Abstract
A phylogenetic study of the Loricariidae with emphasis on the Neoplecostominae is presented based on a maximum parsimony analysis of 268 phenotypic characters encompassing osteology, arthrology, and external morphology. Results support previous hypotheses of the monophyly of the Neoplecostominae and each of the included genera: Hirtella, Isbrueckerichthys, Kronichthys, Neoplecostomus, Pareiorhaphis, and Pareiorhina. In addition, previously undiscovered diversity was revealed within the subfamily as an additional genus-level taxon, herein described as Euryochus. Relationships among neoplecostomine genera are: (Kronichthys (Euryochus ((Hirtella + Pareiorhaphis) (Pareiorhina (Isbrueckerichthys + Neoplecostomus))))). Additional undescribed diversity was also detected among most neoplecostomine genera and the Hypoptopomatinae. In addition, recently discovered genera Nannoplecostomus and Microplecostomus were included in the analysis, and were identified as sequential sister-taxa to Neoplecostominae + Hypoptopomatinae, which are currently not included in any subfamily and regarded as incertae sedis in Loricariidae. The three species of Lithogenes were included in an encompassing phylogenetic analysis for the first time, and were identified as a monophyletic unit and sister group to all remaining loricariids. The other loricariid subfamilies were also corroborated as monophyletic, and presented the following interrelationships (Lithogeninae (Delturinae (Loricariinae (Hypostominae (Nannoplecostomus (Microplecostomus (Hypoptopomatinae + Neoplecostominae). The Neoplecostominae and its genera are phylogenetically diagnosed, and hypothesized relationships are compared to those of previous morphological and molecular phylogenetic studies.
References
Aquino, A. & Schaefer, S.A. (2002) Revision of Oxyropsis Eigenmann and Eigenmann, 1889 (Siluriformes, Loricariidae). Copeia, 2002, 374–390.
https://doi.org/10.1643/0045-8511(2002)002[0374:ROOEAE]2.0.CO;2Armbruster, J.W. (1997) Phylogenetic relationships of the sucker-mouth armored catfishes (Loricariidae) with particular emphasis on the Ancistrinae, Hypoptopomatinae and Neoplecostominae. Unpublished Ph.D. Dissertation, University of Ilinois, Urbana-Champaign, 409 pp.
Armbruster, J.W. (2004) Phylogenetic relationships of the suckermouth armoured catfishes (Loricariidae) with emphasis on the Hypostominae and the Ancistrinae. Zoological Journal of the Linnean Society, 141, 1–80.
https://doi.org/10.1111/j.1096-3642.2004.00109.xArmbruster, J.W. (2008) The genus Peckoltia with the description of two new species and a reanalysis of the phylogeny of the genera of the Hypostominae (Siluriformes: Loricariidae). Zootaxa, 1822, 1–76.
Armbruster, J.W., Sabaj, M.H., Hardman, M., Page, L.M. & Knouft, J.H. (2000) Catfish genus Corymbophanes (Loricariidae: Hypostominae) with description of one new species: Corymbophanes kaiei. Copeia, 2000, 997–1006.
https://doi.org/10.1643/0045-8511(2000)000[0997:CGCLHW]2.0.CO;2Arratia, G. (1987) Description of the primitive family Diplomystidae (Siluriformes, Teleostei, Pisces) morphology, taxonomy and phylogenetic implications. Bonner Zoologische Monographien, 24, 1–120.
Arratia, G. (2003) Catfish head skeleton and the siluriform postcranial skeleton. In: Arratia G., Kapoor, B.G., Chardon, M., Diogo. R. (Eds.), Catfishes. Science Publisher, Enfield, pp. 121–157.
Bailey, R.M. & Baskin, J.N. (1976) Scoloplax dicra, a new armored catfish from the Bolivian Amazon. Occasional Papers of the Museum of Zoology University of Michigan, 674, 1–14.
Baskin, J.N. (1973) Structure and relationships of the Trichomycteridae. Unpublished Phil. Thesis, City University of New York, New York City, 389 pp.
Bhatti, H.K. (1938) The integument and dermal skeleton of Siluroidea. Transactions of the Zoological Society of London, 24, 1–102.
https://doi.org/10.1111/j.1096-3642.1938.tb00395.xBizzeril, C.R.S.F. (1995) Description of a new species of Hemipsilichthys (Loricariidae: Hypostominae) from the state of Santa Catarina. Acta Biologica Leopoldensia, 17, 115–122.
Bremer, K. (1994) Branch support and tree stability. Cladistics, 10, 295–304.
https://doi.org/10.1111/j.1096-0031.1994.tb00179.xBritto, M.R. (2002) Análise filogenética da ordem Siluriformes com ênfase nas relações da superfamília Loricarioidea (Teleostei: Ostariophysi). Unpublished Ph.D. Dissertation, University of São Paulo, São Paulo, 512 pp.
Buckup, P.A., Menezes, N.A. & Ghazzi, M.S. (Eds.) (2007) Catálogo das espécies de peixes de água doce do Brasil. Museu Nacional, Rio de Janeiro, 195 pp.
Chardon, M. 1968. Anatomie comparée de l'appareil de Weber et des structures connexes chez les Siluriformes. Annales du Musée Royal de l'Afrique Centrale, Série 8, Sciences Zoologiques, 169, 1–277.
Chiachio, M.C., Oliveira, C. & Montoya-Burgos. J.I. (2008) Molecular systematic and historical biogeography of the armored Neotropical catfishes Hypoptopomatinae and Neoplecostominae (Siluriformes: Loricariidae). Molecular Phylogenetics and Evolution, 49, 606–617.
https://doi.org/10.1016/j.ympev.2008.08.013Cramer, C.A., Bonatto, S.L. & Reis, R.E. (2011) Molecular phylogeny of the Neoplecostominae and Hypoptopomatinae (Siluriformes: Loricariidae) using multiple genes. Molecular Phylogenetics and Evolution, 59, 43–52.
https://doi.org/10.1016/j.ympev.2011.01.002Cramer, C.A., Liedke, A.M.R., Bonatto, S.L. & Reis, R.E. (2008) The phylogenetic relationships of the Hypoptopomatinae and Neoplecostominae (Siluriformes: Loricariidae) as inferred from mitochondrial Cytochrome c Oxidase I sequences. Bulletin of Fish Biology, 9, 51–59.
Delariva, R.L. & Agostinho, A.A. (2001) Relationship between morphology and diets of six neotropical loricariids. Journal of Fish Biology, 58, 832–847.
https://doi.org/10.1111/j.1095-8649.2001.tb00534.xDerijst, E. (1996) Note on the type species of the mailed catfish genus Pareiorhaphis Miranda Ribeiro, 1918 (Pisces: Siluriformes: Loricariidae), with the introduction of Isbrueckerichthys nom. nov. Aquarium Wereld, 49, 62–64.
Eigenmann, C.H. (1910) Catalogue of the fresh-water fishes of tropical and south temperate America. In: Reports of the Princeton University expeditions to Patagonia 1896–1899. Zoology, 3, 375–511.
Eigenmann, C.H. & Eigenmann, R. (1890) A revision of the South American Nemathognathi, or cat-fishes. Occasional Papers of the California Academy of Sciences, 1, 1–508.
Eschmeyer, W.N. & Fong, J.D. (2016) Species by family/subfamily. Available from http://researcharchive.calacademy.org/research/ichthyology/catalog/SpeciesByFamily.asp. (accessed 30 September 2016)
Ferraris, Jr. C.J. (2003) Subfamily Neoplecostominae (Armored catfishes). In: Reis, R.E., Kullander, S.O. & Ferraris, Jr. C.J., (Eds.), Check List of the Freshwater Fishes of South and Central America. Edipucrs, Porto Alegre, pp. 319–320.
Ferraris, Jr. C.J. (2007) Checklist of catfishes, recent and fossil (Osteichthyes: Siluriformes), and catalogue of siluriform primary types. Zootaxa, 1418 (1), 1–628.
https://doi.org/10.11646/zootaxa.1418.1.1Fink, S.V. & Fink, W.L. (1981) Interrelationships of the ostariophysan fishes (Teleostei). Zoological Journal of the Linnean Society, 72, 297–353.
https://doi.org/10.1111/j.1096-3642.1981.tb01575.xGauger, M.F.W. & Buckup, P.A. (2005) Two new species of Hypoptopomatinae from the rio Paraíba do Sul basin, with comments on the monophyly of Parotocinclus and the Otothyrini (Siluriformes: Loricariidae). Neotropical Ichthyology, 3, 509–518.
https://doi.org/10.1590/S1679-62252005000400008Geerinckx, T. & Adriens, D. (2006) The erectile cheek-spine apparatus in bristlenose catfish Ancistrus (Loricariidae, Siluriformes), and its relation to the formation of a secondary skull roof. Zoology, 109, 287–299.
https://doi.org/10.1016/j.zool.2006.04.001Geerinckx, T., Poorter, J. & Adriens, D. (2007) Morphology and development of teeth and epidermal brushes in loricariid catfish. Journal of morphology, 268, 805–814.
https://doi.org/10.1002/jmor.10547Ghazzi, M.S. (2003) Relações filogenéticas e revisão do gênero Sturisoma Swainson, 1838 (Siluriformes: Loricariinae). Unpublished Ph.D. Dissertation, University of São Paulo, São Paulo, 340 pp.
Goloboff, P.A. (1999) Analyzing large data sets in reasonable times: Solutions for composite optima. Cladistics, 15, 415–418.
https://doi.org/10.1111/j.1096-0031.1999.tb00278.xGoloboff, P.A., Farris, J. & Nixon, K. (2008) TNT, a free program for phylogenetic analysis. Cladistics, 24, 774–786.
Gosline, W.A. (1945) Catálogo dos Nematognatos de Água Doce da América do Sul e Central. Boletim do Museu Nacional, Zoologia, 33, 1–138.
Gosline, W.A. (1947) Contributions to the classification of the loricariid catfishes. Archivos do Museu Nacional, 49, 79–134.
Grande, L. (1987). Redescription of †Hypsidoris farsonensis (Teleostei: Siluriformes), with a reassessment of its phylogenetic relationships. Journal of Vertebrate Paleontology, 7, 24–54.
https://doi.org/10.1080/02724634.1987.10011636Grant, T. & Kluge, A.G. (2008a) Credit where credit is due: The Goodman-Bremer support metric. Molecular Phylogenetics and Evolution, 49, 405–406.
https://doi.org/10.1016/j.ympev.2008.04.023Grant, T. & Kluge, A.G. (2008b) Clade support measures and their adequacy. Cladistics, 24, 1051–1064.
https://doi.org/10.1111/j.1096-0031.2008.00231.xHowes, G.J. (1983) The cranial muscles of loricarioid catfish, their homologies and value as taxonomic characters (Teleostei: Siluroidei). Bulletin of the British Museum (Natural History) Zoological Series, 45, 309–345.
https://doi.org/10.5962/bhl.part.28004Isbrücker, I.J.H. (1978) Descriptions préliminaries de nouveaux taxa de la famille des Loricariidae, poissons-chats cuirassés néotropicaux, avec un catalogue critique de la sous-famille nominale (Pisces, Siluriformes). Revue Française d'Aquariologie Herpétologie, 5, 86–117.
Isbrücker, I.J.H. (1980) Classification and catalogue of the mailed Loricariidae (Pisces, Siluriformes). Verslagen en Technische Gegevens, 22, 1–181.
International Union for Conservation of Nature [IUCN] (2016) Standards and Petitions Subcommittee. (2016) Guidelines for using the IUCN Red List Categories and Criteria. Version 12. Prepared by the Standards and Petitions Subcommittee. Available from: http://cmsdocs.s3.amazonaws.com/RedListGuidelines.pdf (accessed 30 September 2016)
Jerep, F.C., Shibatta, O.A., Pereira, E.H.L. & Oyakawa, O.T. (2006) Two new species of Isbrueckerichthys Derijst, 1996 (Siluriformes: Loricariidae) from the rio Paranapanema basin, Brazil. Zootaxa, 1372, 53–68.
Kluge, A.G. & Grant, T. (2006) From conviction to anti-superfluity: old and new justifications of parsimony in phylogenetic inference. Cladistics, 22, 276–288.
https://doi.org/10.1111/j.1096-0031.2006.00100.xLangeani, F. (1990) Revisão do gênero Neoplecostomus, com a descrição de quatro espécies novas do sudeste brasileiro (Ostariophysi, Siluriformes, Loricariidae). Comunicações do Museu de Ciências da PUCRS, Série Zoologia, 3, 3–31.
Lehmann, P. (2006) Anatomia e relações filogenéticas de Loricariidae (Ostariophysi: Siluriformes) com ênfase na subfamília Hypoptopomatinae. Unpublished Ph.D. Dissertation, Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre, 420 pp.
Lujan, N.K., Armbruster, J.W., Lovejoy, N.R. & López-Fernández, H. (2015) Multilocus molecular phylogenyof the suckermouth armored catfishes (Siluriformes: Loricariidae) with a focus on subfamily Hypostominae. Molecular Phylogenetics and Evolution, 82, 269–288.
https://doi.org/10.1016/j.ympev.2014.08.020Lundberg, J.G. & Baskin, J.N. (1969) The caudal skeleton of the catfishes, order Siluriformes. American Museum Novitatates, 2398, 1–49.
Maddison, W.P. & Maddison, D.R. (2011) Mesquite: a modular system for evolutionary analysis. Version 2.75. Available from: http://mesquiteproject.org (accessed 13 March 2017)
Martins, F.O., Britski, H.A. & Langeani, F. (2014) Systematics of Pseudotothyris (Loricariidae: Hypoptopomatinae). Zoological Journal of the Linnean Society, 170, 822–874.
https://doi.org/10.1111/zoj.12107Miranda Ribeiro, A. (1918) Três gêneros e dezessete espécies novas de peixes brasileiros. Revista do Museu Paulista, 10, 631–646.
Miranda Ribeiro, A. (1924) Ainda Hemipsilichthys e gêneros alliados. Boletim do Museu Nacional, 1, 365–366.
Mo, T. (1991) Anatomy, relationships and systematics of the Bagridae (Teleostei: Siluroidei) with a hypothesis of siluroid phylogeny. Unpublished Ph.D. Dissertation. Koeltz Scientific Books, Königstein, vii + 216 pp.
Montoya-Burgos, J.I. (2001) Phylogenetic relationships of the Hypostominae (Siluriformes: Loricariidae) with investigations on the phylogeny and evolution of the catfishes. Unpublished Ph.D. Dissertation, University of Genève, Genève, 198 pp.
Montoya-Burgos, J.I., Muller, S., Weber, C. & Pawlowski, J. (1998) Phylogenetic relationships of the Loricariidae (Siluriformes) based on mitochondrial rRNA gene sequences. In: Malabarba, L.R., Reis, R.E., Vari, R.P., Lucena, Z.M.S. & Lucena, C.A.S. (Eds.), Phylogeny and classification of Neotropical fishes. Edipucrs, Porto Alegre, pp. 363–374.
Muller, S. & Weber, C. (1992) Les dents des sous-familles Hypostominae et Ancistrinae (Pisces, Siluriformes, Loricariidae) et leur valeur taxonomique. Revue Suisse de Zoologie, 99, 747–754.
https://doi.org/10.5962/bhl.part.79851Nixon, K.C. (1999a) The Parsimony Ratchet, a new method for rapid parsimony analysis. Cladistics, 15, 407–414.
https://doi.org/10.1111/j.1096-0031.1999.tb00277.xNixon, K.C. (1999b) Winclada. Version 1.0. Computer software distributed by the author, Ithaca, New York.
Ørvig, T. (1977) A survey of odontodes (“dermal teeth”) from developmental, structural, funcional, and phyletic points of view. In: Andrews, S.M., Miles, R.S., Walker, A.D. (Eds.). Linnean Society Symposium Series, 4, 53–75.
Paixão, A.C. & Toledo-Piza, M. (2009) Systematics of Lamontichthys Miranda-Ribeiro (Siluriformes: Loricariidae), with descriptions of two new species. Neotropical Ichthyology, 7, 519–568.
https://doi.org/10.1590/S1679-62252009000400002Pereira, E.H.L. (2005) Ressurrection of Pareiorhaphis Miranda Ribeiro, 1918 (Teleostei: Siluriformes: Loricariidae), and description of a new species from the rio Iguaçu basin, Brazil. Neotropical Ichthyology, 3, 271–279.
https://doi.org/10.1590/S1679-62252005000200004Pereira, E.H.L., Oliveira, J.C. & Oyakawa, O.T. (2000) Hemipsilichthys papillatus, a new species of loricariid catfish (Teleostei: Siluriformes) from Minas Gerais, Brazil. Ichthyological Exploration of Freshwaters, 11, 377–383.
Pereira, E.H.L. & Oyakawa, O.T. (2003) Isbrueckerichthys epakmos, a new species of loricariid catfish from the rio Ribeira de Iguape basin, Brazil (Teleostei: Siluriformes). Neotropical Ichthyology, 1, 3–9.
https://doi.org/10.1590/S1679-62252003000100001Pereira, E.H.L. & Reis, R.E. (2002) Revision of the loricariid genera Hemipsilichthys and Isbrueckerichthys (Teleostei, Siluriformes), with descriptions of five new species of Hemipsilichthys. Ichthyological Exploration of Freshwaters, 13, 97–146.
Pereira, E.H.L., Vieira, F. & Reis, R.E. (2007) A new species of sexually dimorphic Pareiorhaphis Miranda Ribeiro, 1918 (Siluriformes: Loricariidae) from the rio Doce basin, Brazil. Neotropical Ichthyology, 5, 443–448.
https://doi.org/10.1590/S1679-62252007000400003Pereira, E.H.L., Zanata, A., Cetra, M. & Reis, R.E. (2014) A remarkable sexually dimorphic new genus and species of neoplecostomine catfish (Siluriformes, Loricariidae) from a coastal drainage of eastern Brazil. Copeia, 2014, 673–681.
https://doi.org/10.1643/CI-14-075de Pinna, M.C.C. (1992) A new subfamily of the Trichomycteridae (Teleostei, Siluriformes), lower loricarioid relationships and a discussion on the impact of additional taxa for phylogenetics analysis. Zoological Journal of the Linnean Society, 106, 175–229.
https://doi.org/10.1111/j.1096-3642.1992.tb01247.xde Pinna, M.C.C. (1993) Higher-level phylogeny of Siluriformes (Teleostei, Ostariophysi), with a new classification of the order. Unpublished Ph.D. Dissertation, City University of New York, New York City, 474 pp.
de Pinna, M.C.C. (1996) A phylogenetic analysis of the Asian catfish families Sisoridae, Akisidae and Amblycipitidae with a hypothesis on the relationships of the neotropical Aspredinidae (Teleostei, Ostariophysi). Fieldiana, 1478, 1–83.
de Pinna, M.C.C. (1998) Phylogenetic relationships of Neotropical Siluriformes: Historical overview and synthesis of hypothesis. In: Malabarba, L.R., Reis, R.E., Vari, R.P., Lucena, Z.M.S. & Lucena, C.A.S. (Eds.), Phylogeny and classification of Neotropical fishes. Edipucrs, Porto Alegre, pp. 279–330.
de Pinna, M.C.C, Ferraris, C.J. & Vari, R.P. (2007) A phylogenetic study of the neotropical catfish family Cetopsidae (Osteichthyes, Ostariophysi, Siluriformes), with a new classification. Zoological Journal of the Linnean Society, 150, 755–813.
https://doi.org/10.1111/j.1096-3642.2007.00306.xPy-Daniel, L.H.R. (1997) Phylogeny of the Neotropical armored catfishes of the subfamily Loricariinae (Siluriformes: Loricariinae). Unpublished Ph.D. Dissertation, University of Arizona, 280 pp.
Py-Daniel, L.H.R. & Fernandes, C.C. (2005) Dimorfismo sexual em Siluriformes e Gymnotiformes (Ostariophysi) da Amazônia. Acta Amazonica, 35, 97–110.
https://doi.org/10.1590/S0044-59672005000100015Quevedo, R. & Reis, R.E. (2002) Pogonopoma obsurum: A new species of loricariid catfish (Siluriformes: Loricariidae) from Southern Brazil, with comments on the genus Pogonopoma. Copeia, 2002, 402–420.
https://doi.org/10.1643/0045-8511(2002)002[0402:POANSO]2.0.CO;2Regan, C.T. (1904) A monograph of the fishes of the family Loricariidae. Transactions of the Zoological Society of London, 17, 191–350.
https://doi.org/10.1111/j.1096-3642.1904.tb00040.xRegan, C.T. (1920) XV—Pisces. Zoological Record [1918], 55, 1–19.
Reis, R.E. (1998) Anatomy and phylogenetic analysis of the neotropical callichthyid catfishes (Ostariophysi, Siluriformes). Zoological Journal of the Linnean Society, 124, 105–168.
https://doi.org/10.1111/j.1096-3642.1998.tb00571.xReis, R.E. (2013) Conserving the freshwater fishes of South America. International Zoo Yearbook, 47, 65–70.
https://doi.org/10.1111/izy.12000Reis, R.E., Albert, J.S., Di Dario, F., Mincarone, M.M., Petry, P. & Rocha, L.A. (2016) Fish biodiversity and conservation in South America. Journal of Fish Biology, 89, 12–47.
https://doi.org/10.1111/jfb.13016Reis, R.E., Kullander, S.O., Ferraris, Jr. C.J. (Eds.) (2003) Check list of the freshwater fishes of South and Central America. Edipucrs, Porto Alegre, 729 pp.
Reis, R.E., Pereira, E.H.L. & Armbruster, J.A.W. (2006) Delturinae, a new loricariid catfish subfamily (Teleostei, Siluriformes), with revisions of Delturus and Hemipsilichthys. Zoological Journal of the Linnean Society, 147, 277–299.
https://doi.org/10.1111/j.1096-3642.2006.00229.xRibeiro, A.C., Lima, F.C.T. & Pereira, E.H.L. (2012) A new genus and species of a minute suckermouth armored catfish (Siluriformes: Loricariidae) from the Rio Tocantins drainage, central Brazil: The smallest known loricariid catfish. Copeia, 2012, 637–647.
https://doi.org/10.1643/CI-11-137Roxo, F.F., Oliveira, C. & Zawadzki, C.H. (2012b) Three new species of Neoplecostomus (Teleostei: Siluriformes: Loricariidae) from the upper Rio Paraná basin of southeastern Brazil. Zootaxa, 3233, 1–21.
Roxo, F.F., Zawadzki, C.H., Silva, G.J.C., Chiachio, M.C., Foresti, F. & Oliveira, C. (2012a) Molecular systematics of the armored neotropical catfish subfamily Neoplecostominae (Siluriformes: Loricariidae). Zootaxa, 3390, 33–42.
Schaefer, S.A. (1987) Osteology of Hypostomus plecostomus (Linnaeus), with a phylogenetic analysis of the loricariid subfamilies (Pisces: Siluroidei). Contributions in Science of the Natural History Museum of Los Angeles County, 394, 1–31.
Schaefer, S.A. (1988) Homology and evolution of the opercular series in the loricarioid catfishes (Pisces: Siluroidei). Journal of Zoology, 214, 81–93.
https://doi.org/10.1111/j.1469-7998.1988.tb04988.xSchaefer, S.A. (1990) Anatomy and relationships of the scoloplacid catfishes. Proceedings of the Academy of Natural Sciences of Philadelphia, 142, 167–210.
Schaefer, S.A. (1991) Phylogenetic analysis of the loricariid subfamily Hypoptopomatinae (Pisces: Siluroidei: Loricariidae), with comments on generic diagnoses and geographic distribution. Zoological Journal of the Linnean Society, 102, 1–41.
https://doi.org/10.1111/j.1096-3642.1991.tb01535.xSchaefer, S.A. (1997) The Neotropical cascudinhos: systematics and biogeography of the Otocinclus catfishes (Siluriformes: Loricariidae). Proceedings of the Academy of Natural Sciences of Philadelphia, 148, 1–120.
Schaefer, S.A. (1998) Conflict and resolution: Impact of new taxa on phylogenetic studies of the neotropical cascudinhos (Siluroidei: Loricariidae). In: Malabarba, L.R., Reis, R.E., Vari, R.P., Lucena, Z.M.S. & Lucena, C.A.S. (Eds.), Phylogeny and classification of Neotropical fishes. Edipucrs, Porto Alegre, pp. 375–400.
Schaefer, S.A. (2003) Relationships of Lithogenes villosus Eigenmann, 1909 (Siluriformes, Loricariidae): Evidence from high-resolution computed microtomography. American Museum Novitates, 3401, 1–55.
https://doi.org/10.1206/0003-0082(2003)401<0001:ROLVES>2.0.CO;2Schaefer, S.A. & Buitrago-Suarez, U.A. (2002) Odontode morphology and skin surface features of Andean astroblepid catfishes (Siluriformes, Astroblepidae). Journal of Morphology, 254, 139–148.
https://doi.org/10.1002/jmor.10024Schaefer, S.A. & Lauder, G.V. (1986) Historical transformation of functional design: evolutionary morphology of feeding mechanisms in loricariioid catfishes. Systematic Zoology, 35, 489–508.
https://doi.org/10.2307/2413111Schaefer, S.A. & Lauder, G.V. (1996) Testing historical hypothesis of morphological change: Biomechanical decoupling in loricarioid catfishes. Evolution, 50, 1661–1675.
https://doi.org/10.2307/2410902Schaefer, S.A. & Provenzano, F. (2008) The Lithogeninae (Siluriformes, Loricariidae): Anatomy, interrelationships, and description of a new species. American Museum Novitates, 3637, 1–49.
https://doi.org/10.1206/625.1Schaefer, S.A. & Stewart, D.J. (1993) Systematics of the Panaque dentex species group (Siluriformes: Loricariidae), wood-eating armored catfishes from tropical South America. Ichthyological Exploration of Freshwaters, 4, 309–342.
Shelden, F.F. (1937) Osteology, myology and probable evolution of the nematognath pelvic girdle. Annals of the New York Academy of Sciences, 37, 1–96.
https://doi.org/10.1111/j.1749-6632.1937.tb55481.xSilva, G.S.C., Roxo, F.F., Ochoa, L.E. & Oliveira, C. (2016) Description of a new catfish genus (Siluriformes, Loricariidae) from the Tocantins River basin in central Brazil, with comments on the historical zoogeography of the new taxon. ZooKeys, 598, 129–157.
https://doi.org/10.3897/zookeys.598.7400Sullivan, J.P., Lundberg, J.G. & Hardman, M. (2006) A phylogenetic analysis of the major groups of catfishes (Teleostei: Siluriformes) using rag1 and rag2 nuclear gene sequences. Molecular Phylogenetics and Evolution, 41, 636–662.
https://doi.org/10.1016/j.ympev.2006.05.044Taylor, W.R. & Van Dyke, G.C. (1985) Revised procedores for staining and clearning smal fishes and other vertebrates for bone and cartilage study. Cybium, 9, 107–119.
Weitzman, S.H. (2004) Hearing in catfishes. Especially that of the family Loricariidae. In: Evers H.-G., Seidel, I. (Eds.), Catfish Atlas. Vol 1. Mergus Verlag, Melle, pp. 31–39.
Zawadzki, C.H., Pavanelli, C.S. & Langeani, F. (2008) Neoplecostomus (Teleostei: Loricariidae) from the upper Rio Paraná basin, Brazil, with description of three new species. Zootaxa, 1757, 31–48.