Hyphessobrycon panamensis daguae Eigenmann, 1922: 141 Eigenmann & Myers, 1929: 534–535 Ibarra & Stewart, 1987: 48 Zarske & Géry, 2002: 26 Hyphessobrycon condotensis Hyphessobrycon panamensis Hyphessobrycon panamensis Eigenmann, 1918: 186 Eigenmann, 1920a: 17 Barriga, 1994: 73 Eigenmann, 1922: 141 Weitzman & Palmer, 1997: 228 H. panamensis Hyphessobrycon ecuadoriensis Barriga, 2012: 107 García-Alzate et al. , 2013: 184 Hyphessobrycon chocoensis Garcia-Alzate, Román-Valencia & Taphorn, 2013: 183 Maldonado-Ocampo et al ., 2012: 115 DoNascimiento et al. , 2017: 49 Hyphessobrycon Jiménez-Prado et al ., 2015: 139 Taxonomic review of the Hyphessobrycon panamensis species-group (Characiformes: Characidae) Ota, Renata R. Carvalho, Fernando R. Pavanelli, Carla S. Zootaxa 2020 2020-03-18 4751 3 401 436 5J328 Eigenmann, 1922 Eigenmann 1922 [151,671,1521,1548] Actinopterygii Characidae Hyphessobrycon Animalia Characiformes 14 415 Chordata species daguae      Hyphessobrycon panamensis daguae Eigenmann, 1922: 141[original description, typelocality: “Dagua and Patía basins”, Colombia]—.  Eigenmann & Myers, 1929: 534–535[comments on the taxon]—.  Ibarra & Stewart, 1987: 48[ typecatalog]—.  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, typelocality: “ 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.    Hyphessobryconsp. nov.  Jiménez-Prado et al., 2015: 139[brief description; biological data; distribution].   Diagnosis.  Hyphessobrycon daguaediffers from all congeners, except from  H. compressus,  H. diancistrus,  H. otrynus, and those belonging to the  H. panamensisspecies-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 daguaecan be distinguished from  H. compressusby 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. diancistrusand  H. otrynusby 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. panamensisgroup 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. columbianusby 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. savageiby 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. panamensisby 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. bussingiand  H. condotensisby 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,   FIGURE 12.  Hyphessobrycon daguae, IUQ 3035, holotype of  Hyphessobrycon chocoensis, 35.3 mm SL, male: Colombia, Nariño, Barbacoas, stream la Tundera.   TABLE 3.Morphometric data of  Hyphessobrycon condotensisand  H. sebastiani. SD = standard deviation; N = number of specimens.  Hyphessobrycon condotensis, n= 4 for Standard lengh, other measures n= 3.      H. condotensis– types   H. sebastiani– Holotype río Atrato basin – non types río San Juan basin – non types  Range Mean SD N Range Mean SD N Range Mean SD  Standard length 14.4–33.1 26.3 – 38.4 42 18.4–40.0 29.7 – 9 13.9–38.4 30.1 –  Body depth 38.1–41.1 39.2 1.9 43.2 18 34.7–46.5 41.5 3.0 7 38.9–43.2 41.0 1.7  Predorsal distance 51.8–52.8 52.2 0.5 52.9 18 50.9–56.6 54.4 1.5 7 52.7–55.3 53.8 0.9  Prepectoral distance – – – 27.9 18 29.1–32.8 31.0 1.1 7 27.3–31.3 29.2 1.4  Prepelvic distance 45.7–47.9 46.7 1.1 49.0 18 48.1–52.4 50.3 1.2 7 47.8–50.8 49.3 1.0  Preanal distance – – – 62.5 18 61.0–69.4 64.7 2.1 7 61.7–65.7 63.3 1.4  Caudal-peduncle depth 10.8–11.9 11.2 0.6 13.1 18 8.7–12.2 10.6 0.9 7 8.6–13.1 11.6 1.6  Caudal-peduncle length 11.9–12.7 12.2 0.4 9.4 18 6.5–11.9 9.6 1.2 7 8.9–11.2 9.8 0.8  Dorsal-fin base length 14.1–15.1 14.5 0.5 16.4 18 14.5–18.7 16.3 1.1 7 14.1–16.4 15.1 0.8  Dorsal-fin length 34.0–34.4 34.2 0.3 29.7 18 25.2–32.3 29.1 2.0 7 26.0–32.7 29.3 2.0  Pectoral-fin length 23.9–26.6 25.2 1.3 25.8 18 19.6–25.9 24.0 1.7 7 23.0–27.0 24.8 1.6  Pelvic-fin length 17.7–19.5 18.5 0.9 19.3 18 15.9–21.2 19.1 1.6 7 16.8–20.5 18.6 1.4  Anal-fin base length 34.7–38.0 36.4 1.6 39.6 18 35.0–40.0 37.1 1.3 7 35.4–39.6 37.7 1.6  Anal-fin length 24.0–25.5 24.8 0.7 19.8 18 17.9–23.7 21.2 1.5 7 18.2–25.9 21.5 2.9  Orbit to dorsal-fin origin 34.9–36.1 35.6 0.6 39.1 18 38.0–41.1 39.5 0.9 7 38.3–40.1 39.0 0.6  Dorsal-fin origin to caudal peduncle 53.9–56.0 55.0 1.1 51.6 18 49.1–56.7 52.1 2.0 7 38.9–52.6 50.0 4.9  Head depth 33.1–35.6 34.1 1.4 32.6 18 29.8–35.5 32.8 1.7 7 30.6–32.7 31.7 0.8  Head length 28.0–29.2 28.7 0.6 27.9 18 26.8–30.9 29.1 1.0 7 27.5–29.3 28.3 0.6  Dorsal-fin origin to adipose-fin origin 37.2–39.1 37.9 1.0 39.1 18 35.2–42.2 37.9 1.8 7 37.1–39.1 38.0 0.7  Pelvic- to anal-fin origin – – – 14.6 18 12.7–17.4 14.5 1.2 7 14.1–15.9 15.0 0.7  Orbital diameter 43.2–46.0 44.3 1.4 44.9 18 40.4–47.6 43.4 2.0 7 43.3–47.1 45.0 1.5  Snout length 20.9–22.5 21.7 0.8 26.2 18 23.5–27.6 25.5 1.1 7 23.5–27.4 25.6 1.3  Interorbital width 29.7–33.9 31.7 2.1 32.7 18 28.7–35.2 31.9 1.7 7 30.6–34.5 32.6 1.5  Upper-jaw length 45.2–48.6 47.3 1.9 46.7 18 46.6–53.8 50.5 2.3 7 46.7–51.2 49.0 1.8   FIGURE 13.Distribution map of  Hyphessobrycon daguae(blue dots; blue star represents type locality) and  H. ecuadoriensis(orange dots; orange star represent type locality).  Color in life.(Based in Jimenéz-Prado et al.2015: 141, as  Hyphessobryconsp. 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 daguaeis known from the río Dagua, río Patía and río Telembí basins in Colombia, and río Mira (at border between Colombiaand 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 daguaeoccur in watercourses with slow flowing, clear water, and abundant riparian vegetation ( Jiménez-Prado et al. 2015, as  Hyphessobryconsp. nov.).   Remarks. Eigenmann (1922)described  Hyphessobrycon daguaeas 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 daguaehas been considered as a junior synonym of  H. condotensisor of  H. panamensis. However,  H. condotensisand  H. panamensispresent 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. daguaecan 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).   TABLE 4.Morphometric data of  Hyphessobrycon daguae. SD = standard deviation; N = number of specimens.    Holotype of  H. chocoensis Paratypes of  H. chocoensis Non-types  N Range Mean SD N Range Mean SD  Standard length 35.3 14 15.6–40.6 30.2 – 17 21.5–43.4 36.4 –   Percentages of standard length  Body depth 43.6 10 39.8–43.0 41.5 1.1 16 39.0–45.7 41.0 1.8  Predorsal distance 54.4 10 51.0–53.8 52.5 1.0 16 48.0–53.1 50.7 1.4  Prepectoral distance 29.2 10 27.8–28.8 28.4 0.4 16 27.7–29.9 28.7 0.6  Prepelvic distance 48.7 10 45.8–48.1 47.3 0.9 16 44.7–48.5 46.5 1.1  Preanal distance 61.5 10 59.5–62.8 61.2 1.2 16 57.7–60.9 59.7 0.9  Caudal-peduncle depth 10.8 10 9.9–11.5 10.8 0.5 16 9.7–11.2 10.5 0.5  Caudal-peduncle length 11.0 10 9.5–10.8 10.1 0.5 16 9.7–12.0 10.6 0.6  Dorsal-fin base length 16.4 10 14.5–17.7 16.1 1.2 16 14.5–19.9 17.2 1.3  Dorsal-fin length 34.6 10 28.3–36.3 31.7 2.6 16 29.1–45.7 34.5 4.3  Pectoral-fin length 25.8 10 22.9–25.6 24.3 0.9 16 22.7–25.8 24.7 0.9  Pelvic-fin length 21.5 10 19.1–22.1 20.0 0.9 16 17.8–21.4 19.9 1.0  Anal-fin base length 39.9 10 35.6–42.5 38.6 1.9 16 36.0–41.3 39.0 1.3  Anal-fin length 17.0 10 17.3–24.6 20.4 2.3 16 16.7–22.1 18.6 1.7  Orbit to dorsal-fin origin 41.6 10 38.0–40.7 38.9 0.9 16 35.4–39.2 37.8 1.1  Dorsal-fin origin to caudal peduncle 53.5 10 50.9–53.7 52.4 1.0 15 51.0–54.6 52.7 1.1  Head depth 32.9 10 31.0–32.6 31.9 0.6 16 29.3–34.3 31.7 1.4  Head length 27.8 10 26.3–29.0 27.7 0.9 16 25.9–28.6 27.0 0.8  Dorsal-fin origin to adipose- fin origin 40.8 10 36.8–40.6 38.3 1.4 16 36.8–41.9 39.3 1.4  Pelvic- to anal fin origin 13.3 10 13.7–16.0 14.8 0.8 16 11.8–16.3 14.5 1.2   Percentages of head length  Orbital diameter 45.9 10 41.1–46.7 44.4 1.8 16 39.8–47.9 43.1 2.3  Snout length 25.5 10 22.2–26.9 25.2 1.3 16 23.7–26.9 25.4 1.1  Interorbital width 31.6 10 30.8–33.3 31.8 0.8 16 30.5–34.5 32.7 1.3  Upper-jaw length 49.0 10 44.1–51.8 48.3 2.1 15 44.2–49.6 47.4 1.6   Hyphessobrycon daguaepresents polymorphism on the lateral line (which can be incompletely, disrupted, or completely pored) similar to that found in some other Characidae, such as  H. balbusMyers ( Burger et al., 2019), and  Hemigrammus ataktosMarinho, 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 syntypesof  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. daguaeis herein revalidated and  H. chocoensisis considered as its junior synonym. The type series of  Hyphessobrycon panamensis daguaeis composed by nine syntypes(FMNH 56156, FMNH 56526, FMNH 56527, FMNH 56528, FMNH 56529, and FMNH 58316). The lectotypeherein designated is the 26.0 mm SL specimen of FMNH 58316 (Fig. 11a), and the remaining syntypesbecome thus paralectoytpes, as ruled by the article 74 of ICZN (1999).   FIGURE 14.  Hyphessobrycon daguae, ZUEC 11330: Ecuador, Esmeraldas, Estero la Chiquita, a) 37.8 mm SL, male, b) 34.8 mm SL, female. Another issue involving  Hyphessobrycon daguaeconcerns the specimens from the province of Esmeraldas, Northern Ecuador. Weitzman & Palmer (1997: 228-229)identified the specimens from the río Esmeraldasas  H. panamensis, while Barriga (2012)recorded  H. ecuadoriensisfrom the río Santiago (a river drainage very close to río Esmeraldas). On the other hand, García-Alzate et al.(2013)recorded  H. ecuadoriensisfrom the río Mira (at the border between Colombiaand Ecuador) to the drainages of Southern Ecuador. However, Jiménez-Prado et al.(2015)restricted the distribution of  H. ecuadoriensisto the río Guayasbasin (southern Ecuador), and identified the specimens from the río Mataje and río Santiago basins as “  Hyphessobryconsp. 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 “  Hyphessobryconsp. nov.” is herein considered to be best interpreted as a population variation within  H. daguae. Therefore, the distribution of  Hyphessobrycon daguaeis extended herein to the río Mira, río Cayapas, río Mataje, and río Santiago basins, in Ecuador.   Material examined.   34 specimens( 15.6–43.4 mmSL). Type material. FMNH 58316, lectotypeof  Hyphessobrycon daguae(by present designation), 26.0 mm SL: Colombia, río Patíabasin; A. W. Henn, 1913. FMNH 144330, paralectotypeof  Hyphessobrycon daguae, 28.0 mm SL, same data as lectotype. IUQ 3035, holotypeof  Hyphessobrycon chocoensis, 35.3 mmSL: 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, paratypesof  Hyphessobrycon chocoensis, 6, 15.6–40.6 mmSL: 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, paratypesof  Hyphessobrycon chocoensis, 7, 24.9–33.4 mmSL, same data as holotype. Non-type material. Colombia, Nariño, río Telembibasin: IUQ 2311, 3, 21.5–34.4 mmSL, 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 mmSL, tributary of the río Bravo,  8 kmabove río Bravofalls, 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 mmSL, same data as MEPN3697; ZUEC 11330, 5, 30.5– 27.8 mmSL, Estero la Chiquita, tributary of the río Santiago, 1°13’50”N, 78°45’17”W; R. Barriga& C. Mina,  21 Oct 1985. 2577369981 [437,1092,1806,1833] FMNH 20 421 34 2577369997 FMNH A. W. Henn Colombia rio Patia 20 421 1 lectotype 2577369985 IUQ A. W. Henn Colombia rio Patia 20 421 1 paralectotype 2577369996 2008-07-16 R, V la Tundera & C. Roman-Valencia & C. Garcia-Alzate & Tovar Sr. & Tovar Jr. & H. Cortes Colombia 1.6608334 rio Quigualpi 21 -78.151115 Barbacoas 20 421 1 Narino holotype 2577369999 2008-07-16 R, V C. Roman-Valencia & Alzate & Tovar Sr. & Tovar Jr. & H. Cortes Colombia 21 422 1.6608334 rio Telembi 21 -78.151115 upper rio Yamunde 20 421 1 Narino paratype 2577369993 2008-07-16 IUQ la Tundera & C. Roman-Valencia & C. Garcia-Alzate & Tovar Sr. & Tovar Jr. & H. Cortes Colombia 1.6608334 rio Telembi 21 -78.151115 Barbacoas 21 422 2 Narino paratype 2577369980 [209,1433,260,287] 2008-07-16 IUQ Colombia 1.5 rio Telembi 1308 -77.96667 Hacienda 21 422 3 1 Narino paratype 2577369975 [151,479,297,323] MEPN Ecuador Esmeraldas 21 422 1 Esmeraldas paratype 2577369987 1985-07-18 R Barriga, C. & W. Nazareno Ecuador 0.6394444 rio Cayapas 21 -78.92333 rio Bravo 21 422 4 1 Esmeraldas paratype 2577369982 1985-07-18 MEPN Barriga, C. & W. Nazareno Ecuador 0.6394444 rio Cayapas 21 -78.92333 rio Bravo 21 422 6 1 Esmeraldas paratype 2577369979 1985-10-21 R Barriga Ecuador 1.2305555 rio Santiago 21 -78.75472 Estero la Chiquita 21 422 5, 30.5 1 Esmeraldas paratype