Overview

Comprehensive Description

[[ Genus Hypostomus Lacépède ZBK ]]

Hypostomus ZBK , the type genus of the subfamily Hypostominae, constitutes an assemblage of bottom-dwelling loricariid fishes widely distributed throughout South America, occurring in a variety of freshwater ecosystems such as small mountain streams and large lowland river areas. Maximum diversity in number of species of Hypostomus ZBK occurs in rivers of the Paraná-Paraguay system (Weber, 2003). As presently defined the genus consists of more than 130 nominal species (107 species considered valid by Weber, 2003 and expanded by Armbruster, 2004).

Species level taxonomy of Hypostomus ZBK is still poorly known as well as the phylogenetic relationships within and of the genus. Hypostomus ZBK was established by Lacépède (1803), who included in the genus only a single species, H. guacari ZBK . Old descriptions, similar to Lacépède ’s, and available for various subsequent species added to the genus, are very often scanty, with incomplete or poor characterization of species. This fact, together with a number of Hypostomus ZBK species yet to be described, makes the recognition of most taxa of the genus difficult. Hypothesis of phylogenetic relationships of Hypostomus ZBK based on molecular data (Montoya-Burgos et al., 1997, 2002) revealed that the genus does not form a natural group. According to these authors, some species currently assigned to Hypostomus ZBK should be removed from the genus, while the genus Cochliodon ZBK should be treated as synonym of Hypostomus ZBK , an action effectuated by Weber & Montoya-Burgos (2002: 265). Morphological based phylogenetic studies presented by Armbruster (2004) reinforce the conclusions above and place Aphanotorulus ZBK and Isorineloricaria ZBK in the synonymy of Hypostomus ZBK . In spite of the cited studies, more detailed phylogenetic analysis of the relationships within Hypostomus ZBK awaits for future studies.

The elevated number of species currently included in Hypostomus ZBK , allied to the broad variability in morphology and color pattern of those taxa, poses difficulties for its entire revision. Various authors acquainted with these problems provided regional revisions of the genus, as exemplified by Boeseman (1968) with a study of the Surinam representatives of the genus, Weber (1985, 1986, 1987) that deals with Hypostomus ZBK from Paraguay, more recently Reis et al. (1990) reviews and describes three new species from Southern Brazil, Mazzoni et al. (1994) provides a taxonomical revision of Hypostomus ZBK from Lower rio Paraíba do Sul, State of Rio de Janeiro, Brazil, and finally Hollanda Carvalho & Weber (2004) provided a revision of Hypostomus cochliodon group ZBK from middle and lower Amazon system.

  • Osvaldo T. Oyakawa, Alberto Akama, Angela M. Zanata (2005): Review of the genus Hypostomus Lacépède, 1803 from rio Ribeira de Iguape basin, with description of a new species (Pisces, Siluriformes, Loricariidae). Zootaxa 921, 1-27: 2-2, URL:http://www.zoobank.org/urn:lsid:zoobank.org:pub:913A8172-1A2E-4784-96DB-50BACBEC7C25
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[[Genus Hypostomus Lacepede ZBK ]]

Discussion

Four nominal species of Hypostomus ZBK are cited in the literature for rio Ribeira de Iguape basin: Hypostomus agna (Miranda-Ribeiro, 1907) , Hypostomus commersonii Valenciennes, 1840 , Hypostomus interruptus (Miranda-Ribeiro, 1918) , and Hypostomus lacerta (Nichols, 1919) . Hypostomus lacerta was recently transferred to the genus Kronichthys ZBK by Armbruster & Page (1997). Hypostomus commersonii was described for the La Plata river basin, Uruguay, and to rio São Francisco, Brazil. Subsequent authors registered the species in other Brazilian drainages (e.g., rio Paraíba do Sul (Steindachner, 1876; MirandaRibeiro, 1911, 1918), rio Paraná (Miranda-Ribeiro, 1911), and rio Ribeira de Iguape (Miranda-Ribeiro, 1907, 1908a, b, 1918)), broadening the distribution of H. commersonii . Weber (1986), on the contrary, restricted the area of occurrence of H. commersonii to the La Plata river system and Mazzoni et al. (1994) corroborated the absence of this species in the rio Paraíba do Sul drainage. Similarly, we herein confirm the absence of H. commersonii in the Ribeira de Iguape river system. Our analysis revealed that the loricariids collected in the Ribeira do Iguape basin and previously identified in the literature as H. commersonii , have several distinctive features when compared to the La Plata river form. In fact, this material represents a new species and is described herein as H. tapijara ZBK , sp. n. Besides having a relatively distinct overall body form, H. tapijara ZBK can be also distinguished from H. commersonii on the basis of several characters including the presence of larger and more numerous dark spots distributed over body and fins, and absence of oblique inconspicuous bands over flanks. These two species can be further distinguished from each other by the relatively higher degree of development of ridges and keels in H. commersonii .

As noted in the ‘Introduction’ some authors provided regional revisions of the genus Hypostomus ZBK . From those revisions, only Mazzoni et al. (1994) examined species from eastern Brazilian basins, recognizing only two valid species for lower portion of rio Paraíba do Sul basin: Hypostomus affinis (Steindachner, 1877) and Hypostomus auroguttatus Kner, 1854 ZBK , senior synonym of H. luetkeni (Steindachner, 1877) (see Weber, 2003:356). According to Mazzoni et al. (1994) both species are only in the rio Paraíba do Sul or northern Brazilian drainages. Furthermore, those species were not cited in the literature for the rio Ribeira de Iguape basin. Examination of specimens of H. affinis in the present study revealed some similarities in coloration with H. interruptus and H. tapijara ZBK . However, H. affinis have body comparatively more elongated than H. interruptus and also have larger and more conspicuous dark spots distributed over body and fins (compare Fig. 6 with fig. 2 of Mazzoni et al., 1994). Hypostomus affinis differs from H. tapijara ZBK by the absence of broad dorsal fin and absence of strongly developed keels characteristic of the later species (compare Fig. 7 with fig. 2 of Mazzoni et al, 1994). According to Bizerril (1994), there is another species from rio Paraíba do Sul basin, H. punctatus ZBK , which belongs to the H. commersoni group, and can be readily distinguished from H. tapijara ZBK by having smaller spots over body and fins, four dark brown inconspicuous oblique bands, absence of a broad dorsal fin and body not so elongated.

In the case of Hypostomus auroguttatus ZBK , comparisons with species of Hypostomus ZBK from rio Ribeira de Iguape basin revealed several similarities with H. agna , mainly related to the body shape, alignment of series of plates in anterior and posterior portion of trunk, absence of keels, and coloration (compare Fig. 2 with fig. 3 of Mazzoni et al, 1994).

AlthoughH. agna is endemic to the rio Ribeira de Iguape system (Weber, 2003:355) and H. auroguttatus ZBK was never reported in the literature to occur in this river basin, the distinction between the two species based on the examination of the available material of both species is apparently problematic. In fact, taxonomic difficulties involving H. auroguttatus ZBK were previously emphasized by Weber (2003:364), under his comments of Hypostomus vermicularis (Eigenmann & Eigenmann) . According to the author, H. vermicularis needs a revision, a task beyond the scope of this study. Considering the known geographic distribution of H. auroguttatus ZBK , the undoubted presence of H. agna in rio Ribeira de Iguape basin and the limited focus of the present study we defer to propose a synonymization of these two species and maintain H. agna as the species present in the rio Ribeira de Iguape basin.

Thus the four species found to occur in rio Ribeira do Iguape basin includes those two previously cited in the literature and confirmed to be present in this study, recognized as H. agna and H. interruptus , one species recorded for the first time in this system that is Hypostomus ancistroides , and the new species H. tapijara ZBK .

  • Osvaldo T. Oyakawa, Alberto Akama, Angela M. Zanata (2005): Review of the genus Hypostomus Lacépède, 1803 from rio Ribeira de Iguape basin, with description of a new species (Pisces, Siluriformes, Loricariidae). Zootaxa 921, 1-27: 23-25, URL:http://www.zoobank.org/urn:lsid:zoobank.org:pub:913A8172-1A2E-4784-96DB-50BACBEC7C25
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Key to the species of Hypostomus ZBK from rio Ribeira de Iguape basin

1. Keels absent or poorly developed; anterior plates of mid-dorsal series longitudinally aligned with those of posterior portion of trunk, not interrupted by the first plate of dorsal series (Fig. 1A); caudal peduncle trapezoid in cross-section; 3 to 5 plates around supraoccipital (Fig. 2)............................................................ Hypostomus agna

1’. Keels well developed; anterior plates of mid-dorsal series not aligned with those of posterior portion of trunk, interrupted by the first plate of dorsal series (Fig. 1B); caudal peduncle ovoid in cross-section; 1 to 2 plates around supraoccipital.....................2

2. Plates on ventral surface of head absent in adults; spots on posterior half of body less conspicuous and more sparsely distributed than on anterior portion of body; spots on fins usually aligned, sometimes forming dark bands (Fig. 4) .................. Hypostomus ancistroides

2’. Ventral region of head completely covered with dermal ossifications, except for small naked areas; pattern of distribution of spots on body and fins relatively uniform, never forming dark bands.......................................................................................................3

3. Presence of oblique inconspicuous dark bands on flanks; small spots over body and fins, smaller over head; dorsal fin comparatively smaller (dorsal-fin spine length 21.6 -28.4% SL) (Fig. 6) ................................................................. Hypostomus interruptus

3’. Dark bands on flanks absent; well defined large, dark, and roundish spots somewhat homogeneously distributed over body and fins; relatively broad dorsal-fin (dorsal-fin spine length 27.6-34.3% SL) (Fig. 7).................................. Hypostomus tapijara ZBK , sp.n.

  • Osvaldo T. Oyakawa, Alberto Akama, Angela M. Zanata (2005): Review of the genus Hypostomus Lacépède, 1803 from rio Ribeira de Iguape basin, with description of a new species (Pisces, Siluriformes, Loricariidae). Zootaxa 921, 1-27: 4-4, URL:http://www.zoobank.org/urn:lsid:zoobank.org:pub:913A8172-1A2E-4784-96DB-50BACBEC7C25
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[[ Hypostomus ZBK ]]

The genus Hypostomus ZBK has 138 species currently recognized as valid making it the largest genus in the Loricariidae (suckermouth armored catfishes; Armbruster, 2004). Few of the species are well defined, and a comprehensive review of the genus is lacking. Other than individual species descriptions, the taxonomic reviews that have been done on Hypostomus ZBK are limited to geographic reviews (Boeseman, 1968; Reis et al., 1990) or to small monophyletic groups such as the H. unicolor group (Armbruster and Page, 1996; Armbruster, 1998) and the H. cochliodon ZBK group (Armbruster, 2003b; Hollanda Carvalho and Weber, 2004).

The type species of Hypostomus ZBK is H. plecostomus , a species from the coastal Guyanas (Boeseman, 1968). It is characterized by dentaries angled just greater than 90° (Armbruster, 2004), viliform teeth, and a generalized morphology consisting of a fairly broad, fairly short body, that is mildly dorsoventrally flattened and brown with dark spots (Boeseman, 1968). There are species of similar morphology in lowlands of most major river basins of tropical South America.

Armbruster (2004) recognized very few subdivisions within Hypostomus ZBK , recognizing only an H. emarginatus ZBK group and an H. cochliodon ZBK group. For the purposes of this paper, Hypostomus ZBK is recognized in two main groups the H. emarginatus ZBK group (as defined in Armbruster, 2004) and the H. plecostomus group (all the other species). The H. plecostomus group additionally has the H. cochliodon ZBK subgroup as defined in Armbruster (2003b, 2004), Hollanda Carvalho and Weber (2004), and Armbruster and de Souza (2005). Few species of the H. plecostomus group (excepting species of the H. cochliodon ZBK subgroup) have been described from the Orinoco River basin (just H. argus ) and none from the Negro (Weber, 2003, pers. obs.). In addition, there is a species of Hypostomus ZBK in the Orinoco that is very similar to H. robinii ZBK (described from Trinidad, pers. obs.). Some specimens of a Hypostomus ZBK with extremely small spots have been found in the Orinoco River drainage of southern Venezuela, and Armbruster (2004) referred these specimens to H. micromaculatus ZBK (a species described from Suriname, Boeseman, 1968). Examination of types of H. micromaculatus ZBK reveals that the Venezuelan specimens do not belong to this species, and comparison of the specimens to types or original descriptions of all other species of Hypostomus ZBK reveals that the specimens represent a new species.

  • Jonathan W. Armbruster, Leigh A. Tansey, Nathan K. Lujan (2007): Hypostomus rhantos (Siluriformes: Loricariidae), a new species from southern Venezuela. Zootaxa 1553, 59-68: 59-60, URL:http://www.zoobank.org/urn:lsid:zoobank.org:pub:F87041AF-3B5A-4F02-9D98-886814090969
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Ecology

Habitat

Depth range based on 42 specimens in 5 taxa.
Water temperature and chemistry ranges based on 1 sample.

Environmental ranges
  Depth range (m): 0.3 - 7
  Temperature range (°C): 18.742 - 18.742
  Nitrate (umol/L): 1.406 - 1.406
  Salinity (PPS): 22.907 - 22.907
  Oxygen (ml/l): 5.526 - 5.526
  Phosphate (umol/l): 0.707 - 0.707
  Silicate (umol/l): 9.028 - 9.028

Graphical representation

Depth range (m): 0.3 - 7
 
Note: this information has not been validated. Check this *note*. Your feedback is most welcome.

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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats
                                        
Specimen Records:462Public Records:155
Specimens with Sequences:402Public Species:23
Specimens with Barcodes:395Public BINs:7
Species:38         
Species With Barcodes:38         
          
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Barcode data: Hypostomus sp1

The following is a representative barcode sequence, the centroid of all available sequences for this species.


No available public DNA sequences.

Download FASTA File
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Statistics of barcoding coverage: Hypostomus sp1

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 1
Specimens with Barcodes: 1
Species With Barcodes: 1
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Statistics of barcoding coverage: Hypostomus cf. uruguayensis

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 0
Specimens with Barcodes: 1
Species With Barcodes: 1
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Statistics of barcoding coverage: Hypostomus cf. affinis

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 0
Specimens with Barcodes: 8
Species With Barcodes: 1
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Barcode data

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Locations of barcode samples

Collection Sites: world map showing specimen collection locations for Hypostomus

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Wikipedia

Hypostomus

Hypostomus is the largest genus of armored catfish. They are native to tropical and subtropical South America, with a single species, H. plecostomus, also found in Panama. This species is the popular freshwater aquarium fish formerly known as Plecostomus plecostomus. There is a lot of confusion as to the precise taxonomic structure of the Loricariidae.


Taxonomy and phylogeny[edit]

Hypostomus is the only genus in the tribe Hypostomini of the subfamily Hypostominae, and is the type genus of Hypostominae.[1][2] The type species of Hypostomus is H. guacari.[3]

With the inclusion of Aphanotorulus, Cheiridodus, Cochliodon, Isorineloricaria, Squaliforma, and Watawata, the genus Hypostomus is by far the largest genus of Loricariidae, and the second largest genus of catfish.[4][5] However, a good way to split up the genus further is unclear.[1] 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.[2] However, small monophyletic groups are known to exist within the genus Hypostomus, such as the H. unicolor group and the H. cochliodon group.[4] Aphanotorulus, Isorineloricaria, and Squaliforma are considered separate genera by some authors based on molecular information.[5]

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.[6]

Distribution and habitat[edit]

Hypostomus species 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. They are essentially ubiquitous throughout their range.[1] Maximum diversity in number of species of Hypostomus occurs in rivers of the Paraná-Paraguay system.[2]

Hypostomus species are found in almost all aquatic habitats, from large rivers to small rivulets.[7] 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 Hypostomus may be found over a range of substrates such mud, detritus, gravel, and sand.[1] Many species spawn in hollows dug into mud banks or within hollow logs.[1]

Description and ecology[edit]

As in all loricariids, Hypostomus 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 the 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.[8]

Also like other loricariids, Hypostomus species exhibit plates of armor over their body.[1] 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.[1] The caudal fin is forked with the lower lobe longer than upper. Most species have stout bodies, but some do have thinner, elongate bodies.[1]

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.[1]

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.[9]

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.[10]

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.[10]

Species[edit]

There are currently 141 recognized species in this genus:[11][12][13][14]

References[edit]

  1. ^ a b c d e f g h i Armbruster, Jonathan W. "Hypostomus". Retrieved 2007-05-06. 
  2. ^ a b c 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. Retrieved 2009-06-23. 
  3. ^ Froese, Rainer, and Daniel Pauly, eds. (2007). Species of Hypostomus in FishBase. May 2007 version.
  4. ^ a b c 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. Retrieved 2009-06-23. 
  5. ^ a b c 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. 
  6. ^ Alves, Anderson Luís; Oliveira, Claudio; Foresti (2005). "Comparative cytogenetic analysis of eleven species of subfamilies Neoplecostominae and Hypostominae (Siluriformes: Loricariidae)". Genetica 124 (2–3): 127–136. doi:10.1007/s10709-004-7561-4. PMID 16134327. 
  7. ^ a b c 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. 
  8. ^ Garavello, J. C.; Garavello, J. P. (August 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). Braz. J. Biol. 64 (3b): 103–141. doi:10.1590/S1519-69842004000400006. Retrieved 2009-06-23. 
  9. ^ "Modifications of the Digestive Tract for Holding Air in Loricariid and Scoloplacid Catfishes" (PDF). Copeia (3): 663–675. 1998. Retrieved 2009-06-23. 
  10. ^ a b Armbruster, Jonathan W. (2003). "The species of the Hypostomus cochliodon group (Siluriformes: Loricariidae)" (PDF). Zootaxa 249: 1–60. Retrieved 2009-06-23. 
  11. ^ Froese, Rainer, and Daniel Pauly, eds. (2013). Species of Hypostomus in FishBase. April 2013 version.
  12. ^ a b Zawadzki, C.H., Birindelli, J.L.O. & Lima, F.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.
  13. ^ a b Zawadzki, C.H. & Carvalho, P.H. (2014): A new species of the Hypostomus cochliodon group (Siluriformes: Loricariidae) from the rio Aripuanã basin in Brazil. Neotropical Ichthyology, 12 (1): 43-52.
  14. ^ a b Zawadzki, C.H., Tencatt, L.F.C. & Froehlich, O. (2014): A new unicuspid-toothed species of Hypostomus Lacépède, 1803 (Siluriformes: Loricariidae) from the rio Paraguai basin. Neotropical Ichthyology, 12 (1): 97-104.
  15. ^ Cardoso, Y.P., Almirón, A., Casciotta, J., Aichino, D., Lizarralde, M.S. & Montoya-Burgos, J.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.
  16. ^ Martins, F.O., Marinho, M.M.F., Langeani, F. & Serra, J.P. (2012): A New Species of Hypostomus (Siluriformes: Loricariidae) from the Upper Rio Paraguay Basin, Brazil. Copeia, 2012 (3): 494-500.
  17. ^ Birindelli, Jose 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. Retrieved 2009-06-23. 
  18. ^ Zawadzki, C.H., de Oliveira, R.R. & Debona, T. (2013): A new species of Hypostomus Lacépède, 1803 (Siluriformes: Loricariidae) from the rio Tocantins-Araguaia basin, Brazil. Neotropical Ichthyology, 11 (1): 73-80.
  19. ^ Zanata, A.M., Sardeiro, B. & Zawadzki, C.H. (2013): A new dark-dotted species of Hypostomus Lacépède (Siluriformes: Loricariidae) from rio Paraguaçu, Bahia State, Brazil. Neotropical Ichthyology, 11 (2): 247-256.
  20. ^ Jerep, Fernando C.; Shibatta & Zawadzki (2007). "A new species of Hypostomus Lacepede, 1803 (Siluriformes: Loricariidae) from the upper rio Parana basin, southern Brazil". Neotropical Ichthyology 5 (4): 435–442. doi:10.1590/S1679-62252007000400002. 
  21. ^ Garavello, J.C., Britski, H.A. & Zawadzki, C.H. (2012): The cascudos of the genus Hypostomus Lacépède (Ostariophysi: Loricariidae) from the rio Iguaçu basin. Neotropical Ichthyology, 10 (2): 263-283.
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