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''Megapiranha'' is an extinct
serrasalmid The Serrasalmidae (serrasalmids) are a family of characiform fishes, recently elevated to family status. It includes more than 90 species. The name means "serrated salmon family", which refers to the serrated keel running along the belly of these ...
characin Characiformes is an order of ray-finned fish, comprising the characins and their allies. Grouped in 18 recognized families, more than 2000 different species are described, including the well-known piranha and tetras.; Buckup P.A.: "Relationshi ...
fish from the
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene Epoch made up of two stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million years ago) to 5.333 Ma. The e ...
(8–10
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago ...
)
Ituzaingó Formation The Ituzaingó Formation ( es, Formación Ituzaingó), in older literature also described as Entre Ríos or Entrerriana Formation, is an extensive geological formation of Late Miocene ( Tortonian, or Huayquerian in the SALMA classification) age in ...
of
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
, described in 2009. The type species is ''M. paranensis''.''Megapiranha''
at
Fossilworks Fossilworks is a portal which provides query, download, and analysis tools to facilitate access to the Paleobiology Database, a large relational database assembled by hundreds of paleontologists from around the world. History Fossilworks was crea ...
.org
It is thought to have been about in length and in weight. The holotype consists only of
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
e and a zigzag tooth row; the rest of its body is unknown.
Summary of the paper
.
This dentition is reminiscent of both the double-row seen in
pacu Pacu () is a common name used to refer to several species of omnivorous South American freshwater serrasalmid fish that are related to the piranha. Pacu and piranha do not have similar teeth, the main difference being jaw alignment; piranha h ...
s, and the single row seen in the teeth of modern
piranha A piranha or piraña (, , or ; or , ) is one of a number of freshwater fish in the family Serrasalmidae, or the subfamily Serrasalminae within the tetra family, Characidae in order Characiformes. These fish inhabit South American rivers, f ...
s, suggesting that ''M. paranensis'' is a
transitional form A transitional fossil is any fossilized remains of a life form that exhibits traits common to both an ancestral group and its derived descendant group. This is especially important where the descendant group is sharply differentiated by gross a ...
. Its bite force is estimated between .


History and naming

The holotype of ''Megapiranha'' was discovered in an unknown locality of the
Ituzaingó Formation The Ituzaingó Formation ( es, Formación Ituzaingó), in older literature also described as Entre Ríos or Entrerriana Formation, is an extensive geological formation of Late Miocene ( Tortonian, or Huayquerian in the SALMA classification) age in ...
,
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
, in the early 20th century near the towns of Paraná and
Villa Urquiza Villa Urquiza is a barrio or neighborhood of Buenos Aires, Argentina. It is located between the barrios of Villa Pueyrredón, Belgrano, Villa Ortúzar, Coghlan, Saavedra and Agronomía. Its limits are the streets and avenues Constituyentes, ...
. The specimen, a fragment of the animal's premaxilla containing several teeth, was later rediscovered by Alberto Luis Cione in the collection of the
Museo de La Plata The La Plata Museum ( es, Museo de la Plata) is a natural history museum in La Plata, Argentina. It is part of the (Natural Sciences School) of the UNLP (National University of La Plata). The building, long, today houses 3 million fossils and ...
. An isolated tooth discovered in 1999 has also been referred to this genus. The name ''Megapiranha'' is a combination of the word "mega" in reference to the animal's large size and piranha, a common name for typically carnivorous members of
Serrasalmidae The Serrasalmidae (serrasalmids) are a family of characiform fishes, recently elevated to family status. It includes more than 90 species. The name means "serrated salmon family", which refers to the serrated keel running along the belly of the ...
. The word piranha itself is a Portuguese merging of words originating in the Tupi language and may have several meanings including "tooth fish", "cutting fish", "devil fish" or "biting fish". The
species name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
was chosen to reflect ''Megapiranha''s place of origin near the city of Paraná.


Description

Although only three teeth are fully preserved, the holotype specimen shows the presence of seven premaxillary teeth which are arranged in a zig-zag pattern with some overlap. The third, fourth and fifth tooth are preserved and all share very similar morphology with one another. Of these teeth the first and third formed the inner row, while the rest formed the outer row. In other serrasalmids the teeth are arranged either two rows of seven teeth with a morphologically distinct second tooth of the inner row (third tooth), such an arrangement is seen in the Tambaqui for instance, a zig-zag pattern of five teeth (like in '' Catoprion'') or a single row of six teeth, which is typical for the carnivorous species. Each teeth only shows a single tooth crown the shape of an almost equilateral triangle, which sits atop a constriction of the tooth. Towards the apex of the crown the teeth take on a more sloping edge which is finely serrated in addition to slight labio-lingual compression. Much like in the arrangement of the teeth, ''Megapiranha'' differs significantly from its modern relatives. Pacus generally have more complex and broad teeth while true piranhas have teeth with multiple cusps, well developed serration and strong compression, making them thin and well suited for cutting. Between the three preserved teeth the size varies greatly, with the third being the largest and the fourth the smallest. The attachment scars likewise differ in size, showing a similar size distribution. The preserved
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
is almost straight and the teeth are all positioned on the same horizontal plane. The dorsal surface of the bone is slightly concave and slopes upwards towards the front as it transitions to the ascending process, which is barely tapering. Here too ''Megapiranha'' provides a unique combination of features amongst its family, with serrasalmids that share the straight axis of the premaxilla typically having a straight dorsal margin and two different planes on which the teeth are placed, while those with a single horizontal plane and concave dorsal surface lack a straight axis. The entire premaxilla is long with a rugose outer surface that most likely housed nerves and blood vessels. The symphyseal joint is interlocking. Based on the size of the holotype, ''Megapiranha'' has originally been estimated to have reached a length of and a weight of , larger than any other member of the family, living or extinct. Later research using ''
Serrasalmus rhombeus The redeye piranha (''Serrasalmus rhombeus''), also known as the black piranha, white piranha, spotted piranha or yellow piranha, is a species of freshwater ray-finned fish, a piranha from the family Serrasalmidae. It is found in northern South Am ...
'' as a basis arrived at a more conservative size estimate of long and heavy.


Phylogeny

''Megapiranha'' combines several traits known from more basal serrasalmids with those of derived members. Both the subcircular tooth attachment scars as well as the presence of seven, not six, teeth are in line with what is known from most members of the group, while the triangular shape of the crown, fine serration and slight labiolingual compression are more in line with the morphology seen in the teeth of true piranhas. The interdigitating symphyseal joint meanwhile draws parallels to basal pacus like ''
Colossoma The tambaqui (''Colossoma macropomum'') is a large species of freshwater fish in the family Serrasalmidae. It is native to tropical South America, but kept in aquaculture and introduced elsewhere. It is also known by the names black pacu, black- ...
'' and '' Mylossoma'', however differences in the structure of the joint between ''Megapiranha'' and extant forms suggests that this trait developed independently from one another. This unique combination of characters supports the idea that within serrasalmids an evolutionary trend led to the shift from double-rowed dentition with broad teeth to the single row of flattened teeth observed in piranhas. ''Megapiranha'' represents an intermediate form between the two, with triangular, slightly compressed teeth but maintaining two rows of teeth that are still relatively broad.


Paleobiology

In the 2009 description of the genus, Cione and colleagues suggest that the dentition of ''Megapiranha'' may not have been a direct adaptation towards carnivory and likely helped the animal with a wide range of food sources. Part of their reasoning for this is the broad range of diets found within serrasalmids, including many herbivorous and omnivorous forms in addition to carnivores, as well as the highly specialised wimple piranha which feeds primarily on the scales of other fish. In 2012 Justin R. Grubich and colleagues suggest that the dentition of ''Megapiranha'' may have been a transitional form between feeding on hard prey and specialising in slicing flesh. To arrive at this conclusion, they conducted extensive measurements of the bite force of the extant ''
Serrasalmus rhombeus The redeye piranha (''Serrasalmus rhombeus''), also known as the black piranha, white piranha, spotted piranha or yellow piranha, is a species of freshwater ray-finned fish, a piranha from the family Serrasalmidae. It is found in northern South Am ...
'' as a standin for its Miocene relative. With this method they calculated a bite force of 1240 Newton for the smaller estimates and 4749 Newton for the older, larger size estimates. Even the more conservative estimates would put the biteforce of ''Megapiranha'' four times higher than that of the largest extant piranha species. The authors additionally note that the fact that the measurements were taken on live animals may lead to understimates caused by fatigue and stress. Furthermore, the measurements were restricted to the anterior biteforce, not including the potential of doubled biteforce along the lower jaw. This may result in forces between 2480 and 9498 Newton. Tests using a bronze-alloy replica of ''Megapiranha''s dentition showed that it would be able to penetrate the thick outer layer of a bovine femur, the shell of a turtle and the armor of certain catfish species. They conclude by suggesting that ''Megapiranha'' could have hypothetically fed on hard-shelled animals such as turtles, armored catfish and even attacked larger mammals. The shape of ''Megapiranha''s teeth is infered to effectively focus stress at the tip of the teeth while piercing flesh before distributing it impact stresses throughout the base of the tooth while crushing hard material such as bones in a similar fashion to how Pacus crack hard-shelled fruits and nuts. However, this ecology is only hypothetical in the absence of any fossil material bearing the bite marks of ''Megapiranha''.


References


External links

*
New fossil tells how piranhas got their teeth
. ''eurekalert.org''. 2009. {{Taxonbar, from=Q21447457 †Megapiranha Prehistoric ray-finned fish genera Transitional fossils Miocene fish Miocene animals of South America Neogene Argentina Fossils of Argentina Ituzaingó Formation Paraná Basin Fossil taxa described in 2009