Phocavis
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''Phocavis'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
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
flightless Flightless birds are birds that cannot fly, as they have, through evolution, lost the ability to. There are over 60 extant species, including the well-known ratites ( ostriches, emus, cassowaries, rheas, and kiwis) and penguins. The smal ...
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 ...
, belonging to the family
Plotopteridae Plotopteridae is an extinct family of flightless seabirds with uncertain placement, generally considered as member of order Suliformes. They exhibited remarkable convergent evolution with the penguins, particularly with the now extinct giant peng ...
, and distantly related with modern
cormorant Phalacrocoracidae is a family of approximately 40 species of aquatic birds commonly known as cormorants and shags. Several different classifications of the family have been proposed, but in 2021 the International Ornithologists' Union (IOU) ado ...
s. Its fossils, found in the
Keasey Formation The Keasey Formation is a geologic formation in northwestern Oregon. It preserves fossils dating back to the Paleogene The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million year ...
in
Oregon Oregon ( , ) is a U.S. state, state in the Pacific Northwest region of the United States. It is a part of the Western U.S., with the Columbia River delineating much of Oregon's northern boundary with Washington (state), Washington, while t ...
, are dated from the Late
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 ...
.


History and Etymology

The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
of ''Phocavis'', LACM 123897, an isolated 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 ...
, was collected in Late
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 ...
rocks belonging to the
Keasey Formation The Keasey Formation is a geologic formation in northwestern Oregon. It preserves fossils dating back to the Paleogene The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million year ...
, near
Vernonia, Oregon Vernonia is a city in Columbia County, Oregon, United States. It is located on the Nehalem River, in a valley on the eastern side of the Northern Oregon Coast Range that is the heart of one of the most important timber-producing areas of the ...
, by James L. Goedert in 1979. The fossil was only described in 1988 as a new genus and species of
Plotopteridae Plotopteridae is an extinct family of flightless seabirds with uncertain placement, generally considered as member of order Suliformes. They exhibited remarkable convergent evolution with the penguins, particularly with the now extinct giant peng ...
, ''Phocavis maritimus''. Due to the lack of existing material on the two described genera of
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
n plotopterids, ''
Tonsala ''Tonsala'' is an extinct genus of Plotopteridae, a family of flightless seabird similar in biology with penguins, but more closely related to modern cormorants. The genus is known from terrains dated from the Late Oligocene of the State of Washi ...
'' and '' Plotopterum'', the validity of the genus was only assumed due to the assumed size of the living bird, estimated to be intermediate between the two later genera, and its much older geological age. Comparison could however be made with the then undescribed remains of ''
Copepteryx ''Copepteryx'' is an extinct genus of flightless bird, flightless bird of the family Plotopteridae, endemic to Japan during the Oligocene living from 28.4 to 23 Annum, mya, meaning it existed for approximately . History and Etymology Remains of ...
'' from
Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...
to verify the identity of the bone as a fossil of plotopterid. In 2004, in an heavily criticized article in which he considered the
Spheniscidae Penguins are a group of Water bird, aquatic flightless birds from the family (biology), family Spheniscidae () of the order (biology), order Sphenisciformes (). They live almost exclusively in the Southern Hemisphere. Only one species, the G ...
to be related to
Suliformes The order Suliformes (, dubbed "Phalacrocoraciformes" by ''Christidis & Boles 2008'') is an order of birds recognised by the International Ornithological Congress, International Ornithologist's Union. Regarding the recent evidence that the tradit ...
based on their shared similarities with plotopterids,
Gerald Mayr Gerald Mayr is a German palaeontologist who is Curator of Ornithology at the Senckenberg Research Institute in Frankfurt am Main, Hesse. He has published extensively on fossil birds, especially the Paleogene avifauna of Europe.Mayr, Gerald (2016). ...
noted that the tarsometatarsus of ''Phocavis'' shared similarities with that of the Eocene
frigatebird Frigatebirds are a Family (biology), family of seabirds called Fregatidae which are found across all tropical and subtropical oceans. The five extant species are classified in a single genus, ''Fregata''. All have predominantly black plumage, l ...
''
Limnofregata ''Limnofregata'' ("Freshwater frigatebird") is an extinct genus of primitive frigatebird. The two known species were described after fossils from the Early Eocene Green River Formation (c. 49 million years ago) of Wyoming. A number of good comple ...
'', and tentatively assigned ''Phocavis'' as the sister taxon of a clade including plotopterids and modern-day
penguins Penguins are a group of aquatic flightless birds from the family Spheniscidae () of the order Sphenisciformes (). They live almost exclusively in the Southern Hemisphere. Only one species, the Galápagos penguin, is equatorial, with a sm ...
. These affirmations have later be rebutted, on the basis of the more robust and larger shape of the bone, as well as similarities with more derived plotopterids. In 2016,
Gerald Mayr Gerald Mayr is a German palaeontologist who is Curator of Ornithology at the Senckenberg Research Institute in Frankfurt am Main, Hesse. He has published extensively on fossil birds, especially the Paleogene avifauna of Europe.Mayr, Gerald (2016). ...
and James L. Goedert suggested that some of the remains attributed to ''Phocavis'' were virtually indifferentiable from those of its later relative, '' Tonsala hildegardae''.


Etymology

The genus name, ''Phocavis'', is constructed with the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
prefix “''Phoca''-”, meaning
seal Seal may refer to any of the following: Common uses * Pinniped, a diverse group of semi-aquatic marine mammals, many of which are commonly called seals, particularly: ** Earless seal, also called "true seal" ** Fur seal ** Eared seal * Seal ( ...
, and the suffix “''-avis''”, meaning bird, as a reference of its supposed adaptation towards swimming. The
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
name, ''maritimus'', means in Latin “from the sea”.


Description

''Phocavis'' is only known from its holotype tarsometatarsus, which could be identified as belonging to a new genus of plotopterid thanks to comparison with fossil remains of large plotopterids from
Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...
. The 60 mm large tarsometatarsus was much smaller than its Japanese counterparts, but larger than its later relative '' Plotopterum maritimus''. It was also more elongated, and anatomically distinct from the Japanese plotopterids. The primitiveness of the tarsometatarsus compared to that of the latter ''
Copepteryx ''Copepteryx'' is an extinct genus of flightless bird, flightless bird of the family Plotopteridae, endemic to Japan during the Oligocene living from 28.4 to 23 Annum, mya, meaning it existed for approximately . History and Etymology Remains of ...
'' indicates that ''Phocavis'' represented a considerably primitive form compared to its
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
counterparts.


Paleoenvironment

''Phocavis'' is the earliest and presumably most basal known genus of plotopterid, dating from the Late Eocene. The Keasey Formation, in which it was found, represents deep sea deposits, at an approximate deposition depth of 500 to 1000 m, possibly indicating that plotopterids were able to venture far from the coast early in their evolution. In the Eocene of this formation are also represented a pseudodontorn, a
marine turtle Sea turtles (superfamily Chelonioidea), sometimes called marine turtles, are reptiles of the order Testudines and of the suborder Cryptodira. The seven existing species of sea turtles are the flatback, green, hawksbill, leatherback, loggerhe ...
, a
cetacean Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...
,
teleost fish Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
and
shark Sharks are a group of elasmobranch cartilaginous fish characterized by a ribless endoskeleton, dermal denticles, five to seven gill slits on each side, and pectoral fins that are not fused to the head. Modern sharks are classified within the ...
s. The cohabitation between plotopterids and early cetaceans since the Eocene goes against an earlier theory justifying the extinction of the plotopterids during the
Miocene The Miocene ( ) is the first epoch (geology), 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 mea ...
by an unequal concurrence with
porpoise Porpoises () are small Oceanic dolphin, dolphin-like cetaceans classified under the family Phocoenidae. Although similar in appearance to dolphins, they are more closely related to narwhals and Beluga whale, belugas than to the Oceanic dolphi ...
s, and may designate other culprits, such as the dramatic
climatic Climate is the long-term weather pattern in a region, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorolog ...
and oceanographic changes in the North Pacific during the Early and Middle Miocene, affecting
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater Aquatic ecosystem, ecosystems. The name comes from the Greek language, Greek words (), meaning 'plant', and (), mea ...
ic communities and causing the extinction of the specialized plotopterids, in favor of other group such as the enaliarctine
pinniped Pinnipeds (pronounced ), commonly known as seals, are a widely range (biology), distributed and diverse clade of carnivorous, fin-footed, semiaquatic, mostly marine mammals. They comprise the extant taxon, extant families Odobenidae (whose onl ...
s. ''Phocavis'' was contemporaneous of at least one other distinct species of plotopterid, colloquially known as the "Whiskey Creek plotopterid", from the Late Eocene of the Makah Formation.


References

{{Taxonbar, from=Q87705879 Fossil taxa described in 1988 Plotopteridae Extinct flightless birds Prehistoric bird genera