Apistogramma Eleutheria and A. Flavipedunculata, Two New Species of Dwarf Cichlids from the Rio Curuá on Serra Do Cachimbo, Brazil (Teleostei: Cichlidae)

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Apistogramma eleutheria and A. Flavipedunculata, two new species of dwarf cichlids from the rio Curuá on Serra do Cachimbo, Brazil (Teleostei: Cichlidae)

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Henrique Varella Ricardo Britzke Universidade Federal da Bahia São Paulo State University

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Apistogramma eleutheria and A. flavipedunculata, two new species of dwarf cichlids from the rio Curuá on Serra do Cachimbo, Brazil (Teleostei: Cichlidae)

Henrique R. Varella* and Ricardo Britzke**

Apistogramma eleutheria and Apistogramma flavipedunculata are described from the upper portion of two adjacent tributaries of the rio Curuá on the Serra do Cachimbo. Apistogramma eleutheria, new species, is isolated from the lower portion of rio Treze de Maio (a tributary of the middle rio Curuá), by a waterfall with same name. It is similar to several species of the A. regani species group, but differs by the coloration of brooding females. Brood- ing females of A. eleutheria have a very dark, blackish pigmentation on the median area of gular region occupying the ventral portion of opercular series and branchiostegal membranes, which is most similar to that found in females of A. kullanderi. In A. kullanderi, however, the dark gular pigmentation is more expansive, extending onto the ventral half of the head, and continuing posteriorly onto belly and along anal-fin base. Apistogramma flavipe dunculata, new species, is only known from the rio Escorpião, and is probably isolated in the upper rio Curuá by a series of three waterfalls. It is distinguished by the unique color pattern of adult males, which have the posterior part of the flank and caudal peduncle dominated by a light yellow pigmentation, which results in a dark midlateral band fragmented posteriorly and dark bars masked or absent (though present in juveniles); also differing from all congeners except A. luelingi and A. eleutheria by having exclusively four anal-fin spines instead of three.

Introduction much of tropical and subtropical South America, but most of the species of the genus are thought Apistogramma Regan, 1913 and Crenicichla Heckel, to have distributions restricted to a particular 1840 are the most species-rich genera among region or hydrographic basin (Kullander, 1986). Neotropical cichlids, the former with 89 species Only few species of the genus have wide distri- and the latter with 90 species (Kullander, 2003; butions, as Apistogramma agassizi (Steindachner, Kullander & Varella, 2015; Römer et al., 2015). The 1875), A. bitaeniata Pellegrin, 1936, A. cacatuoides geographic distribution of Apistogramma includes Hoedeman, 1951, A. commbrae (Regan, 1906) and

* Instituto de Biociências da Universidade de São Paulo, Museu de Zoologia da Universidade de São Paulo, Caixa Postal 42494, 04218-970 São Paulo, SP, Brazil. E-mail: [email protected] ** Universidade Estadual Paulista, Instituto de Biociências, Departamento de Morfologia, Rubião Jr. s/n., 18618- 970 Botucatu, SP, Brazil; Universidad Técnica de Machala, Facultad de Ciencias Agropecuarias, Laboratorio 1 de Acuacultura, Km. 4 /2 Vía Machala-Pasaje, E583, Machala, El Oro, Ecuador. E-mail: [email protected]

A. borellii (Regan, 1906). The highest diversity and include the last half-centrum and were taken from degree of endemism of Apistogramma is found radiographs made with the system Faxitron LX-60 in the Western Amazon, notably in Peru (with (DC 12, version 1.0). 24 valid species, 21 of them endemic), in the río Institutional abbreviations are: ANSP, Aca- Orinoco basin (19 endemic species) and in the rio demy of Natural Sciences of Philadelphia; LBP, Negro basin (13 endemic species). The diversity Laboratório de Biologia e Genética de Peixes, of Apistogramma in the Brazilian Shield remains Botucatu, São Paulo; MZUSP, Museu de Zoolo- little studied with many putative undescribed gia da Universidade de São Paulo; MCP, Museu species indicated for some drainages (e. g. Kos de Ciências e Tecnologia PUCRS, Porto Alegre; lowski, 2002). MUSM, Museo de Historia Natural, Universidad Apistogramma kullanderi Varella & Sabaj Pérez, Nacional Mayor de San Marcos, Lima; and ROM, 2014, from the upper rio Curuá on the Serra do Royal Ontario Museum, Toronto. Comparative Cachimbo, is the only nominal species described material examined is as listed by Varella & Sabaj for the rio Xingu drainage. The Serra do Cachimbo Pérez (2014) and Britzke et al. (2014), with addition is the highest portion of the Brazilian Shield in of the material listed at the end of this paper. the Amazon basin that includes headwaters from both middle Xingu (i. e., rio Curuá) and rio Tapajós (rio Jamanxim and rio Teles Pires) river basins Results (Birindelli et al., 2009). Inventories and taxonomic studies of the region have shown high degrees Apistogramma eleutheria, new species of endemism for various fishes. For instance, the (Fig. 1) upper rio Curuá presents six endemic species described only recently: Leporinus guttatus Birindelli Apistogramma sp. ‘upper Treze de Maio’: Varella & Britski, 2009, Jupiaba kurua Birindelli et al., 2009, & Sabaj Pérez, 2014: 255. Moenkhausia petymbuaba Lima & Birindelli, 2006, Lebiasina marilynae Netto-Ferreira, 2012, L. melano Holotype. MZUSP 115293, 39.5 mm SL, adult guttata Netto-Ferreira, 2012 and Apistogramma male; Brazil: Pará: Altamira: rio Xingu basin: rio kullanderi Varella & Sabaj Pérez, 2014. Iriri drainage: rio Treze de Maio, a tributary of Varella & Sabaj Pérez (2014) mentioned two rio Curuá, upstream of PCH Salto Treze de Maio undescribed species of Apistogramma, one syntopic dam; 8°45'06" S 55°02'05" W; A. Netto-Ferreira, J. with A. kullanderi in the upper rio Curuá and Birindelli, L. Sousa & P. Hollanda-Carvalho, 23 another from the rio Treze de Maio, a tributary Jan 2009. that joins the rio Curuá downstream of the major waterfalls (Salto do Curuá) that mark its depar- Paratypes. MZUSP 97095, 1 unsexed, 25.1 mm SL, ture from the Serra do Cachimbo. The aim of this 1 female, 38.9 mm SL; same locality as holotype; paper is to provide formal descriptions of those J. Birindelli, A. Netto-Ferreira, L. Sousa, M. Sabaj species and to discuss aspects of the diversity of Pérez & N. Lujan, 22 Oct 2007. – MZUSP 101422, 2 Apistogramma recently discovered on the Serra do unsexed or juveniles, 23.2-29.0 mm SL, 2 females, Cachimbo. 31.3-40.1 mm SL; collected with holotype.

Diagnosis. Apistogramma eleutheria is similar in Material and methods general counts and body shape to several species of the A. regani species group sensu Kullander Measurements, counts and color pattern termi- (1980), but differs from them by details of col- nology follow Kullander (1980, 1986); counts for oration of mature males and females. The color holotype are denoted by asterisks. Scale rows pattern of mature females includes a very dark, are numbered according to Kullander (1990). blackish pigmentation on the median area of gular 1 A half-scale row (/2 scale) typically occurs adjacent region, occupying the ventral portion of opercular to a fin base and is formed by scales approxi- region and branchiostegal membranes, expressed mately half the size of a normal flank scale. The as a ventral continuation of the suborbital stripe correspondence between coloration and sex was (Fig. 5a). Among species of the A. regani group, determined by examination of gonads or genital only females of A. kullanderi show similar gular papillae under stereomicroscope. Vertebral counts pigmentation, but it differs from A. eleutheria by

Fig. 1. Apistogramma eleutheria: a, MZUSP 115293, holotype, 39.5 mm SL, adult brooding male; b, MZUSP 97095, paratype, 38.9 mm SL, adult brooding female; c, MZUSP 101422, paratype, 31.3 mm SL, unsexed subadult; Brazil, Pará, rio Xingu basin, rio Curuá drainage, rio Treze de Maio above the waterfalls. having this dark pigmentation more expansive, interspaces whereas females of A. kullanderi have extending onto the ventral half of the head, and the basal portion of the dorsal fin intensely and continuing posteriorly onto the belly and along continuously dark pigmented. Besides, males of the anal-fin base (Fig. 5b); females of A. eleutheria A. eleutheria do not have the reticulate pattern have dark pigments on basal portion of the dorsal of narrow dark stripes on flank and the coarse fin discontinuous, forming blotches with paler pattern of dark vermiculations on head present

Ichthyol. Explor. Freshwaters, Vol. 27, No. 1 Copyright © Verlag Dr. Friedrich Pfeil 84 in males of A. kullanderi. Apistogramma eleutheria species) and adult males do not present the first apparently does not reach a large size (around dorsal-fin lappets prolonged (vs. first dorsal-fin 40 mm SL) while A. kullanderi reaches 79.7 mm SL. lappets prolonged in A. hongsloi, as in the other Additionally, A. eleutheria differs from oth- A. macmasteri-group species). ers species of the A. regani species group, like A. cinilabra, A. commbrae, A. gosssei, A. inconspicua, Description. Largest male 39.5 mm SL, largest A. ortegai, A. ortmanni, A. paulmuelleri, A. regani female 40.1 mm SL. Body deep (35.1-37.4 % SL, and A. rubrolineata, by the presence of two ab- mean 36.4), greatest depth between verticals dominal stripes only in juveniles (vs. three or through dorsal-fin spines V and VII. Dorsal and four abdominal stripes invariably present). Apisto ventral head profiles convex in juveniles and gramma eleutheria differs from A. eunotus, A. gossei adults, except one male with a soft nuchal hump. and A. ortegai by the presence of small dark dots Snout relatively short (10.7-12.6 % SL, mean forming narrow vertical bands on caudal fin (vs. 11.5); posterior border of maxilla reaching verti- caudal fin immaculate). It differs from A. kullan cal through anterior margin of orbit or slightly deri, A. rubrolineata and A. tucurui by the absence beyond it. Orbit dorsolateral and entirely situated of the reticulate pattern with narrows dark stripes in anterior half of head. Posterior margins of pre- on flanks in males (vs. present) and the presence opercle and supracleithrum entire; posttemporal of abdominal stripes in juveniles (vs. abdominal serrations lacking. stripes invariably absent). Almost all body scales ctenoid, except for Apistogramma eleutheria may also be com- cycloid scales on anterior portion of cheek, prepared to A. hongsloi, in the A. macmasteri species opercle, interopercle, subopercle, anteromedian group, by having a similar pattern of blotches on portion of predorsal and interpelvic area and abdominal area. The pattern of irregular dark ab- eventually around urogenital papilla. Scales dominal blotches in A. eleutheria, however, starts in E1 row 22* (5) or 23 (2). Cheek fully scaled, at pectoral fin and ends at the 7th bar whereas 4 horizontal scale rows. Predorsal scales 8-9. Pre- in A. hongsloi this pattern of blotches starts more pelvic area scaled; scales anterior to tip of cleithra posteriorly (at the 3rd bar) and ends at the 7th absent or 2-3, and 6-8 scales from tip of cleithra to 1 bar. Also, females of A. eleutheria do not have the pelvic fin. Transverse row with 1 /2 scales above 1 dark blotch on chest (present in mature females and /2 scales below anterior ramus of lateral line. of A. hongsloi, as in all other A. macmasteri-group Circumpeduncular scale rows 16; 2 lateral line

Table 1. Proportional measurements of holotype and six paratypes of Apistogramma eleutheria and A. flavipedun culata. Pectoral-fin length of A. flavipedunculata obtained from 5 paratypes only. Holotype measurements included in sample range. SD, standard deviation.

A. eleutheria A. flavipedunculata holotype range mean SD holotype range mean SD Standard length (mm) 39.5 23.2-40.1 – – 45.5 26.3-45.5 –– Percents of standard length Body depth 37.4 35.1-37.4 36.4 1.0 37.9 35.9-38.5 37.3 1.0 Head length 35.2 35.2-36.7 36.0 0.6 34.3 34.3-35.7 34.8 0.5 Head depth 25.8 25.5-27.4 26.4 0.8 28.6 26.5-28.6 27.4 0.7 Head width 15.1 15.1-16.1 15.6 0.4 15.8 15.2-16.6 15.8 0.4 Orbital diameter 9.9 9.9-11.8 10.7 0.7 10.1 10.1-11.5 10.6 0.5 Interorbital width 8.9 8.7-9.5 9.1 0.4 9.0 8.5-9.9 9.1 0.4 Snout length 11.8 10.7-12.6 11.5 0.6 11.0 10.0-11.4 10.9 0.5 Preorbital depth 4.0 3.3-4.6 4.1 0.4 4.4 3.4-4.4 3.9 0.3 Caudal peduncle depth 16.9 15.8-16.9 16.4 0.4 16.1 16.1-18.0 16.7 0.6 Caudal peduncle length 10.5 10.0-11.0 10.5 0.3 9.8 9.3-10.4 9.8 0.4 Pectoral-fin length 26.3 26.3-28.7 27.1 0.9 29.0 28.9-30.7 29.8 0.8 Pelvic-fin length 26.2 25.9-29.2 27.6 1.2 32.6 29.8-34.1 32.3 1.7 Upper jaw length 10.6 9.2-10.9 10.0 0.7 11.8 10.0-11.8 10.7 0.6 Lower jaw length 14.3 13.4-14.3 13.9 0.3 14.1 13.6-14.4 14.0 0.3

Varella & Britzke: Apistogramma eleutheria and A. flavipedunculata Copyright © Verlag Dr. Friedrich Pfeil 85 scales plus 7 dorsally and 7 ventrally. Lateral-line additional, short and irregular, symphyseal row scales range from 14-17/6-8. Tube-bearing scales on upper and lower jaws. on upper branch of lateral line 11 (2), 12 (2), 13 (1), Gill rakers on first epibranchial lobe 2 (1), 14 (1), 15* (1), plus 2-5 pore-bearing scales. Tube- 3* (4); one gill raker in angle and 4 (5) gill rakers bearing scales on lower branch of lateral line 3 (3), on ceratobranchial. Gill rakers laterally on lower 5 (2), 6* (2), plus 1-4 pore-bearing scales. Scales pharyngeal toothplate 9 (1), 10 (2), 11 (2). between upper lateral line and first dorsal-fin Vertebrae 11 + 13 = 24 (3), 11 + 14 = 25 (2), 12 + 12 1 spine 4. Only /2 scale between anterior ramus of = 24 (1) or 12 + 13 = 25* (1), including last half- lateral line and last dorsal-fin spine. Fins naked centrum. Hypurals 1-5 separate. except caudal fin, which is scaled basally for one- third of its length or slightly beyond. Color pattern in alcohol. Juveniles and imma- Infraorbital series composed of three bony ture specimens (three specimens 23.2-29.0 mm elements: lachrymal, infraorbital one and infraor- SL) with tan ground colour, dorsal half of head bital 2 + 3 (co-ossification interpreted by presence and body darker than ventral half; markings of median pore). Infraorbital canal with four dark brown or blackish. Brown preorbital stripe pores on lachrymal, ventro-posterior one shared running from anterior margin of orbit to post- by anterior opening of canal through infraorbi- labial skin fold. Postorbital stripe aligned with tal one; postlachrymal infraorbital pores three: preorbital stripe, running from posterior margin anteriormost pore shared between infraorbital of orbit to posterior edge of opercle, divided in one and infraorbital 2 + 3, second pore median, a blotch just posterior to orbit and a stripe on and posteriormost pore close to posterior end dorsal border of opercle. Dark supraorbital stripe of infraorbital 2 + 3. Dentary with five pores; conspicuous only closer to dorsoposterior mar- anguloarticular canal present with simple pore. gin of orbit, running more faintly onto median Preopercular pores six; nasal with pores at each area of nape. Suborbital stripe evident, oblique end, posterior one shared with anterior opening from just beneath ventral margin of orbit to of frontal canal. Frontal with four pores plus end of preopercle, with another dark marking coronalis pore. expressed as a posteroventral continuation of Dorsal fin XVI.6 (3), XVII.5 (1), XVII.6* (3). suborbital stripe occupying area of articulation Lappets of dorsal-fin spines not conspicuously between subopercle, interopercle and opercle. prolonged in mature males. Posterior margin About seven to eight irregularly pigmented dark of soft dorsal fin pointed with 3rd ray longest; brown blotches from nape, along dorsal-fin base, tip of 3rd ray reaching approximately one-third to caudal peduncle. Anteriormost blotch on nape, length of caudal fin. Anal fin III.6 (2), IV.5* (4), followed by one blotch at dorsal-fin origin, four or IV.6 (1). Posterior margin of soft anal fin pointed five blotches along dorsal fin and one on caudal in males and females; third ray longest, reach- peduncle. Dorsal blotches corresponding in po- ing approximately one-third length of caudal sition to six or seven vertical bars on flank, only fin. Caudal fin rounded with 3 procurrent and 8 nuchal blotch not corresponding to any bar. Bars principal each lobe. Pectoral fin rounded, nearly 1-2 fainter or almost indistinct, situated anterior symmetrical, with 13* (7) rays; 6th ray longest, to vertical through dorsal-fin spine V. Bars 3-7 tip reaching vertical through genital papilla. more evident; bar 3 between dorsal-fin spines Pelvic fin pointed, first ray elongated; tip of first VII and IX, bars 4-6 between verticals through pelvic-fin ray usually reaching base of 1st anal-fin anal-fin origin and end of dorsal fin, and bar 7 spine, extending beyond urogenital papilla but on middle of caudal peduncle. Midlateral dark not reaching anal-fin origin in few specimens. band aligned with postorbital stripe, finishing just Jaw teeth unicuspid, erect, cusp slightly ret- before caudal flexure and separate from darker rorse; outer row teeth similar in size or slightly caudal-fin blotch. Band occupying entire depth larger than inner row teeth; 24-25 teeth in upper of E1 scale row plus adjacent half of scales in row right jaw outer hemi-series, 23-24 in upper left, E0, but fragmented between bars 5-6, between 26-28 in lower right jaw outer hemi-series, 26-27 bars 6-7, and less frequently between bars 2-3. in lower left. Outer row of teeth occupying almost Additional two narrow abdominal stripes formed entire premaxilla and dentary margins; inner row by a series of dark spots, fainter than midlateral of premaxilla and dentary extending to approxi- band, running from end of bar 1 to end of bar 5 1 mately /3 of jaw margin; few teeth forming one and occupying two horizontal scale rows below

Fig. 2. Apistogramma flavipedunculata: a, MZUSP 115292, holotype, 45.5 mm SL, adult male; b, MZUSP 97594, paratype, 30.5 mm SL, unsexed subadult; Brazil, Pará, rio Xingu basin, upper rio Curuá drainage, rio Escorpião.

E0 row. Caudal fin with a large blotch some- than females. Juvenile pattern of two abdominal what rectangular but with poorly defined edges, stripes replaced in adult male by irregular, faint tangent to caudal flexure and situated between pigmentation on abdomen, occupying most of caudal-fin rays D4-V4; two or three faint dark, area of four abdominal horizontal scale rows narrow bands posterior to caudal blotch. Dorsal below E0 series, from pectoral-fin base to bar 7. fin dusky, with more concentration of dark pig- Dorsal and anal fins of male more conspicuously ment in basal portion of spines, corresponding ornamented; dorsal fin darker with three series of with position of flank bars, and with two or light spots on soft portion (besides basal blotches three irregular lighter spaces on soft portion of also present in juveniles), soft portion of anal fin fin. Anal fin dusky, with two paler spaces on soft with lighter and darker stripes interspaced by portion. A specimen measuring 31.3 mm SL, ten- dusky background. tatively considered an immature female, may be Ground colour of adult females brown dor- interpreted as showing intermediate colouration sally, pale yellowish on ventral half. Juvenile pat- between juvenile and adult patterns, with anal fin tern of abdominal stripes replaced by an irregular dark margined. Pectoral and pelvic fins hyaline, pattern of dark pigmentation on abdomen; differ- with no markings except pelvic fin with a very ent from males, this pigmentation is conspicuous faint pigmentation on its spine. and expressed approximately in same position of Adult male and females retaining most of vertical bars, resulting in an overall barred pat- juvenile colour pattern, with some differences tern. Midlateral band appearing more fragmented attributed mainly to sexual dichromatism. than in males and juveniles, forming a series of Adult male ground colour brown, darker six blotches on midlateral area. Dark blackish

Apistogramma eleutheria Apistogramma flavipedunculata acete da erra do Cachio Apistogramma kullanderi Apistogramma p. Peioto oad Apistogramma p. iddle Cr Apistogramma p. aai olie ca. Fig. 3. Serra do Cachimbo, showing known distribution of Apistogramma eleutheria, A. flavipedunculata, A. kullan deri and other putative new species occurring in the rio Curuá and adjacent rivers. 1, waterfalls of the rio Treze de Maio; 2, series of waterfalls of the rio Curuá. pigmentation of pelvic fin in adult females more Etymology. The specific epithet eleutheria conspicuous than in adult males, occupying spine (λευθερα) is a Greek noun that means freedom. It and first soft ray. Suborbital stripe continuing onto makes reference to the name of the river, because median area of gular region, occupying ventral Treze de Maio is the date (13 May 1888) on which portion of opercular region and branchiostegal Princess Isabel signed the ‘Áurea’ law, marking membranes. Anterior to gular pigmented area, the official end of legal slavery in Brazil. A noun another concentration of dark pigmentation in opposition. ventrally on dentary, separated from gular pigmentation by a small pale area (Fig. 5a). Apistogramma flavipedunculata, new species Distribution and habitat. Apistogramma eleutheria (Fig. 2) is known only from the rio Treze de Maio above its major waterfalls (see photo in Sousa et al., Apistogramma sp. ‘upper Curuá’: Varella & Sabaj 2010: fig. 5a), rio Xingu basin, Brazil. Although Pérez, 2014: 255. some maps and the ‘Índice de Nomes Geográ- Holotype. MZUSP 115292, 45.5 mm SL, adult ficos’ (IBGE, 2011) give the name “rio Três de male; Brazil: Pará: Altamira: rio Xingu basin: rio Maio”, Treze de Maio is the correct name based Iriri drainage: rio Escorpião, tributary of the up- on communications with locals and road signs per rio Curuá, at route BR163 bridge; 8°53'54" S along BR163. 54°59'20" W; A. Netto-Ferreira, J. Birindelli, L. Sousa, M. Sabaj-Pérez & N. Lujan, 29 Oct 2007.

Paratypes. MZUSP 97594, 5 immature specimens specimens, falling short of orbital margin in larg- identified as males, 26.3-39.6 mm SL, 1 adult male, est specimens. Orbit dorsolateral, mostly but not 45.3 mm SL; collected with holotype. entirely situated in anterior half of head. Posterior margins of preopercle and supracleithrum entire; Diagnosis. Adult males of A. flavipedunculata posttemporal serrations lacking. are distinguished from males of all congeners by Almost all body scales ctenoid, except for having the posterior part of the flank and caudal cycloid scales on cheek, preopercle, interopercle, peduncle dominated by a light yellow pigmenta- subopercle, anteromedian portion of predorsal tion, resulting in the dark midlateral band frag- and interpelvic area and eventually around uro- mented posteriorly and the dark bars (present in genital papilla. Scales in E1 row 22 (2) or 23* (5). juveniles) masked or absent in the males. Cheek fully scaled with 3 horizontal scale rows. Apistogramma flavipedunculata can be distin- Predorsal scales 9-10. Prepelvic area scaled; scales guished from all congeners except A. luelingi, anterior to tip of cleithra absent or 3-4, and 6-7 A. eleutheria and A. pleurotaenia by having exclu- scales from tip of cleithra to pelvic fin. Transverse 1 1 sively four anal-fin spines instead of three (four row with 1 scale above and 6 /2 or 7 /2 scales below spines occurring very rarely in few species; see anterior ramus of lateral line. Circumpeduncular discussion). It differs from A. luelingi by the ab- scale rows 16; 2 lateral line scales plus 7 dorsally sence of midlateral spot, the lack of accentuated and 7 ventrally. Lateral-line scales range from elongation of lappets on spines of dorsal fin and 14-17/7-8. Tube-bearing scales on upper branch lyrate caudal fin in adult males, the absence of of lateral line 12 (1), 13* (3), 14 (3), plus 2-4 pore- conspicuous abdominal stripes (vs. three dark bearing scales. Tube-bearing scales on lower stripes evident on abdomen), and by the presence branch of lateral line 2 (1), 5 (1), 6* (2), 7 (3) plus of numerous dark narrow bands on the caudal 1-4 pore-bearing scales. Scales between upper 1 fin (vs. dark markings on caudal fin absent or lateral line and first dorsal-fin spine 3 or 3 /2. Only 1 inconspicuous). /2 scale between anterior ramus of lateral line and Among species of the A. regani group, A. acr last dorsal-fin spine. Fins naked except caudal fin, ensis, A. caetei, A. ortmanni, A. paulmuelleri and scaled basally on approximately one-third of its A. regani resemble A. flavipedunculata with respect length. to body shape and presence of narrow vertical Configuration of infraorbital bones and later- bands on the caudal fin. Apistogramma flavipe alis pores of head as in A. eleutheria. dunculata is distinguished from A. acrensis and Dorsal fin XVII.5 (5), XVII.6* (2). Lappets A. caetei by having more dorsal-fin spines (16- of anteriormost eight dorsal-fin spines slightly 17 vs. 15), from A. ortmanni, A. paulmuelleri and prolonged in mature males. Posterior margin of A. regani by having two or three series of faint soft dorsal fin pointed with second and third rays dark markings on abdominal area not forming longest, with tips reaching approximately one- well-defined stripes (vs. presence of three or third length of caudal fin in smaller specimens four well-defined dark abdominal stripes) and and half-length of caudal fin in largest specimens. caudal-fin blotch rounded and restricted to the Anal fin IV.6* (7). Posterior margin of soft anal fin centralmost rays of the caudal fin (vs. caudal-fin pointed; 3rd and/or 4th ray longest, extending blotch vertically elongated in A. ortmanni, A. paul slightly beyond half-length of caudal fin. Caudal muelleri and A. regani, occupying mostly or totally fin rounded with 3 procurrent and 8 principal the depth of caudal-fin base). rays each lobe. Pectoral fin rounded, nearly symmetrical, with 11 (1), 12* (6) rays; 6th ray longest, Description. Largest male 45.5 mm SL, no female tip reaching genital papilla or extending slightly specimens confirmed. Body deep (35.9-38.5 % beyond vertical through genital papilla. Pelvic fin SL, mean 37.3), greatest depth between verticals pointed, first ray elongated; tip of first pelvic-fin through dorsal-fin spines IV and V. Dorsal and ray usually reaching base of 1st anal-fin spine. ventral head profiles convex in juveniles and Jaw teeth unicuspid, erect, cusp slightly retrorse; adults, with interorbital concavity in largest outer row teeth similar in size or slightly larger males. Snout relatively short (10.1-11.4 % SL, than inner row teeth; 20-24 teeth in upper right mean 10.9); posterior border of maxilla reaching jaw outer hemi-series, 21-24 in upper left, 28-31 vertical through anterior margin of orbit in most in lower right jaw outer hemi-series, 29-30 in

2 lower left. Outer row of teeth occupying /3 length postorbital stripe, ending approximately at verti- of premaxilla and entire dentary margins; inner cal through caudal peduncle blotch and separate row of premaxilla and dentary extending to ap- from darker caudal-fin blotch in five specimens proximately middle of jaw margin; additional (26.3-39.6 mm SL); midlateral band fragmented irregular, symphyseal row of teeth present on posteriorly and faded past vertical through bar upper and lower jaws. 5 in two largest males. Band occupying entire Gill rakers on first epibranchial lobe 4 (1), 5 (1), depth of E1 scale row plus adjacent half of scales 6* (5); one gill raker in angle and 1 (1), 2 (2), 3 (3), in row E0 at least anteriorly (i. e., band fragmented 4* (1) gill rakers on ceratobranchial. Gill rakers posteriorly in largest specimens). Two or three laterally on lower pharyngeal toothplate 10 (1), faded abdominal series of irregular dark mark- 11 (2), 12* (4). ings, composed of pigmentation on distal margin Vertebrae 11 + 13 = 24 (5), 12 + 12 = 24* (1), of some scales (present only in four specimens 12 + 13 = 25, including last half-centrum. Hypurals and more visible in holotype); abdominal series 1-5 separate. occupying two or three horizontal scale rows below E0 row. Dominance of light pigmentation Color pattern in alcohol. Type material com- posteriorly to vertical through first soft dorsal-fin posed of large males and smaller specimens ray onto base of caudal fin, masking melanin- tentatively determined as males (by examination based colour markings except caudal fin blotch, of shape of genital papilla); same color pattern for as most remarkable difference between largest all specimens excepting some aspects as otherwise adult males and smaller specimens. noted. Ground colour yellowish tan, slightly Caudal fin with a rounded blotch tangent to darker on dorsum closer to dorsal-fin base than caudal flexure and situated between caudal-fin ventrally; colour markings on head and flanks rays D3-4 and V4; posterior to caudal blotch, dark brown and blackish on distal margin of fins. eight or nine well-defined narrow bands to dis- Brown preorbital stripe from anterior margin tal margin of fin. Dorsal fin dusky, with strong of orbit to postlabial skin fold, extending onto blackish pigmentation on lappets of dorsal-fin tip of upper jaw. Postorbital stripe aligned with spines occupying most of first dorsal-fin spine and preorbital stripe, running from posterior margin distal half of second and third dorsal-fin spines; of orbit to posterior edge of opercle. Dark supraor- distal portion of dorsal fin slightly darker than bital stripe evident from dorsoposterior margin basal portion. Dorsal fin with four slightly darker of orbit obliquely onto median area of nape in vertical bands on distal portion of soft rays. Anal largest males; reduced to a rounded blotch near fin dusky with dark gray pigmentation on distal orbit in few specimens. Suborbital stripe running portion of spines and soft rays; four dark vertical obliquely from just beneath ventral margin of bars on two posteriormost soft rays. Pectoral fin orbit to anterior margin of subopercle, often with hyaline. Pelvic fin hyaline except black blotch a brief pale hiatus across posteroventral margin covering its spine and one or two adjacent soft of preopercle; suborbital stripe more evident in rays, much more evident in largest males than in two largest males. Interorbital stripes distinct smaller specimens. only in holotype, fragmented into two paired dark vermiculations from anterodorsal margin Distribution. Apistogramma flavipedunculata is of orbit onto median area. known only from the rio Escorpião, tributary of About seven or eight irregularly pigmented the upper rio Curuá, rio Xingu basin, Brazil. The dark brown blotches from nape, along dorsal-fin species is syntopic with A. kullanderi in this river. base, to caudal peduncle; anteriormost blotch on nape, followed by one blotch on dorsal-fin origin, Etymology. The specific epithet flavipedunculata, five blotches along dorsal fin and one on caudal an adjective, refers to the yellow pigmentation peduncle. Blotches restricted to dorsum, not that dominates the posterior part of the body in extending ventrally to form bars and more evi- adult males; from the Latin flavus, yellow, and dent in smallest specimens (i. e., three specimens pedunculatus, with a peduncle. 26.3-35.8 mm SL); posteriormost blotch, on caudal peduncle, partially or totally indistinct in larger specimens. Midlateral dark band aligned with

Fig. 4. Apistogramma sp. ‘middle Curuá’, ca. 25 mm SL; Brazil, Pará, rio Xingu basin, rio Curuá basin, rio Treze de Maio below the waterfalls; alive just after capture. Photograph by William Ohara.

Discussion portion of the rio Curuá (A. flavipedunculata and A. kullanderi) and the rio Treze Maio (A. eleutheria). The number of nominal species of Apistogramma Apistogramma eleutheria is isolated from the lower reported from the rivers of the Brazilian Shield portion rio Treze de Maio by a waterfall with same is relatively low when compared to that of the name (Souza et al., 2010: 259, fig. 5c). Similarly, Western Amazon, Orinoco basin and Guiana A. kullanderi and A. flavipedunculata are isolated Shield. Only the rio Madeira basin, along the from the middle portion of the Curuá drainage western border of the Brazilian Shield, presents by a series of three waterfalls (indicated in Fig. 3 a high known diversity with 12 valid species, ten by arrow 2), as previously described by Birindelli of them endemic. This may be correlated with et al. (2009: 16, fig. 6) and Varella & Sabaj Pérez the early scientific expeditions to this area (e. g. (2014: 252, fig. 7). Johann Natterer in 1817-1835). Other tributaries Besides those three species, an expedition to to the right margin of the rio Amazonas were the Serra do Cachimbo in 2014 brought fishes of systematically explored later, more intensively in another new species from the rio Curuá and rio the last fifty years. Only a few species have been Treze de Maio below the waterfalls, provisionally described from these drainages, as A. taeniata denominated here as Apistogramma sp. ‘middle (Günther, 1862) described from the lower rio Curuá’ (Fig. 4). This undescribed Apistogramma Tapajós (distributed also in the lower portion rio is known only from juveniles and small adults Xingu) and A. tucurui Staeck, 2003 from the area of that resemble other forms encountered in the Turucui reservoir in the rio Tocantins. Other spe- lower portions of the rio Iriri and rio Xingu, and cies of Apistogramma reported from these drain- shares many similarities with several species of ages are tentatively identified as species widely the A. regani group. Although the lack of mature distributed in the main channel or lower portion males and females of this species precludes a of tributaries of the Amazon river, such as A. cf. definitive identification, Apistogramma sp. ‘middle pertensis (Haseman, 1911) in the rio Tapajós (RB, Curuá’ can be easily distinguished from A. eleuthe pers. obs.) and A. agassizi in the lower rio Xingu, or ria and A. flavipedunculata by the presence of a are putatively undescribed (e. g. Koslowski, 2002). black blotch just below the posterior edge of the Apistogramma eleutheria and A. flavipedunculata, mouth (arrow in Fig. 4; vs. absent in A. eleutheria together with the recently described A. kullan and A. flavipedunculata) and three anal-fin spines deri, are the only formally named species of the (vs. modally or always four spines in A. eleutheria genus from the middle and upper portions of and A. flavipedunculata, respectively). the rio Xingu basin. Their known distribution is Rivers adjacent to the rio Curuá (tributaries restricted to the Serra do Cachimbo in the upper of the rio Teles Pires and rio Jamanxim) present

a b c Fig. 5. Schematic drawing of ventral body in species of Apistogramma showing extent and condition of dark markings present on brooding females. a, A. eleutheria, MZUSP 97095, 38.9 mm SL; b, Apistogramma sp. ‘Peixoto’, MZUSP 116959, 38.7 mm SL; and c, A. kullanderi, MZUSP 97597, 58.7 mm SL. putative undescribed species deserving of com- pigmentation, and the presence of dark markings ments. Apistogramma sp. ‘Jamanxim’ is most simi- on the chest and belly (Fig. 5). In A. eleutheria, the lar to A. flavipedunculata with respect to general conspicuous ventral marking is restricted to the body shape, dorsal and anal fins comparatively gular area, but there are fainter markings on mid- high and with their posterior margin extending ventral area, on spine of pelvic fin and laterally half the length of the caudal fin, and elements of on chest close to opercle (Fig. 5a). Apistogramma the color pattern such as caudal fin with numer- sp. ‘Peixoto’ presents conspicuous black marking ous dark narrow bands and conspicuous black on gular region, extending posteriorly onto the marking on the spine and first ray of pelvic fin in median portion of chest, the spine and lateralmost males. Apistogramma flavipedunculata differs from rays of pelvic fin; separate from that, another it by having four instead of three anal-fin spines, strong black marking on midventral area extends by the peculiar color pattern on the posterior posteriory onto anal fin (Fig. 5b). Apistogramma portion of the flank and caudal peduncle (i. e., kullanderi has the sexually dimorphic colora- dominated by yellow pigmentation that masks tion in females over-expressed, resulting in the the melanin-based markings), and by not having ventral body almost totally covered by intense dark-margined scales on dorsum (vs. distal mar- black pigmentation (Fig. 5c). Thus, in A. eleuthe gin of scales on dorsum blackish or dark brown). ria, A. sp. ‘Peixoto’ and A. kullanderi, which are Apistogramma sp. ‘Peixoto’ is an undescribed spe- allopatrically isolated in rivers of the Serra do cies from the rio Braço Norte, tributary of the rio Cachimbo, females develop sexual dimorphism Peixoto de Azevedo (rio Teles Pires basin), similar based on more or less the same color pattern ele- in several aspects to A. eleutheria and A. kullanderi. ments, differing from each other by the intensity Brooding females of A. eleutheria, A. sp. ‘Peixoto’ and distribution of the color expression. On the and A. kullanderi share the possibly apomorphic other hand, males of A. kullanderi are much larger states of the gular region covered by a blackish than females and seem to show very distinctive

Ichthyol. Explor. Freshwaters, Vol. 27, No. 1 Copyright © Verlag Dr. Friedrich Pfeil 92 dimorphic color pattern consisting of many dark Graça, 2008; Musilová et al., 2009), a condition stripes on flanks and a coarse pattern of dark of three to four anal-fin spines was codified as vermiculations on the head. Males of A. eleutheria plesiomorphic among cichlids. Reduction to two are slightly darker than females, and males of anal-fin spines is an autapomorphy of Biotoecus Apistogramma sp. ‘Peixoto’ have few faint dark dicentrarchus Kullander, 1989, and the condition markings on the head and some pale spots on of five or more spines is synapomorphic for the the flank, but neither species present the strong tribe Heroini (sensu Kullander, 1998). sexually dimorphic coloration as in A. kullanderi. The only geophagine with five or more anal-fin Varella & Sabaj Pérez (2014) inferred that large spines is Apistogrammoides pucallpaensis Meinken, body in A. kullanderi may be due to its isolation in 1965 (6-9 spines), which has been considered the upper part of rio Curuá, which is depauperate closely related to, if not congeneric with Apisto of cichlids and potential predators (i. e., phylo- gramma (e. g. Kullander, 1998; López-Fernández et genetic constraints on body size reduced when al., 2005, 2010). Almost all species of Apistogramma competition and predatory pressure are limited). have three anal-fin spines. Exceptions with four Ready et al. (2006) studied allopatric populations anal-fin spines include a few specimens of A. agas from the Guamá and adjacent rivers in Pará, sizi and A. cacatuoides (1 specimen each; Kullander, previously identified as A. caetei Kullander, 1980, 1980), A. cruzi (1 specimen; Kullander, 1986) and but with slight differences in color pattern. They A. kullanderi (2 of 44 specimens). Among species found three closely related lineages with strong typically with three anal-fin spines, A. commbrae is prezygotic isolation and sexual selection through the only one that frequently presents four anal-fin female mate choice involved in the diversification spines (15 of 56 specimens examined here; also in Apistogramma. Following the same idea, the about 15 % of the material examined by Kullander, evolution of large size and sexually dimorphic 1982). Apistogramma flavipedunculata exclusively coloration in A. kullanderi would be possibly and A. luelingi Kullander, 1976 almost exclusively correlated: an unrestrained male-biased sexual (40 of 42 specimens, according to Kullander, 1986) dimorphism in an isolated area with reduced en- have four-anal fin spines. Apistogramma eleutheria vironmental constraints resulting in a large body. presented 5 of 7 specimens with four spines, Occurring syntopically with Apistogramma sp. perhaps an indication that it is the most common ‘Peixoto’ in the rio Braço Norte, there are other count for the species. Finally, the holotype (only cichlids (e. g. species of Satanoperca, Crenicichla specimen known) of A. pleurotaenia (Regan, 1909) and Cichla) and other fish groups (e. g. species also possesses four spines. of Serrasalmus and Pimelodus), which can be con- Kullander (1988) and Varella & Moreira sidered as potential competitors and predators. (2013) described two species of Teleocichla having They may represent environmental constraints exclusively four anal-fin spines (T. centrarchus for the development of a very flashy dimorphic and T. wajapi, respectively). These are the only coloration and large body like that found in A. kul species with four instead of three anal-fin spines landeri. However, A. eleutheria occurs above large among species of the Crenicichla-Teleocichla clade, waterfalls in the upper rio Treze de Maio, which is although four spines are rarely recorded for a few depauperate like the upper rio Curuá. Body size species typically having three (Varella & Moreira, and sexual dimorphism in A. eleutheria, however, 2013). For this reason, the authors considered this have apparently not evolved like in A. kullanderi. condition probably apomorphic and indicative of a closer-relationship between T. centrarchus and Anal-fin spines. The number of anal-fin spines T. wajapi, which would demand the division of is traditionally useful in the taxonomy of species the character state if in a cladistic analysis. Also and genera of Cichlidae, as well as to reconstruc- Kullander (1996) suggested considering modal tions of their phylogenetic relationships, notably anal-fin counts rather than ranges in phylogenetic within the Neotropical clade. Kullander (1983a, analyses, if there are adequate data available 1989, 1996) discussed in detail this character and (i. e. large amount of specimens of the terminal its implications for the delimitation of supraspe- taxa). In the particular case of a comprehensive cific groups. In many published or unpublished phylogeny on Apistogramma, to follow this sug- cladistics analysis (e. g. Cichocki, 1976; Kul- gestion might avoid missing important variation lander, 1998; Landim, 2006; Chakrabarty, 2007; present between the included species.

Comparative material. Apistogramma sp. ‘middle Cu- A. luelingi: ROM 82894, 2, 21.6-28.5 mm SL, Peru, ruá’: Brazil: Pará: Altamira: rio Xingu drainage: MZUSP Madre de Dios, Tambopata, Tambopata River. 97617, 2 juveniles, 9.6-10.2 mm SL; flooding at brigde A. ortegai: MUSM 48898, holotype, 38.4 mm SL; of road BR-163, close to district of Castelo dos Sonhos; Peru, Loreto, Distrito Pebas: small stream tributary of 8°15'17" S 55°06'40" W. – MZUSP 116007, 4, 19.4-25.0 mm Ampiyacu River. SL; tributary of rio Treze de Maio below the waterfalls, A. ortmanni: ROM 91316, 8, 35.1-55.7 mm SL, Guy- rio Curuá drainage; 8°39'08" S 55°02'08" W. – MZUSP ana, Imbaima creek, tributary of Kuribong River. 116025, 5, 17.0-25.2 mm SL; tributary of rio Curuá, close A. rubrolineata: ROM 63879, 8, 20.5-34.5 mm SL, to district of Cachoeira da Serra; 8°36'18.51" S 55°05' Peru, Madre de Dios, Tambopata, Planchon creek. 28.67" W. – MZUSP 116620, 3, 17.6-24.4 mm SL; tributary A. taeniata: ANSP 195006 (examined from photo- of rio Curuá, close to district of Cachoeira da Serra; graph); Brazil, Pará, lower rio Xingu close to Porto de 8°36'41.80" S 55°07'25.00" W. Moz. Apistogramma sp. ‘Jamanxim’: Brazil: Pará, Novo Progresso: rio Tapajós drainage: MZUSP 97261, 1, 22.7 mm SL; rio Jamanxim, approximately 30 km from Acknowledgements Castelo dos Sonhos; 8°15'17" S 55°06'40" W. – MZUSP 97561, 18, 12.4-37.9 mm SL; rio Jamanxim, flooded Thanks to André Netto-Ferreira, José Luís Birindelli, shoreline at BR163 bridge, south of Vila Mil; 7°51'47.00" S Leandro Sousa, Mark Sabaj Pérez, Nathan Lujan and 55°10'47.00" W. Pedro Hollanda-Carvalho for collecting the type spec- Apistogramma sp. ‘Peixoto’: Brazil: Pará: Novo imens. Fieldwork was supported in part by the “All Progresso: rio Teles Pires drainage (rio Tapajós basin): Catfish Species Inventory” (NSF DEB-0315963). Informa- MZUSP 96770, 1, 21.1 mm SL; right margin tributary of tion on Apistogramma from the lower Xingu and sug- rio Peixoto de Azevedo, at BR163 bridge, close to Força gestions for improving the manuscript were provided Aérea Brasileira (FAB) base: 10°17'14" S 54°50'57" W. – by Mark Sabaj Pérez (supported by the “iXingu Project”, MZUSP 96828, 43, 13.7-48.3 mm SL; MZUSP 116052, NSF DEB-1257813). Flávio Bockmann and Ricardo 10, 28.0-61.7 mm SL; tributary of rio Braço Norte (rio Castro allowed us to take radiographs in the Laborató- Peixoto de Azevedo drainage), at BR163 bridge, close rio de Ictiologia de Ribeirão Preto (LIRP), using the to FAB base; 9°25'55" S 54°52'11" W. – MZUSP 96846, machine obtained by the project “Melhoria e Capacita- 141, 13.0-46.7 mm SL; MZUSP 101428, 3, 28.1-30.0 mm ção das Coleções Científicas do Departamento de Bio- SL; MZUSP 116053, 20: 26.6-53.3 mm SL; tributary of logia da Faculdade de Filosofia, Ciências e Letras de rio Braço Norte (rio Peixoto de Azevedo drainage), at Ribeirão Preto-USP” (FAPESP 09/54931-0). Visit of the BR163 bridge, close to FAB base; 9°28'20" S 54°52'11" W. authors to MCP collection was supported by the project – MZUSP 101434, 1, 40.5 mm SL; tributary of rio Braço “Qualificação taxonômica da base de dados da coleção Norte, rio Peixoto de Azevedo, at BR163 bridge, close de peixes do Museu de Ciências e Tecnologia – PUCRS to FAB base; 9°19'17" S 54°50'26" W. – MZUSP 116595, para uso on-line” (MCT/CNPq: 45/2012: 504177/2012- 7, 13.9-38.7 mm SL; tributary of rio Braço Norte (rio 5). H. Varella is financially supported by FAPESP (grant Peixoto de Azevedo drainage), on road BR163 close to 2011/14630-0) and R. Britzke by FAPESP (grant border between States of Pará and Mato Grosso, 2011/00269-4) and PROPe (004/2014). 09°28'20" S 54°51'32" W. A. acrensis: MCP 29419, 2, 30-32 mm SL; Brazil: Acre: rio Purus basin, rio Branco. A. agassizii: ANSP 197892 (examined from photo- Literature cited graph); Brazil: Pará: lower rio Xingu close to Porto do Moz. Birindelli, J. L. O. & H. A. Britski. 2009. New species of A. commbrae: MCP 22849, 1, 25 mm SL; Brazil: Rio the genus Leporinus Agassiz (Characiformes: Ano Grande do Sul: Uruguaiana: rio Uruguai basin. – MZUSP stomidae) from the rio Curuá, rio Xingu basin, 59972, 22 (with 3 anal-fin spines), 13.5-23,3 mm SL, 13 Serra do Cachimbo, Brazil, with comments on (with 4 anal-fin spines), 15.7-21.6 mm SL; Brazil: Mato Leporinus reticulatus. Neotropical Ichthyology, 7: Grosso do Sul: Corumbá: rio Paraguai basin: rio Ver- 1-10. melho. – MZUSP 100185, 2 (with 4 anal-fin spines), Birindelli, J. L. O., A. M. Zanata, L. M. Sousa & A. L. 20.5-22.4 mm SL, 12 (with three anal-fin spines), 14.1- Netto-Ferreira. 2009. New species of Jupiaba Zana- 21.3 mm SL: Brazil: Mato Grosso: Corumbá: rio Paraguai ta (Characiformes: Characidae) from Serra do basin: rio Miranda. Cachimbo, with comments on the endemism of A. hongsloi: MZUSP 96446, 15, 13.0-46.3 mm SL, upper rio Curuá, rio Xingu basin, Brazil. Neo- Venezuela, Bolivar, Caicara del Orinoco. – LBP 18704, tropical Ichthyology, 7: 11-18. 18, 21.2-47.1 mm SL, Colombia Meta, Canõ Iraca, Río Britzke, R., C. Oliveira & S. O. Kullander, 2014. Apisto Guaviare basin. gramma ortegai (Teleostei: Cichlidae), a new species A. inconspicua: ROM 69320, 2, 17.6-24.3 mm SL, of cichlid fish from the Ampyiacu River in the Pe- Bolivia, Santa Cruz, Río Negro, Vers Perseverancia. ruvian Amazon basin. Zootaxa, 3869: 409-419.

Chakrabarty, P. 2007. A morphological phylogenetic — 1983a. A revision of the South American cichlid analysis of Middle American cichlids with special genus Cichlasoma (Teleostei: Cichlidae). Naturhis- emphasis on the section “Nandopsis” sensu Regan. toriska Riksmuseet, Stockholm, 296 pp. Miscellaneous Publications, Museum of Zoology, — 1983b. Cichlid fishes from the La Plata basin. Part University of Michigan, 198: 1-31. IV. Review of the Apistogramma species, with de- Cichocki, F. P. 1976. Cladistic history of cichlid fishes scription of a new species (Teleostei, Cichlidae). and reproductive strategies of the American genera Zoologica Scripta, 11: 307-313. Acarichthys, Biotodoma and Geophagus. Volume I. — 1986. Cichlid fishes of the Amazon river drainage Unpublished PhD dissertation, University of Mich- of Peru. Swedish Museum of Natural History, igan, Ann Arbor. Stockholm, 431 pp. Eigenmann, C. H. 1912. The freshwater fishes of British — 1988. Teleocichla, a new genus of South American Guiana, including a study of the ecological group- rheophilic cichlid fishes with six new species (Tele ing of species, and the relation of the fauna of the ostei: Cichlidae). Copeia, 1988: 196-230. plateau to that of the lowlands. Memoirs of the — 1989. Biotoecus Eigenmann and Kennedy (Teleostei: Carnegie Museum, 5: i-xxii + 1-578. Cichlidae): description of a new species from the Graça, W. J. 2008. Sistemática da tribo Heroini Kullan- Orinoco Basin and revised generic diagnosis. Jour- der, 1998 (Perciformes, Cichlidae). Unpublished nal of Natural History, 23: 225-260. doctoral dissertation, Universidade Estadual de — 1990. Mazarunia mazarunii (Teleostei: Cichlidae), a Maringá, Maringá, 217 pp. new genus and species from Guyana, South America. Günther, A. 1862. Catalogue of the fishes in the British Ichthyological Exploration of Freshwaters, 1: 3-14. Museum. Catalogue of the Acanthopterygii, Pha- — 1996. Heroina isonycterina, a new genus and species ryngognathi and Anacanthini in the collection of of cichlid fish from western Amazonia, with com- the British Museum. Volume 4. British Museum, ments on cichlasomine systematics. Ichthyological London, xxi + 534 pp. Exploration of Freshwaters, 7: 149-172. Haseman, J. D. 1911. An annotated catalog of the cich- — 1998. A phylogeny and classification of the South lid fishes collected by the expedition of the Carnegie American Cichlidae (Teleostei: Perciformes). Pp. Museum to central South America, 1907-10. Annals 461-498 in: L. R. Malabarba et al. (eds.), Phylogeny of the Carnegie Museum, 7: 329-373. and classification of neotropical fishes. Edipucrs, Heckel, J. J. 1840. Johann Natterer’s neue Flussfische Porto Alegre. Brasilien’s nach den Beobachtungen und Mitthei- — 2003. Family Cichlidae. Pp. 605-654 in: R. E. Reis lungen des Entdeckers beschrieben (Erste Abthei- et al. (eds.), Check list of the freshwater fishes of lung, Die Labroiden). Annalen des Wiener Museums South and Central America. Edipucrs, Porto Alegre. der Naturgeschichte, 2: 325-471. Kullander, S. O. & H. R. Varella. 2015. Wallace’s pike Hein, G., A. Zarske & J. Zapata. 2002. Apistogramma rubro cichlid gets a name after 160 years: a new species lineata sp. n., ein neuer Buntbarsch (Teleostei: Perci- of cichlid fish (Teleostei: Cichlidae) from the upper formes: Cichlidae) aus dem Rio Manuripi, Departa- rio Negro in Brazil. Copeia, 103: 512-519. mento Pando, in Bolivien. Das Aquarium, 402: 15-19. Landim, M. I. P. F. 2006. Relações filogenéticas na fa- Hoedeman, J. J. 1951. Notes on the fishes of the cichlid mília Cichlidae Bonaparte, 1840 (Teleostei: Perci- family I. Apistogramma cacatuoides sp. n. Beaufortia, formes). Unpublished doctoral dissertation, Insti- 4: 1-4. tuto de Biociências da Universidade de São Paulo, IBGE, 2011. Ìndice de nomes geográficos, Volume 1, 445 pp. Carta. Instituto Brasileiro de Geografia e Estatística, Lima, F. C. T. & J. L. O. Birindelli. 2006. Moenkhausia Rio de Janeiro, 400 pp. petymbuaba, a new species of characid from the Koslowski, I. 2002. Die Buntbarsche Amerikas. Volume Serra do Cachimbo, Rio Xingu basin, Brazil (Chara 2. Apistogramma & Co. Ulmer, Stuttgart, 320 pp. ciformes: Characidae). Ichthyological Exploration Kullander, S. O. 1976. Apistogramma luelingi sp. nov., a of Freshwaters, 17: 53-58. new cichlid fish from Bolivia (Teleostei: Cichlidae). López-Fernández, H., R. L. Honeycutt, M. L. J. Stiassny Bonner Zoologische Beiträge, 27: 258-266. & K. O. Winemiller. 2005. Morphology, molecules, — 1979. Species of Apistogramma (Teleostei, Cichlidae) and character congruence in the phylogeny of South from the Orinoco drainage basin, South America, American geophagine cichlids (Perciformes, La- with descriptions of four new species. Zoologica broidei). Zoologica Scripta, 34: 627-651. Scripta, 8: 69-79. López-Fernández, H., K. O. Winemiller & R. L. Honey- — 1980. A taxonomical study of the genus Apisto cutt. 2010. Multilocus phylogeny and rapid radia- gramma Regan, with a revision of Brazilian and tions in Neotropical cichlid fishes (Perciformes: Peruvian species (Teleostei: Percoidei: Cichlidae). Cichlidae: Cichlinae). Molecular Phylogenetics and Bonner Zoologische Monographien, 14: 1-152. Evolution, 55: 1070-1086. — 1982. Description of a new species of Apistogramma Meinken, H. 1965. Über eine neue Gattung und Art der Regan, from the Oyapock and Approuague River Familie Cichlidae aus Peru (Pisces, Percoidea, systems (Teleostei: Cichlidae). Cybium, 6: 65-72. Cichlidae). Senckenbergiana Biologica, 46: 47-53.

Musilová, Z., O. Rícan & J. Novák. 2009. Phylogeny of Römer, U., D. P. Soares, C. R. G. Dávila, F. Duponchelle, the neotropical cichlid fish tribe Cichlasomatini J. F. Renno & I. Hahn. 2015. Re-description of Apis (Teleostei: Cichlidae) based on morphological and togramma payaminonis Kullander, 1986, with descrip- molecular data, with the description of a new genus. tions of two new cichlid species of the genus Journal of Zoological Systematics and Evolutionary Apistogramma (Teleostei, Perciformes, Geophaginae) Research, 47: 234-247. from northern Peru. Vertebrate Zoology, 65: 287- Netto-Ferreira, A. L. 2012. Three new species of Lebia 314. sina (Characiformes: Lebiasinidae) from the Brazil- Sousa, L. M., A. L. Netto-Ferreira & J. L. O Birindelli. ian shield border at Serra do Cachimbo, Pará, 2010. Two new species of Moenkhausia Eigenmann Brazil. Neotropical Ichthyology, 10: 487-498. (Characiformes: Characidae) from Serra do Cachim- Pellegrin, J. 1936. Un poisson d’aquarium nouveau du bo, Pará, Northern Brazil. Neotropical Ichthyology, genre Apistogramma. Bulletin de la Société Nationale 8: 255-264. d’Acclimatation de France, 1936: 56-58. Staeck, W. 2003. Cichliden-Lexikon, Teil 3: Südameri- Ready, J. S., I. Sampaio, H. Schneider, C. Vinson, T. dos kanische Zwergbuntbarsche. Dähne, Ettlingen, Santos & G. F. Turner. 2006. Colour forms of Ama- 219 pp. zonian cichlid fish represent reproductively iso- Steindachner, F. 1875. Beiträge der Kenntniss der Chro- lated species. Journal of Evolutionary Biology, 19: miden des Amazonenstromes. Sitzungsberichte der 1139-1148. kaiserlichen Akademie der Wissenschaften. Mathe- Regan, C. T. 1906. Revision of the South-American matisch-Naturwissenschafliche Classe, 71: 61-137. cichlid genera Retroculus, Geophagus, Heterogramma, Varella, H. R. & C. R. Moreira. 2013. Teleocichla wajapi, and Biotoecus. Annals and Magazine of Natural a new species of cichlid from the rio Jari, Brazil, History, Ser. 7, 17: 49-66. with comments on T. centrarchus Kullander, 1988 — 1909. Description of a new cichlid fish of the genus (Teleostei: Cichlidae). Zootaxa, 3641: 17-187. Heterogramma from La Plata. Annals and Magazine Varella, H. R. & M. H. Sabaj Pérez. 2014. A titan among of Natural History, Ser. 8, 3: 270. dwarfs: Apistogramma kullanderi, new species (Tele — 1913. Fishes from the River Ucayali, Peru, collected ostei: Cichlidae). Ichthyological Exploration of by Mr. Mounsey. Annals and Magazine of Natural Freshwaters, 25: 243-258. History, Ser. 8, 12: 281-283.

Received 18 June 2015 Revised 17 February 2016 Accepted 13 March 2016

Varella & Britzke: Apistogramma eleutheria and A. flavipedunculata Ichthyological Exploration of Freshwaters An international journal for field-orientated ichthyology

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Volume 27 • Number 1 • April 2016

C O N T E N T S

Chen, Yan-Qiao, Chang-Lian Peng and E Zhang: Sinocyclocheilus guanyangensis, a new species of cavefish from the Li-Jiang basin of Guangxi, China (Teleostei: Cyprinidae).. 1 Cardoso, Yamila P., Florencia Brancolini, Ariel Paracampo, Marta Lizarralde, Raphael Covain and Juan I. Montoya-Burgos: Hypostomus formosae, a new catfish species from the Paraguay River Basin with redescription of H. boulengeri (Siluriformes: Loricari- idae)...... 9 Britski, Heraldo A. and José L. O. Birindelli: Redescription of Leporinus altipinnis, a senior synonym of L. falcipinnis, and comments on L. holostictus (Characiformes: Anostomi- dae)...... 25 Britz, Ralf: Pillaiabrachia siniae, a new species of earthworm eel from northern Myanmar (Teleostei: Synbranchiformes: Chaudhuriidae)...... 41 Decru, Eva, Emmanuel Vreven, Oumar Sadio and Jos Snoeks: Brycinus epuluensis, a new species from the Epulu River (Congo basin), Africa (Teleostei: Alestidae)...... 49 Liu, Shu-Wei, Jun-Xing Yang and Xiao-Yong Chen: Paralepidocephalus translucens, a new species of loach from a cave in eastern Yunnan, China (Teleostei: Cobitidae)...... 61 de Oliveira, Renildo R., Lucia Rapp Py-Daniel, Claudio H. Zawadzki and Jansen Zuanon: Two new Amazonian species of Ancistrus with vestigial adipose fin, with an appraisal on adipose fin loss in neotropical armoured catfishes (Teleostei: Loricariidae)...... 67 Varella, Henrique R. and Ricardo Britzke: Apistogramma eleutheria and A. flavipedunculata, two new species of dwarf cichlids from the rio Curuá on Serra do Cachimbo, Brazil (Teleostei: Cichlidae)...... 81

Cover photograph Pillaiabrachia siniae (photograph by Ralf Britz) Ralf Britz (this volume pp. 41-47)

Articles appearing in this journal are indexed in:

AQUATIC SCIENCES and FISHERIES ABSTRACTS BIOLIS - BIOLOGISCHE LITERATUR INFORMATION SENCKENBERG CAMBRIDGE SCIENTIFIC ABSTRACTS CURRENT CONTENTS/AGRICULTURE, BIOLOGY & ENVIRONMENTAL SCIENCES and SCIE FISHLIT ZOOLOGICAL RECORD

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