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      Two new species and a new subgenus of toothed Brachyhypopomus electric knifefishes (Gymnotiformes, Hypopomidae) from the central Amazon and considerations pertaining to the evolution of a monophasic electric organ discharge

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          Abstract

          Abstract

          We describe two new, closely related species of toothed Brachyhypopomus ( Hypopomidae: Gymnotiformes: Teleostei) from the central Amazon basin and create a new subgenus for them. Odontohypopomus , new subgenus of Brachyhypopomus , is diagnosed by (1) small teeth present on premaxillae; (2) medialmost two branchiostegal rays thin with blades oriented more vertically than remaining three rays; (3) background color in life (and to lesser extent in preservation) distinctly yellowish with head and sides peppered with small, widely spaced, very dark brown stellate chromatophores that greatly contrast with light background coloration; (4) a dark blotch or bar of subcutaneous pigment below the eye; (5) electric organ discharge waveform of very long duration (head-positive phase approx. 2 milliseconds or longer, head-negative phase shorter or absent) and slow pulse repetition rate (3–16 Hz). The type species of the new subgenus, Brachyhypopomus (Odontohypopomus) walteri sp. n., is diagnosed by the following additional character states: (1) subcutaneous dark pigment at base of orbit particularly prominent, (2) body semi-translucent and nearly bright yellow background coloration in life, (3) a biphasic electric organ discharge (EOD) waveform of very long duration (between 3.5 and 4 milliseconds at 25° C) with head-positive first phase significantly longer than second head-negative phase in both sexes. Brachyhypopomus (Odontohypopomus) bennetti sp. n. is diagnosed by two character states in addition to those used to diagnose the subgenus Odontohypopomus : (1) a deep electric organ, visible as large semi-transparent area, occupying approximately 14–17% body depth directly posterior to the abdominal cavity in combination with a short, but deep, caudal filament, and (2) a monophasic, head-positive EOD waveform, approximately 2.1 milliseconds in duration in both sexes. These are the only described rhamphichthyoid gymnotiforms with oral teeth, and Brachyhypopomus bennetti is the first Brachyhypopomus reported to have a monophasic (head-positive) EOD waveform. Unlike biphasic species, the waveform of its EOD is largely unaffected by tail damage from predators. Such injuries are common among specimens in our collections. This species’ preference for floating meadow habitat along the major channels of the Amazon River basin may put it at particularly high risk of predation and “tail grazing.”

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          Nós descrevemos duas novas espécies proximamente relacionadas de Brachyhypopomus ( Hypopomidae: Gymnotiformes: Teleostei) da porção central da bacia Amazônica e criamos um novo subgênero para elas. Odontohypopomus , novo subgênero de Brachyhypopomus , é diagnosticado por (1) apresentar dentes pequenos no pré-maxilar; (2) pela forma dos dois raios branquiostegais mais internos, com expansão laminar fina orientada mais verticalmente do que nos três raios remanescentes; (3) coloração de fundo em vida (e, em menor grau, após preservação) distintamente amarelada, com a cabeça e flancos marcados por pequenos cromatóforos marrom escuro de formato estelar e bem espaçados entre si, contrastando fortemente com a tonalidade clara de fundo; (4) uma mancha ou barra escura de pigmento subcutâneo abaixo do olho; (5) forma da onda de descarga do órgão elétrico (DOE) de duração muito longa (fase positiva com aproximadamente 2 milisegundos ou mais, fase negativa mais curta ou ausente) e uma baixa taxa de repetição de pulsos (3–16 Hz). A espécie-tipo do novo subgênero, Brachyhypopomus (Odontohypopomus) walteri sp. nov., é adicionalmente diagnosticada pelos seguintes estados de caráter: (1) pigmentação subcutânea na base da órbita particularmente conspícua; (2) corpo semi-translúcido e com coloração geral em vida amarelo forte, (3) DOE com onda bifásica e de duração muito longa (entre 3,5 e 4 milisegundos a 25° C), com a fase inicial positiva significativamente mais longa do que a segunda fase (negativa) em ambos os sexos. Brachyhypopomus (Odontohypopopmus) bennetti sp. nov. é diagnosticada por dois estados de caráter, além daqueles utilizados para diagnosticar o subgênero Odontohypopomus : (1) órgão elétrico largo em vista lateral, visível como uma grande área semitransparente ocupando aproximadamente 14–17% da altura do corpo, medida imediatamente posterior à cavidade abdominal, combinada com um filamento caudal curto e alto; e (2) DOE monofásica positiva, com duração de aproximadamente 2,1 milisegundos em ambos os sexos. Esses são os únicos Gymnotiformes Rhamphichthyoidea com dentes orais e Brachyhypopomus bennetti é a primeira espécie de Brachyhypopomus conhecida por apresentar uma DOE monofásica positiva. Diferentemente das espécies com DOE bifásica, a forma de onda/pulso dessa espécie não é significantemente afetada por danos ao filamento caudal decorrentes de predação. Esse tipo de dano é comum entre os espécimes examinados em nossas coleções. A ocorrência predominante dessa espécie em bancos flutuantes de macrófitas ao longo de alguns dos principais rios formadores da Bacia Amazônica pode representar um risco particularmente alto de predação e mutilação do filamento caudal.

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          Amazonian ecology: tributaries enhance the diversity of electric fishes.

          Neotropical rivers support a diverse array of endemic taxa, including electric fishes of the order Gymnotiformes. A comprehensive survey of the main channels of the Amazon River and its major tributaries (>2000-kilometer transect) yielded 43 electric fish species. Biogeographical analyses suggest that local mainstem electric fish diversity is enhanced by tributaries. Mainstem species richness tends to increase downstream of tributary confluences, and species composition is most similar between tributaries and adjacent downstream mainstem locations. These findings support a "nodal" or heterogeneous model of riverine community organization across a particularly extensive and diverse geographical region.
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            Stimulus filtering and electroreception: Tuberous electroreceptors in three species of Gymnotoid fish

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              Predation enhances complexity in the evolution of electric fish signals.

              Theories of sexual selection assume that predation is a restrictive, simplifying force in the evolution of animal display characters and many empirical studies have shown that predation opposes excessive elaboration of sexually selected traits. In an unexpected turnaround, I show here that predation pressure on neotropical, weakly electric fish (order Gymnotiformes) seems to have selected for greater signal complexity, by favouring characters that have enabled further signal elaboration by sexual selection. Most gymnotiform fish demonstrate adaptations that lower detectability of their electrolocation/communication signals by key predators. A second wave phase added to the ancestral monophasic signal shifts the emitted spectrum above the most sensitive frequencies of electroreceptive predators. By using playback trials with the predatory electric eel (Electrophorus electricus), I show that these biphasic signals are less detectable than the primitive monophasic signals. But sexually mature males of many species in the family Hypopomidae extend the duration of the second phase of their electric signal pulses and further amplify this sexual dimorphism nightly during the peak hours of reproduction. Thus a signal element that evolved for crypsis has itself been modified by sexual selection.
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                Author and article information

                Contributors
                Journal
                Zookeys
                Zookeys
                ZooKeys
                ZooKeys
                Pensoft Publishers
                1313-2989
                1313-2970
                2013
                28 August 2013
                : 327
                : 1-34
                Affiliations
                [1 ]Cornell University Museum of Vertebrates, 159 Sapsucker Woods Road, Ithaca, New York 14850 USA
                [2 ]Instituto Nacional de Pesquisas da Amazônia, Coordenação de Biodiversidade, Avenida André Araújo 2936, CEP 69080-971, Manaus, Brazil
                [3 ]Biology Department, Morrill Science Center, University of Massachusetts, Amherst, Massachusetts 01003 USA
                Author notes
                Corresponding author: John P. Sullivan ( jpsullivan@ 123456cornell.edu )

                Academic editor: Carole Baldwin

                Article
                10.3897/zookeys.327.5427
                3807744
                24167419
                8d86f940-c02d-469a-b3f5-230a690dae9e
                John P. Sullivan, Jansen Zuanon, Cristina Cox Fernandes

                This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 8 May 2013
                : 9 August 2013
                Categories
                Article

                Animal science & Zoology
                gymnotiform,weakly electric fish,electric organ,electric organ discharge,eod,neotropical freshwater fishes,evolution

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