Brycon atricaudatus Brycon oligolepis Brycon coxeyi Fowler, 1943 : 3 Géry, 1972b : 7 A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae) Lima, Flávio C. T. Zootaxa 2017 4222 1 1 189 NH75 Fowler, 1943 Fowler 1943 [151,489,1628,1654] Actinopterygii Bryconidae Brycon Animalia Characiformes 20 21 Chordata species coxeyi    Brycon atricaudatus(sic) (non Kner & Steindachner, 1863): Boulenger, 1898a: 3 (part;  Ecuador, “ RioZamora”).  Brycon oligolepis(non Regan): Tortonese, 1939: 49 (  Ecuador, “ RioZamora”; same specimen examined by Boulenger, 1898a).  Brycon coxeyi Fowler, 1943: 3–4, fig.2 ( Typelocality, “Hacienda La Mascota, mouth of the Rio Pastaza, basin of the Rio   Marañon, Ecuador”).   Lima, 2003: 175–176 (correction of typelocality to “ Ecuador, Napo-Paztaza, Hacienda Mascota,  Río    Topo, a tributary of RíoPaztaza, 1°25’S, 78°11’W”).  ?  Bryconspecies:  Géry, 1972b: 7–8 ( RíoVillano, tributary of RíoCururay, Amazon basin).   Diagnosis.  Brycon coxeyican be distinguished from all remaining cis-andean  Bryconspecies, except  Brycon stolzmanni, by possessing a distinct patch of dark pigmentation on the opercle (vs. patch of dark pigmentation on the opercle absent in the remaining cis-andean  Bryconspecies) and infraorbital series not closed, with the orbital margin partially delimited by the frontal bone (vs. infraorbital series closed, frontal excluded from the orbital series by the contact between the supraorbital and the sixth infraorbital bone).  Brycon coxeyican be distinguished from  Brycon stolzmanniby possessing the dark patch of pigmentation on opercle well-defined, typically lying at its center (vs. dark patch of pigmentation diffuse and restricted to the upper half of opercle in  B. stolzmanni).   Description.Morphometric data are presented in Table 2. Relatively small-sized species, largest examined specimen 225.0 mm SL. Body moderately high. Largest body height slightly ahead of dorsal-fin origin. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, slightly concave from latter point to basis of supraoccipital process, and moderately convex from latter point to dorsal-fin origin, straight along dorsal-fin basis, straight to slightly convex from dorsal-fin terminus to adipose-fin origin. Dorsal profile of caudal peduncle slightly concave. Ventral profile slightly convex from lower lip to pelvic-fin insertion, straight to slightly convex from this point to anal-fin origin and approximately straight along anal-fin base. Ventral profile of caudal peduncle slightly concave.   TABLE 2.Morphometric data of  Brycon coxeyi(A: holotype, ANSP 70156).    A n Range Mean  Standard length (SL) 131.6 58 50.2–218.0 -  Percentages of standard length  Depth at dorsal-fin origin - 57 26.2–34.0 30.4  Snout to dorsal-fin origin 50.1 58 49.1–54.3 51.5  Dorsal-fin base length 11.0 58 10.6–14.1 12.2  Posterior terminus of dorsal fin to adipose fin 23.8 58 18.9–27.9 25.0  Posterior terminus of dorsal fin to hypural joint 35.6 58 35.6–50.2 40.8  Snout to pelvic-fin insertion - 56 46.2–53.3 50.3  Snout to anal-fin origin 70.5 58 62.6–71.3 66.7  Anal-fin base length 19.8 58 19.8–25.1 22.6  Caudal peduncle length 15.4 58 12.6–18.0 16.1  Dorsal-fin height 20.1 57 16.6–22.2 19.0  Pectoral-fin length 19.4 58 17.5–23.6 19.9  Pelvic-fin length - 58 12.1–17.3 14.7  Caudal peduncle depth 9.3 58 8.6–11.2 9.9  Head length - 57 25.4–30.0 28.4  Percentages of head length  Head height - 57 72.9–85.6 79.2  Snout length - 57 22.1–32.8 28.4  Upper jaw length - 57 40.9–49.5 45.2  Horizontal eye diameter - 57 19.7–31.5 23.8  Post-orbital length - 57 45.6–55.7 52.4  Least interorbital width - 57 27.5–36.2 32.4 Head profile slightly acute anteriorly, mouth terminal, jaws approximately isognathous. Maxillary extending posteriorly to vertical slightly anterior to middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Five (2), 6 (24), 7 (25), 8(5), or 9(1) tricuspidate teeth in outer series. Two (4), 3 (42), or 4 (7) tri- to tetracuspidate teeth in second, inner premaxillary row, plus 2 (4), 3 (42), or 4(7) tricuspidate teeth between the first and third rows. Two teeth in third premaxillary row, medial teeth largest, symphyseal teeth smaller, both pentacuspidate. Maxillary with distal portion expanded and rounded in profile. Nine to 21 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 6(2), 7(1), 9(1), 11 (1), or 13 (1) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, tetra- to pentacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, tri- to unicuspidate. Inner (lingual) series consisting of a single unicuspid symphyseal tooth, considerably smaller than teeth of main series anterior to it, plus row of 6 small, unicuspidate teeth, originating behind last tooth of the main series.   FIGURE 11.  Brycon coxeyi, holotype, ANSP 70156, 131.6 mm SL: Ecuador, Tungurahua, Río Paztaza. Photo by M.W. Littmann.   FIGURE 12.  Brycon coxeyi, MEPN uncatalogued, 192.9 mm SL: Ecuador, Zamora-Chinchipe, Río Nangaritza.   FIGURE 13.  Brycon coxeyi, MEPN uncatalogued, 148.5 mm SL: Ecuador, Zamora-Chinchipe, Río Nupatakaine.   FIGURE 14.  Brycon coxeyi, MUSM 6136, 131.5 mm SL: Peru, Amazonas, Río Comainas.   FIGURE 15.  Brycon coxeyi, MUSM 6136, 135.8 mm SL: Peru, Amazonas, Río Comainas. Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-two (1), 43 (1), 44(3), 45 (15), 46 (17), 47 (8), 48 (9), 49 (2), 50 (1), or 52 (1) scales in lateral line series; laterosensory tube simple, straight or slightly deflected downwards. Horizontal scale rows between dorsal-fin origin and lateral line 8 (22), 9 (34), or 10 (2). Horizontal scale rows between lateral line and pelvic-fin 4 (9), 5(43), or 6(5). Circumpeduncular scales 14 (4), 15 (4), 16 (30), 17 (16), 18 (1), or 19 (1). Dorsal-fin rays ii, 9, one specimen ii, 10. Dorsal fin origin at middle of SL. Dorsal fin displaying numerous (c. 15–30 per fin-ray main branch) small hooks on last unbranched and all branches (more numerous on last branch) of all branched rays in 13 specimens(MEPN 11101, 205.0 mm SL; MEPN 11108, 155.4 mmSL; MEPN 11115, 144.8 mmSL; MEPN 11109, 150.9 mmSL; MEPN 11103, 187.0 mm SL; MEPN 11105, 168.1 mmSL; MEPN 11114, 2, 153.0– 157.2 mmSL; MEPN 11112, 122.6 mmSL; MUSM 6127, 2, 126.6– 135.4 mmSL; MUSM 6136, 2, 131.5– 135.8 mmSL). Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 21 (6), 22 (17), 23 (33), or 24 (10). Last unbranched and anterior 2–3 branched anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin displaying numerous (c. 20–30 per fin-ray main branch) small hooks associated with dense, gelatinous tissue on last unbranched and posterior main branch of all branched rays except posteriomost ray in 15 specimens(MEPN 11101, 205.0 mm SL; MEPN 11108, 155.4 mmSL; MEPN 11115, 144.8 mmSL; MEPN 11109, 2, 149.6– 150.9 mmSL; MEPN 11103, 187.0 mm SL; MEPN 11105, 168.1 mmSL; MEPN 11102, 218.0 mm SL; MEPN 11114, 2, 153.0– 157.2 mmSL; MEPN 11112, 122.6 mmSL; MUSM 6127, 2, 126.6– 135.4 mmSL; MUSM 6136, 2, 131.5– 135.8 mmSL). Small hooks also present in the anterior main anal-fin branch in the two anteriormost anal-fin rays. Sheath of scales composed of a single series composed by 17–20 rectangular scales, covering basis of anal-fin rays. Pectoral-fin rays i, 12 (28), 13 (24), or 14 (4). Pelvic-fin rays i,7, one specimen i,6, and another single specimen i,8. Branched pelvic-fin rays with minutes hooks (c.10 per branch) on distal portion of posterior ray branches in 14 specimens(MEPN 11101, 205.0 mm SL; MEPN 11108, 155.4 mmSL; MEPN 11115, 144.8 mmSL; MEPN 11109, 2, 149.6– 150.9 mmSL; MEPN 11103, 187.0 mm SL; MEPN 11105, 168.1 mmSL; MEPN 11102, 218.0 mm SL; MEPN 11114, 2, 153.0– 157.2 mmSL; MEPN 11112, 122.6 mmSL; MUSM 6127, 2, 126.6– 135.4 mmSL; MUSM 6136, 2, 131.5– 135.8 mmSL). Main caudal-fin rays 10/9. Caudal fin forked, lobes rounded. Four branchiostegal rays. First gill arch with 10 (1), 11 (2), or 14 (1) lower gill rakers, 9 (3) or 10 (1) upper gill rakers and 1 at angle. Vertebrae 44 (1).  Coloration in alcohol.Top of head, snout, supraorbital, first, sixth, and fifth infraorbital bones, and dorsal portion of body dark grey to dark brown. Dentary, maxillary, gular area, and ventral portion of body cream-colored. Second through fourth infraorbital bones silvery in specimens retaining guanine, with a brownish tinge. Opercle silvery, with a large, rounded conspicuous dark patch of pigmentation typically lying at its central area, with the exception of darkened large specimens, described below. Humeral blotch present, little conspicuous, more evident in smaller specimens or specimens lacking scales on humeral region, split in two halves. Humeral blotch vertically elongated, lower half more conspicuous, crossed at its middle by the lateral line, extending horizontally from the third to the fourth lateral line scales. Upper half of humeral blotch less conspicuous, separated from lower half of humeral blotch by a narrow clear-colored area, two scales rows high. Lateral portions of body clear, with a silvery hue in specimens retaining guanine. Body with multiple moderately conspicuous longitudinal rows of wavy dark stripes, formed by pigmentation concentrated along scale junctions, more discernible dorsally. Caudal peduncle blotch present, moderately to highly conspicuous, rounded, extending throughout 4–5 last lateral line scales into basis of 5–6 innermost caudal-fin rays. Caudal, anal and dorsal-fins with numerous small dark chromatophores scattered over interradial membranes. Pectoral and pelvic-fins with relatively few dark chromatophores on interradial membranes. Adipose fin light grey, with a well-defined clear area at its basis. Diffuse dark pigmentantion present at lateral portion of body in several middle to large sized specimens ( 120–220 mmSL, e.g., MEPN 11101, 1, 205.0 mm SL, MEPN 11112, 122.6 mmSL). Some specimens (e.g., MEPN 11115, 3, 164.6–168.0 mm SL; MEPN 11105, 2, 168.1– 173.4 mmSL; MEPN 11103, 187.0 mm SL; MUSM 6141, 145.1 mmSL; MUSM 6136, 3, 131.5– 135.8 mmSL) develop a intense dark pigmentantion on body sides, presenting a much enlarged dark blotch on opercle, covering the entire middle and lower surface of that bone, and sides of body with a marbled pattern, with intense dark and white/silvery pigmentation alternating (see item “Sexual Dimorphism”, below). Asingle specimen (MEPN 11103, 187.0 mm SL) with a large amount of dark pigmentation on the third to the fifth infraorbital bones.  Color in life.Description based on pictures of several freshly-collected specimens from the RíoMachinaza (upper RíoZamora/ Río Santiagobasin), taken by R. Barriga and collaborators. Snout, top of head, and dorsum plumbeous-grey to brown; sides of body silvery, with a plumbeous tinge. Lower portion of head and branchiostegal rays clear, tinged with orangish in larger specimens. Dark patch of pigmentation in opercle and caudal-peduncle blotch conspicuous. Longitudinal wavy stripes moderately conspicuous, more discernible dorsally. Pectoral and pelvic-fins reddish. Anal and caudal fin with red pigmentation concentrated at its basis.  Sexual dimorphism.Hooks were observed at the anal-fin rays (and in most cases, also dorsal-fin rays and distal portion of pelvic-fin rays) of 18 specimens (MEPN 11101, 205.0 mm SL; MEPN 11108, 155.4 mmSL; MEPN 11115, 144.8 mmSL; MEPN 11109, 2, 149.6– 150.9 mmSL; MEPN 11103, 187.0 mm SL; MEPN 11105, 168.1 mmSL; MEPN 11102, 218.0 mm SL; MEPN 11114, 2, 153.0– 157.2 mmSL; MEPN 11112, 122.6 mmSL; MUSM 6141, 145.1 mmSL; MUSM 6127, 2, 126.6– 135.4 mmSL; MUSM 6144, 108.2 mmSL; MUSM 6136, 3, 131.5– 135.8 mmSL). Specimen MEPN 11115 ( 144.8 mmSL) was dissected and is a male, with well-developed testes. On other hand, a specimen lacking any hooks but collected alongside two specimens possessing hooks, MEPN 11114 (182.0 mm SL), proved to be female on dissection, with ripe oocytes. A moderate to intense dark pigmentation on body sides and a enlarged dark blotch on opercle are present in some specimens and seem to be an additional dimorphic feature, since most specimens presenting these color features also present hooks on fins. However, some specimens presenting an enlarged dark blotch on opercle (MEPN 11115, 3, 164.6–168.0 mm SL; MEPN 11117, 2, 147.9– 177.1 mmSL; MUSM 6144, 1, 108.2 mmSL) does not possess any fin hooks, and conversely, some specimens presenting hooks (MEPN 11115, 144.8 mmSL; MUSM 6127, 2, 126.6– 135.4 mmSL) present instead the “normal” color pattern.  Brycon guatemalensisis known to possess mature males with a marbled darkened pattern on sides (e.g., Bussing, 1987) and males of  Bryconspecies from all river systems of Pacific versant of the Andes in Ecuador also display a darkened color pattern when mature (R. Barriga, pers. comm..).  Common names.“Blanco”, “sábalo” (Zamora-Chinchipe, Ecuador; R. Barriga, pers. comm.).   Distribution.  Brycon coxeyiis only known from upper tributaries of the Río Marañonbasin in  Ecuadorand Peru, at altitudes ranging from 500–1392 meters a.s.l. ( Fig. 10).  Ecological notes.  Brycon coxeyiis found in fast-running, rocky-bottomed rivers of the Andean piedmont in Ecuador. Two of the the rivers known to harbor  Brycon coxeyipopulations, the Río Nangaritza and the Río Machinaza, both tributaries of the upper Río Zamora/Río Santiago basin, are fast-running, rocky-bottomed blackwater rivers (R. Barriga, pers. comm..). The Río Comainas in Cordillera del Condor, Peru, where the species also occurs, is a shallow (0.3–1.9 meters deep), relatively small (8–20 meters wide) clear-water river, with a rocky/ gravel/sandy/clay substrate from most of the year, except after heavy rains, when it becomes torrential, with turbid waters (Vari et al., 1995: 294). Crushed vegetal matter (leaves and apparently also fruit remains) were found in two specimens (MEPN 11115, 163.1 mmSL; MEPN 11114, 182.0 mm SL) examined for gut contents. Specimens presenting fin hooks and as such presumably in breeding period were found in March, April, July, September, and November. A female with ripe oocytes (MEPN 11114, 182.0 mm SL) was collected in April and a male with ripe testes (MEPN 11115, 144.8 mmSL) was collected in September. Specimens with a large amount of fat in the abdominal cavitiy were collected in February (MEPN 11119).   Remarks. Fowler (1943: 3)described  Brycon coxeyibased on a single specimen collected at the Río Topo in the upper Río Paztaza basin, Ecuador. He noticed that the species was “apparently not closely related to any of the many known forms distributed in the Atlantic drainage of the eastern Andes”, and compared it with to  Brycon henniEigenmann, a species occurring in several trans-andean river drainages of Colombia, and  B. stolzmanni. Fowler (1943)incorrectly assumed that the latter could be differentiated from  Brycon coxeyiby supposedly lacking the opercular dark patch of pigmentation, when in fact  B. stolzmannipossess it. Both species can be differentiated by the characters mentioned in the item “Diagnosis”, above. The holotype (ANSP 70156) is in a very poor condition of preservation, being very soft, with no scales left, most fins broken and the opercle bone almost disarticulated from the rest of the opercular apparatus (see Fig. 11). Still, it displays the rounded dark patch of pigmentation which is characteristic from the remaining  Bryconspecimens from the upland areas of the Amazon basin in Ecuador herein assigned to the species. As noticed by Böhlke (1958: 24–25)and Lima (2003: 175–176), the type-locality given by Fowler (1943)for  Brycon coxeyiis incorrect, and was amended by the latter as “Ecuador, Napo-Paztaza, Hacienda Mascota, Río Topo, a tributary of Río Paztaza, 1°25’S, 78°11’W”. However, according to the old label present in the jar of the holotype of  Brycon coxeyi, the specimen was not collected at the Río Topo itself but rather at the Río Paztaza near the mouth of the Río Topo, and should be again corrected, this time to “Ecuador, Província Tungurahua, Hacienda Mascota, Río Paztaza, near mouth of Río Topo, c. 1°25’S, 78°11’W”.  Boulenger (1898a: 3)examined seven  Bryconspecimens from the Río Mira and Río Peripa (Río Guayas basin), plus one specimen from the Río Zamora, all of which he identified as  Brycon atricaudatus(=  B. atrocaudatus). We have not re-examined the specimen from the Río Zamora (which is presumably deposited at the MSNG), but judging both from the locality and its meristic data (lateral line scales 46, scales above lateral line 9, and anal-fin rays 27—i.e., iii, 24— Boulenger, 1898a: 3), this record is with all likelihood referable to  Brycon coxeyi. Tortonese (1939: 49)re-examined this same specimen and incorrectly re-identified it as  Brycon oligolepis. Géry (1972b: 7–8)cited an unidentified species of  Bryconbased in specimens collected in the Río Curaray basin (not “Cururay”), a tributary of the Río Napo, Ecuador. Judging by the color pattern and locality data from these specimens, this record might refer to  Brycon coxeyi, though anal-fin rays counts are too high (iv, 26–27, according to Géry, 1972: 7). Howes (1982: 21), without presenting any evidence to support such assertion, suggested that  Brycon coxeyimight represent a synonym of  B. melanopterus. However, both species are very distinct from each other and does not even seem to be closely related (see the account for  Brycon melanopterus, below).    Material examined. Typematerial:  ANSP70156( 1, 131.6 mmSL): Ecuador, Província Tungurahua, Hacienda Mascota,  RíoPaztaza, near mouth of RíoTopo, c. 1°25’S, 78°11’W; W. Judson Coxey,  April 1931. Holotypeof  Brycon coxeyiFowler.    Non types. Ecuador: Província Zamora-Chinchipe, Río Zamora/ Río Santiagobasin: MEPNuncataloged ( 1, 148.5 mmSL): Río Nupatakaine,  500 mfrom Destacamento Shaine, at confluence with Río Nangaritza, c. 4°28'S, 78°39'W; R. Barriga,  27 July 1993.   MEPN17484(1, 94.6 mmSL): Quebrada Mayayacu. 2 kmfrom its mouth at RíoNangaritza, 3°59’9’’S, 78°38’27’’W; R. Barriga,  18 July 1993.  MEPNuncataloged ( 1, 192.9 mmSL): RíoNangaritza, near sitio El Dorado, 3°46'S, 78°38'W; R. Barriga& M. Ibarra,  23 Sept 1978. MEPN 17494 (8, 50.4–123.6 mmSL);   ZUEC11325( 1, 106.3 mmSL): Quebrada Yunganza, trib. Río Santiago, 3°24’00’’S, 78°33’25’’W; R. Barriga, M. Ibarra& R. Loma,  27 March 1979.   MEPN11116( 1, 111.2 mmSL): RíoNangaritza; R. Barriga& M. Ibarra,  18 March 1979.   MEPN11114(5, 153.8–182.0 mm SL): RíoMachinaza, 3°45’43’’S, 78°29’53’’W; R. Barriga et al.,  14 Apr 2009.   MEPN11115(6, 152.3– 167.6 mmSL): RíoMachinaza, 3°45’43’’S, 78°29’46’’W; R. Barriga et al.,  13 Sept 2009.   MEPN11102(1, 218.0 mm SL): Zamora-Chinchipe, Río Machinaza Grande  0.5 kmN of Garita de Aurelean, 3°45’34’’S, 78°29’49’’W; R. Barriga, J. Sandoval& J. Sharupi,  19 Nov 2008.   MEPN11103(1, 187.0 mm SL): Zamora-Chinchipe, Río Machinaza, Peñas 3°47'1''S, 78°29'35''W; M. Andrade,  23 Jul 2010.   MEPN11104(2, 149.5– 179.8 mmSL): Zamora-Chinchipe, Río Machinaza,  500 mbelow confluence with Río La Zanza, 3°45’46’’S, 78°29’46’’W; R. Barriga& M. Andrade,  4 Oct 2009.   MEPN11105(3, 168.1– 180.9 mmSL): Zamora-Chinchipe, meeting of Río Zarzawith Río Blanco, 3°45’44’’S, 78°30’39’’W; R. Barriga, J. Sandoval& S. Charupe,  9 Nov 2008.   MEPN11106(4, 130.7– 177.1 mmSL): Zamora-Chinchipe, Río Machinaza, 3°45’21’’S, 78°29’34’’W; R. Barriga et al.,  13 Sept 2009.   MEPN11106(4, 130.7– 177.1 mmSL): Zamora-Chinchipe, Río Machinaza, 3°45’21’’S, 78°29’34’’W; R. Barriga et al.,  13 Sept 2009.   MEPN11108(4, 155.4– 177.1 mmSL): Zamora-Chinchipe, Río Machinaza, 0,5 km above confluence with Río La Zalza, 3°45’45’’S, 78°29’46’’W; R. Barriga& M. Andrade,  4 Nov 2009.   MEPN11109(13, 59.6–150.0 mm SL): Zamora- Chinchipe, Sachavaca Alta, RíoMachinaza, 3°41’13’’S, 78°29’27’’W; M. Andrade,  29–30 Jul 2011.   MEPN11110(4, 79.3–135.8 mmSL): Zamora-Chinchipe,  500 mabove confluence of Río Zorzawith Río Blanco, 3°45’43’’S, 78°30’38’’W; R. Barriga, J. Sharupe& J. Sandoval,  9 Nov 2008.   MEPN11111(7, 55.2–124.9 mmSL): Zamora- Chinchipe, Sacha Vaca Alta, RíoMachinaza, 3°40’53’’S, 78°29’25’’W; M. Andrade,  3 Aug 2011.   MEPN11113(6, 147.3– 191.7 mmSL): Zamora-Chinchipe, Río Machinaza, 3°45’46’’S, 78°29’46’’W; R. Barriga et al.,  13 Sept 2009.   MEPN11119(20, 148.3–225.0 mm SL): Zamora-Chinchipe, Rio Machinaza, Peñas; R. Barriga et al.,  15 Feb 2010.   MEPN11120(2, 158.1– 185.9 mmSL): Zamora-Chinchipe, Río Machinaza, Sachavaca, c. 3°42'S, 78°29'W; M. Andrade,  29 Jul–3 Aug 2011.   MEPN11121(3, 132.8– 158.5 mmSL): Zamora-Chinchipe, Río Blanco, below the bridge, San Antonio, 3°50'49''S, 78°41'41''W; R. Barriga et al.,  18 July 2010.   MEPN11117(4, 39.8–121.2 mmSL): RíoTundayme, trib.  Río Santiago, 3°35’29’’S, 78°28’10’’W; no collector specified,  7 Aug 2010.   MEPN11118(1, 48.0 mm SL): RíoTundayme, trib.  Río Santiago, 3°34’35’’S, 78°28’47’’W; no collector specified,  6 Aug 2010.  Província Morona-Santiago:  MEPN1979(4, 119.1– 132.9 mmSL): Morona-Santiago, Río Chuchumbletza, 3°32’20’’S, 78°30’20’’W; R. Barriga,  13 Sept 1978.  MEPNuncataloged (1, 50.2 mmSL): Quebrada Chuza, 3 kmfrom Tayuza, c. 2°43’S, 78°15’W; R. Barriga,  7 May 1991.   MEPN11117(2, 147.9– 177.1 mmSL): RíoChiguaza, trib.  RíoPaztaza, 2°1'30''S, 77°59'38''W; M. Olalla,  8 Aug 1956.   MEPN5239(22, 27.8– 47.2 mmSL): RioChupianza, 3 kmfrom Chupianza, 2°43’31’’S, 78°18’48’’W; R. Barriga,  6 March 1991.   MEPN5190(1, 56.0 mm SL): RíoPaute at the mouth of río Napiñanza, 2°39’20’’S, 78°23’18’’W; R. Barriga,  7 May 1991.   Peru, Depto. Amazonas, Provincia Condorcanqui:  MUSM6136(10, 52.8–135.8 mmSL); MUSM 6153 (3, 97.3–117.8 mmSL); MUSM 6163 (1, 104.0 mm SL); MUSM 6144 (4, 75.0– 108.2 mmSL); MUSM 6141 ( 1, 145.1 mmSL);   MUSM6123(7, 53.4–103.4 mmSL): RíoComainas, tributary of RíoCenepa, at  Posto de Vigilancia22, c. 3°57’40’’S, 78°24’40’’W; H. Ortega et al.,  24–28 Jul 1994. MUSM 6127 (8, 82.8–135.4 mmSL);   ZUEC6851(2, 96.5–105.4 mmSL): RíoComainas, quebrada 3,  Posto de Vigilancia22, 3°57’40’’S, 78°24’40’’W; H. Ortega et al.,  21 Jul 1994. 1425222670 [199,293,1827,1848] Maranon 20 21 1 Maranon 1425222667 [418,1389,1826,1848] Ecuador Lima 20 21 1 Lima holotype 1425222659 [1399,1435,1826,1848] Ecuador Rio 20 21 1 Rio holotype 1425222658 [199,736,1857,1879] -1.4166666 Topo 1308 -78.183334 20 21 1 Rio 1425222657 ANSP Ecuador Hacienda Mascota 25 26 ANSP 70156 1 Tungurahua holotype 1425222641 1931-04 W. Judson Coxey Ecuador -1.4166666 Rio 1308 -78.183334 25 26 1 Rio holotype 1425222675 1993-07-27 MEPN R. Barriga Ecuador Non 500 -4.4666667 Rio Santiago 1306 -78.65 Rio Zamora 25 26 1 Zamora-Chinchipe holotype 1425222651 1993-07-18 MEPN R. Barriga Ecuador -3.9858334 Rio 21 -78.64083 25 26 MEPN 17484 1 Rio holotype 1425222672 1978-09-23 MEPN R. Barriga & M. Ibarra Ecuador -3.7666667 Rio 1307 -78.63333 25 26 1 Rio holotype 1425222644 1979-03-27 ZUEC R. Barriga & M. Ibarra & R. Loma Ecuador -3.4 Quebrada Yunganza 21 -78.556946 26 27 ZUEC 11325 1 Rio Santiago holotype 1425222662 1979-03-18 MEPN R. Barriga & M. Ibarra Ecuador Rio 26 27 MEPN 11116 1 Rio holotype 1425222654 2009-04-14 MEPN R. Barriga Ecuador -3.7619445 Rio 21 -78.498055 26 27 MEPN 11114 1 Rio holotype 1425222652 2009-09-13 MEPN R. Barriga Ecuador -3.7619445 Rio 21 -78.49611 26 27 MEPN 11115 1 Rio holotype 1425222674 2008-11-19 MEPN R. Barriga & J. Sandoval & J. Sharupi Ecuador -3.7594445 Rio Machinaza Grande 21 -78.49694 26 27 MEPN 11102 1 Zamora-Chinchipe holotype 1425222660 2010-07-23 MEPN M. Andrade Ecuador -3.783611 Penas 21 -78.49305 Rio Machinaza 26 27 MEPN 11103 1 Zamora-Chinchipe holotype 1425222677 2009-10-04 MEPN R. Barriga & M. Andrade Ecuador 500 -3.7627778 Rio La Zanza 21 -78.49611 Rio Machinaza 26 27 MEPN 11104 1 Zamora-Chinchipe holotype 1425222655 2008-11-09 MEPN R. Barriga & J. Sandoval & S. Charupe Ecuador -3.7622223 Rio Blanco 21 -78.51083 Rio Zarza 26 27 MEPN 11105 1 Zamora-Chinchipe holotype 1425222647 2009-09-13 MEPN R. Barriga Ecuador -3.7558334 Rio Machinaza 21 -78.492775 26 27 MEPN 11106 1 Zamora-Chinchipe holotype 1425222648 2009-09-13 MEPN R. Barriga Ecuador -3.7558334 Rio Machinaza 21 -78.492775 26 27 MEPN 11106 1 Zamora-Chinchipe holotype 1425222663 2009-11-04 MEPN R. Barriga & M. Andrade Ecuador -3.7625 Rio La Zalza 21 -78.49611 Rio Machinaza 26 27 MEPN 11108 1 Zamora-Chinchipe holotype 1425222682 2011-07-29 2011-07-30 2011-07-29 MEPN M. Andrade Ecuador -3.6869445 Sachavaca Alta 21 -78.49083 Chinchipe 26 27 MEPN 11109 1 Rio holotype 1425222666 2008-11-09 MEPN R. Barriga & J. Sharupe & J. Sandoval Ecuador 500 -3.7619445 Rio Blanco 21 -78.51056 Rio Zorza 26 27 MEPN 11110 1 Zamora-Chinchipe holotype 1425222661 2011-08-03 MEPN M. Andrade Ecuador -3.6813889 Sacha Vaca Alta 21 -78.49027 Chinchipe 26 27 MEPN 11111 1 Rio holotype 1425222643 2009-09-13 MEPN R. Barriga Ecuador -3.7627778 Rio Machinaza 21 -78.49611 26 27 MEPN 11113 1 Zamora-Chinchipe holotype 1425222665 2010-02-15 MEPN Penas & R. Barriga Ecuador Rio Machinaza 26 27 MEPN 11119 1 Zamora-Chinchipe holotype 1425222697 2011-07-29 2011-08-03 2011-07-29 MEPN M. Andrade Ecuador -3.7 Sachavaca 1307 -78.48333 Rio Machinaza 26 27 MEPN 11120 1 Zamora-Chinchipe holotype 1425222690 2010-07-18 MEPN R. Barriga Ecuador -3.8469443 San Antonio 21 -78.694725 Rio Blanco 26 27 MEPN 11121 1 Zamora-Chinchipe holotype 1425222683 MEPN Ecuador Rio 26 27 MEPN 11117 1 Rio holotype 1425222686 [458,1344,1015,1040] 2010-08-07 Ecuador -3.5913887 Rio Santiago 21 -78.469444 26 27 1 Rio Santiago holotype 1425222681 MEPN Ecuador Rio 26 27 MEPN 11118 1 Rio holotype 1425222669 2010-08-06 Ecuador -3.5763888 Rio Santiago 21 -78.47972 26 27 1 Rio Santiago holotype 1425222696 1978-09-13 MEPN R. Barriga Ecuador -3.538889 Rio Chuchumbletza 21 -78.505554 26 27 MEPN 1979 1 Morona-Santiago holotype 1425222673 1991-05-07 MEPN R. Barriga Ecuador -2.7166667 Tayuza 1307 -78.25 Quebrada Chuza 26 27 1 Morona-Santiago holotype 1425222698 MEPN Ecuador Rio 26 27 MEPN 11117 1 Rio holotype 1425222664 [489,1160,1196,1221] 1956-08-08 M. Olalla Ecuador -2.025 Rio 21 -77.99389 26 27 1 Rio holotype 1425222691 1991-03-06 MEPN R. Barriga Ecuador -2.725278 Chupianza 21 -78.31334 26 27 MEPN 5239 1 Rio holotype 1425222680 1991-05-07 MEPN R. Barriga Ecuador -2.6555557 rio Napinanza 21 -78.38833 26 27 MEPN 5190 1 Rio holotype 1425222679 [151,1228,1303,1328] MUSM Peru Amazonas 26 27 MUSM 6136 1 Amazonas holotype 1425222689 1994-07-24 1994-07-28 1994-07-24 MUSM Posto de Vigilancia & H. Ortega Peru -3.961111 Posto de Vigilancia 21 -78.41111 26 27 MUSM 6123 1 Rio holotype 1425222685 1994-07-21 ZUEC Posto de Vigilancia & H. Ortega Peru -3.961111 Posto de Vigilancia 21 -78.41111 26 27 ZUEC 6851 1 Rio holotype