Received: 10 May 2019 Accepted: 6 July 2019 DOI: 10.1111/jfb.14091 FISH REGULAR PAPER A new species of Hemigrammus (Characiformes: Characidae) from western Amazon River basin, Colombia Juan G. Albornoz-Garzón1 | Alejandro Méndez-López1 | Carlos DoNascimiento1 | 2 Flávio C. T. Lima 1 Colecciones Biológicas, Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Boyacá, Colombia 2 Museu de Zoologia da Universidade Estadual de Campinas “Adão José Cardoso”, São Paulo, Brazil Correspondence Carlos DoNascimiento, Colecciones Biológicas, Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Carrera 8, No. 15-08, Villa de Leyva, Boyacá, Colombia. Email: cdonascimiento@humboldt.org.co Abstract A new species of Hemigrammus is described from the Amazon Basin near Leticia, Departamento Amazonas, Colombia. In common with some congeners and some Hyphessobrycon spp., the new species colour pattern lacks a humeral blotch but has a caudal-peduncle blotch. It can be distinguished from congeners with a similar colour pattern by having: a relatively deep body (30.2%–39.0% LS), 20–24 branched anal-fin rays, 6–8 perforated scales of the lateral line, anal-fin base without a conspicuous black stripe, 3–5 maxillary teeth, maxillary teeth with 1–3 cusps, a thin longitudinal midlateral line and a well-defined oval shaped caudal blotch, extending from caudal peduncle into the lower caudal-fin rays. Comparisons with congeners and with Funding information FAPESP, Grant/Award Number: 2011/51532-7 and 2013/20936-0 Hyphessobrycon species sharing the same general colour pattern are presented. KEYWORDS freshwater fishes, Hemigrammus tridens group, Río Amazonas, Stethaprioninae, taxonomy 1 | I N T RO D U C T I O N Colombia. This new species is similar to some congeners assigned by Géry (1977) to the Hemigrammus tridens group by presenting a colour The genus Hemigrammus Gill 1858 currently includes 61 valid species pattern that includes a conspicuous caudal peduncle blotch but lacks a and is widely distributed throughout the major cis-Andean river basins humeral blotch. A description of the new species is given here. across South America (Lima et al., 2016; Ota et al., 2019; Reia & Benine, 2019). Traditionally, Hemigrammus species have been defined by a combination of non-exclusive characters, first advanced by Durbin (in Eigenmann, 1918): incomplete lateral line, teeth in the premaxilla arranged in two rows, inner premaxillary tooth row composed by five teeth, few or none maxillary teeth and scales extending to the basal region of caudal-fin lobes. Recent phylogenies of Characidae using either morphological (Mirande, 2010), or molecular data (Javonillo et al., 2010; Oliveira 2 | MATERIALS AND METHODS Counts and measurements follow Fink and Weitzman (1974), except for the number of horizontal scale rows below the lateral line (counted to the pelvic-fin insertion) and the number of post-anal scales (median scales between the posterior rim of the anus and the base of the first anal-fin ray) and the addition of three measurements: postorbital head et al., 2011) have failed to demonstrate the monophyly of the genus. length (measured from posterior bony orbital margin to posterior bony More recently, Mirande (2018), using a total-evidence approach, also margin of opercle), dorsal-fin base length and anal-fin base length failed to recover the genus as monophyletic. There are, however, (measured between the bases of the anterior and posteriormost rays some propositions of monophyletic groups within the genus (Lima of each fin). All measurements were taken with digital callipers to the et al., 2016; Ota et al., 2019). nearest 0.1 mm on the left side of specimens. Measurements are Recently, a new species of Hemigrammus was identified during expressed as percentage of standard length (LS), except subunits of searches in the IAvH-P collection, from the western Amazon Basin in the head, which are expressed as percentage of head length (LH). 932 © 2019 The Fisheries Society of the British Isles wileyonlinelibrary.com/journal/jfb J Fish Biol. 2019;95:932–939. ALBORNOZ-GARZÓN ET AL. FISH 933 Counts are followed by their absolute frequency in parentheses. Asterisks indicate counts of the holotype. Circuli and radii were counted on scales from the row immediately dorsal to the lateral line at the vertical through the dorsal-fin origin. Counts of supra-neurals, branchiostegal rays, gill-rakers of first branchial arch, teeth cusps, diminutive posteriormost dentary teeth, unbranched anal-fin rays, procurrent caudal-fin rays, uroneurals and position of pterygiophores were taken from cleared and stained (c&s) specimens prepared according to Taylor and Van Dyke (1985). Vertebrae of the Weberian apparatus were counted as four elements and the compound caudal centrum (PU1 + U1) as a single element. Precaudal vertebral counts include the Weberian apparatus and the vertebrae lacking haemal spines. Caudal vertebral counts include all vertebrae with haemal spines. Catalogue numbers are followed by the total number of specimens and their LS range. The number of c&s specimens is given in parentheses, followed by their respective LS range. Institutional abbreviations follow Sabaj (2019). Morphometric data for Hemigrammus boesemani Géry 1959 are based on the original description by Géry (1959). F I G U R E 2 Hemigrammus amacayacu sp. nov. paratypes, Colombia, Amazonas, Amazon Basin, Leticia, Parque Nacional Natural Amacayacu, unnamed tributary of quebrada Matamatá: a IAvH-P 8285, 24.9 mm LS, b IAvH-P 8283, 24.4 mm LS 3 | RESULTS 3.1 | Hemigrammus amacayacu sp. nov. (Figures 1–3) urn:lsid:zoobank.org:act:F7BBFFA6-312D-46B7-BD68-D7BEE1CD4083. Hemigrammus levis (not Durbin): Arbeláez et al., 2008: 514 (abundance, quebrada Matamatá, Departamento Amazonas, Colombia). 3.1.1 | Holotype IAvH-P 21889, 23.2 mm LS , Colombia, Amazonas, Leticia, Parque Nacional Natural Amacayacu, unnamed tributary of quebrada Matamatá, 03
47 0 53.3” S, 70
15’ 58” W, F. Arbeláez, 20 Mar 2006. 3.1.2 | Paratypes All from Colombia, Amazonas, Amazon Basin, Leticia, Parque Nacional Natural Amacayacu, unnamed tributary of quebrada Matamatá. IAvHP 8282, 12, 17.9–26.3 mm LS, collected with the holotype. IAvH-P F I G U R E 3 Hemigrammus amacayacu sp. nov. IAvH-P 8283, paratype, 24.5 mm LS, medial view of left jaws: a maxilla, b premaxilla, c dentary 8280, 4, 23.1–25.2 mm LS, same locality as holotype, F. Arbeláez, F I G U R E 1 Hemigrammus amacayacu sp. nov. holotype, IAvH-P 21889, 23.2 mm LS, Colombia, Amazonas, Amazon Basin, Leticia, Parque Nacional Natural Amacayacu, unnamed tributary of quebrada Matamatá 17 Mar 2006. IAvH-P 8281, 18, 21.6–26.4 mm LS (1 c&s, 24.0 mm LS), same locality as holotype, F. Arbeláez, 16 Mar 2006. ANSP 206738, 2, 24.0–24.8 mm LS; IAvH-P 8283, 14, 12.0–24.5 mm LS (2 c&s, 934 ALBORNOZ-GARZÓN ET AL. FISH 24.2–24.5 mm LS); ZUEC 17117, 2, 23.1–23.8 mm LS, same locality as dorsal-fin origin. Dorsal profile of head slightly convex from tip of holotype, F. Arbeláez, 18 Mar 2006. IAvH-P 8284, 1, 19.7 mm LS, 03
snout to vertical through anterior margin of orbit; straight from latter 480 23” S, 70
15’ 59.8” W, F. Arbeláez, 15 Mar 2006. IAvH-P 8285, point to end of supraoccipital spine; convex from that point to dorsal- 1, 25.2 mm LS, same locality as IAvH-P 8284, F. Arbeláez, fin origin. Dorsal-fin base slightly convex and posteroventrally 13 Mar 2006. inclined; straight and slanted from dorsal-fin terminus to adipose-fin origin; slightly concave to straight from that point to anteriormost 3.1.3 | Diagnosis dorsal procurrent caudal-fin ray. Ventral profile convex from anterior tip of dentary to pelvic-fin origin, straight and dorsally slanted from Hemigrammus amacayacu sp. nov. differs from most congeners (except from H. boesemani, Hemigrammus brevis Ellis 1911, Hemigrammus mahnerti Uj & Géry 1989, Hemigrammus parana Marinho, Carvalho, Langeani & Tatsumi 2008, Hemigrammus rodwayi Durbin 1909 and Hemigrammus tridens Eigenmann 1907) by having a conspicuous caudal-peduncle blotch and humeral blotch lacking (v. either humeral blotch or blotches present, caudal-peduncle blotch present, or presence of some other colour feature as dark pigmentation on caudal-fin that point to anal-fin origin. Ventral profile along anal-fin base straight and posterodorsally inclined. Ventral profile of caudal peduncle slightly concave. Mouth terminal. Posterior tip of maxilla crossing vertical through anterior margin of pupil. Premaxillary teeth in two rows: outer row with 2* (31) or 3 (9), uni to tricuspid teeth; inner row with 5 (40) tri to pentacuspid teeth. Maxilla with 3 (12), 4* (20) or 5 (7) teeth, lobes, or a conspicuous midlateral dark stripe). Hemigrammus amacayacu differs from H. boesemani by having a deeper body (2.6–3.1 times in LS v. 3.1–3.6 times), anal-fin base without a conspic- T A B L E 1 Morphometric data for holotype and paratypes of Hemigrammus amacayacu sp. nov. (n = 37) uous black stripe (v. anal-fin base with a conspicuous black stripe) and Paratypes caudal blotch roughly oval in shape (v. caudal blotch roughly rhomboiHolotype Mean Range SD 23.2 23.6 19.7–26.4 1.4 Depth at dorsal-fin origin 39.0 34.6 30.2–38.1 2.1 Snout to dorsal-fin origin 53.4 51.5 48.3–54.9 1.8 Snout to pectoral-fin origin 27.4 28.2 25.0–30.8 1.4 Snout to pelvic-fin origin 46.8 46.4 43.9–48.4 1.2 extending from caudal peduncle into lower caudal-fin rays (v. roughly Snout to anal-fin origin 62.6 59.9 56.7–63.3 1.5 triangular or rectangular, restricted to caudal fin, not extending into Caudal-peduncle depth 11.3 10.4 7.7–12.1 1.1 lower caudal-fin rays) and largest dentary teeth tricuspid (v. pen- Caudal-peduncle length 11.9 12.5 9.8–15.2 1.2 tacuspid). Hemigrammus amacayacu can be distinguished from Pectoral-fin length 20.4 19.8 17.1–24.6 1.9 dal in shape). Hemigrammus amacayacu can be distinguished from H. brevis by presenting a broad, roughly oval caudal blotch (v. caudal Standard Length (LS, mm) blotch as an elongated midlateral stripe) and 20–24 branched anal-fin Percentage of LS rays (v. 17–18 branched anal-fin rays). Hemigrammus amacayacu can be distinguished from H. mahnerti by narrow line above anal-fin base absent (v. present), darkened area on sides of body immediately above urogenital opening absent (v. present), maxillary teeth uni to tricuspid (v. pentacuspid), larger dentary teeth tricuspid (v. penta to heptacuspid). Hemigrammus amacayacu can be distinguished from H. parana by having a thin longitudinal midlateral line (v. relatively broad longitudinal midlateral band) and an oval shaped caudal blotch, H. rodwayi by a lower number of lateral-line scales (6–8 v. 9–22), Pelvic-fin length 16.3 16.5 15.0–23.0 1.7 larger dentary teeth tricuspid (v. pentacuspid) and by having a well- Dorsal-fin length 31.9 30.0 26.4–33.3 1.4 defined caudal-peduncle blotch, slightly asymmetrical, with pigmenta- Dorsal-fin base length 14.4 14.3 11.7–16.3 0.8 53.8 54.9 50.8–57.7 1.4 relatively diffuse, roughly symmetrical, dark pigmentation at caudal fin Dorsal-fin origin to caudal-fin origin restricted to middle caudal-fin rays). Hemigrammus amacayacu can be Anal-fin base 30.2 29.9 25.8–33.2 1.9 distinguished from H. tridens by presenting maxillary teeth uni to tri- Anal-fin length 21.8 20.2 17.2–22.6 1.5 37.6 37.3 35.0–39.9 1.1 fin blotch, without an extension along middle caudal-fin rays (v. caudal Posterior margin of eye to dorsal-fin origin peduncle blotch with posterior portion elongated along middle Head length (LH) 27.7 26.6 25.2–28.6 0.9 Postorbital head length 35.7 37.1 32.2–43.1 2.6 Snout length 21.3 23.4 19.9–27.9 1.8 Least interorbital width 34.4 32.6 27.7–36.6 2.3 Upper jaw length 44.2 44.7 41.3–50.8 2.1 Eye diameter 36.1 37.9 35.3–44.4 1.9 tion extending into lower caudal-fin rays (v. caudal-peduncle blotch cuspid (v. pentacuspid), by presenting a roughly oval-shaped caudal- caudal-fin rays) and by the higher number of branched anal-fin rays (20–24 v. 15–17). 3.1.4 | Description Morphometric data of holotype and paratypes are given in Table 1. Body compressed and moderately deep; greatest body depth at Percentage of LH ALBORNOZ-GARZÓN ET AL. FISH 935 anteriormost one or two teeth tricuspid and remaining posterior teeth darkened area present in humeral region due to thinning of hypaxial conic, five from 15 examined specimens with all maxillary teeth conic. musculature. A longitudinal band of chromatophores originating Dentary with 11 (1), 12–13 (1) or 13–15 (1) teeth. Anteriormost four immediately posterior to opercle, becoming more slender posteriorly teeth larger, mostly tricuspid (except one specimen with first three and with few chromatophores reaching caudal peduncle blotch. Lat- teeth on right side and second teeth on left side tetracuspid), eral septum with dark chromatophores forming a thin longitudinal remaining teeth unicuspid, gradually decreasing in size posteriorly midlateral line, beginning slightly anterior to vertical through middle of (Figure 3). Central cusp of multicuspid teeth oriented slightly back- pectoral-fins, becoming more darkly intense posteriorly and extending wards. First gill arch with 1 (1) or 2 (2) gill rakers on hypobranchial, to caudal peduncle blotch. Dark chromatophores aligned with margins 8 (2) or 9 (1) on ceratobranchial, 1 (1) on cartilage between of myomeres, starting slightly anterior to anal-fin origin. Abdominal ceratobranchial and epibranchial and 5 (2) or 7 (1) on epibranchial. region light brown, without chromatophores. Caudal peduncle with Total number of gill rakers on first gill arch 15 to 17. Branchiostegal large, conspicuous oval shaped dark blotch, slightly asymmetrical, rays four (3). occupying most of posterior half of caudal peduncle and extending Scales cycloid, with three to five strongly marked radii, diverging from focus to posterior portion of scale; circuli well-marked anteriorly. Lateral line incomplete, with 6 (13), 7* (8) or 8 (2) pored scales. Longitudinal series of scales including lateral-line scales 30 (1), 31* (4), 32 (5), or 33 (3). Scale rows between dorsal-fin origin and lateral line along middle caudal-fin rays and ventrally to base of lower caudal-fin rays. Remaining fins with dark chromatophores mainly scattered over inter-radial membrane. Dorsal and anal fins with chromatophores concentrated along distal half. Adipose fin without chromatophores. 5 (19). Scale rows between lateral line and pelvic-fin insertion 3 (1) or 4* (20). Scale rows between lateral line and anal-fin insertion three* (15) or four (2). Predorsal scale row with 9 (5), 10* (24) or 11 (6). Circumpeduncular scales 10 (2), 11 (3) or 12* (13). Postanal scales 2* (23) or 3 (5). Caudal fin with one to four scales on basal one-third of each lobe, similar in size to adjacent body scales. Pectoral-fin rays i,9 (1), 10* (22), 11 (13) or 12 (1). Tip of pectoral fin reaching pelvic-fin insertion. Pelvic-fin rays i,6 (2) or 7*(35). Tip of pelvic fin reaching anal-fin origin in some specimens. Dorsal-fin rays 3.1.6 | Distribution Hemigrammus amacayacu sp. nov. is only known from two localities of a short unnamed tributary of the quebrada Matamatá. The quebrada Matamatá is a forest stream draining an upland area of the Parque Nacional Natural Amacayacu and is a direct tributary of the main channel of the Amazon River, departamento Amazonas, Colombia (Figure 4). ii,9* (37); first unbranched ray about one-half length of second unbranched ray; second unbranched and first branched rays longest. Dorsal-fin origin posterior to vertical through pelvic-fin origin; base of last dorsal-fin ray slightly posterior to vertical through anal-fin origin. Anteriormost dorsal-fin pterygiophore inserted posterior to neural spine of 9th (3) vertebra. Anal-fin rays iv* (3), 20 (2), 21 (9), 22* (11), 23 (13) or 24 (2). Anteriormost anal-fin pterygiophore inserted posterior to neural spine of 15th (3) vertebra. Anal fin with anterior lobe formed by last unbranched and five anteriormost branched-fin rays, remaining rays decreasing gradually in size. Adipose fin present. Caudal fin forked, lobes similar in size. Principal caudal-fin rays i,10 + 9,i. 3.1.7 | Ecological notes The localities where the specimens of Hemigrammus amacayacu sp. nov. were found are streams draining pristine terra firma forest. Some physical and chemical variables from these localities are: temperature 24.7–24.9
C, dissolved oxygen 3.7–4.0 mg O2 l−1, pH 5.7–5.9, conductivity 13.5–15.3 μS cm−1, phosphates 0.08 mg l−1 and total bases 0.13–0.15 mg l−1 (Arbeláez et al., 2008). The gut content of one cleared and stained specimen (IAvH-P 8281, 24.0 mm LS) was filled with insect remains, from which several ants and a small Dorsal caudal-fin procurrent rays 8 (2); ventral caudal-fin procurrent cockroach could be identified. Hemigrammus amacayacu was collected rays 6 (2) or 8 (1); one pair of uroneurals. Supraneurals 4 (3). Vertebrae with Characidium sp., Hoplias malabaricus (Bloch 1794), Serrasalmus 33 (3); precaudal vertebrae 13 (1) or 14 (2), caudal vertebrae sp., Cyphocharax pantostictos Vari & Barriga Salazar 1990, Copella vil- 19 (2) or 20 (1). mae Géry 1963, Nannostomus marginatus Eigenmann 1909, Carnegiella strigata (Günther 1864), Gasteropelecus sternicla (L. 1758), Ace- 3.1.5 | Colour in alcohol strorhynchus falcatus (Bloch 1794), Astyanax anterior Eigenmann 1908, Chrysobrycon mojicai Vanegas-Ríos & Urbano-Bonilla, 2017, Charax Overall body colour uniformly light brown. Small, dark chromato- tectifer (Cope 1870), Gymnocorymbus thayeri Eigenmann 1908, Knodus phores densely concentrated on dorsal surface of head, premaxilla, sp., Moenkhausia comma Eigenmann 1908, Moenkhausia oligolepis anterior portion of maxilla and dentary. Infraorbitals 3–6, opercle and (Günther 1864), Tyttocharax cochui (Ladiges 1949) and Bryconops inpai preopercle with scattered dark chromatophores. Ventral portion of Knöppel, Junk & Géry 1968. head clear, except by presence of scattered dark chromatophores restricted to anteroventral region of dentary. Dark chromatophores densely concentrated on predorsal area. Scales from laterodorsal por- 3.1.8 | Sexual dimorphism tion of body with reticulated pattern formed by dark chromatophores No secondary sexually dimorphic characters were observed in exam- concentrated along scale borders. Humeral blotch absent, but ined specimens of Hemigrammus amacayacu sp. nov. However, it 936 ALBORNOZ-GARZÓN ET AL. FISH 72° 71° 70° 69° N 3° Putumayo River F I G U R E 4 Map of southern portion of Colombia showing type locality of Hemigrammus amacayacu sp.nov. The symbol represent more than one lot and locality 4° r ive va Ya ri R Am o Ri r ve Ita qu ai Ri ve r 5° az s na 50 0 50 100 150 200 km should be noted that all specimens were collected in the same month 498–499) as lacking a humeral blotch, actually possess a relatively (March 2006). inconspicuous humeral blotch (Lima et al., 2013, pp. 260–263, for pictures of specimens of H. hyanuary and H. levis). A comparison with 3.1.9 | Etymology these species is relevant given the fact that they are superficially similar to H. amacayacu and indeed, the species was mistakenly identified The specific name amacayacu comes from the Parque Nacional Natu- as H. levis by Arbeláez et al. (2008). In addition to the lack of a humeral ral Amacayacu where the specimens of this species were found. The blotch, H. amacayacu differs from H. hyanuary and H. levis by having a word amacayacu is a quechua word meaning el río de las hamacas (i.e., deeper body (32.7%–39.0% LS v. 24.4%–29.6% LS and 22.9%–29.8% river of the hammocks). The Parque Nacional Natural Amacayacu is an LS, respectively), a higher number of branched anal-fin rays (20–24 v. important national park because represents around 40% of the area 11–14 and 16–18, respectively). Hemigrammus amacayacu further dif- known in Colombia as the Amazonian trapeze, conserving vast areas fers from H. levis by a fewer number of cusps in the largest teeth of of both terra firme and seasonally flooded forests. A noun in inner row of premaxilla and dentary (3–5 v. 7–9) and is distinguished apposition. from H. micropterus by the lower number of perforated scales of the lateral line (6–8 v. 8–12), largest tooth of inner row of premaxilla pen- 4 | DISCUSSION tacuspid (v. heptacuspid), number of maxillary teeth and cusps (3–5 Hemigrammus amacayacu sp. nov. shares with H. boesemani, H. brevis, blotch oval shaped, slightly asymmetrical, with pigmentation exten- H. mahnerti, H. micropterus, H. parana, H. rodwayi and H. tridens the ding into lower caudal-fin rays (v. caudal-peduncle blotch roughly lack of a humeral blotch and the presence of a caudal-peduncle rhomboidal in shape, extending into middle caudal-fin rays). conical or tricuspid teeth v. 1 heptacuspid tooth) and caudal-peduncle blotch. Some of these species (the ones already known at the time, Faria et al. (2018) discussed that some Hyphessobrycon species H. boesemani, H. brevis, H. micropterus, H. rodwayi and H. tridens) were (namely, Hyphessobrycon diastatos Dagosta, Marinho & Camelier included by Géry (1977, pp. 495) in his “He. tridens-group”, which was 2014, Hyphessobrycon juruna Faria, Lima & Bastos 2018, Hyphe- defined by the combination of these characters. There is no evidence ssobrycon negodagua Lima & Gerhard 2001 and Hyphessobrycon par- that this group is a monophyletic unit and as other species groups pro- vellus Ellis 1911) closely resembles He. rodwayi by sharing a similar posed by Géry (1977) for species-rich characiform genera, was a mere colour pattern; i.e., the lack of a humeral blotch and the presence of a convenient tool for aiding species identifications rather than a propo- diffuse caudal peduncle blotch. The traditional diagnosis between sition of a possible monophyletic group. Among the species included Hyphessobrycon Durbin 1908 and Hemigrammus is based on the by Géry (1977) in the “He. tridens-group”, Hemigrammus hyanuary absence of scales on the caudal peduncle (v. its presence in the latter Durbin 1918, Hemigrammus levis Durbin 1908 and Hemigrammus genus), micropterus Meek 1907, although reported both by Durbin Hemigrammus spp. possess just a few scales at the caudal-fin base, (in Eigenmann, 1918, pp. 136–137) and Géry (1977, pp. 495, often missing in preserved specimens. Hemigrammus amacayacu is an unreliable diagnostic character, given that most ALBORNOZ-GARZÓN ET AL. 937 FISH distinguished from Hy. diastatos by presenting caudal peduncle blotch possible that the occurrence of H. amacayacu may be connected with asymmetrical, extending into lower portion of caudal peduncle and the nutrient-richer water from streams draining the Pebas formation. base of lower caudal-fin rays (v. caudal-peduncle blotch lying at the Hemigrammus amacayacu is another recent discovery from the centre of caudal peduncle and base of middle caudal-fin rays), dorsal Leticia area, a relatively well-sampled region in the Amazon Basin for fin not elongated and anal-fin lobe only moderately developed (v. dor- fishes, but which yielded some recently described fish species (Lima sal fin and anal-fin lobe considerably elongated in mature males) and a et al., 2016; Valdesalici, 2016; Vanega-Ríos & Urbano-Bonilla, 2017; higher number of branched anal-fin rays (20–24 v. 15–18). DoNascimiento et al., 2019). These recent discoveries emphasizes the Hemigrammus amacayacu can be distinguished from Hy. juruna, need of additional, broader ichthyological surveys in the Amazonian H. negodagua and H. parvellus by presenting a well-defined caudal Trapeze of Colombia. peduncle blotch (v. caudal-peduncle blotch blurred, with ill-defined contours) and larger dentary teeth with only three cusps (v. larger 5 | ADDITIONAL SPECIMENS EXAMINED dentary teeth with five cusps in H. negodagua and H. parvellus and with 7–9 cusps in H. juruna). It can be additionally distinguished from 5.1 | Hemigrammus boesemani H. negodagua by possessing 20–24 branched anal-fin rays (v. 14–17). The area surrounding Leticia was intensely surveyed ichthyologi- MHNG 2180.009, 10, 25.1–30.4 mm LS, no locality data. MHNG cally by Galvis et al. (2006), who recorded seven Hemigrammus spp. 2180.010, 2, 24.0–25.4 mm LS, Suriname, Coropina Creek. MHNG from the area; i.e., Hemigrammus analis Durbin 1909, Hemigrammus 2180.011, 6, 24.5–30.8 mm LS, Suriname, Marowijne. MHNG bellottii (Steindachner 1882), Hemigrammus cf. erythrozonus Durbin 2180.012, 3, 33.4–36.7 mm LS, no locality data. MHNG 2180.016, 1909, Hemigrammus luelingi Géry 1964, Hemigrammus ocellifer (Steindachner 1882), Hemigrammus pulcher Ladiges 1938 and Hemigrammus schmardae (Steindachner 1882). However, as noticed respectively by Lima et al. (2013), p. 233) and Lima et al. (2016, pp. 124–125), the record of H. analis in Galvis et al. (2006) actually refers to Bryconella pallidifrons (Fowler 1946), while the record of H. ocellifer refers instead to Hemigrammus aguaruna Lima, Correa & 1, 27.1 mm LS, Suriname, Marowijne. MHNG 2180.015, 7, 21.4–25.5 mm LS, Suriname, Paramaribo. MHNG 2181.077, 4, 21.0–28.4 mm LS, French Guyana, Cayenne. MHNG 2181.078, 2, 27.1–28.9 mm LS, French Guyana, Maroni River. MHNG 2616.053, 7, 19.6–25.3 mm LS, French Guyana, Sinnamary River. NRM 70353, 5, 25.7–29.8 mm LS, French Guyana, Kourou. NRM 70684, 1, 26.5 mm LS, French Guyana, Sinnamary Basin. Ota 2016. Additionally, the record of Hemigrammus cf. erythrozonus by Galvis et al. (2006) is a misidentification, the examination of the voucher material at ICN-MHN by us showed it to belong to an unidentified Hemigrammus species. Hemigrammus erythrozonus, the popular glowlight tetra of aquarists, is only known from some tributaries of 5.2 | Hemigrammus brevis ZUEC 13494, 6, 19.0–26.7 mm LS, Brazil, Sergipe, Rio São Francisco basin. the Essequibo River in Guyana and Venezuela (F.C.T.L., unpublished data). Among the sympatric Hemigrammus spp., H. amacayacu is more similar to H. schmardae, with which it shares a conspicuous caudalpeduncle blotch. However, H. schmardae possess a conspicuous, narrow, vertically elongated humeral blotch, while H. amacayacu lacks any hint of the presence of a humeral blotch and for presenting a 5.3 | Hemigrammus hyanuary ZUEC 8879, 28, 24.1–31.0 mm LS, Brazil, Pará, Santarém, Rio Tapajós drainage. ZUEC 13813, 141, 25.4–31.5 mm LS, Brazil, Amazonas, Guajará, Rio Juruá drainage. ZUEC 15314, 20, 31.2–36.5 mm LS, Brazil, Amazonas, Tefé, Rio Tefé. lower number of branched anal-fin rays (15–20 v. 20–24, respectively), among other differences. Galvis et al. (2006) probably did not have access to any specimen 5.4 | Hemigrammus levis of Hemigrammus amacayacu because they have examined relatively ZUEC 7971, 30, 39.1–47.8 mm LS, Brazil, Amazonas, Barcelos, Rio little material from the Parque Nacional Natural Amacayacu. Most of Negro drainage. ZUEC 8310, 136, 24.6–41.3 mm LS, Brazil, Amazonas, the material used in their study from upland streams was collected in Iranduba, lago Janauari. ZUEC 11813, 23, 23.5–47.4 mm LS, Brazil, the quebradas crossing the road to Taracua, to the north-west of the Pará, town of Leticia (Galvis et al., 2006, p. 74). The quebrada Matamatá 62, 25.0–40.3 mm LS, Brazil, Amapá, Calçoene, Rio Amapá Grande. drains the Pebas formation, while streams to the east sampled by ZUEC 16746, 166, 23.4–35.4 mm LS, Brazil, Amapá, Tartarugalzinho, Galvis et al. (2006) drain the Terciario Superior Amazónico formation, Rio Araguari drainage. Santarém, Rio Arapiuns drainage. ZUEC 16529, which presents more leached and coarser soils than soils from the Pebas formation (Arbeláez et al., 2008). Arbeláez et al. (2008) reported considerable differences in fish composition between streams draining 5.5 | Hemigrammus mahnerti the Pebas formation v. streams draining the Terciario Superior ZUEC 3216, 1, 17.0 mm LS; ZUEC 5476, 1, 20.4 mm LS, Brazil, Mato Amazónico formation, with the first presenting a considerable larger Grosso, Poconé, Rio Bento Gomes. ZUEC 6656, 48, 15.1–19.0 mm LS, biomass and species richness when compared with the latter. It is Brazil, Mato Grosso, Poconé, Rio Piraputanga. 938 ALBORNOZ-GARZÓN ET AL. FISH 5.6 | Hemigrammus micropterus FUNDING INF ORMATI ON IAvH-P 9509, 592, 12.5–34.2 mm LS, Colombia, Casanare, Aguazul, Financial support was given by FAPESP, project “Systematics of the laguna El Tinije, río Meta drainage. IAvH-P 20232, 1, 33.6 mm LS, tetras (genera Hemigrammus, Hyphessobrycon, Thayeria, Parapristella Colombia, Vichada, Cumaribo, caño Fruta, Caño Matavén drainage, and Bryconella), with emphasis on the species from cis-Andean South Orinoco River basin. IAvH-P 20250, 72, 8.7–34.5 mm LS, Colombia, America”, grants # 2011/51532-7 and 2013/20936-0. Meta, Puerto Gaitán, laguna Mururito, río Meta drainage. ZUEC 16918, 50, 20.3–28.1 mm LS, Colombia, Meta, río Meta drainage, OriOR CID noco River basin. Carlos DoNascimiento https://orcid.org/0000-0002-8680-7942 5.7 | Hemigrammus parana ZUEC 7505, 10, 24.8–29.8 mm LS, Brazil, Minas Gerais, Conceição das Alagoas, Rio Grande drainage. 5.8 | Hemigrammus rodwayi ZUEC 12885, 85, 20.2–34.0 mm LS, Brazil, Pará, Primavera, Rio Primavera drainage. ZUEC 14437, 14, 26.1–40.5 mm LS, Brazil, Pará, Igarapé Açú, Rio Maracanã drainage. ZUEC 16062, 100, 22.2–32.3 mm LS, Brazil, Amapá, Porto Grande, Rio Araguari drainage. 5.9 | Hemigrammus schmardae CIACOL 2050, 13, 18.5–28.5 mm LS, Colombia, Amazonas, Amazon River basin. IAvH-P 9024, 22, 24.7–30.1 mm LS, Colombia, Amazonas, Leticia, quebrada Sufragio, Amazon River basin. ZUEC 15091, 38, 21.0–31.1 mm LS, Brazil, Amazonas, Tabatinga, Rio Solimões drainage. ZUEC 15149, 61, 26.0–33.1 mm LS, Brazil, Amazonas, Tefé, Rio Tefé drainage. ZUEC 16989, 733, 12.7–26.6 mm LS, Brazil, Amazonas, Tabatinga, Rio Solimões drainage. 5.10 | Hemigrammus tridens ZUEC 6683, 1, 17.8 mm LS, Brazil, Mato Grosso, Poconé, Rio Piraputanga. UMMZ 206594, 147, 15.2–28.1 mm LS, Paraguay, San Pedro, Río Aguaray-Mi. ACKNOWLEDGEMEN TS We thank Jorge E. García-Melo and Eduardo G. Baena for the photographs of the paratypes (Figures 2a and 2b, respectively). The fourth author (FCTL) is grateful to A. Acosta-Santos and E. Agudelo Cordoba (CIACOL), J. I. Mojica, H. D. Agudelo-Zamora and G. Serrato (ICNMHN), D. W. Nelson (UMMZ), M. H. Sabaj, J. G. Lundberg and M. Arce Hernández (ANSP), R. Covain (MHNG), S. O. Kullander, B. Delling and E. Ahlander (NRM) for access to comparative material used in this study. RE FE RE NCE S Arbeláez, F., Duivenvoorden, J. F., & Maldonado-Ocampo, J. A. (2008). Geological differentiation explains diversity and composition of fish communities in upland streams in the southern Amazon of Colombia. 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