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''Priscomyzon riniensis'' is an extinct
lamprey Lampreys (sometimes inaccurately called lamprey eels) are a group of Agnatha, jawless fish comprising the order (biology), order Petromyzontiformes , sole order in the Class (biology), class Petromyzontida. The adult lamprey is characterize ...
that lived some 360 million years ago during the
Famennian The Famennian is the later of two faunal stages in the Late Devonian epoch. The most recent estimate for its duration is that it lasted from around 371.1 to 359.3 million years ago. An earlier 2012 estimate, still used by the International Commis ...
(
Late Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era during the Phanerozoic eon, spanning 60.3 million years from the end of the preceding Silurian period at million years ago ( Ma), to the beginning of the succeeding ...
) in a marine or estuarine environment in
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. This small
agnathan Agnatha (; ) or jawless fish is a paraphyletic infraphylum of animals in the subphylum Vertebrata of the phylum Chordata, characterized by the lack of jaws. The group consists of both living ( cyclostomes such as hagfishes and lampreys) and ext ...
is anatomically similar to the
Mazon Creek The Mazon Creek fossil beds are a conservation ' found near Morris, in Grundy County, Illinois. The fossils are preserved in ironstone concretions, formed approximately in the mid- Pennsylvanian epoch of the Carboniferous period. These concr ...
lampreys, but is some 35 million years older. Its key developments included the first known large oral disc, circumoral teeth and a branchial basket.


Context

Though common and diverse during the
Silurian The Silurian ( ) is a geologic period and system spanning 23.5 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the third and shortest period of t ...
and
Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
, jawless fish are today represented only by lampreys and
hagfish Hagfish, of the Class (biology), class Myxini (also known as Hyperotreti) and Order (biology), order Myxiniformes , are eel-shaped Agnatha, jawless fish (occasionally called slime eels). Hagfish are the only known living Animal, animals that h ...
, both groups being quite specialized. Lampreys have seven gill pouches (whereas jawed fish have only five), no paired fins, and a rudimentary skeleton of cartilage. They also have a sucker disc of cartilage surrounded by a soft lip and a central small mouth set about with simple hooked teeth. They attach to the bodies of other vertebrates by suction, securing their grip with the hooked teeth, after which a rasped tongue scrapes a hole providing access to the host's softer tissues.


Implications of the find

''Priscomyzon'' provides evidence that agnathans close to modern lampreys had existed before the end of the
Devonian period The Devonian ( ) is a geologic period and system of the Paleozoic era during the Phanerozoic eon, spanning 60.3 million years from the end of the preceding Silurian period at million years ago ( Ma), to the beginning of the succeeding C ...
. Lampreys have ancient origins, have endured for 360 million years and lived through four major
extinction events An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp fall in the diversity and abundance of multicellular organisms. It occ ...
- even so, their evolutionary history is obscure. In ancient seas jawless vertebrate fish led to jawed fish, and thence to all other
vertebrates Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
, including humans. Being the only extant jawless vertebrates and a virtual time capsule of embryology and
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
, lampreys and hagfish are of immense scientific interest."Discovery of the oldest fossil lamprey in the world"
Knowing the period during which they became parasitic would shed light on how typical they are of ancient fish. Having no
bone tissue A bone is a Stiffness, rigid Organ (biology), organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red blood cell, red and white blood cells, store minerals, provi ...
, effectively being essentially boneless, their cartilaginous framework leaves almost no fossil record - a mere three fossil species had previously been described, and in none of these was a sucker disc detected.


Description

The exceptionally well-preserved fossil of ''P. riniensis'' is only long and reveals details of its fin, gill basket and mouth region. A striking feature of ''Priscomyzon'' is its relatively large oral disk, inside a soft outer lip, supported by annular cartilage. The circular mouth at the center of the oral disk is surrounded by 14 small, evenly spaced, simple teeth, with no associated radiating series or plates of supplementary teeth, but otherwise quite similar to present-day lampreys, suggesting that their blood-sucking lifestyle was developed in ancient seas. The oral disk of the
Late Carboniferous Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * Late (The 77s album), ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudo ...
lamprey ''
Mayomyzon pieckoensis ''Mayomyzon pieckoensis'' is an extinct species of lamprey that lived during the Late Carboniferous period, about 300 million years ago. It is the only known species of the genus ''Mayomyzon'', which belongs to the family Mayomyzontidae. It is kno ...
'', if present, is much smaller, while there is no evidence of a disk in the
Early Carboniferous Early may refer to: Places in the United States * Early, Iowa, a city * Early, Texas, a city * Early Branch, a stream in Missouri * Early County, Georgia * Fort Early, Georgia, an early 19th century fort Music * Early B, stage name of Jamaican d ...
''
Hardistiella montanensis ''Hardistiella montanensis'' is an extinct species of lamprey, found, dating from the Carboniferous period, at the Bear Gulch Limestone site in the U.S. state of Montana. P. Janvier and R. Lund. 1983. Hardistiella montanensis n. gen. et sp. (Petr ...
''. ''Priscomyzons are the earliest teeth found in any fossil lamprey, and have a similar arrangement to that of the 19 or more teeth found in modern forms such as ''
Ichthyomyzon ''Ichthyomyzon'' is a genus of lampreys in the family Petromyzontidae, native to North America. Species There are currently six recognized species in this genus: * '' Ichthyomyzon bdellium'' ( D. S. Jordan, 1885) (Ohio lamprey) * '' Ichthyomyzon ...
'', ''
Petromyzon The sea lamprey (''Petromyzon marinus'') is a parasitic lamprey native to the Northern Hemisphere. It is sometimes referred to as the "vampire fish". It was likely introduced to the Great Lakes region through the Erie Canal in 1825 and the Well ...
'', ''
Caspiomyzon ''Caspiomyzon'' is a genus of lamprey in the family Petromyzontidae. They are native to Eastern Europe and parts of Western and Central Asia. Two of the three species in the genus are endemic to Greece. Taxonomy Species There are currently ...
'' and ''
Geotria ''Geotria'' is the only genus in the lamprey family Geotriidae. It has 2 known species: ''Geotria australis'' (pouched lamprey) and ''Geotria macrostoma'' (Argentinian lamprey). Both species were considered conspecific until ''G. macrostoma'' was ...
''. Its posterior teeth are more elongate than the rest, while in modern species lateral or anterior teeth tend to be largest. Its teeth are also quite simple in shape compared to those of modern species, and in this respect are probably primitive. The position of the orbits is not clear as there are no darkened areas suggesting eye locations. The branchial imprint is preserved in great detail, parts of both the right and left baskets having been preserved, and the posterior five branchial arches being well defined. Anterior to these the presence of seven branchial pouches is evident. The dorsal fin originates immediately behind the branchiae and continues to the caudal extremity, resembling a modern ammocoete or lamprey larva rather than an adult, in which separate anterior and posterior dorsal fins are to be found. Fossils of juvenile specimens indicate that it did not have the filter feeding "ammocoete" juvenile stage found in modern lampreys, with juveniles morphologically similar to adults.


Discovery

The lamprey fossil was discovered on Waterloo Farm in rocks of the Witteberg Group near
Grahamstown Makhanda, formerly known as Grahamstown, is a town of about 75,000 people in the Eastern Cape province of South Africa. It is situated about northeast of Gqeberha and southwest of East London. It is the largest town in the Makana Local Mun ...
in
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. The species was described by Robert W. Gess & Bruce S. Rubidge of the Bernard Price Institute (Palaeontology)
Wits University The University of the Witwatersrand, Johannesburg (), commonly known as Wits University or Wits, is a multi-campus public research university situated in the northern areas of central Johannesburg, South Africa. The university has its roots in ...
, and Michael I. Coates of the Department of Organismal Biology
University of Chicago The University of Chicago (UChicago, Chicago, or UChi) is a Private university, private research university in Chicago, Illinois, United States. Its main campus is in the Hyde Park, Chicago, Hyde Park neighborhood on Chicago's South Side, Chic ...
in the journal ''
Nature Nature is an inherent character or constitution, particularly of the Ecosphere (planetary), ecosphere or the universe as a whole. In this general sense nature refers to the Scientific law, laws, elements and phenomenon, phenomena of the physic ...
'' of 26 October 2006. The shale containing the fossil was discovered as far back as 1985 during construction of the N2 bypass outside Grahamstown. The site consists of black carbonaceous shale formed from anaerobic mud deposited in a marine estuary on the Agulhas Sea. A variety of organic remains are found in this setting, including algae, terrestrial plants and fish. Invertebrate remains are of small bivalves,
ostracod Ostracods, or ostracodes, are a Class (biology), class of the crustacean, Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 33,000 species (only 13,000 of which are extant taxon, extant) have been identified,Brandão, S.N.; Antoni ...
s,
clam shrimp Clam shrimp are a group of bivalved branchiopod crustaceans that resemble the unrelated bivalved molluscs. They are extant and also known from the fossil record, from at least the Devonian period and perhaps before. They were originally classif ...
, and a
eurypterid Eurypterids, often informally called sea scorpions, are a group of extinct marine arthropods that form the Order (biology), order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period, 467.3 Myr, mil ...
. Fossil material has been greatly compressed and original tissue replaced by metamorphic mica altered to chlorite during uplift. To prevent rockfalls onto the road from the unstable formation, the steep slopes were cut back again in 1999 and once more in 2007/8. During these upgrades Gess managed to obtain a large sample of rock blocks with the help of the road construction company, and worked intermittently on exposing their contents.


Etymology

The scientific name derives from the Latin ''
priscus Priscus of Panium (; ; 410s/420s AD – after 472 AD) was an Eastern Roman diplomat and Greek historian and rhetorician (or sophist)...: "For information about Attila, his court and the organization of life generally in his realm we have the ...
'' "ancient", and Greek '' myzon'' "sucking". 'Rini' is the
isiXhosa Xhosa ( , ), formerly spelled ''Xosa'' and also known by its local name ''isiXhosa'', is a Bantu language, indigenous to Southern Africa and one of the official languages of South Africa and Zimbabwe. Xhosa is spoken as a first language ...
name for
Grahamstown Makhanda, formerly known as Grahamstown, is a town of about 75,000 people in the Eastern Cape province of South Africa. It is situated about northeast of Gqeberha and southwest of East London. It is the largest town in the Makana Local Mun ...
and its surrounding valley.


Holotype

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 ...
is held in the
Albany Museum The Albany Museum, South Africa, situated in Makhanda, South Africa, is affiliated to Rhodes University and dates back to 1855,Chinsamy, Anusuya. (1997). "Albany Museum, Grahamstown, South Africa." ''Encyclopedia of Dinosaurs''. Edited by Philli ...
, Grahamstown, Eastern Cape, with the catalogue number AM5750. The locality and horizon is the Waterloo Farm, Grahamstown, South Africa; Witpoort Formation, Witteberg Group, Famennian, Late Devonian.


See also

*''
Gondwanascorpio emzantsiensis ''Gondwanascorpio emzantsiensis'' is an extinct Gondwanan scorpion that lived 360 million years ago in the Devonian. Its fossil remains, clearly showing pincer and sting, were discovered in rocks of the Witteberg Group near Grahamstown in South A ...
'' *''
Serenichthys kowiensis ''Serenichthys kowiensis'' is a fossil species of coelacanth described in 2015 from near Grahamstown in South Africa. Some 30 complete specimens of this new species were found in the black shale lagerstätte on Waterloo Farm, preserved by th ...
''


References

* {{Taxonbar, from=Q15325723
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
Devonian jawless fish Late Devonian fish Devonian fish of Africa Famennian genera Fossil taxa described in 2006