Palaeochenoides Mioceanus
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''Palaeochenoides'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of the
prehistoric Prehistory, also known as pre-literary history, is the period of human history between the use of the first stone tools by hominins 3.3 million years ago and the beginning of recorded history with the invention of writing systems. The use of ...
pseudotooth birds of somewhat doubtful validity. 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 ...
s and
stork Storks are large, long-legged, long-necked wading birds with long, stout bills. They belong to the family called Ciconiidae, and make up the order Ciconiiformes . Ciconiiformes previously included a number of other families, such as herons an ...
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 in ...
, and are here placed in the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
Odontopterygiformes to account for this uncertainty. Only a single
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, ''Palaeochenoides mioceanus'', is known to date. The first
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 preserved ...
assigned to it – a
distal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
right
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
piece – was found near the source of the
Stono River The Stono River or Creek is a tidal channel in southeast South Carolina, located southwest of Charleston. The channel runs southwest to northeast between the mainland and Wadmalaw Island and Johns Island, from north Edisto River between Johns ...
in
Charleston County, South Carolina Charleston County is located in the U.S. state of South Carolina along the Atlantic coast. As of the 2020 census, its population was 408,235, making it the third most populous county in South Carolina (behind Greenville and Richland counties). ...
(United States). At first it was believed to be from the Early Miocene
Hawthorne Formation Hawthorne often refers to the American writer Nathaniel Hawthorne. Hawthorne may also refer to: Places Australia *Hawthorne, Queensland, a suburb of Brisbane Canada *Hawthorne Village, Ontario, a suburb of Milton, Ontario United States *Hawtho ...
– its
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 ...
, seemingly referring to the "
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
ocean The ocean (also the sea or the world ocean) is the body of salt water that covers approximately 70.8% of the surface of Earth and contains 97% of Earth's water. An ocean can also refer to any of the large bodies of water into which the wo ...
" as presumed
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
but actually a simple spelling error for "miocaenus", "from the Miocene", that was never corrected and hence became valid – alludes to this. But in fact no Hawthorne Formation rocks were known in the Charleston region when the fossil was found, and consequently modern authors consider a
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) age more likely and suggest the fossil came from 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), alias of Dutch producers Klubbheads * Cooper (video game character), in ...
or
Chandler Bridge Formation The Chandler Bridge Formation is a geologic formation in South Carolina. It preserves fossils dating back to the Chattian (Late Oligocene) of the Paleogene period, corresponding to the Arikareean in the NALMA classification.
. Specimen MCZ 2514, a distal left
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 bones found in other types of animals, and homologous to the mammalian tarsus (ankle bones) and meta ...
fragment from the
Ashley River The Ashley River is a blackwater and tidal river in South Carolina, rising from the Wassamassaw and Great Cypress Swamps in western Berkeley County. It consolidates its main channel about five miles west of Summerville, widening into a ti ...
, was more tentatively assigned to ''P. mioceanus'' later on; it was also erroneously believed to be from the Hawthorne Formation.


Description and systematics

The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
femur's classification mirrors the recently renewed uncertainties about the pseudotooth birds' placement. At the time of its description, when it was still much surrounded by
matrix Matrix most commonly refers to: * ''The Matrix'' (franchise), an American media franchise ** ''The Matrix'', a 1999 science-fiction action film ** "The Matrix", a fictional setting, a virtual reality environment, within ''The Matrix'' (franchis ...
, it was believed to be from a giant
goose A goose ( : geese) is a bird of any of several waterfowl species in the family Anatidae. This group comprises the genera ''Anser'' (the grey geese and white geese) and '' Branta'' (the black geese). Some other birds, mostly related to the ...
or
swan Swans are birds of the family (biology), family Anatidae within the genus ''Cygnus''. The swans' closest relatives include the goose, geese and ducks. Swans are grouped with the closely related geese in the subfamily Anserinae where they form t ...
. This is referred to in the genus name, which means "ancient goose-like
ird IRD or Ird may refer to the following: * Ird (Bedouin), a Bedouin honor code for women * Ird, alternate name of Arad, Iran, a city in Fars Province * Ishwardi Airport (IATA airport code) * Kaarel Ird (1909–1986), Estonian theatre leader, director ...
. But only one year later, the bone had been prepared from the matrix and was submitted to an improvised phenetic analysis of its details. It was compared to that of
Anserinae The Anserinae are a subfamily in the waterfowl family Anatidae. It includes the swans and true geese. Under alternative systematical concepts (see e.g., Terres & NAS, 1991), it is split into two subfamilies, the Anserinae contain the geese and ...
and
Dendrocygninae The whistling ducks or tree ducks are a subfamily, Dendrocygninae, of the duck, goose and swan family of birds, Anatidae. In other taxonomic schemes, they are considered a separate family, Dendrocygnidae. Some taxonomists list only one genus, ...
(other
Anseriformes 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 in ...
were either similar to these or too unlike ''P. mioceanus''), as well as with Pelecanidae,
Phaethontidae Tropicbirds are a family, Phaethontidae, of tropical pelagic seabirds. They are the sole living representatives of the order Phaethontiformes. For many years they were considered part of the Pelecaniformes, but genetics indicates they are most cl ...
and
Phalacrocoraciformes The order Suliformes (, dubbed "Phalacrocoraciformes" by ''Christidis & Boles 2008'') is an order recognised by the International Ornithologist's Union. In regard to the recent evidence that the traditional Pelecaniformes is polyphyletic, it has ...
of the "higher waterbird" radiation, and found to resemble the former in one, the latter in 4 out of 5 traits. Some minor details of the femur were also unlike in the Anseriformes. The study proposed the fossil bird to have been "a large steganopod ..somewhat larger than '' Pelecanus erythrorhynchos'' or '' P. onocrotalus''," and not securely assignable to a known
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
. But as cladistic analysis had not yet been developed, it was of course not studied whether the similarities between ''P. mioceanus'' and the "higher waterbirds" are
apomorph In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
ies. It has been noted, however, that
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...
to oceanic habits, has induced in pseudotooth bird legs and feet features convergent to other
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
s. Still, the fossil is a rather good specimen, considering how little humerus material known from pseudotooth birds; a recent cladistic analysis (which did not include ''P. mioceanus'') did not put much emphasis on humerus traits, for example, as so few good fossils are known. When
Witmer Stone Witmer Stone (September 22, 1866 – May 24, 1939) was an American ornithologist, botanist, and mammalogist, and was considered one of the last of the “great naturalists.” Stone is remembered principally as an ornithologist. He was president ...
, then editor of ''
The Auk ''Ornithology'', formerly ''The Auk'' and ''The Auk: Ornithological Advances'', is a peer-reviewed scientific journal and the official publication of the American Ornithological Society (AOS). It was established in 1884 and is published quarterly. ...
'', commented on the
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
confusion that the bird caused during less than 25 months after its description, he noted:
"It would seem desirable that those who name fossil birds should not fashion their generic names on those of existing birds as it is embarrassing to find them shifted into other families or orders where the name becomes somewhat of a misnomer!"
Of the
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 bones found in other types of animals, and homologous to the mammalian tarsus (ankle bones) and meta ...
, only the end remains, but this is complete and even not very much abraded. Only on the edges of the outer trochleae does significant material seem to be lost. Altogether, the bone is very similar to that of the
sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species sh ...
and probably contemporary '' Tympanonesiotes wetmorei'', only appearing a bit less
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 Pacifi ...
-like. The spread of the toes must have resembled that found in a fulmar quite a lot, by contrast. The thin-walled bone has a second
toe Toes are the digits (fingers) 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 ''plant ...
trochlea that attaches notably kneewards from the others and is angled slightly outwards while the
hallux Toes are the digits (fingers) 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 '' pl ...
was vestigial or missing, as is typical for the pseudotooth birds. The fossil is about one-quarter larger than that of ''Tympanonesiotes'', with a maximum end width of – probably a bit more in life, as the trochlea rims are eroded away. The shaft is wide at the point where it flares into the trochleae. It was thus more than twice 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 or lowest stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by the Eocene Lutetian Age. The Ypresian i ...
of the Ouled Abdoul Basin (
Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
), and – like '' Osteodontornis orri'' – thus belonged to the large pseudotooth birds, with a wingspan of more than 5, perhaps as much as 6 m (16–20 ft). Its bill was presumably long and massive as in its relatives, and in length the bird exceeded the largest
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 ...
s. The early systematic uncertainties have been noted again in recent times, in the scope of the debate whether the pseudotooth birds were
Galloanseres Fowl are birds belonging to one of two biological orders, namely the gamefowl or landfowl (Galliformes) and the waterfowl ( Anseriformes). Anatomical and molecular similarities suggest these two groups are close evolutionary relatives; togethe ...
closely related to waterfowl or
Neoaves Neoaves is a clade that consists of all modern birds (Neornithes or Aves) with the exception of Paleognathae (ratites and kin) and Galloanserae (ducks, chickens and kin). Almost 95% of the roughly 10,000 known species of extant birds belong to ...
related to pelicans. Meanwhile, as a presumed pelican relative, ''P. mioceanus'' was allied with ''
Cyphornis ''Cyphornis'' is a genus of the prehistoric pseudotooth birds. These were probably rather close relatives of either pelicans and storks, or of waterfowl, and are here placed in the order Odontopterygiformes to account for this uncertainty. Descr ...
'' (another pseudotooth bird, known only from a
proximal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
left
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 bones found in other types of animals, and homologous to the mammalian tarsus (ankle bones) and meta ...
) and placed in a
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Cyphornithidae. This was subsequently assigned to a "pelecaniform"
suborder Order ( la, ordo) 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 ...
Cladornithes. But the enigmatic Late Oligocene ''
Cladornis The Pelecaniformes are an order of medium-sized and large waterbirds found worldwide. As traditionally—but erroneously—defined, they encompass all birds that have feet with all four toes webbed. Hence, they were formerly also known by such n ...
'' – the type genus of that supposed suborder – from the
Argentinian Argentines (mistakenly translated Argentineans in the past; in Spanish (masculine) or ( feminine)) are people identified with the country of Argentina. This connection may be residential, legal, historical or cultural. For most Argentines, ...
part of Patagonia is known from a
distal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
right tarsometatarsus only, and thus was not directly comparable to ''Palaeochenoides'' and ''Cyphornis''. '' Tympanonesiotes wetmorei'', yet another pseudotooth bird, known from a distal right tarsometatarsus fragment, was initially presumed to be similar to ''Cladornis'' however, and also to ''Cyphornis''; consequently, the three genera were allied simply because of their size and some similarities to the corresponding bones of pelicans. Today however, ''Cladornis'' is more generally held to be a terrestrial bird rather than a
seabird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
. As regards the supposed Cyphornithidae, most if not all pseudotooth birds placed there are probably closely related to the better-known ''
Pelagornis ''Pelagornis'' is a widespread genus of prehistoric pseudotooth birds. These were probably rather close relatives of either pelicans and storks, or of waterfowl, and are here placed in the order Odontopterygiformes to account for this uncertain ...
'', type genus of the family Pelagornithidae. And even if ''Cyphornis'' is the senior synonym of ''Palaeochenoides'' and ''Tympanoneisiotes'' (which is not overly likely due to size differences), according to the rules of zoological nomenclature the family name Pelagornithidae would not be affected. Cyphornithidae would almost certainly be a
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...
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 the smaller ''Tympanoneisiotes'' appear to be very closely related and are probably part of a
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
lineage of (usually) giant pseudotooth birds. As at the time of its description no adequate comparative material was known, it was explicitly cautioned that ''P. mioceanus'' might include ''Pseudodontornis'' – which may actually be synonymous with ''Pelagornis''.Olson (1985: pp.195-197), Mlíkovský (2002: p.81), Mayr (2009: pp.58-59)


Footnotes


References


Supplement
* * Christidis, Les & Boles, Walter E. (2008): ''Systematics and Taxonomy of Australian Birds''. CSIRO Publishing, CollingwoodVictoria, Australia. * *Mayr, Gerald (2009): ''Paleogene Fossil Birds''. Springer-Verlag, Heidelberg & New York. * *Mlíkovský, Jirí (2002)
''Cenozoic Birds of the World, Part 1: Europe''
Ninox Press, Prague. *Olson, Storrs L. (1985)
The Fossil Record of Birds
''In:'' Farner, D.S.; King, J.R. & Parkes, Kenneth C. (eds.): ''Avian Biology'' 8: 79-252. * * * {{Taxonbar, from1=Q42351553, from2=Q7126399 Pelagornithidae Oligocene birds Extinct animals of North America Prehistoric bird genera