Brachyhypopomus bombilla Loureiro & Silva, 2006

Crampton, William G. R., de Santana, Carlos D., Waddell, Joseph C. & Lovejoy, Nathan R., 2016, A taxonomic revision of the Neotropical electric fish genus Brachyhypopomus (Ostariophysi: Gymnotiformes: Hypopomidae), with descriptions of 15 new species, Neotropical Ichthyology 14 (4) : -

Brachyhypopomus bombilla Loureiro & Silva, 2006

( Fig. 15; Tables 2-5, 9)

Rhamphichthys brevirostris . - Boulenger, 1896: 38 ( Brazil, Mato Grosso, Descalvados, list of fishes collected in the Paraguay river by C. Ternetz, record corresponds to BMNH 1895.5.17.265-268; identified as B. bombilla [part, n = 3], B. gauderio [part, n = 1]).

Hypopomidae indet. -Hagedorn & Keller, 1996: 491, fig. 2b, drawing of adult specimen ( Peru, Pakitza, Manu Reserve; diversity of gymnotiform species and EODs).

Brachyhypopomus electropomus . - Sullivan, 1997: 122, 322, fig. 54, line drawing of MZUSP 45032, Paraguay drainage (description in unpublished thesis with disclaimer stating nomenclatural acts therein not available, a nomen nudum; name therein refers also to B. menezesi and B. regani ).

Brachyhypopomus sp. N. - Crampton & Albert, 2006: 672, fig. 23.8, position in phylogenetic tree; 681, notes on EODs (species and EOD diversity).

Brachyhypopomus bombilla Loureiro & Silva, 2006: 666 , fig. 1, black and white photograph of holotype (original description, type locality – Uruguay, Rocha Department, Cuatro Palmas stream, Patos-Mirim dr.). - Almirón et al., 2010: 573, fig. 2, photograph of live individual ( Argentina, Entre Ríos, Pre-Delta National Park, Paraná dr., listing of Brachyhypopomus ). -Crampton, 2011: 176, table 10.2, species list; 179, figs. 10.2- 10.3, phylogeny, geographical and ecological distributions (gymnotiform biology). -Giora et al., 2011 (reproductive and trophic ecology). - Koerber, 2011: 7 ( Argentina, species list). - Carvalho, 2013: 177, fig. 37B color photograph of head showing position of accessory electric organ; 472, fig. 1, color photographs of head; 181-185, figs. 41-43, position in phylogeny (phylogenetic systematics of Rhamphichthyoidea ). -Mirande & Koerber, 2015: 48 ( Argentina, catalog of fishes). - Crampton et al., 2016: 1-66, table 1, 3-4, figs. 1-7, 18-20 (phylogeny, biogeography and ecology of Brachyhypopomus ).

Diagnosis. Brachyhypopomus bombilla is diagnosed from congeners by the following combination of characters: presence of accessory electric organ over the opercular region, vs. absence in all congeners except B. menezesi and B. regani ; dorsal surface uniformly speckled with small brown chromatophores on a pale background, vs. presence of large dark blotches against a lighter background in B. menezesi and B. regani .

Description. Head and body shape, and pigmentation illustrated in Fig. 15 and in the original description. Meristic and morphometric data for examined specimens are presented in Tables 2-5 and 9. Body shallow to moderate in depth. Head short to moderate in length and shallow to moderate in depth. Dorsal profile of head approximately straight from occiput to snout, concave above eye, ventral profile of head straight, snout truncated to rounded. Eye small to moderate in size. Upper jaw with slight sigmoidal angle between premaxillary and maxillary portions in lateral view. Accessory electric organ (AEO) over operculum present; skin over AEO with similar pattern of chromatophores to surrounding area, obscuring margin of AEO and rendering AEO difficult to see in most specimens. Gill filaments on first gill arch 29-34 (median 32, n = 6). Pectoral fin narrow to moderate in breadth, pectoral-fin rays 14-17 (mode of 15 in populations from Patos-Mirim lagoon system and upper Madeira, mode of 14 in populations from Uruguay drainages). Precaudal vertebrae 15-18 (mode of 18 from Patos-Mirim, 16 from río Paraguay drainages of Brazil and Paraguay, and 17 from upper Madeira), including 1-2 transitional vertebrae. Anal-fin origin slightly (<0.25 HL distance) anterior or posterior to, or near, tip of pectoral fin. Anal-fin rays 155-193 (median of 183 in populations from Patos-Mirim, 171 from río Paraguay, and 176 from upper Madeira). Dorsal rami of recurrent branch of anterior lateral line nerve not visible. Middorsal region of body scaled. Rows of scales above lateral line 5-7 (mode 6). Lateral line continuous. Multiple groove-like depigmented epidermal canals, some with dark borders, form parallel striations mostly dorsal to lateral line in posterior portion of body, beginning ca. two thirds of LEA; striations reaching 1-4 scales (including lateral line series) dorsally, and occasionally (with much lower density) 1-2 scales ventrally. Epidermal canals absent in anterior portion of body, never form a pair of long irregular lines either side of dorsal surface in dorsal portion of flank, and do not form isolated horizontal wavy line midway from lateral line to dorsal midline in midbody or anterior portion of body. Three bilateral horizontal columns of electrocytes at anal-fin terminus, and at mid-point between anal-fin terminus and tip of caudal filament in immature, mature female, and mature male specimens. Electrocytes of hypaxial electric organ extend anterior to anal and urogenital pores and appear to extend into opercular area to form paired accessory electric organ. Caudal filament short to moderate in length.

Coloration. ( Fig. 15). Background pale straw. Dorsal region with more or less homogenous speckling of small brown chromatophores, declining in density ventrally. Dorsal region without pale stripe extending along midline from occiput to base of caudal filament. Flank ventral to lateral line only sparsely pigmented with irregular small clusters of brown stellate chromatophores. Caudal filament darker. Hypaxial electric organ conspicuous in posterior half to third of body. Head with evenly scattered dark chromatophores, darker dorsally. Eye usually with suborbital patch of subcutaneous pigmentation, of varying darkness and prominence. Region around lips usually depigmented. Presence of chromatophores over entire operculum disrupts borders of AEO, rendering it difficult to see in many specimens. This may account for why the authors of the original description failed to report the presence of this important character (although we inspected the holotype and found the AEO conspicuous). Pectoral and anal-fin membranes hyaline. Pectoral-fin rays hyaline. Anal-fin rays with uniform light brown pigmentation near bases, hyaline distally. Color in life faintly yellow in individuals from sediment-rich waters, with opercular region usually rosy due to underlying gills.

Size. Small to moderate adult size, largest specimen examined 161 mm TL, 140 mm LEA (n = 537). Largest male specimen examined 161 mm TL, 140 mm LEA (n = 13). Largest female specimen examined 147 mm TL, 124 mm LEA (n = 43).

Sexual dimorphism. No known secondary sexual dimorphism.

Geographic distribution. Argentina, Bolivia, Brazil, Paraguay, Peru, and Uruguay ( Fig. 16). Known from the Patos-Mirim lagoon system, Uruguay drainage, Paraná below the former Guaíra Falls, lower and upper Paraguay, and upper Madeira (Amazon drainage) above its major series of rapids beginning at the Salto Santo Antônio (near Porto Velho, Rondônia, Brazil).

Population variation: We found complete overlap in the range of meristics ( Tables 2-5) and morphometric proportions ( Table 9), and observed similar pigmentation among populations of B. bombilla in the Patos-Mirim system, Paraguay dr., and upper rio Madeira. These observations, in combination with molecular data ( Crampton et al., 2016), support the hypothesis that all populations we have assigned to B. bombilla are members of a single geographically widespread species, which is morphologically and genetically distinct from all congeners (although we were unable to obtain specimens for genetic analyses from the type locality in the Laguna Merín drainage).

adapted to a wide range of temperature, conductivity, and other parameters. In high latitudes it exhibits a period of dormancy in the austral winter, which is apparently unique among congeners (Giora et al., 2011). At this time it burrows in muddy substrates, slows its EOD, ceases to forage, and exhibits arrested gonadal development (Giora et al., 2011). Stomach contents in populations from southern Brazil (Giora et al., 2011) and from the upper Madeira (WGRC pers. obs.) comprise mostly aquatic insect larvae, microcrustacea, and other small aquatic invertebrates.

Co-occurring congeners: In the upper Madeira B. bombilla occurs in geographical sympatry and ecological syntopy with B. arrayae , B. brevirostris , B. pinnicaudatus , and B. walteri . It also exhibits an allotopic distribution with B. alberti and B. sullivani . In the upper portions of the Paraguay, B. bombilla co-occurs in sympatry and syntopy with B. brevirostris , B. draco , B. gauderio , and B. walteri . In the lower Paraná, B. bombilla co-occurs in sympatry and syntopy with B. draco , B. gauderio and B. walteri . In the lower portions of the Paraguay, and in the Patos-Mirim system, B. bombilla co-occurs in sympatry and syntopy with B. draco and B. gauderio .

Local names. Argentina, Paraguay: morenita; Bolivia: Cuchillo; Brazil: tuvira, moreninha (southern Brazil); Peru: macana; Uruguay: morenita, limpiavidrio ( Nion et al., 2002).

Ecological notes. Brachyhypopomus bombilla occurs in floodplains, swamps and along the edges of slow-moving creeks and rivers – always in association with dense root mats of floating aquatic vegetation ( Almirón et al., 2010; Giora et al., 2011; Koerber, 2011; Loureiro & Silva, 2006). Giora et al. (2011) document the reproductive ecology of B. bombilla from southern Brazil. Breeding occurs during the austral spring and summer. In the upper Madeira, WGRC collected B. bombilla during the low water period from floating aquatic vegetation in floodplain pools and channelmargins of the río Beni floodplain (conductivity ca. 150 μScm-1, pH 7.3-7.6, dissolved oxygen 5-6 mgl-1, temperature ca. 28-32°C). Hagedorn & Keller (1996) report B. bombilla (identifications confirmed from voucher specimens at the USNM) from creeks of upper Madeira drainages in Peru. In the Paraná-Paraguay drainage, collection records indicate that B. bombilla is found in a variety of habitats, including swamps, riverine floodplains and slow-flowing creeks. Throughout its range B. bombilla appears to be Material examined. 527 specimens. Argentina. Corrientes. AI 267, 10, 91-132 mm, Esteros del Iberá, Laguna Galarza, Paraná dr., 28°04′56″S, 056°42′04″W. MACN-ict 9457, 2, 91-96 mm, Iberá wetlands, río Corrientes , Paraná dr., 28°42′45″S, 058°06′40″W . Entre Ríos. MACN-ict 9458, 1, 65 mm, Arroyo Las Mangas, Pre-Delta National Park , Paraná dr., 32°07′59″S, 060°39′37″W . Bolivia. Beni (localities are from the rio Madeira dr., Amazonas dr.). CAS 81632, 1, 100 mm, Reyes, 24 mi. NE Rurrenabaque, río Beni , 14°17′42″S, 067°20′27″W . CBF 10271 , 1 , immature, 93 mm , UMSS 7035 , 1 , female, 106 mm, mun. Riberalta, stream nr. Hormiga, on Riberalta-Guayaramerín rd., affl. río Beni , 11°01′34″S, 065°52′58″W . CBF 10272 , 1 , immature, 87 mm , CBF 10273 , 1 , immature ( CS), 91 mm , CBF 10274 , 1 , male ( CS), 96 mm , CBF 10275 , 1 , immature, 105 mm , CBF 10276 , 1 , female, 101 mm , UF 177344 , 2 , immature, 82-90 mm , UMSS 7036 , 1 , female, 99 mm , UMSS 7037 , 1 , immature, 90 mm , UMSS 7038 , 1 , female, 90 mm , UMSS 7039 , 1 , immature, 87 mm , UMSS 7040 , 1 , female, 83 mm, mun. Riberalta, arroyo Tres Cuchillos, río Beni floodplain, 10°51′44″S, 066°04′50″W . INHS 37243, 3, 74-87 mm, 41km E Estación Biológica del Beni, on rd. to San Ignacio [de Moxos], río Chevejecure, affl. río Apere , affl. río Mamoré , 14°52′S, 065°57′W . INHS 50011, 2, 65-85 mm, 46 km E Estación Ecológica del Beni on rd. to San Ignacio [de Moxos], río Moseruna, affl. río Apere , affl. río Mamoré , ca. 14°55′S, 065°54′W . UF 81921 , 2 , 102-104 mm, Bella Vista, rio Blanco, affl. rio Guaporé , 13°16′S, 063°41′W . UMMZ 66420, 1, 89 mm, lago Rogoagua, lake and nearby lagoons, río Beni floodplain, ca. 13°58′S, 066°58′W . La Paz. CAS 72232, 1, 124 mm, Ixiamas, 60 mi. NW Rurrenbaque, río Beni , ca. 13°45′S, 068°03′W . Pando. FMNH 106643, 1, 78 mm, FMNH 106644, 28, 57-75 mm, FMNH 106645, 4, 57-72 mm, río Manuripi, ca. 20 km upstream Puerto Rico, río Manuripi, affl. río Orthon, affl. río Madre de Dios, affl. río Beni , ca. 11°09′06″S, 067°33′41″W . FMNH 106646, 11, 53-129 mm, FMNH 106647, 17, 41-68 mm, río Manuripi, ca. 13 km upstream Puerto Rico, río Manuripi, affl. río Orthon, affl. río Madre de Dios, affl. río Beni , ca. 11°13′S, 067°34′W . Santa Cruz. UF 82190 , 3 , 65-79 mm, 71 km N San José de Chiquitos, río San Diablo, affl. río Itonama, affl. río San Martín, affl. río Guaporé , affl. río Mamoré , 17°18′S, 060°35′W . UF 82309 , 1 , 65 mm, creek 22 km W San Javierito and 49 km W San Ignacio, río Paragua, affl. río Guaporé , affl. río Mamoré , ca. 16°21′S, 061°05′W . Brazil. Acre. MCP 41034 , 2 , 66-107 mm, Fazenda Uberaba, ca. 6 km hwy. BR-317, rio Iná, affl. rio Xipamanu, affl. rio Abunã , 10°44′13″S, 068°11′16″W . Mato Grosso (localities in rio Paraguai dr., Paraná dr., unless otherwise stated). ANSP 53894, 2, 57-78 mm, BMNH 1895.5.17.265-268 (part), 3, 71-107 mm, mun. Cácares, Descalvados, 16°43′59″S, 057°44′54″W . BMNH 1900.4.14.91-94 (part), 1, 72 mm, mun. Carandàzinho, rio Cuiabá , ca. 16°27′S, 056°08′W . FMNH 54551, 10, 54-127 mm, MCP 15629 , 9 (9 immature, 76-110 mm), mun. Cáceres, nr. Cáceres, ca. 16°04′S, 057°39′W . FMNH 54553, 1, nr. Conceição [Porto Conceição], ca. 15°57′S, 060°10′W. FMNH 70397, 1, 88 mm, nr. Conceição, ca. 17°09′S, 057°23′W. MCP 15646 , 1 , female, 69 mm, Porto Esperidião, arroio Zé do Cacete , km 70 on Cáceres-Porto Esperidião hwy. BR-174, 15°49′S, 058°17′W . MCP 41030 , 1 , 80 mm, mun. Vila Bela da Santíssima Trindade, stream 10 km rd. between Pontes e Lacerda and Vila Bela da Santíssima Trindade, affl. rio Guaporé, affl. rio Mamoré , affl. rio Madeira , Amazonas dr., 15°13′58″S, 059°24′30″W . MNRJ 12242 , 6 , 93-115 mm, mun. Cáceres, Fazenda Pantanalzinho, district of Porto Espiridião, Córrego Carrapato, rio Jauru , ca. 15°51′S, 058°27′W . MZUSP 19243 , 1 , male, 98 mm, Poconé, ca. 16°14′S, 056°37′W . MZUSP 25137 , 3 (1 immature, 94 mm, 2 female, 87-97 mm), mun. Cáceres, Ilha da Taiamã [Estação Ecológica Taiamã], rio Paraguai , 16°53′S, 057°27′W [coordinates from center of island]. MZUSP 25257 , 1 , 93 mm, Fazenda Santo Antônio do Paraíso, rio Piquiri-Itiquira , 17°12′S, 054°09′W . MZUSP 43681 , 26 , MZUSP 45032 , 1, 101 mm, mun. Cáceres, Descalvados, ca. 16°44′S, 057°44′W . MZUSP 44261 , 9 , mun. Cácares, Cácares and vicinity, ca. 16°04′S, 057°40′W . USNM 326426 , 10 , 61-106 mm, nr. Cáceres, ca. 16°04′S, 057°39′W . ZUEC 4291, 1, 76 mm, ZUEC 4329, 1, 72 mm, mun. Poconé, rio Piraputanga, affl. rio Vermelho, affl. rio Cuiabá , ca. 16°20′S, 053°56′W . ZUEC 4351, 1, 94 mm, ZUEC 5449, 3, 73-90 mm, mun. Poconé, km 19, Transpantaneira hwy., rio Bentes Gomes , ca. 16°33′S, 56°42′W . Mato Grosso do Sul (localities in rio Paraguai dr., Paraná dr.). FMNH 54553, 1, 105 mm, FMNH 108540, 16, 38- 88 mm, Corumbá, rio Vermelho , 19°37′13″S, 56°57′16″W . FMNH 108539, 3, 70-80 mm, rio Vermelho at Fazenda Xarães, 19°36′44″S, 56°55′25″W . MUSM 16955 , 1 , immature, 99 mm, Taquari, rio Bonito , 18°16′23″S, 053°27′27″W . MUSM 17021 , 4 , 71-81 mm, Brazo de Santana Sofia , 19°35′59″S, 056°20′47″W . MZUSP 36341 , 2 , Ladário, nr. Corumbá, 19°00′S, 057°36′W . MZUSP 48334 , 3 , female, 69-80 mm, Pantanal do Paiaguás, Baía da Sede , 18°30′S, 054°45′W . MZUSP 59406 , 1 , female, 87 mm, Córrego da Furna , riacho Água Clara, 18°15′45″S, 053°27′59″W . MZUSP 59411 , 1 , male, 97 mm, Corumbá, rio Negro on rd. between Nhecolândia and hwy. BR-262, 19°17′16″S, 057°03′39″W . MZUSP 59412 , 1 , immature, 49 mm, Corumbá,rioAbobral, 19°27′37″S, 057°01′14″W . MZUSP 59441 , 33 , (15 measured: 2, immature, 61-78 mm, 13 female, 61-100 mm), Corumbá, rio Miranda, bay on left bank, rio Vermelho , 19°37′22″S, 056°57′27″W . MZUSP 59444 , 4 (1 immature, 99 mm, 2 female, 72-93 mm, 1 male, 100 mm), Aquidauana, Fazenda Taboco, dead end of rio Taboco , 20°04′38″S, 055°38′51″W . MZUSP 59445 , 2 , female, 68-79 mm, rio Verde de Mato Grosso, Anhuma, on rd., rio Negro , 19°09′57″S, 055°17′56″W . MZUSP 60051 (part), 2, immature, 76-99 mm, rio Verde de Mato Grosso, rio Negro , 19°18′17″S, 055°10′20″W . MZUSP 95081 , 12 (10 immature, 52-83 mm, 2 female, 89-102 mm), mun. Barão de Melgaço, rio Mutum, between Vila de Mimoso and Joselândia , Pantanal de Paiaguás , 16°19′30″S, 055°49′59″W . Rio Grande do Sul. BMNH 1889.8.24.8, 1, 124 mm, mun. Camaquã, Camaquã, arroio Duro, affl. rio Camaquã , Patos-Mirim dr., ca. 30°51′S, 051°48′W . BMNH 1891.3.16.103-105, 3, 89-140 mm, “ Rio Grande do Sul ”, drainage unknown, no coordinates. MCP 15326 , 1 , female, 73 mm, canals along hwy. BR-290 between 1 and 6 km junction with hwy. BR-116, rio Guaíba , Patos-Mirim dr., between 30°02′39″S, 051°21′20″W and 30°02′48″S, 051°23′27″W . MCP 19545 , 1 , male, 149 mm, 13.7 km, from São Gabriel, rio Vacacaí, affl. rio Jacuí , affl. rio Guaíba , Patos-Mirim dr., 30°27′18″S, 054°22′26″W . MCP 19807 , 4 (1 immature, 88 mm, 2 female, 81-98 mm, 1 male, 117 mm), river at Barra do Ribeiro, affl. rio Guaíba , Patos-Mirim dr., 30°17′03″S, 051°18′04″W . MCP 19847 (part), 6 (3 immature, 127-140 mm [2 CS], 2 female, 100-147 mm, 1 male, 156 mm), bridge over Banhado do Inhatium, hwy. BR- 290, 21 km São Gabriel, affl. rio Ibicuí , Uruguay dr., 30°15′43″S, 054°31′33″W . MCP 19869 , 1 , male, 161 mm , MCP 45355 , 3 , 51-65 mm, bridge on São Gabriel-Tiaraju hwy., affl. rio Vacacaí, affl. rio Jacuí , affl. rio Guaíba , Patos-Mirim dr., 30°17′29″S, 054°20′18″W . UFRGS 6495 , 1 , male, 131 mm , UFGRS 6496 , 1 , male, 133 mm , UFRGS 6498 , 1 , male, 136 mm, mun. Rosário do Sul, stream at km 82 on hwy. BR-290, affl. rio Santa Maria, affl. rio Ibicuí , Uruguay dr., 30°12′42″S, 055°03′17″W . UFRGS 6741 , 1, 138 mm , UFRGS 6742 , 1 , 63 mm , UFRGS 6743 , 1 , 65 mm , UFRGS 6744 , 1 , 59 mm , UFRGS 6745 , 1 , 88 mm , UFRGS 6746 , 1 , UFRGS 6747 , 1 , 49 mm , UFRGS 9273 , 18 , 57-126 mm , UFRGS 9282 , 15 , 58-99 mm , UFRGS 9284 , 17 , 61-132 mm, Rosário do Sul, Sanga do Jacaré, stream at BR-290, affl. rio Ibicuí , Uruguay dr., 30°12′42″S, 055°03′19″W . UFRGS 10561 , 2 , 94-108 mm, mun. Rosário do Sul, arroio do Jacaré, affl. rio Ibicuí , Uruguay dr., 30°12′43″S, 055°03′18″W . Rondônia (localities from rio Madeira, dr., Amazonas dr.). INPA 19944 , 5 , 71- 97 mm, rio Mamoré , upstream Surprêsa, ca. 11°53′S, 064°59′W . MCP 41031 (part), 2, 87-88 mm, mun. Nova Mamoré, rio Ribeirão at hwy. BR-425, ca. 62 km S hwy. BR-364, affl. rio Guaporé, affl. rio Mamoré , 10°13′55″S, 065°16′55″W . UFRO-I 6478, 6, 61-76 mm, rio Madeira, no specific locality, no coordinates. UFRO-I 6507, 2, 112- 112 mm, UFRO-I uncat. CAU2010010701, 1, 85 mm, mouth of the rio Cautário, affl. rio Guaporé , affl. rio Mamoré , 12°10′51″S, 064°34′00″W . Paraguay (localities from río Paraguay dr., Paraná dr.). Alto Paraguay. FMNH 108057, 1, 49 mm , MZUSP 54330 , 1 , 86 mm, Estancia Voluntad, at Puerto Voluntad, 20°42′S, 057°57′W . FMNH 108058, 1, 103 mm, rio Negro , 20°09′S, 058°10′W . Caaguazú. ANSP 175179, 1, 97 mm, stream crossing rd. N Coronel Oviedo, 24°52′59″S, 056°20′10″W . Canindeyú. NRM 32529, 1, 116 mm, río Jejui-Guazú , 76 km, rd. to Curuguaty, 24°19′42″S, 055°05′13″W . NRM 32530, 1, female, 111 mm, stream nr. rd. 3km before Curuguaty, 24°29′29″S, 055°39′12″W. NRM 32531, 1, male, 115 mm, small stream, ca. 12 km on Curuguaty- Mbutuy rd. , 24°29′47″S, 055°47′08″W . UMMZ 206285, 2, male, 115-142 mm, Arroyo Carimbatay, ca. 15.6 km WSW Curuguaty, Arroyo Curuguaty, río Jejui-Guazú , 24°31′06″S, 056°43′30″W . UMMZ 206470, 1, 84 mm, small stream, affl. río Jegui-Guazú , ca. 79 km by dirt rd. ENE turnoff to Curuguaty, nr. Santo Domingo, ca. 24°21′S, 055°06′W [coordinates for Santo Domingo]. UMMZ 206536, 1, 96 mm, small stream, affl. Arroyo Curuguaty, ca. 5.3 km NNW Curuguaty, río Jejui-Guazu , 24°25′06″S, 055°40′48″W . Central. BMNH 1935.6.4.370-372, 2, 75-85 mm, riacho Caracara, W Asunción, ca. 25°18′S, 057°38′W. Concepción. MZUSP 54331 , 1 , female, 86 mm, riacho Blandengue, nr. confl. with río Apa , 22°14′S, 057°21′W . Guairá. NRM 41902, 1, male, 118 mm, stream crossing rd. from Mauricio J. Troche to Independencia, 5 km S río Tebicuary bridge, 25°40′59″S, 056°18′40″W . Itapúa. NRM 42291, 1, female, 136 mm, NRM 42294, 1, 75 mm, Bella Vista, Arroyo Posore at 12 km before Pirapó on rd. from Obligado, 26°54′34″S, 055°36′37″W . Paraguarí. NRM 32533, 1, 98 mm, lago Ypoá, Estancia Ypoá, 25°56′43″S, 057°27′03″W . San Pedro. NRM 23195, 1, female, 122 mm, Estancia Laguna Blanca, Arroyo Mborebi from outlet from Laguna Blanca, 23°48′48″S, 056°17′44″W . Peru. Madre de Dios (localities from río Madre de Dios dr., affl. río Beni, affl. río Madeira , Amazonas dr.). MUSM 1388 , 1, 132 mm , MUSM 1428 , 1, 126 mm , MUSM 3947 , 2 , 118-122 mm , MUSM 4524 , 4 , 120-136 mm , MUSM 4526 , 2 , 113-118 mm , MUSM 5068 , 1, 109 mm, Parque Nacional ( PN) Manú, Pakitza Biological Station, nr. Aguajal , ca. 12°52′S, 069°16′W . MUSM 2553 , 1, 105 mm, PN Manú, Pakitza Biological Station , nr. Aguajal, quebrada Fortaleza, ca. 11°45′S, 070°17′W . MUSM 4520 , 8 , 84-122 mm , MUSM 14015 , 73 , 56-133 mm, PN Manú, Pakitza Biological Station , nr. Aguajal, quebrada Carpintero, ca. 12°52′S, 069°16′W . MUSM 4564 , 1, 129 mm , MUSM 14016 , 11 , 79-140 mm, PN Manú, Pakitza Biological Station , nr. Aguajal, quebrada Trompetero, ca. 12°52′S, 069°16′W . MUSM 21691 , 1 , Huitoto, no coordinates . MUSM 25041 , 1 , Planchón las Piedras, stream at km. 50, río Manuripe , no coordinates . MUSM 37016 , 2 , 120-155 mm, stream nr. Soledad Lodge, río Las Piedras , 12°02′03″S, 069°47′47″W . ROM 62201, 5, 74-120 mm, PN Manú, Pakitza Biological Station , stream in aguajal, ca. 12°52′S, 069°16′W . Puno. MUSM 3692 , 2 , Zona Reservada ( ZR) Tambopata-Candamo, río Tambopata , ca. 13°30′S, 069°34′W . MUSM 11786 , 1, 111 mm, ZR Tambopata-Candamo, río Pacal, affl. río Candamo , ca. 13°31′S, 069°41′W . MUSM 11186 , 2 , 106-128 mm, ZR Tambopata-Candamo, río Candamo , ca. 13°31′S, 069°41′W . Uruguay. Artigas. ZVC-P 5687, 1, 90 mm, 30º26′20″S, 056º48′14″W , ZVC-P 5916, 2, 133- 133 mm, Tres Cruces stream, Uruguay dr., 30º18′09″S, 057º23′21″W. Durazno. UF183773 , 2 , immature (1 CS), 105-130 mm, Laguna Lavalle, río Negro, Uruguay dr., 33°01′16″S, 055°22′30″W . Rocha. CU 77188, 1, Velasquez stream, Patos-Mirim dr., 34°03′S, 054°28′W . CU 77189, 1, Sauces stream, Patos-Mirim dr., ca. 34°00′S, 053°51′W. ZVC-P 6287, 1 (holotype), 154 mm, Arroyo Cuatro Palmas, Route 15, km 268, nr. Cebollatí, affl. Arroyo Pelotas, affl. Laguna Merín, Patos-Mirim dr., 33º23′43″S, 053º51′53″W. Tacuarembó. ZVC-P 5685, 1, 137 mm, Batoví stream, Uruguay dr., 31º54′28″S, 056º01′03″W .