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Academy of Natural Sciences Academy of Natural Sciences The Neotropical Cascudinhos: Systematics and Biogeography of the Otocinclus Catfishes (Siluriformes: Loricariidae) Author(s): Scott A. Schaefer Source: Proceedings of the Academy of Natural Sciences of Philadelphia, Vol. 148 (Oct. 31, 1997), pp. 1-120 Published by: Academy of Natural Sciences Stable URL: http://www.jstor.org/stable/4065046 Accessed: 26-03-2015 15:15 UTC REFERENCES Linked references are available on JSTOR for this article: http://www.jstor.org/stable/4065046?seq=1&cid=pdf-reference#references_tab_contents You may need to log in to JSTOR to access the linked references. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Academy of Natural Sciences is collaborating with JSTOR to digitize, preserve and extend access to Proceedings of the Academy of Natural Sciences of Philadelphia. http://www.jstor.org This content downloaded from 192.134.151.170 on Thu, 26 Mar 2015 15:15:03 UTC All use subject to JSTOR Terms and Conditions PROCEEDINGS OF THE ACADEMY OF NATURAL SCIENCES OF PHILADELPIA 148: 1-120. 31 OCTOBER 1997 The Neotropical cascudinhos:Systematics and biogeography of the Otocinclus catfishes (Siluriformes:Loricariidae) SCOTT A. SCHAEFER Department of Ichthyology,American Museumof Natural History, Central Park Westat 79th Street,New York, NY 10024-5192, USA ABSTRACT - The genus OtocinclusCope (1872) of the siluriform family Loricariidaeis diagnosed as monophyletic on the basis of shared derived characters of the cranial and hyobranchial skeleton, dorsal gill arch musculature, and gut. Otocinclus are relatively small herbivorous catfishes restrictedto small streams and quiet slow-flowing margins of larger rivers, most frequently living in close association with aquatic macrophytes and terrestrial marginal grasses extending into the water column. Otocinclusspecies share a novel modification of the distal esophageal wall which is developed into an accessory blind diverticulum that may function in aerial respiration and for providing additional modulatory positive buoyancy for remaining in the upper water column at stream margins. Otocinclus has no junior synonyms, however several nominal species originally described in Otocinclus are here formally re-assignedto other genera in the subfamily Hypoptopomatinae. Otocinclus cephalacanthusRibeiro 1911, 0. depressicaudaRibeiro 1918, 0. francirochai Ihering 1928, 0. laevior Cope 1894, 0. leptochilus Cope 1894, 0. maculipinnis Regan 1904, 0. nigricauda Boulenger 1891, and 0. paulinus Regan 1908 are all placed in the genus MicrolepidogasterEigenmann & Eigenmann 1889; 0. obtusos Ribeiro 1911 was placed in PseudotothyrisBritski & Garavello 1984; the genus Nannoptopoma Schaefer 1996 was erected for 0. spectabilisEigenmann 1914 in the tribe Hypoptopomatini; 0. gibbosusRibeiro 1908 is removed from Otocinclus,yet remains of undetermined generic status. Thirteen species are recognized in Otocinclus: 0. affinis Steindachner 1877 of the lower Parana/Paraguayand Uruguay basins and coastal streams of southeastern Brazil; 0. bororon. sp. of the upper Rio Paraguay;0. caxararin. sp. of the middle Rio Guapore/Mamore system; 0. flexilis Cope 1894 of the lower Parana/Paraguay and Uruguay basins and coastal streams of southeastern Brazil; 0. hasemaniSteindachner 1915 of northern Brazil; 0. hoppeiRibeiro 1939 of the upper Amazon, Tocantins and Paraguay basins and coastal streams of northeastern Brazil; 0. huaorani n. sp. of the upper Amazon and Orinoco basins; 0. macrospilusEigenmann & Allen 1942 of the upper Amazon basin of Colombia, Ecuador, and Peru; 0. mariae Fowler 1940 of the lower Amazon, upper Madeira and Paraguay basins; 0. mura n. sp. of the middle Amazon River; 0. vestitus Cope 1872 of the upper Amazon and lower Parana basins; 0. vittatus Regan 1904 of the Amazon, Orinoco, Parana/Paraguay,and Tocantins basins; and 0. xakriaba n. sp. of the rio Sao Fransisco basin. Two species are placed in synonymy: Otocinclusarnoldi Regan 1909 and 0. fimbriatus Cope 1894 are junior synonyms of 0. flezilis. Keys to the species of Otocinclus and genera of the Hypoptopomatinae are provided. A descriptive treatment of the osteology and cranial myology is provided for 0. vittatus. Detailed analysis of meristic and morphometric variation based on geometric morphometric procedures is provided for the phenetically similar species pairs 0. mariae and 0. vittatus, 0. bororo and 0. huaorani in an a posteriori evaluation of separate species status. The phylogenetic relationships among Otocinclusspecies, and the phylogenetic position of Otocinclus among genera of the Hypoptopomatinae, are determined based on analysis of 27 morphological featuresusing cladistic parsimony. Monophyly of Otocincluswas confirmed; within Otocinclus,a clade comprised of 0. affinis and 0. flexilis is the sister-group to the remainder of the genus. Within that latter clade, 0. hasemani and 0. xakriaba are the first and second-level sister-groups to the remainder of the genus, within which relationships among species are not fully resolved with available data. The phylogenetic biogeography of Otocinclusis informative regardingthe historical relationshipsamong major river drainage basins, particularly of those river systems of the Brazilian Shield. A biogeographic hypothesis is proposed based on the area cladogram derived from the species-level phylogenetic relationships, which suggests successive vicariance and speciation in the non-Amazonian regions of endemism of southeastern and eastern South America, followed by speciation and dispersal within the Amazon, Orinoco and upper Paraguay basins. The pattern of vicariance revealed by the Otocinclusspecies-level phylogeny is congruent with the geologic history of the major river drainage basins of the Brazilian Shield. This result suggests that, for Otocinclus and perhaps other loricariid catfishes, much of their generic and species-level diversification occurred prior to the formation of the Amazon basin. SPECIESNOVAE: Otocinclus bororo, 0. caxarari, 0. huaorani, 0. mura, 0. xakriaba. RESUMO - 0 genero Otocinclus Cope (1872) da famflia de siluriformes Loricariidae e diagnosticado como monofiletico baseado em caracteres derivados do cranio e do esqueleto hipobranquial, musculatura dorsal dos arcos branquiais, e v'sceras. Otocinclus sao bagres herbivoros, relativamente pequenos, restritos a pequenos riachos e a 1 This content downloaded from 192.134.151.170 on Thu, 26 Mar 2015 15:15:03 UTC All use subject to JSTOR Terms and Conditions 2 SCOTT A. SCHAEFER margens de baixa correnteza de rios largos, vivendo mais frequentemente em associac,ao 'a macrofitas aquaticas e gramineas terrestres que atingem a coluna d'agua. As especies de Otocinclus compartilham uma nova e exclusiva modificacao da parede distal do esofago que se desenvolveu em um divertlculo acessorio de fundo cego que talvez sirva para respiragao aerea e para prover modulagao positiva adicional na flutua,ao para permanencia na parte superior da coluna d'agua nas margens dos rios. Otocinclusnao possui sinonimos junior, no entanto, diversasespecses nominais originalmente descritasem Otocinclus, sao aqui transferidaspara outros generos da subfamfliaHypoptopomatinae. Otocincluscephalacanthus Ribeiro 1911, 0. depressicaudaRibeiro, 1918, 0. francirochai Ihering 1928, 0. laevior Cope 1894, 0. leptochilus Cope 1894, 0. maculipinnis Regan 1904, 0. nigricauda Boulenger 1891, e 0. paulinus Regan 1908 sao colocados no genero MicrolepidogasterEigenmann & Eigenmann 1899; 0. obtususRibeiro 1911 foi colocado em PseudotothyrisBritski & Garavello 1984; 0. spectabilisEigenmann 1914 representaum genero nao descrito da tribo Hypoptopomini; 0. gibbosus Ribeiro 1908 e removido de Otocinclus, mas seu status generico permanece indeterminado. Treze especies sao reconhecidas em Otocinclus:0. affinis Steindachner 1877 das bacias do baixo Paranai/Paraguaie Uruguai e rios costeiros do sudeste do Brasil; 0. bororosp. n. do alto rio Paraguai;0. caxaraisp. n. do sistema do medio rio Guapore/Mamore; 0. flexilis Cope 1894 das bacias do baixo Paranat/Paraguaie Uruguai e rios costeiros do sudeste do Brasil; 0. hasemani Steindachner 1915 do norte do Brasil; 0. hoppeiRibeiro 1939 das bacias do alto Amazonas, Tocantins e Paraguaie rios costeiros do nordeste do Brasil; 0. huaorani sp. n. das bacias do alto Amazonas e Orinoco; 0. macrospilusEigenmann & Allen 1942 do alto Amazonas da Colombia, Equador e Peru; 0. mariaeFowler 1940 das bacias do baixo Amazonas, alto Madeira e Paraguai;0. mura sp. n. do merdiorio Solimrnes;0. vestitus Cope 1872 das bacias do alto Amazonas e baixo Parana';0. vittatus Regan 1904 das bacias do Amazonas, Orinoco, Paranai/Paraguaie Tocantins; 0. xakriaba sp. n. da bacia do rio Sao Francisco. Duas especies sao colocadas em sinonirmia:Otocinclus arnoldi Regan 1909 e 0. fimbriatus Cope 1894 sao sinonimos junior de 0. flexilis. Sao fornecidas chaves de identificagao das especies de Otocinclus e dos generos de Hypoptopomatinae. Uma descrigao da osteologia e musculaturacraniana de 0. vittatus e fornecida. Uma an'alisedetalhada da variaIao mer stica e morfomertricabaseada em procedimentos
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