Geophagus abalios, López-Fernández, Hernán & Taphorn, Donald C., 2004

López-Fernández, Hernán & Taphorn, Donald C., 2004, Geophagus abalios, G. dicrozoster and G. winemilleri (Perciformes: Cichlidae), three new species from Venezuela, Zootaxa 439, pp. 1-27 : 6-11

Geophagus abalios n. sp.

( Figs. 2 b, 3–5, 9)

Holotype. MCNG 47600, 163.0 mm SL; Venezuela: Apure: Río Cinaruco: Laguna Larga (6.5339 N 67.4150 W); K. Winemiller, H. López­Fernández, D.A. Arrington, L. Kelso­ Winemiller, H. López­Chirico and J. Arrington, 1­3 Jan 1999.

Paratypes. MCNG 30939, 3, 86.0–115.0 mm SL; Venezuela: Anzoátegui: Río Orinoco: Laguna Tineo (8.1903°N 63.4722°W); M.A. Rodríguez, 0 4 April 1987. — MCNG 33723, 3, 54.3–132.0; Venezuela: Bolívar: Río Orinoco: Laguna Bartolico (7.6417°N 66.1167°W); M.A. Rodríguez, 13 Jan 1987. — MCNG 35035, 1, 74.4 mm SL; Venezuela: Amazonas: Río Casiquiare: Playa Macanilla (2.4331 N 66.4547 W); K. O. Winemiller and D. Jepsen, 31 Jan 1997. — MCNG 40878, 1, 112.1 mm SL; Venezuela: Apure: Río Cinaruco: Laguna Guayaba; D.A. Arrington and J. Arrington, 12 April 1999. — MCNG 41124, 2, 45.5–55.5 mm SL; Venezuela: Apure: Río Cinaruco; D.A. Arrington and J. Arrington 14 April 1999. — AMNH 233634 (ex­MCNG 44865), 1, 96.3 mm SL; Venezuela: Apure: Río Cinaruco (6.5333 N 67.4164 W); D.A. Arrington and J.A. Arrington, 16 March 1999. — MCNG 47602 (ex­MCNG 6278), 1, 151.0 mm SL; Venezuela: Apure: Rio Cinaruco: Hato Las Delicias (6.5750°N 67.2361°W); D.C. Taphorn, C. Lilyestrom and B. Stergios, 11 Jan 1982. — MCNG 47601, 2, 96.3–160.0 mm SL; collected with holotype. — AMNH 93052, 2, 132.9–150.0 mm SL; Venezuela: Amazonas: Río Mavaca: small tributary on left bank; C.J. Ferraris and R. Royero, 10 March 1989.

Diagnosis. The lack of head markings distinguishes G. abalios n. sp. from Geophagus grammepareius , G. taeniopareius , G. harreri and G. argyrostictus , which have a complete infraorbital stripe, and from G. dicrozoster n. sp., G. winemilleri n. sp., G. brachybranchus and G. proximus , which have a black preopercular marking. Preserved specimens of Geophagus abalios can be distinguished from all other Geophagus species without head markings except G. brokopondo by the possession of six vertical, parallel bars on the flank ( Fig.1 , 2 ); it can be distinguished from G. brokopondo by the anterior three bars, which are medially bisected by a clearer area, giving the impression of two thinner bars, whereas in the latter species all bars are solid; additionally, the sixth bar in G. abalios is elongate and restricted to the dorsal half of the caudal peduncle, above the lower lateral line, and in G. brokopondo the line covers the entire caudal peduncle ( Fig. 2 b and 2d).

Description. Based on holotype (163.0 mm SL) and 16 paratypes 45.5–192.0 mm SL with notes on variation among smaller specimens. Measurements and counts are summarized in Table 1 . Sexes appear to be isomorphic.

Shape. Moderately elongate; dorsal outline more convex than ventral outline; head slightly broader ventrally than dorsally, chest flat; specimens 45.0 mm SL and smaller more elongate, with rounder nape; interorbital area moderately concave. Dorsal head profile straight, slightly concave in front of orbit, straight or slightly convex in specimens smaller than 112.0 mm SL, then sloping to dorsal­fin origin; dorsal­fin base descending, slightly convex to last ray, dorsal caudal peduncle forming a moderately concave curve to caudal­fin base. Ventral head profile straight, slightly descending to pelvic­fin insertion; chest slightly convex in one specimen 192.0 mm SL; straight, horizontal from pelvic­fin insertion to origin of anal fin; anal­fin base straight, ascending; ventral caudal peduncle straight to slightly concave, slightly ascending or horizontal in specimens 45.0 mm SL and smaller; ventral caudal peduncle 1.5–1.6 times in dorsal. Lips moderately wide, lower without caudally expanded fold (see Kullander et al., 1992, Fig. 3 ). Maxilla reaching at most one third of the distance between nostril and orbit; ascending premaxillary process reaching slightly above midline of orbit. Opercule, preopercule, cleithrum, postcleithrum, and post­temporal lacking serration.

Scales. E1 33(4), 34(10), 35(3); scales between upper lateral line and dorsal fin 5.5– 7.5 anteriorly, 2.5 posteriorly. Scales between lateral lines 2. Scales on upper lateral line 21(1), 22(4), 23(9), 24(1) and lower lateral line 13(1), 14(3), 15(6), 16(5). Anterior 1/3–1/ 2 of cheek naked, remainder with ctenoid scales; cheek scale rows 8–9. Opercule and subopercule covered with ctenoid scales. Interopercule with ctenoid scales caudally, otherwise naked. Single postorbital column of cycloid scales. Occipital and flank scales ctenoid. Circumpeduncular scale rows 7 above upper, 9 below lower lateral lines, ctenoid.

Fin scales. Pectoral and pelvic fins naked. Dorsal fin with double or triple columns of ctenoid scales along interradial membranes to one third to one half of fin height. Scaly pad at base of dorsal fin formed by irregularly arranged small, ctenoid scales extending from first spine to fifth to seventh soft ray; specimens 55.5 mm SL or smaller, pad scales are cycloid or moderately ctenoid. Anal fin scaled on anterior section of soft portion, scales ctenoid, arranged in a single column along interradial membranes to one quarter to one third of fin height; anal fin naked in specimens 55.5 mm SL or less. Scaly pad on base of anal fin, scales small, ctenoid. Caudal fin entirely scaled except the tip of rays, and membranes between D3 and V3, scales ctenoid. Accessory caudal fin extension of lateral line between V4–V5, absent on dorsal lobe.

Fins. Dorsal XVII­11(1), XVII­12(1), XVIII­10(2), XVIII­11(6), XVIII­12(5), XIX­ 11(2); anal III­8 (14), III­9 (3). Dorsal spines increasing in length from first to sixth, equal length to ninth, then slightly shorter; lose membranes behind spine tips (lappets) acutely pointed, up to 1/3 the length of spines. Soft portion moderately expanded and pointed, reaching about 1/3 of caudal­fin length, rays 3–6 longest but not produced into filaments; specimens 56.0 mm SL and smaller with rounded soft portion, not quite reaching caudalfin base. Anal fin pointed, with 2nd and 3rd soft rays slightly produced, not reaching caudal fin or barely beyond its base in specimens 90.6 and 192.0 mm SL. Caudal fin emarginate with lobes of approximately the same length and without filaments; one specimen 112.1 mm SL with slightly produced rays D8 and V8. Pectoral fin elongate, more or less triangular, longest at 4th ray, reaching 1st or 2nd anal­fin soft rays, then progressively shorter ventrally. Pelvic fin triangular, first ray produced into a filament reaching 5th anal­fin soft ray; in one specimen 112.1 mm SL reaching over 1/2 of caudal­fin length; specimens 45.5 mm SL or less with rays only slightly produced, reaching at most 1st spine of anal fin.

Teeth. Outer row of upper jaw with 10–28, blunt, slightly recurved unicuspid teeth; much larger than in inner rows, extending along most of premaxillary length. 2–3 inner rows, separated by a clear gap from outer row; teeth very thin, pointed, straight or slightly recurved unicuspids. Inner rows parallel to outer over its length, not forming a tight pad. Outer row of lower jaw with 6–25 unicuspid, blunt, slightly recurved unicuspids; medial 4–5 teeth larger than rest on outer row, cylindrical, slightly recurved, blunt and more labially positioned than rest of row. Inner rows 3–4, only on medial third of dentary, separated from outer row by distinct gap; teeth long, thin, straight or slightly recurved, much smaller than outer row.

Gills. External rakers on first gill arch; 9(5), 10(3) on epibranchial lobe, 1 in angle and 12(7), 13(2) on ceratobranchial, none on hypobranchial. Microbranchiospines on the outer face of second to fourth arches. Gill filaments with narrow basal skin cover.

Tooth plates. Lower pharyngeal tooth plate elongate; width of bone 80% of length; dentigerous area 80% of width; 30 teeth in posterior row, 10 in median row. Anteriormost teeth subconical or subcylindrical, erect; most teeth laterally compressed and with small, low ridge rostrally, cusps on caudal half of teeth; lateral marginal teeth on anterior half like anteriormost, on caudal half smaller and thinner; posteromedial teeth much larger, nearly round in circumference, posterior cusps, almost blunt ( Fig. 5 ). Ceratobranchial 4 with 4 toothplates with 11, 28, 6 and 4 teeth.

Vertebrae. 14+18=32(1), 14+19=33(1), 15+18=33(3), 15+19=34(5), 15+20=35(1), 16+18=34(4); 11–13 epihemal ribs.

Color pattern in alcohol ( Fig. 3 ). Base color grayish yellow; nape, snout and upper lip darker gray, fading caudally to base color towards cheek; lower lip yellowish white. No markings on the head, preopercule immaculate. Opercule darker on dorsal third; lower half of opercule and subopercule dusky yellow; silvery white in some specimens, probably depending on preservation. Ventrally, gill cover yellowish white; white in some specimens; branchiostegal membrane grayish. Chest white laterally and ventrally; in best preserved specimens white extends ventrally to base of caudal fin and to scale row H3 on caudal peduncle ( Fig. 1 ). Flanks with 6, dorso­ventrally directed, yellowish­gray bars fading or disappearing ventrally ( Fig. 2 b). Bar 1 expands from the 4th or 5th predorsal scale to the base of the 4th dorsal­fin spine; its anterior edge delimited by the extrascapular and its posterior edge descending vertically and disappearing ventrally at the pectoral­fin insertion. Bar 2 extends between the 6th and 8th dorsal­fin spines, and runs vertically to H7.

Bar 3 extends between the 10th and 13th dorsal­fin spines, and runs parallel to bar 2, fading ventrally at H6–H7. Bars 1–3 are generally bisected dorso­ventrally by a lighter column about 1 scale wide, giving the appearance of being two narrow bars in some specimens; this feature may be lost on poorly preserved specimens. A diffuse, blackish medial spot coincides with bar 3, extending rostro­caudally between scales 11–12 and 14–15 of E3 and dorso­ventrally between E3 and E1, such that the upper lateral line traverses the uppermost row of scales of the spot. Bar 4 extends between the bases of dorsal­fin spines 13–14 to 16–18, descends vertically and fades at H4–H5. Bar 5 extends between the first soft ray and ray 4–5 of dorsal fin, it descends vertically and disappears at H3–H4; in other specimens the bar is located between the last dorsal­fin spine and ray 3. Bar 6 extends from the base of the 6–7 (4–5 in some specimens) dorsal­fin rays and extends to the base of the caudal fin; bar is restricted to dorsal portion of caudal peduncle, above lower lateral line, and is longer horizontally than vertically ( Fig. 2 b).

Dorsal fin dusky, lappets dark gray or blackish, forming a dark edge along fin; soft and posterior third of spinous portion white­spotted on interradial membranes; four distinguishable longitudinal, parallel, grayish stripes alternate with light stripes along most of fin, turning almost hyaline rostrally; number of stripes increases with size to 6 in a 192.0 mm SL specimen. Anal fin hyaline to slightly dusky; 4 longitudinal, parallel gray stripes along soft portion of fin (5 in largest specimen). Caudal fin dusky, with round, whitish spots increasing in size towards dorsal edge; spots develop into horizontal stripes in larger specimens and a 192.0 mm SL specimen shows virtually no spots; specimens 55.5 mm SL and smaller with 4 dark, vertical bars. Pectoral fin immaculate. Pelvic fin whitish gray, dusky distally; dusky in largest specimen (192.0 mm SL), spine and first ray whitish gray to dusky.

Live colors ( Fig. 4 ). Background color greenish gray, breeding specimens more metallic gray. Head without markings except for iridescent blue on the upper lip, continued as a stripe extending to the corner of the preopercule, and a slight marking of the same color on the ventral edge of orbit. A variable number of iridescent blue spots on the preopercule apparently limited to breeding specimens. Six yellow stripes extend between the base of dorsal and H4–5; in adult, breeding specimens, dorsalmost stripes appear as brownish­orange vermiculations and spots. Ventrum distinctly white; breeding adults with bright orange or red chest. Dorsal and anal fins reddish with faint iridescent blue horizontal banding that turns brighter during breeding; caudal brownish red with iridescent blue spots and bands in no clear pattern; pelvic reddish orange with iridescent blue banding, first ray white or very light blue. An aquarium photograph in Weidner (2000: 148, Fig. 1 ) of an unidentified Geophagus from Venezuela is undoubtedly of a mature adult of G. abalios .

Distribution and habitat. ( Fig. 9 ) Geophagus abalios is commonly found in black or clear water rivers in the llanos, and is known from the Apure, Cinaruco­Capanaparo, and Aguaro­Guariquito drainages. Its current northern­most collection locality is “Las Majaguas” dam in the Río Cojedes, where it was probably introduced by recommendation of the Venezuelan ichthyologist A. Fernández­Yépez. According to his account (Fernández­Yépez and Anton, 1966), Geophagus species were not naturally present in the reservoir, and he recommended the introduction of " Geophagus surinamensis " along with some other species, presumably for sport fishing purposes. G. abalios reaches the Andean piedmont to the west, and is the only Geophagus found in clear to white water seasonal lagoons along the main­stem of the Orinoco to the east ( Rodríguez and Lewis Jr. 1990; 1994). The species appears restricted to the Caura drainage on the Guyana Shield, but it extends into the tributaries of the middle and upper Orinoco, including the Ventuari, Mavaca, and along the Río Casiquiare, nearly to the headwaters of the Río Negro.

Etymology. From the Greek a, not or without and balios, spotted. In reference to the lack of preopercular markings. To be regarded as an adjective in masculine form.

Figures

FIGURE 1. Diagrammatic representation of scale nomenclature, head markings and lateral bar patterns of Geophagus as used in this paper. Abbreviations are as follows: E 1, first epaxial longitudinal series of scales, used to count number of longitudinal scales; E 6, last epaxial longitudinal series of scales; H 1 – H 5, hypaxial longitudinal series of scales; IOS, infraorbital stripe; MLS, midlateral spot; POM, preopercular mark; ULL, upper lateral line; LLL, lower lateral line; 1 – 7, lateral bars. Scale nomenclature after Kullander (1992, 1996). Infraorbital, preopercular and mid­lateral black markings are observable both in live and preserved specimens; lateral bar patterns are gener­

FIGURE 2. Diagrammatic representation of preopercular markings and lateral bars distinguishing Geophagus species within the G. surinamensis complex. a, G. dicrozoster, n. sp.; b, G. abalios n. sp.; c, G. winemilleri n. sp.; d, G. brokopondo; e, G. brachybranchus; f, G. s u r i n a m e n s i s; g, G. proximus; h, G. megasema; i, G. camopiensis, and j, G. altifrons. Preopercular markings are visible in both live and preserved individuals; lateral bar patterns are generally visible only in preserved specimens.

FIGURE 3. Geophagus abalios. Holotype, MCNG 47600, 163.0 mm SL; Venezuela: Apure: Río Cinaruco: Laguna Larga (6.5339 N 67.4150 W).

FIGURE 4. Geophagus abalios. Uncatalogued specimen, adult in breeding coloration immediately after capture at the type locality: Laguna Larga, Río Cinaruco, Apure State, Venezuela. 6.3335 N, 67.2471 W.

FIGURE 5. Geophagus abalios. Lower pharyngeal toothplate on occlusal view; from MCNG 40636, 70.5 mm SL; scale bar 1 mm.

FIGURE 9. Known distribution area of Geophagus abalios n. sp. (), G. dicrozoster n. sp. (), and G. brachybranchus () in Venezuela. One dot may represent more than one collection locality.