Brachyhypopomus (Odontohypopomus) walteri, Sullivan, John P., Zuanon, Jansen & Cox Fernandes, Cristina, 2013

Sullivan, John P., Zuanon, Jansen & Cox Fernandes, Cristina, 2013, 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, ZooKeys 327, pp. 1-34 : 5-9

Brachyhypopomus (Odontohypopomus) walteri sp. n. Figs 3, 4, Appendix I;Tables 1,3

Holotype.

INPA 8941, tag no. 93-219, 163 mm TL, 126 mm LEA, sex undetermined, Amazonas, Brazil: floating meadow alongside of lake in the Paraná do Paracuúba, near mouth of Rio Negro and entrance to Lago Janauari, approx. 15 km due south of Manaus, 03°12.6'S, 059°59.4'W, J.P. Sullivan and J. Zuanon. 23 April 1993.

Paratypes

(20). Brazil: Amazonas: INPA 8926 (3 cs, tag nos. 93-58, 93-140, 93-156), collection data same as for holotype, 24 March– 18 April 1993; INPA 8896 (4 alc, tag nos. 93-18, 93-19, 93-22, 93-23, 102-122 mm LEA), Ilha da Marchantaria, Rio Solimões, emergent grasses on shore of island, approx. 15 km upstream from confluence with Rio Negro, near Manaus, approx. 03°14'S, 059°59'W, J.P. Sullivan and J. Zuanon, 9 March 1993; INPA 8880 (3alc, tag nos. 93-55, 93-56, 93-57, 108-116 mm LEA), locality same as for holotype, 24 March 1993; INPA 8939 (1 alc, specimen number 93-114, 125 mm LEA), locality same as for holotype, 10 April 1993; ANSP 194031 (1 alc, tag no. JPS11-1-93/13, 84 mm LEA), channel between Rio Solimões and Lago Tefé, approx. 03°21'S, 064°40'W, J.P. Sullivan et al., 1 November 1993; ANSP 194032 (1 alc, 110 mm LEA, tag no. JPS11-20-93/1), Rio Içá in roots of water hyacinth along margin of inlet,15 km upstream of the mouth of the Içá, approx. 03°06'S, 068°05'W, J.P. Sullivan et al., 20 November 1993; ANSP 194033 (1 alc, tag no. CALH-11-20-93/2, 97 mm LEA), near Santo Antonio do Içá and mouth of Rio Içá in floating vegetation, approx. 03°07'S, 067°57'W, J.P. Sullivan et al., 20 November 1993; CUMV 97641 (1 alc, damaged, 80 mm TL, tag no. JPF-93-187/1), Rio Negro above Manaus, 03°05.38'S, 060°27.02'W, J.P. Sullivan and J.P. Friel, 14 December 1993; CUMV 97642 (5 alc, 2 damaged: tag nos. JPF-93-188/3, 188/4, 104, 100 mm LEA, 3 intact: tag nos. JPF-93-188/2, 188/5, 188/6, 95-102 mm LEA), Rio Negro above Manaus, 03°05.59'S, 60°26.83'W, J.P. Sullivan and J.P. Friel, 14 December 1993.

Non-types.

Brazil: Amazonas: Rio Solimões drainage: INHS 70542 (4 of 10, alc, 71-162 mm LEA), Ilha da Marchantaria, approx. 03°14'S, 059°59'W, P. Bayley, 14 March 1978; MZUSP 30061 (1 alc, 70 mm LEA), Rio Tefé, Lago Mucura, M. Goulding, 5 August 1979; USNM 306874 (2 alc, 91 & 100 mm LEA), Paraná da Ilha da Marchantaria, approx. 03°14'S, 059°59'W, depth 0-1.3 meters P. Bayley, 25 April 1978; USNM 306919 (1 alc, 75 mm LEA), Lago Camaleão, Ilha da Marchantaria, approx. 03°14'S, 059°59' W, P. Bayley, 29 March 1977; INPA 33268 (3 alc), Coari, 03°51.17'S, 063°28.12'W, L. Rapp Py-Daniel et al., 13 September 2013; INPA 33253 (3 alc), Manacapuru, Canaboca III, 03°35.55'S, 060°50.15'W, L. Rapp Py-Daniel et al., 17 September 2003; INPA 30241 (8 alc, 63.65-113.62 mm LEA), São Paulo de Olivença, Rio Camatiã, comunidade Monte Sinai, approx. 03°27.57'S, 068°56.00'W, L. Rapp Py-Daniel et al., 4 April 2008; ANSP 194025 (5 alc, 59.9-126.2 mm LEA, 1 cs, 117.4 mm), same data previous. Rio Uatumã: INPA 39074 (1 alc), São Sebastião do Uatumã, right bank in front of São José do Jabote, 01°56.20'S, 058°17.78'W, L. Rapp Py-Daniel et al., 1 October 2011. Rio Japurá: INPA 9945 (30 alc), Lago Caetano ( várzea lake) 1 km N of Jarauá, 02°50.97'S, 064°55.70'W, W.G.R. Crampton, 8 January 1995. Rio Purus: INPA 17112 (20 alc), Paraná do Seixo, Lago Jari, 04°54.67'S, 062°21.42'W, L. Rapp Py-Daniel et al., 8 June 2001; INPA 29259 (12 alc), Beruri, Lago Ayapuá, Igarapé Ajará, 04°25.12'S, 062°15.60'W, L. Rapp Py-Daniel et al., 15 November 2007; INPA 17192 (27 alc), Igarapé das duas bocas, Paraná do Jari, 04°53.12'S, 062°20.18'W, L. Rapp Py-Daniel et al., 7 June 2001. Pará: Rio Tapajós drainage: INPA 32703 (1 alc) Rio Crepori, Jacareacanga, Igarapé do Cocho, 06°44.90'S, 056°54.72'W, W.S. Pedroza, 31 July 2008. Rio Amazonas drainage: INPA 33192 (1 alc), Almeirim, Paranaguara, 01°44.48'S, 053°10.25'W, J. Zuanon et al., 5 October 2003. Rondônia: Rio Guaporé-Madeira drainage: INPA 9721 (19 alc), 15 km above Guajará-Mirim on Rio Pacaás Novos, approx. 10°56.8'S, 065°14.3'W, G.M. dos Santos, 26 November 1983; INPA 9727 (78 alc), mouth of Rio Pacaás Novos into Rio Guaporé, G.M. dos Santos, 26 November 1983. Roraima: Rio Branco drainage: INPA 30748 (11 alc), near Boa Vista, 02°47.51'N, 060°40.15'W, L.N. Carvalho, 29 September 2006; INPA 30749 (8 alc), near Boa Vista, 02°47.51'N, 060°40.15'W, L.N. Carvalho, 29 September 2006. Ecuador: Río Napo drainage: FMNH 102276 (8 alc, 69-107 mm LEA), Río Napo, along edge of Lago Anangucocha, approx. 01°28.48'S, 077°33.73'W, D. Stewart et al., 13 October 1981.

Diagnosis.

Brachyhypopomus (Odontohypopomus) walteri sp. n. is diagnosed by the following three character states in combination with the character states listed above for Odontohypopomus : (1) patch of brown pigment below skin at base of orbit distinct; (2) body yellow and semi-translucent in living specimens; (3) caudal filament long and fine, greater than 20% of LEA in intact specimens; (4) EOD pulse duration very long (between 3.5 and 4 milliseconds at 25° C) with head-positive first phase of longer duration than second head-negative phase in both sexes (Fig. 2A, B).

In all other Brachyhypopomus with biphasic EODs, the second, head-negative phase of the EOD is nearly equal in amplitude or of greater duration than the head-positive first phase. Microsternarchus bilineatus has an EOD waveform of similar duration, but the second phase is roughly equal or longer than the first (and the repetition rate is far faster). No other species of Brachyhypopomus is as distinctly yellow in color, particularly in life.

This species can be distinguished from the similar Brachyhypopomus bennetti sp. n. by a shorter body (depth quickly tapers posteriorly: depth of body at 40th post-abdominal vertebra 36-41% of depth at first abdominal vertebra vs. 46-57% in Brachyhypopomus bennetti ), fewer anal fin rays (198-216 rays vs. 227-255 rays in Brachyhypopomus bennetti ), a shallower electric organ, and a long, fine caudal filament (length 20-32% of LEA vs. 10-19% of LEA in Brachyhypopomus bennetti ) with three or four bilateral columns of electrocytes at base of caudal filament (vs. six columns in Brachyhypopomus bennetti sp. n.). Subcutaneous pigment below eye is absent in other hypopomids and usually less conspicuous in the sister species Brachyhypopomus bennetti sp. n. The EOD waveform of Brachyhypopomus walteri sp. n. is biphasic in contrast to Brachyhypopomus bennetti 's monophasic EOD waveform. Brachyhypopomus bennetti sp. n. tends to be more darkly pigmented and less yellow and translucent.

Description.

Morphometric and meristic data are presented in Tables 1, 3 and 4. A Brachyhypopomus of moderate to small adult size for a hypopomid; largest specimen examined measures 175 mm TL, 125 mm LEA. Body very compressed, depth at posterior end of abdominal cavity 2.7-3.1 times body width. Body more compressed posteriorly, sides of body with only slight curvature posterior to abdominal cavity. Dorsal profile gently convex. Depth quickly tapers posteriorly: depth of body at 40th post-abdominal vertebra 36-41% depth at first abdominal vertebra. Head short in comparison to body length, deep and wide: HL 11.2-12.6% LEA, head depth at occiput 72-81% HL, head width at opercle 54-63% HL. Head triangular in lateral view, dorsal profile of head straight from occiput to point of downturn of snout, ventral profile of head straight from lower jaw to opercular area with little if any concavity between opercular area and tip of lower jaw. Eye moderate in size, 12.4-14.5% HL. Mouth small, terminal, jaws equal, gape 20-23% HL. Closed lips meet ventral to a horizontal through ventral margin of eye. One to five small needle–like conical teeth present on each premaxilla (Fig. 1), lower jaw edentate. Maxilla moderate in length, thin, with slight curvature. Snout moderate in length, 26-29% HL, edge of upper lip close to farthest anterior extent of snout. Posterior naris close to eye, posterior naris–eye 1.8-3.7% HL. Lateral ethmoid present. Round ossification present in anterior portion of palatine cartilage (Fig. 1). Infraorbital portion of cephalic lateralis system incomplete, lacking recurrent anterodorsal segment and associated pores beneath and anterior to the posterior nares that are present in most other Brachyhypopomus (see fig. 53 in Sullivan 1997); fourth supraorbital pore lying near vertical through posterior nostril, pores inconspicuous. Preopercular lateral-line canal embedded in preopercle, canals radiating out to pores. Pores of lateral-line canal immediately behind head without downward pointing tubes. Discernible lateral scales terminate along caudal filament. Five branchiostegal rays, medialmost two thin with blades oriented nearly vertically compared to outer three (see diagnosis of Odontohypopomus ). Gill rakers robust for genus, some with weakly ossified cores, on anterior faces of first four gill arches. Rakers subtended on ceratohyals one to four by small trough-shaped ossicles. Approximately 40 gill filaments on arch one. Three pectoral radials, all partially fused together at proximal end. Mesocoracoid bridge absent. Pectoral fin broad, 12-15 branched plus unbranched rays, length 5.3-7.0% LEA. 198-216 anal fin rays, longest rays 4.0-4.9% LEA. Precaudal vertebrae 13-16, up to 75 caudal vertebrae in advance of regenerated portion of caudal filament. Body excluding head and fins covered with thin cycloid scales, small dorsally, larger posterolaterally, partially obscured by skin. Twelve scale rows above, 13 scale rows below lateral line at farthest extent of pectoral fin. Anal-fin origin slightly posterior to vertical at midpoint of extended pectoral fin. Caudal filaments long and fine in intact mature specimens, 20-32% of TL. Sexual dimorphism of caudal filaments not observed. Three or four bilateral columns of electrocytes along caudal filament, number often alternating along length of caudal filament; 38-63 rows of electrocytes. Electrocytes do not extend farther anteriorly than base of urogenital pore. No accessory electric organs on head or humeral region.

Electric organ discharge.

TheEOD is biphasic and 3.2-4.5 milliseconds in total duration at 25°C; the first head-positive phase is 1.7-1.9 times duration of second head-negative phase (Fig. 2). Resting EOD repetition rate is slow (3.1-16.3 Hz, mean 9.4 Hz, median 9.8, at 21-25°C, n=23). See Appendix II.

Coloration.

Background color yellow in life, yellowish-tan in preservation. In life, body semi-translucent, with gill filaments appearing cherry red through opercle, gut dark, and swim bladder whitish through abdominal wall. Pigmentation variable: poorly to moderately developed irregular bands along sides, darker and wide above lateral line, often with a spot of darker intensity on lateral line itself. Bands either restricted to anterior portion of body above lateral line or connected to fainter bands below. Some bands connect to eight to 12 irregular saddles across dorsum. Saddles more regular in smaller individuals. Dorsal rami of the anterior lateral line nerve visible when viewed from above as two thin, dark parallel lines running along upper back beginning a short distance behind head and continuing to mid-point of the back. Cheeks, underside of head and sides of body below lateral line peppered with prominent dark brown stellate chromatophores that greatly contrast with background color of skin and that do not form part of a larger pattern. Diffuse pigment below eye resembling a teardrop is more prominent in live specimens as overlying tissue becomes opaque upon preservation. Pectoral and anal fin with irregular brown pigment along rays; interradial membranes hyaline.

Distribution and ecology.

See distribution map (Fig. 5). Brachyhypopomus walteri sp. n. is known only from the Amazon basin where it appears to be common in floating meadow habitats, (mostly composed of the grass Paspalum repens , Poaceae ), on the margins of the Amazonas/ Solimões and its tributaries. It has been collected predominantly in white water, but also in areas near the confluence of black water rivers with the Amazonas/ Solimões ranging from low to medium conductivity. Apart from one collection very near Manaus and the white water Rio Branco, it is absent from collections in the Rio Negro system. It is frequently taken with Brachyhypopomus bennetti sp. n. and sometimes with Brachyhypopomus brevirostris . Species of Eigenmannia , Gymnotus , the apteronotid Parapteronotus hasemani and the electric eel, Electrophorus electricus , frequently co-occur in the floating meadow habitats preferred by this species.

Etymology.

This species is named for Walter Heiligenberg (1938-1994) in honor of his discoveries in electric fish neurophysiology and behavior made at the Scripps Institute of Oceanography. These notably include the "jamming avoidance response" in Eigenmannia , often described as the best-understood vertebrate behavior.