Hyphessobrycon daguae Eigenmann, 1922

Ota, Renata R., Carvalho, Fernando R. & Pavanelli, Carla S., 2020, Taxonomic review of the Hyphessobrycon panamensis species-group (Characiformes: Characidae), Zootaxa 4751 (3), pp. 401-436 : 415-421

Hyphessobrycon daguae Eigenmann, 1922

(Figs. 11–14, Tab. 4)

Hyphessobrycon panamensis daguae Eigenmann, 1922: 141 [original description, type locality: “Dagua and Patía basins”, Colombia]—. Eigenmann & Myers, 1929: 534–535 [comments on the taxon]—. Ibarra & Stewart, 1987: 48 [type catalog]—. Zarske & Géry, 2002: 26 [possible junior synonym of Hyphessobrycon condotensis ]—.Lima et al., 2003: 139 [listed; as a synonym of Hyphessobrycon panamensis ].

Hyphessobrycon panamensis (not Durbin) —. Eigenmann, 1918: 186 [in part; specimens from río Patía]—. Eigenmann, 1920a: 17 [in part; specimens from río Patía]—. Barriga, 1994: 73 [list of species, northwest of Ecuador]—. Eigenmann, 1922: 141 [in part; specimens from Colombia: Raspadura, and Tambo]—. Weitzman & Palmer, 1997: 228 [in part: specimens from río Esmeraldas, río Santiago, río Bogotá, Estero Sabalera, Ecuador; comments on color in life and distribution]—.Lima et al., 2003: 139 [as a synonym of H. panamensis ].

Hyphessobrycon ecuadoriensis (not Eigenmann & Henn) —. Barriga, 2012: 107 [list of species, Ecuador]—. García-Alzate et al., 2013: 184 [in part: MEPN 85-117].

Hyphessobrycon chocoensis Garcia-Alzate, Román-Valencia & Taphorn, 2013: 183 [original description, type locality: “ Colombia, Nariño, Barbacoas, Quebrada la Tundera afluente del río Quigualpí”]—. Maldonado-Ocampo et al., 2012: 115 [ Colombia, Chocó; photograph; short description; distribution] —. DoNascimiento et al., 2017: 49 [listed, Colombia]. NEW SYNONYM.

Hyphessobrycon sp. nov. Jiménez-Prado et al., 2015: 139 [brief description; biological data; distribution].

Diagnosis. Hyphessobrycon daguae differs from all congeners, except from H. compressus , H. diancistrus , H. otrynus , and those belonging to the H. panamensis species-group, by the presence of one large bony hook per fin ray, antero-dorsally oriented and surrounded by a fleshy area, on last unbranched and first branched anal-fin rays of mature males (vs. absence of hooks, or presence of small and numerous hooks by ray). Hyphessobrycon daguae can be distinguished from H. compressus by the absence of a black blotch on the dorsal fin (vs. presence), by the presence of predorsal series (vs. absence), and by presenting 32–35 scales in longitudinal series (vs. 41–48). It can be distinguished from H. diancistrus and H. otrynus by presenting the caudal fin hyaline (vs. dark blotches on caudalfin lobes in H. diancistrus ; distal third of caudal fin darkened in H. otrynus ). It differs from the remaining species of the H. panamensis group by presenting only one diffuse humeral blotch (vs. two humeral blotches, the first one well defined), and by presenting the lateral line frequently disrupted (vs. incompletely pored, or rarely disrupted in H. condotensis ). It can be further distinguished from H. columbianus by the absence of a silvery-blue iridescent coloration on dorsal region of flank in life (vs. presence), and by the lower body depth (39.8–43.0% SL vs. 46.0–50.3% SL). It can be distinguished from H. savagei by presenting one diffuse humeral blotch, extending one longitudinal scale row below the lateral line (vs. two humeral blotches equally pigmented and always well defined, extending two to four longitudinal scale rows below the lateral line). It also differs from H. panamensis by a higher number of branched anal-fin rays (23–27, mode = 24 vs. 19–23, modes = 21 and 22). It can be additionally distinguished from H. bussingi and H. condotensis by presenting 7–8 (mode = 7) scale rows between lateral line and dorsal-fin insertion (vs. 6–7, mode = 6½ in H. bussingi ; and 5–7, mode = 6 in H. condotensis ).

Description. Morphometric data in Table 4. Body compressed, greatest depth at vertical through dorsal-fin origin. Dorsal profile of body convex from upper lip to vertical through anterior nostril; straight or slightly concave from this point to distal tip of supraoccipital process. Slightly convex from tip of supraoccipital process to dorsal-fin origin; straight or slightly convex from this point to adipose-fin origin; and slightly concave along caudal peduncle. Ventral profile of body convex from lower lip to pelvic-fin origin; straight or slightly convex from this point to analfin origin; straight or slightly convex along anal-fin base; and slightly concave along caudal peduncle.

Jaws equal, mouth terminal. Premaxillary teeth in two rows; outer row with 2(2), 3(4), 4(11), or 5(2) tricuspid teeth; inner row with 4(1) or 5(25) tri- to pentacuspid teeth. Posterior margin of maxilla slightly posterior to vertical through anterior margin of pupil. Maxilla with 1(2), 2(18), or 3(6) conical to tricuspid teeth. Dentary with 4(10), 5(14), or 6(4) large penta and tricuspid teeth, gradually decreasing in size, followed by up to 11 conical and abruptly smaller teeth. Central cusp longer than lateral ones.

Scales cycloid. Lateral line incompletely pored with 8(1), 9(1), 10(1), 11(1), 13(1), 14*(1), 16(2), 21(1), 25(1), 31(1), disrupted with total of 15(1), 16(1), 18(1), 20(1), 21(1), 23(1), 26(1), 27(1), 28(1), 29(1), 30(1), 33(2), or completely pored with 35(1) perforate scales. Longitudinal series, including perforate scales, with 32(7), 33(5), 34(3), 35(6). Scale rows between dorsal-fin origin and lateral line 7(19) or 8(2). Scale rows between lateral line and pelvic-fin insertion 5(3), 5½(2), or 6(16). Predorsal series with 10(15) or 11(8) scales. Single row of ten scales covering base of anteriormost anal-fin rays. Circumpeduncular scales 13(6) or 14(16).

Dorsal-fin rays ii,7,ii(4), 8,i(9), 9(9), or 10(1). Adipose fin present. Pectoral-fin rays i,10(2), 11(10), or 12(10), distal tip of rays surpassing pelvic-fin origin. Pelvic-fin rays i,7(21), distal tip surpassing anal-fin origin. Anal-fin rays iii(1) or iv(26), 23(5), 24(11), 25*(4), 26(6), or 27(1). Anal fin of mature males with large bony hooks. Caudal fin bifurcate, lobes approximately of same size, i,9/8,i rays.

First gill arch with 6(3), 7(16), or 8(2) rakers on epibranchial, 1(15) on intermediate cartilage, 11(4), 12(11), or 13(5) on cerato+hipobranchial. Branchiostegal rays 4.

Color in alcohol. Background coloration beige. Dorsal region of head and middorsal region of body with dense concentration of melanophores. Region above pupil darkened. Infraorbitals and opercle beige, with few scattered melanophores. Opercle with some guanine pigmentation. Lower portion of maxilla and gular region with few scattered melanophores, slightly more concentrated on upper half. Single humeral blotch, diffuse and vertically elongat- ed, with scattered melanophores equally distributed, extending three longitudinal scale rows above and one below lateral line. Body sides with melanophores evenly distributed, except on anteroventral half, with few melanophores. Posterior margin of scales on superior third of flanks with concentration of melanophores. Brownish-gray longitudinal midlateral stripe, from humeral blotch to caudal peduncle, more conspicuous on posterior region. Fins hyaline,

Color in life. (Based in Jimenéz-Prado et al. 2015: 141, as Hyphessobrycon sp. nov.). Background color silvery to beige. Dark markings as in preserved specimens. Dorsal region of body brownish-gray. Upper half of head brownish-gray; lower half of head silvery to beige. Humeral blotch absent or inconspicuous. Brownish-gray longitudinal midlateral stripe. Abdominal region whitish-yellow. Fins hyaline; rosy or reddish in living specimens.

Distribution. Hyphessobrycon daguae is known from the río Dagua, río Patía and río Telembí basins in Colombia, and río Mira (at border between Colombia and Ecuador), río Cayapas, río Mataje, and río Santiago basins, in Ecuador ( Figs. 13 , and 21).

Sexual dimorphism. Mature males with one pair of large bony hooks per fin ray, antero-dorsally oriented and surrounded by a fleshy area, from the last unbranched (on the fourth segment) to the third branched (varying on fourth to seventh segments) anal-fin ray of mature males. Very small hooks, difficult to discern, can be present on distal segments of most anterior anal-fin rays. Dorsal-fin length of males much longer than in females.

Ecology and habitat. Hyphessobrycon daguae occur in watercourses with slow flowing, clear water, and abundant riparian vegetation ( Jiménez-Prado et al. 2015, as Hyphessobrycon sp. nov.).

Remarks. Eigenmann (1922) described Hyphessobrycon daguae as a subspecies, H. panamensis daguae . In the original description, S.F. Hildebrand highlighted that its lateral line was “frequently almost completely pored”. Despite that, subsequently to its description, H. panamensis daguae has been considered as a junior synonym of H. condotensis or of H. panamensis . However, H. condotensis and H. panamensis present the lateral line typically incompletely pored (except for one specimen of H. condotensis , ICNMHN 205, with lateral line disrupted; and one specimen of H. panamensis , UCR 1463006, with lateral line completely pored). Additionally, H. daguae can be distinguished from these species by presenting a single diffuse humeral blotch (vs. two humeral blotches, the first one always well defined, even if inconspicuous).

Hyphessobrycon daguae presents polymorphism on the lateral line (which can be incompletely, disrupted, or completely pored) similar to that found in some other Characidae , such as H. balbus Myers ( Burger et al., 2019) , and Hemigrammus ataktos Marinho, Birindelli & Dagosta ( Marinho et al., 2014) . This condition, as well as other characters, are similar to those presented in the description of Hyphessobrycon chocoensis , a species described from the río Telembí basin, which is a tributary of the río Patía (type locality of H. daguae ). Furthermore, two syntypes of H. daguae (FMNH 56527 and FMNH 56528) are exactly from the same municipality and drainage of the type series of H. chocoensis . Therefore, as no differences between these nominal species were found, H. daguae is herein revalidated and H. chocoensis is considered as its junior synonym.

The type series of Hyphessobrycon panamensis daguae is composed by nine syntypes (FMNH 56156, FMNH 56526, FMNH 56527, FMNH 56528, FMNH 56529, and FMNH 58316). The lectotype herein designated is the 26.0 mm SL specimen of FMNH 58316 (Fig. 11a), and the remaining syntypes become thus paralectoytpes, as ruled by the article 74 of ICZN (1999).

Another issue involving Hyphessobrycon daguae concerns the specimens from the province of Esmeraldas, Northern Ecuador. Weitzman & Palmer (1997: 228-229) identified the specimens from the río Esmeraldas as H. panamensis , while Barriga (2012) recorded H. ecuadoriensis from the río Santiago (a river drainage very close to río Esmeraldas). On the other hand, García-Alzate et al. (2013) recorded H. ecuadoriensis from the río Mira (at the border between Colombia and Ecuador) to the drainages of Southern Ecuador. However, Jiménez-Prado et al. (2015) restricted the distribution of H. ecuadoriensis to the río Guayas basin (southern Ecuador), and identified the specimens from the río Mataje and río Santiago basins as “ Hyphessobrycon sp. nov.”, distinguishing it from H. daguae (as H. chocoensis ) by the presence of seven perforated scales on the lateral line (vs. 10 or more). Nonetheless, the specimens from the río Bravo (MEPN 3701) and from the río Santiago basins (ZUEC 11330) can present the lateral line incompletely pored or disrupted ( Fig. 14 ). This emphasizes the polymorphic condition of the character for those populations, and, consequently, the count of perforated scales in “ Hyphessobrycon sp. nov.” is herein considered to be best interpreted as a population variation within H. daguae . Therefore, the distribution of Hyphessobrycon daguae is extended herein to the río Mira, río Cayapas, río Mataje, and río Santiago basins, in Ecuador.

Material examined. 34 specimens (15.6–43.4 mm SL). Type material. FMNH 58316, lectotype of Hyphessobrycon daguae (by present designation), 26.0 mm SL: Colombia, río Patía basin; A. W. Henn, 1913. FMNH 144330, paralectotype of Hyphessobrycon daguae , 28.0 mm SL, same data as lectotype. IUQ 3035, holotype of Hyphessobrycon chocoensis , 35.3 mm SL: Colombia, Nariño, Barbacoas, creek la Tundera, tributary of the río Quigualpí , 01°39’39’’N 78°09’04’’W; C. Román-Valencia, C. García-Alzate, R. Ruiz-C, V. Tovar Sr., V. Tovar Jr. & H. Cortes, 16 Jul 2008 ; IUQ 2274, paratypes of Hyphessobrycon chocoensis , 6, 15.6–40.6 mm SL: Colombia, Nariño, Barba- coas, upper río Yamundé, tributary of the río Telembí , 01°39’39’’N 78°09’04’’W; C. Román-Valencia, C. García- Alzate, R. Ruiz-C, V. Tovar Sr., V. Tovar Jr. & H. Cortes, 16 Jul 2008 ; IUQ 2275, paratypes of Hyphessobrycon chocoensis , 7, 24.9–33.4 mm SL, same data as holotype. Non-type material. Colombia, Nariño, río Telembi basin: IUQ 2311, 3 , 21.5–34.4 mm SL, creek on Hacienda, tributary of the río Telembi , 01°30’N 77°58’W, 16 Jul 2008 . Ecuador, Esmeraldas: MEPN 3697, 4 , 34.1–43.4 mm SL, tributary of the río Bravo, 8 km above río Bravo falls, río Cayapas , 00°38’22’’N 78°55’24’’W; R. Barriga, C. Mina & W. Nazareno, 18 Jul 1985 ; MEPN 3701, 6 , 35.0– 43.3 mm SL, same data as MEPN 3697; ZUEC 11330, 5 , 30.5 – 27.8 mm SL, Estero la Chiquita, tributary of the río Santiago , 1°13’50”N, 78°45’17”W; R. Barriga & C. Mina , 21 Oct 1985 .

Figures

FIGURE 13. Distribution map of Hyphessobrycon daguae (blue dots; blue star represents type locality) and H. ecuadoriensis (orange dots; orange star represent type locality).

FIGURE 14. Hyphessobrycon daguae, ZUEC 11330: Ecuador, Esmeraldas, Estero la Chiquita, a) 37.8 mm SL, male, b) 34.8 mm SL, female.