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Brachyhypopomus Bullocki, a New Species of Electric Knifefish

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Brachyhypopomus Bullocki, a New Species of Electric Knifefish ISSN 0097-3157 PROCEEDINGS OF THE ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA 158: 183-192 APRIL 2009 Brachyhypopomus bullocki, a new species of electric knifefish (Gymnotiformes: Hypopomidae) from northern South America JOHN P. SULLIVAN Department of Ichthyology, The Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103-1195, USA, Email: [email protected] CARL D. HOPKINS Department of Neurobiology and Behavior, Cornell University, Ithaca, NY 14853-2702, USA, Email: [email protected] ABSTRACT.—We describe the new species Brachyhypopomus bullocki (Gymnotiformes: Hypopomidae) from the río Orinoco basin of Colombia and Venezuela as well as the Takutu River/rio Branco-rio Negro basin of Guyana and Brazil. Among its congeners, B. bullocki most closely resembles Brachyhypopomus brevirostris (Steindachner, 1868) in possessing an elongated caudal filament which may reach up to 45% TL in reproductive males, in having wide, evenly spaced saddles of pigment over the dorsum connecting to lateral bands of pigment along the abdominal flanks, and in having a mesocoracoid bone. This new species is distinguished from B. brevirostris and other members of the genus by a large eye (15.5–19.2% of head length), a short abdomen usually with 12 pre-caudal vertebrae, and poorly ossified third and fourth branchiostegal rays that are enlarged distally, producing a bulbous appearance of the opercular region. The electric organ discharge waveform of B. bullocki is biphasic, 0.9–1.6 milliseconds in duration, and the pulse rate varies from 20–80 Hz. RESUMEN.— Se describe la nueva especie Brachyhypopomus bullocki (Gymnotiformes: Hypopomidae) de la cuenca del río Orinoco de Colombia y Venezuela, como así también de la cuenca del Río Takutu/Rio Branco-Rio Negro de Guyana y Brasil. Entre las especies de este género B. bullocki se asemeja a Brachyhypopomus brevirostris (Steindachner, 1868) al poseer un filamento caudal que puede alcanzar hasta un 45% del largo total en los machos reproductivos, en tener pigmentos dorsales en forma de montura conectándose con bandas laterales a lo largo de los flancos abdominales, y en presentar un hueso mesocoracoides. Se distingue esta especie de B. brevirostris y de otros miembros del género por tener un ojo grande (15.5–19.2% del largo de la cabeza), un abdomen corto que presenta por lo general 12 vértebras pre-caudales, y el 3er y 4to radio branquiostego pobremente osificados y agrandados distalmente, dando una apariencia bulbosa a la región opercular. La onda de descarga del órgano eléctrico de B. bullocki es bifásica, de 0.9–1.6 milisegundos de duración, y la tasa de pulsación varía de 20–80 Hz. New taxon: Brachyhypopomus bullocki Sullivan and Hopkins INTRODUCTION forms from the longer-snouted Hypopomus Gill, 1864. In addition to the type species Brachyhypopomus brevirostris, The Family Hypopomidae is one of six nominal fami- Mago-Leccia (1994) recognized five other species in this lies of the Neotropical Order Gymnotiformes, all of which genus: B. occidentalis (Regan, 1914), B. beebei (Schultz, produce weak electric impulses for the purposes of electro- 1944), B. diazi (Fernández Yépez, 1972), B. pinnicaudatus location and communication. Hypopomids are nocturnally (Hopkins, 1991), and B. janeiroensis (Costa and Campos active, small to medium-sized gymnotiforms that are com- da Paz, 1992). So far in the 21st Century, three new mon inhabitants of lentic freshwater habitats from Panama Brachyhypopomus species have been described, all from to Uruguay. Here we describe a species of hypopomid the southern range of this genus in Southeastern Brazil gymnotiform from the Orinoco basin of Colombia and and Uruguay: B. jureiae Triques and Khamis (2003), B. Venezuela and the rio Negro basin of Brazil and Guyana. bombilla Loureiro and Silva (2006), and B. draco Gioria Brachyhypopomus is one of seven genera recog- et al. (2008). nized within the family Hypopomidae (Albert and In a morphology-based phylogenetic study of the gym- Crampton, 2003). Mago-Leccia (1994) created the notiforms, Albert (2001) regarded the Brachyhypopmus genus Brachyhypopomus to distinguish Rhamphichthys species recognized by Mago-Leccia (1994) and several brevirostris Steindachner, 1868 and similar short-snouted undescribed forms as a monophyletic group and listed 184 J.P. SULLIVAN AND C.D. HOPKINS four synapomorphies for the genus: (1) premaxilla gracile fered damage from predators. Specimens that had suffered with a curved anterior margin and forming a distinct angle damage anterior to the terminus of the anal fin were ident- with the maxilla in lateral view, (2) dentary gracile, (3) ified from radiographs, and excluded from measurements body cavity with 16 or 17 precaudal vertebrae, and (4) of SL and caudal filament length (CL). HL = head length, a single transitional vertebrae. In an unpublished dis- is taken from the tip of the snout to the bony end of the sertation, Sullivan (1997) regarded the monophyly of opercle, not to the uppermost limit of the branchial mem- the genus Brachyhypopomus to be poorly supported, brane. Interorbital width is the distance between the up- but placed the species described here into an unnamed per margins of the eyes. The term “branched” pectoral-fin clade with B. brevirostris that is diagnosed by (1) caudal rays refers to all rays posterior to the anterior unbranched filament exceedingly long in reproductive males, up to rays, even if the posterior terminal ray is unbranched. The 45% TL, (2) wide regular saddles of pigment over dorsum abbreviation “alc” is used to indicate specimens that are connecting to lateral bands, and (3) mesocoracoid present. preserved in alcohol, “cs” for those that have been cleared Although Mago-Leccia (1994) included the absence of and stained. Institutional abbreviations follow Ferraris a mesocoracoid bridge as a diagnostic character for this (2007) with the exception of MAC-PAY which is for the genus, Sullivan (1997) noted the presence of this element Ministerio de Agricultura y Cría, Estación Experimental in B. brevirostris and in this new species. Because of Amazonas, Puerto Ayacucho, Venezuela. its apparent close relationship to the type species of Because the lack of external landmarks on these fishes Brachyhypopomus, it seems likely that the species we de- makes measuring body depth from homologous landmarks scribe here will remain within Brachyhypopomus even difficult, radiographs were taken of specimens and distance if a future revision renders this genus less inclusive than from the tip of the snout to the image of the 1st, 20th and at present. Sexual dimorphism in the caudal filaments, 40th caudal vertebrae were measured. These distances were electric organ and electric organ discharge (EOD) of then measured off on the specimen itself and the depth of this species was the subject of a masters’ thesis by N.C. the body measured at each of these three points with digital Comfort (1990). needle-point calipers. Measurements are presented as per- centages of LEA except for measurements within the head MATERIAL AND METHODS that are presented as percentages of HL. Staining protocols for bone and cartilage followed We examined the type material for all described Potthoff (1984). In order to count columns of electrocytes Brachyhypopomus species with the exception of those in preserved specimens, caudal filaments were skinned recently described from southern Brazil and Uruguay (B. and observed with strong transmitted light. “Electrocyte jureiae, B. bombilla, and B. draco) for which we consulted columns” refers to the number of to the bilateral bands the published descriptions. Measurements were taken with of electrocytes along the longitudinal axis of the fish that a digital, needle-point caliper to within 0.1 mm under low begin under the head and continue to the tip of the caudal power magnification. All measurements were taken point- filament. These bands are most visible on the area immedi- to-point, i.e. not orthogonal to the main body axis. Counts ately above the posterior portion of the anal-fin base and on of anal-fin rays and vertebrae were made from radiographs the caudal filament. These columns can often be counted of the specimens, observed under magnification. All ver- with no special preparation by viewing the area with strong tebral counts began with C5, the first vertebra to carry a transmitted light. neural spine. “Precaudal vertebrae” include all anterior ver- To record the EOD waveforms, we transported tebrae bearing neural spines up to the first vertebra to bear freshly captured individuals to a field laboratory site in a hemal spine. Vertebrae bearing hemal spines are termed the home-stream water where we recorded the discharge “caudal vertebrae.” Counts of pectoral-fin rays, made with in a 10 cm x 40 cm x 12 cm aquarium with silver/silver- the aid of dissecting microscope and strong transmitted chloride electrodes positioned at the ends and a ground light, include all elements. Measurements were taken on electrode in the center. The signal was amplified with a the left side unless otherwise specified. CWE Corporation bio-amplifier with filters set to 0.1 Hz Anatomical measurements and abbreviations follow to 50,000 Hz and low gain. The fish always faced the Hubbs & Lagler (1958). Those that require explanation positive electrode. The amplified signals were digitized are: LEA = length from tip of snout to the posterior end of on a custom-made pulse logger (Helpware; Department the anal-fin base. This measurement is generally used as of Psychology, University of Sheffield) at 1 MHz (1µs standard length in descriptions of gymnotiforms since most per point), at 8 bit accuracy (4096 points). The pulse lack caudal fins and are often are often found to have re- logger was also used to collect short samples of interpulse generated a portion of their caudal region after having suf- intervals accurate to 1µS. A NEW SPECIES OF ELECTRIC KNIFEFISH FROM NORTHERN SOUTH AMERICA 185 Fig. 1. Brachyhypopomus bullocki.
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