Cyphornithidae would almost certainly 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 Pelagornithidae even if the pseudotooth birds are (as some have proposed) divided into several families – rather than being all placed in the Pelagornithidae as is usual nowadays – as ''Cyphornis'', ''
Osteodontornis ''Osteodontornis'' is an extinct seabird genus. It contains a single named species, ''Osteodontornis orri'' (Orr's bony-toothed bird, in literal translation of its scientific name), which was described quite exactly one century after the first sp ...
'', ''Palaeochenoides'', ''Pelagornis'' and perhaps ''Tympanoneisiotes'' appear to be very closely related and are probably part of a
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
lineage of (usually) giant pseudotooth birds.Olson (1985: pp. 195-197), Mlíkovský (2002: p.81), Mayr (2009: pp. 58-59)


Footnotes


References

* (2005): Osteological evidence for sister group relationship between pseudo-toothed birds (Aves: Odontopterygiformes) and waterfowls (Anseriformes). ''
Naturwissenschaften ''The Science of Nature'', formerly ''Naturwissenschaften'', is a monthly peer-reviewed scientific journal published by Springer Science+Business Media covering all aspects of the natural sciences relating to questions of biological significance. I ...
'' 92(12): 586–591. (HTML abstract
Electronic supplement
(requires subscription) * (1963): Catalogue of fossil birds. Part 1 (Archaeopterygiformes through Ardeiformes). ''Bulletin of the Florida State Museum, Biological Sciences'' 7(4): 179–293
PDF or JPEG fulltext
* (1964): ''Pseudodontornis'' and other large marine birds from the Miocene of South Carolina. ''Postilla'' 83: 1–19
Fulltext
at the
Internet Archive The Internet Archive is an American 501(c)(3) organization, non-profit organization founded in 1996 by Brewster Kahle that runs a digital library website, archive.org. It provides free access to collections of digitized media including web ...
* (1999): ''International Code of Zoological Nomenclature'' (4th ed.). International Trust for Zoological Nomenclature, London. HTML fulltext
* (2009): ''Paleogene Fossil Birds''. Springer-Verlag, Heidelberg & New York. * (2002): ''Cenozoic Birds of the World, Part 1: Europe''. Ninox Press, Prague. PDF fulltext
!-- This should be treated with extreme caution as regards merging of species. Splits are usually good though. See also critical review in Auk121:623-627 here http://www.findarticles.com/p/articles/mi_qa3793/is_200404/ai_n9396879 --> * (1985): The Fossil Record of Birds. ''In:'' : ''Avian Biology'' 8: 79-252
PDF fulltext
* (1972): Bone of a presumed odontopterygian bird from the Miocene of New Zealand. ''New Zealand Journal of Geology and Geophysics'' 15(2): 269–274.
Fulltext
at
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{{Taxonbar, from=Q7860421 Pelagornithidae Prehistoric birds of North America Cenozoic birds of North America