Hypostomus

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Hypostomus
Hypostomus plecostomus - Raphael Covain.png
Hypostomus plecostomus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Siluriformes
Family: Loricariidae
Subfamily: Hypostominae
Tribe: Hypostomini
Kner, 1853
Genus: Hypostomus
Lacépède, 1803
Type species
Hypostomus guacari
Lacépède, 1803
Synonyms

CheiridodusEigenmann, 1922
CochliodonHeckel, 1854
PlecostomusGronovius, 1754
WatawataIsbrücker & Michels, 2001

Contents

Hypostomus is a genus of catfish in the family Loricariidae. They are native to tropical and subtropical South America. H. plecostomus is the popular freshwater aquarium fish formerly known as Plecostomus plecostomus. The taxonomic structure of the Loricariidae is still being expanded by scientists. Hypostomus is a highly species-rich and widely distributed catfish genus. [1]

Taxonomy and phylogeny

This genus is the type member of its tribe Hypostomini and subfamily Hypostominae. [2] [3] The type species of Hypostomus is H. guacari. [4]

With the inclusion of Aphanotorulus , Isorineloricaria and Squaliforma , the genus Hypostomus is by far the largest genus of Loricariidae and the second largest genus of catfish. [5] [6] However, a good way to split up the genus further is unclear. [2] Species level taxonomy of Hypostomus is still poorly known as well as the phylogenetic relationships within and of the genus. It has been shown that the genus does not form a natural group. [3] However, small monophyletic groups are known to exist within the genus Hypostomus, such as the H. cochliodon group. [5] Aphanotorulus and Isorineloricaria are considered separate genera by some authors based on molecular information. [6]

Studies conducted with representatives of some genera of Hypostominae showed that within this group, the diploid number ranges from 2n = 52 to 2n = 80. However, the supposed wide karyotypic diversity that the family Loricariidae or the subfamily Hypostominae would present is almost exclusively restricted to the genus Hypostomus and the species from the other genera had a conserved diploid number. [7]

Distribution and habitat

These species are originate from freshwater habitats of South America and the Caribbean island of Trinidad. They are found throughout most of the range of loricariids except for drainages west of the Atrato River (northwest Colombia). They are essentially ubiquitous throughout their range. [2] Maximum diversity in number of species of Hypostomus occurs in rivers of the Paraná-Paraguay system. [3]

Hypostomus species are found in almost all aquatic habitats, from large rivers to small rivulets. [8] The typical habitat would be relatively fast moving water over a shallow gravel bed. Most species are lowland, sluggish stream- and lake-dwellers usually found associated with submerged wood; however, many species may be found among rocks in piedmont to mountain streams with moderate to swift flow. Members of this genus may be found over a range of substrates such mud, detritus, gravel, and sand. [2] Many species spawn in hollows dug into mud banks or within hollow logs. [2]

Description and ecology

As in all loricariids, these species have a suckermouth. They use their mouth as well as the pectoral, pelvic and caudal fin areas, to interact with the rocky river-bottom. The comb-toothed inferior mouth of Hypostomus is able to grasp green algae on the river bottom. With the help of the suckermouth, which anchor the fishes, in conjunction with the arched design of pectoral and pelvic fin spines, they are also able to remain upright while exploring this substratum. In addition, their large, concave caudal fin is associated with fast locomotion over short distances. [9]

Also like other loricariids, Hypostomus species exhibit plates of armor over their body. [2] The colouration among Hypostomus species varies. They may have a white ground color and black spots, or brown and spotted, or even black with red, gold, or white spots. Abdomen also varies in color from white to black and may be spotted or not. [2] The caudal fin is forked with the lower lobe longer than upper. Most species have stout bodies, but some do have thinner, elongate bodies. [2]

In most species, males develop hypertrophied odontodes on the leading edge of the pectoral fin spine and the end of the spine may become swollen. In some species, the males may also develop hypertrophied odontodes on the body during the breeding season. [2]

Hypostomus species are well known for the ability to breathe air. This is achieved through a slightly modified stomach that is larger and thinner than in non-air breathers. To breathe air, Hypostomus must orient itself vertically. [10]

Species of the H. cochliodon group are fairly large loricariids that reach about 30 centimetres (12 in) SL. Their colouration is typically dark brown with spots generally developed over most surfaces; however, most species of this group have been observed to have a well-developed ability to alter color according to substrate. Except for H. sculpodon, these species tend to have a deep body at the dorsal fin origin; this makes the body appear humped. [11] Along with Panaque , species of the H. cochliodon group are unique among loricariids for xylophagy, or the ability to digest wood, accomplished through specialized spoon-shaped teeth. In H. hemicochliodon and H. sculpodon appear to be intermediate between other species of Hypostomus and the H. cochilodon group by feeding less on wood and lacking specialized teeth; many other Hypostomus will occasionally eat wood, but wood only amounts to a very small fraction of their diet. [11]

Species

Hypostomus alatus Hypostomus alatus.jpg
Hypostomus alatus
Hypostomus asperatus Hypostomus asperatus.jpg
Hypostomus asperatus
Hypostomus commersoni Hypostomus commersoni.jpg
Hypostomus commersoni
Hypostomus subcarinatus Hypostomus subcarinatus Castelnau.jpg
Hypostomus subcarinatus

There are currently 156 recognized species in this genus:

Lustosa-Costa, Ramos, Zawadzki, Jacobina & Lima, 2024

Related Research Articles

<span class="mw-page-title-main">Loricariidae</span> Largest family of catfish

Loricariidae is the largest family of catfish, with over 90 genera and just over 680 species. Loricariids originate from freshwater habitats of Costa Rica, Panama, and tropical and subtropical South America. These fish are noted for the bony plates covering their bodies and their suckermouths. Several genera are sold as "plecos", notably the suckermouth catfish, Hypostomus plecostomus, and are popular as aquarium fish.

<i>Peckoltia</i> Genus of fishes

Peckoltia is a genus of small South American armored suckermouth catfishes. Many of these fish are popular aquarium fish.

<i>Hemiancistrus</i> Genus of fishes

Hemiancistrus is a genus of suckermouth armored catfishes. These species are native to South America. The taxonomy of this genus is complex and unclear, and major work has to be done. Many of these fish are popular aquarium fish.

<span class="mw-page-title-main">Hypoptopomatinae</span> Subfamily of fishes

The Hypoptopomatinae are a subfamily of catfishes of the family Loricariidae, composed of 17 genera and approximately 80 species. This subfamily represents about one-tenth of all loricariid species.

Neoplecostominae is a subfamily of South American catfishes of the family Loricariidae. Species of this subfamily live in high-mountain and swift-flowing river habitats.

<span class="mw-page-title-main">Hypostominae</span> Subfamily of fishes

The Hypostominae are a subfamily of catfishes of the family Loricariidae. Most members are restricted to tropical and subtropical South America, but there are also several species in southern Central America. Hypostomus plecostomus, which is popular in the aquarium trade, has been introduced to several regions far from its native range.

<span class="mw-page-title-main">Ancistrini</span> Tribe of fishes

Ancistrini is a tribe of catfishes of the family Loricariidae. Most are restricted to tropical and subtropical South America, but there are also several genus in southern Central America.

Neoplecostomus is a genus of fish in the family Loricariidae native to South America. Neoplecostomus can be distinguished from all other loricariids by a modified shield of small plates on the abdomen with posteriorly directed odontodes; the shield appears to act as a holdfast. The color pattern is generally mottled brown with the abdomen white. The head is long, rounded, and shovel-shaped. The fin spines are weak. They range from about 8 to 11 cm (3.1–4.3 in) SL. The species of Neoplecostomus live in fast-flowing water.

Hypostomus cochliodon is a species of armored catfish native to the Paraguay and middle Paraná River basins in northern Argentina, southern Brazil and Paraguay. Initially it did not occur in the upper Paraná basin above the Guaíra Falls, but these disappeared after the construction of the Itaipu Dam, allowing this species to spread. It grows to a standard length of 23 cm (9.1 in).

Hypostomus yaku is a species of South American armoured catfish from the family Loricariidae. It is only known from the Rio Quente, a small thermal stream in the Rio Paranaíba drainage of the upper Paraná basin, Brazil. H. yaku is unusual when compared to other species of Hypostomus in that it tolerates warm water up to 34 °C (93 °F) and only attains a relatively small size, growing to 70.8 mm (2.79 in) SL.

Hypostomus jaguar, sometimes known as the jaguar pleco, is a species of catfish in the family Loricariidae. It is native to South America, where it occurs in the Paraguaçu River basin in the state of Bahia in Brazil.

<i>Hypostomus topavae</i> Species of catfish

Hypostomus topavae is a species of catfish in the family Loricariidae. It is native to South America, where it occurs in the Paraná River drainage, including the Rio Grande basin. The species is very large for a loricariid, reaching 70 cm SL.

Hypostomus dardanelos is a species of catfish in the family Loricariidae. Its type locality is stated to be the Praia Grande River, a tributary of the Aripuanã in the Brazilian state of Mato Grosso.

Hypostomus delimai is a species of catfish in the family Loricariidae. It is native to South America, where it occurs in the basins of the Araguaia River and the Tocantins River in Brazil. It is typically found in areas with turbid water, riparian vegetation, and a substrate composed of rocks and sand. It is known to be sympatric with other species of Hypostomus, including H. faveolus and H. pyrineusi, as well as species of Aphanotorulus. The species reaches 25.3 cm in standard length and is believed to be a facultative air-breather.

Hypostomus angipinnatus is a species of catfish in the family Loricariidae. It is native to South America, where it occurs in the Río de La Plata drainage basin, as well as possibly the Paraguay River basin, although further investigation has found its presence in the Paraguay basin unlikely. The species reaches 15 cm in length and is believed to be a facultative air-breather.

Hypostomus denticulatus is a species of catfish in the family Loricariidae. It is native to South America, where it occurs in the Corumbá River in the Paraná River basin in Brazil. It is typically found in turbid waters with a substrate composed of rocks with some amount of sand. It is known to be syntopic with other loricariid species in the genus Hypostomus, including Hypostomus ancistroides, H. heraldoi, H. iheringii, H. margaritifer, and H. regani. The species reaches 19.1 cm in standard length and is believed to be a facultative air-breather.

Hypostomus heraldoi is a species of catfish in the family Loricariidae. It is native to South America, where it occurs in the Corumbá River and Rio Grande basins in the upper Paraná River drainage in Brazil. It is typically found in turbid waters with a substrate composed of rocks with some amount of sand. It is known to be syntopic with other loricariid species in the genus Hypostomus, including Hypostomus ancistroides, H. denticulatus, H. iheringii, H. margaritifer, and H. regani. The species reaches 23.6 cm in standard length and is believed to be a facultative air-breather.

Hypostomus kopeyaka is a species of catfish in the family Loricariidae. It is native to South America, where it occurs in the basin of the Tiquié River, a tributary of the Vaupés River, which is itself a tributary of the Rio Negro, in Brazil, although it has also been reported from Vaupés Department in Colombia. It generally occurs in rapids, cataract pools, and slow-flowing portions of the Tiquié and some of its large tributaries. The species reaches 22.6 cm in standard length and is believed to be a facultative air-breather. Its specific epithet, kopeyaka, refers to its common name in the Tuyuca and Tucano languages, kope ya’ka, reportedly meaning "pleco from the holes", in reference to the species' tendency to hide in holes in the river bank.

Hypostomus nigropunctatus is a species of catfish in the family Loricariidae. It is native to South America, where it occurs in the Iguazu River in the state of Paraná in Brazil. The species reaches 23.3 cm (9.2 in) in standard length and is believed to be a facultative air-breather.

Hypostomus peckoltoides is a species of catfish in the family Loricariidae. It is native to South America, where it occurs in the Cuiabá River in the upper Paraguay River basin in Brazil. It is typically found in environments with turbid water, a substrate composed of rocks and sand, and various types of riparian vegetation. It is known to be syntopic with other species of Hypostomus, including H. boulengeri, H. cochliodon, H. latifrons, H. latirostris, and H. regani.

References

  1. ↑ Jardim de Queiroz, Luiz; Cardoso, Yamila; Jacot-des-Combes, Cécile; Bahechar, Ilham Anne; Lucena, Carlos Alberto; Rapp Py-Daniel, Lucia; Sarmento Soares, Luisa Maria; Nylinder, Stephan; Oliveira, Claudio; Parente, Thiago Estevam; Torrente-Vilara, Gislene; Covain, Raphaël; Buckup, Paulo; Montoya-Burgos, Juan I. (2020-04-01). "Evolutionary units delimitation and continental multilocus phylogeny of the hyperdiverse catfish genus Hypostomus". Molecular Phylogenetics and Evolution. 145: 106711. doi: 10.1016/j.ympev.2019.106711 . ISSN   1055-7903. PMID   31857199. S2CID   209428047.
  2. 1 2 3 4 5 6 7 8 9 Armbruster, J.W. "Hypostomus Lacépède, 1803".
  3. 1 2 3 Oyakawa, Osvaldo T.; Akama, Alberto; Zanata, Angela M. (2005). "Review of the genus Hypostomus Lacépède, 1803 from rio Ribeira de Iguape basin, with description of a new species (Pisces, Siluriformes, Loricariidae)" (PDF). Zootaxa . 921: 1–27. doi: 10.11646/zootaxa.921.1.1 .
  4. ↑ Froese, Rainer and Pauly, Daniel, eds. (2016). Species of Hypostomus in FishBase . January 2016 version.
  5. 1 2 3 Armbruster, Jonathan W.; Tansey, Leigh A.; Lujan, Nathan K. (2007). "Hypostomus rhantos (Siluriformes: Loricariidae), a new species from southern Venezuela" (PDF). Zootaxa . 1553: 59–68. doi:10.11646/zootaxa.1553.1.3.
  6. 1 2 3 Zawadzki, Cláudio H.; Birindelli, José L. O.; Lima, Flávio C. T. (2008). "A new pale-spotted species of Hypostomus Lacépède (Siluriformes: Loricariidae) from the rio Tocantins and rio Xingu basins in central Brazil". Neotropical Ichthyology. 6 (3): 395–402. doi: 10.1590/s1679-62252008000300012 .
  7. ↑ Luìs Alves, Anderson; Oliveira, Claudio; Foresti, Fausto (2005). "Comparative cytogenetic analysis of eleven species of subfamilies Neoplecostominae and Hypostominae (Siluriformes: Loricariidae)". Genetica. 124 (2): 127–136. doi:10.1007/s10709-004-7561-4. PMID   16134327. S2CID   8239240.
  8. 1 2 3 Zawadzki, Cláudio H.; Weber, Claude; Pavanelli, Carla S. (2008). "Two new species of Hypostomus Lacépède (Teleostei: Loricariidae) from the upper rio Paraná basin, Central Brazil". Neotropical Ichthyology. 6 (3): 403–412. doi: 10.1590/s1679-62252008000300013 .
  9. ↑ Garavello, J. C.; Garavello, J. P. (2004). "Spatial distribution and interaction of four species of the catfish genus Hypostomus Lacépède with bottom of Rio São Francisco, Canindé do São Francisco, Sergipe, Brazil (Pisces, Loricariidae, Hypostominae)" (PDF). Brazilian Journal of Biology. 64 (3B): 591–598. doi: 10.1590/s1519-69842004000400006 . PMID   15619997.
  10. ↑ Armbruster, Jonathan W. (1998). "Modifications of the Digestive Tract for Holding Air in Loricariid and Scoloplacid Catfishes" (PDF). Copeia . 1998 (3): 663–675. doi:10.2307/1447796. JSTOR   1447796.
  11. 1 2 Armbruster, Jonathan W. (2003). "The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae)" (PDF). Zootaxa . 249: 1–60. doi:10.11646/zootaxa.249.1.1.
  12. ↑ Castelnau, Francis comte de (1855). Animaux nouveaux ou rares recueillis pendant l'expedition dans les parties centrales de L'Amerique du Sud, de Rio de Janeiro a Lima, et de Lima au Para. Tome Second (in French). France: Chez P. Bertrand. doi:10.5962/bhl.title.101716.
  13. 1 2 3 4 5 6 7 8 9 Isbrücker, I.J.H. (1980). "Classification and catalogue of the mailed Loricariidae (Pisces, Siluriformes)" (PDF). Verslagen en Technische Gegevens. 22 (1): 1–181.
  14. ↑ Burgess, Warren (1989). An atlas of freshwater and marine catfishes : a preliminary survey of the Siluriformes. Neptune City, NJ: T.F.H. ISBN   0-86622-131-X. OCLC   875582011.
  15. 1 2 3 4 5 6 7 8 Armbruster, Jonathan W.; Werneke, David C.; Tan, Milton (2015). "Three new species of saddled loricariid catfishes, and a review of Hemiancistrus, Peckoltia, and allied genera (Siluriformes)". ZooKeys (480): 97–123. doi: 10.3897/zookeys.480.6540 . PMC   4319111 . PMID   25685026.
  16. ↑ Cardoso, Yamila P.; Almirón, Adriana; Casciotta, Jorge; Aichino, Danilo; Lizarralde, Marta S.; Montoya-Burgos, Juan I. (2012). "Origin of species diversity in the catfish genus Hypostomus (Siluriformes: Loricariidae) inhabiting the Paraná river basin, with the description of a new species". Zootaxa. 3453: 69–83. doi:10.11646/zootaxa.3453.1.5. S2CID   85715759.
  17. ↑ Eigenmann, Carl H.; Eigenmann, Rosa Smith (1890). A revision of the South American Nematognathi or cat-fishes / by Carl H. Eigenmann and Rosa Smith Eigenmann. San Francisco: California Academy of Sciences. doi:10.5962/bhl.title.30137.
  18. ↑ Kner, Rudolf (1854). Die Hypostomiden : zweite Hauptgruppe der Familie der Panzerfische (Loricata v. Goniodontes) / von R. Kner (in German). Wien: Kaiserlich-Königlichen Hof- und Staatsdruckerei. doi:10.5962/bhl.title.12573.
  19. 1 2 3 Tencatt, Luiz F. C.; Zawadzki, Cláudio H.; Froehlich, Otávio (2014). "Two new species of the Hypostomus cochliodon group (Siluriformes: Loricariidae) from the rio Paraguay basin, with a redescription of Hypostomus cochliodon Kner, 1854". Neotropical Ichthyology. 12 (3): 585–602. doi: 10.1590/1982-0224-20130162 .
  20. 1 2 "Hypostomus formosae, a new catfish species from the Paraguay River Basin with redescription of H. boulengeri (Siluriformes: Loricariidae)". Ichthyological Exploration of Freshwaters. 27 (1): 9–24. 2016.
  21. ↑ Günther, Albert C. L. G. (1859). Catalogue of the fishes in the British Museum. Vol. 4. London: British Museum (Natural History), Department of Zoology. doi:10.5962/bhl.title.8809.
  22. ↑ Martins, Fernanda O.; Marinho, Manoela M. F.; Langeani, Francisco; Serra, Jane P. (2012). "A New Species of Hypostomus (Siluriformes: Loricariidae) from the Upper Rio Paraguay Basin, Brazil". Copeia. 2012 (3): 494–500. doi:10.1643/ci-11-011. S2CID   84627045.
  23. ↑ Birindelli, José L. O.; Zanata, Angela M.; Lima, Flávio C. T. (2007). "Hypostomus chrysostiktos, a new species of armored catfish (Siluriformes: Loricariidae) from rio Paraguaçu, Bahia State, Brazil" (PDF). Neotropical Ichthyology . 5 (3): 271–278. doi: 10.1590/s1679-62252007000300006 .
  24. ↑ Zawadzki, Cláudio H.; Carvalho, Pedro Hollanda (2014). "A new species of the Hypostomus cochliodon group (Siluriformes: Loricariidae) from the rio Aripuanã basin in Brazil". Neotropical Ichthyology. 12 (1): 43–51. doi: 10.1590/s1679-62252014000100004 .
  25. ↑ Zawadzki, Cláudio Henrique; Oliveira, Renildo Ribeiro de; Debona, Tiago (2013). "A new species of Hypostomus Lacépède, 1803 (Siluriformes: Loricariidae) from the rio Tocantins-Araguaia basin, Brazil". Neotropical Ichthyology. 11 (1): 73–80. doi: 10.1590/s1679-62252013000100008 .
  26. ↑ Zanata, Angela M.; Sardeiro, Byanca; Zawadzki, Cláudio H. (2013). "A new dark-dotted species of Hypostomus Lacépède (Siluriformes: Loricariidae) from rio Paraguaçu, Bahia State, Brazil". Neotropical Ichthyology. 11 (2): 247–256. doi: 10.1590/s1679-62252013000200002 .
  27. ↑ Zawadzki, Cláudio H.; Birindelli, José L. O.; Lima, Flávio C. T. (2012). "A new armored catfish species of the genus Hypostomus Lacépède, 1803 (Siluriformes: Loricariidae) from the upper rio Xingu basin, Brazil". Neotropical Ichthyology. 10 (2): 245–253. doi: 10.1590/s1679-62252012000200003 .
  28. ↑ Penido, I. de S.; Pessali, T.C. & Zawadzki, C.H., 2023 "A new tiny-spotted species of Hypostomus (Siluriformes: Loricariidae) from the upper Rio Tocantins basin, Goiás State, Brazil" (abstract) (Zootaxa, Vol. 5361 No 1)
  29. ↑ Froese, Rainer; Pauly, Daniel, eds. (2022). "Hypostomus laplatae". FishBase.
  30. ↑ Zanata, Angela M.; Pitanga, Bruno R. (2016). "A new species of Hypostomus Lacépède, 1803 (Siluriformes: Loricariidae) from rio Itapicuru basin, Bahia State, Brazil". Zootaxa. 4137 (2): 223–232. doi:10.11646/zootaxa.4137.2.4. PMID   27470717.
  31. ↑ "Hypostomus melanephelis, a new armored catfish species from the rio Tapajós basin, Brazil (Teleostei: Loricariidae)". Ichthyological Exploration of Freshwaters. 26 (1): 49–58. 2015.
  32. ↑ Jerep, Fernando C.; Shibatta, Oscar A.; Zawadzki, Cláudio H. (2007). "A new species of Hypostomus Lacépède, 1803 (Siluriformes: Loricariidae) from the upper rio Paraná basin, Southern Brazil". Neotropical Ichthyology . 5 (4): 435–442. doi: 10.1590/s1679-62252007000400002 .
  33. ↑ Garavello, Julio César; Britski, Heraldo Antônio; Zawadzki, Claudio Henrique (2012). "The cascudos of the genus Hypostomus Lacépède (Ostariophysi: Loricariidae) from the rio Iguaçu basin". Neotropical Ichthyology. 10 (2): 263–283. doi: 10.1590/s1679-62252012000200005 .
  34. ↑ Zawadzki, Claudio H.; Tencatt, Luiz F. C.; Froehlich, Otávio (2014). "A new unicuspid-toothed species of Hypostomus Lacépède, 1803 (Siluriformes: Loricariidae) from the rio Paraguai basin". Neotropical Ichthyology. 12 (1): 97–104. doi: 10.1590/s1679-62252014000100010 .
  35. ↑ Pfeil, Friedrich & München, • & Zawadzki, Cláudio & Silva, Hugmar & Troy, Waldo. (2018). Redescription of Hypostomus latirostris (Regan, 1904) with the recognition of a new species of Hypostomus (Siluriformes: Loricariidae) from the upper rio Paraguay basin, Brazil. Ichthyological Exploration of Freshwaters. 28. 10.23788/IEF-1079.
  36. ↑ Cardoso, Yamila P.; Brancolini, Florencia; Protogino, Lucila; Paracampo, Ariel; Bogan, Sergio; Posadas, Paula; Montoya-Burgos, Juan I. (2019). "An integrated approach clarifies the cryptic diversity in Hypostomus Lacépède 1803 from the Lower la Plata Basin". Anais da Academia Brasileira de Ciências. 91 (2): e20180131. doi: 10.1590/0001-3765201920180131 . hdl: 11336/118954 . PMID   31038530. S2CID   140248209.
  37. ↑ Ray, C. K.; Armbruster, J. W. (2016). "The genera Isorineloricaria and Aphanotorulus (Siluriformes: Loricariidae) with description of a new species". Zootaxa. 4072 (5): 501–539. doi: 10.11646/zootaxa.4072.5.1 . PMID   27395943.
  38. ↑ Martins, Fernanda O.; Langeani, Francisco; Zawadzki, Cláudio H. (2014). "A new spiny species of Hypostomus Lacépède (Loricariidae: Hypostominae) from thermal waters, upper rio Paraná basin, central Brazil". Neotropical Ichthyology. 12 (4): 729–736. doi: 10.1590/1982-0224-20140035 . hdl: 11449/117915 .
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