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Gayana (Concepción)

On-line version ISSN 0717-6538

Gayana (Concepc.) vol.76 no.2 Concepción  2012 

Gayana 76(2): 153-161, 2012.


Distributional patterns of the South American species of Hyalella (Amphipoda: Hyalellidae)


Patrones de distribución de especies sudamericanas de Hyalella (Amphipoda: Hyalellidae)


Patricio De los Ríos-Escalante1,2, Juan J. Morrone3 & Reinaldo Rivera1,4

1Universidad Católica de Temuco, Facultad de Recursos Naturales, Escuela de Ciencias Ambientales, Laboratorio de Ecología Aplicada y Biodiversidad; Casilla 15-D, Temuco, Chile. and

2Núcleo de Estudios Ambientales UCTemuco, Chile

3Museo de Zoología "Alfonso L. Herrera", Departamento de Biología Evolutiva, Facultad de Ciencias, Universidad Nacional Autónoma de México (UNAM), Apartado Postal 70-399, 04510 Mexico, D.F., Mexico.

4Departamento de Zoología, Universidad de Concepción, Casilla 160-C, Concepción, Chile.


Distributional patterns of the South American species of the freshwater amphipod genus Hyalella were analysed using a panbiogeographic approach. Five generalized tracks were found: (1) northern Andes to Lake Titicaca (H. dielaii, H. meinerti, H. dybowskii, H.jelskii, H. lubominsky, and H. pauperocavae; (2) lake Titicaca (H. armata, H. cuprea, H. latinamus, H. lucifugax, H. montforti, H. neveulemairei, H. robusta, H. tiwanaku, H. simplex simplex, and H. solida); (3) central Andes (H. fossamancinii and H. kochi); (4) Pampas (H. bonariensis, H. caeca, H. castroi, H. longispina, H. montenegrinae, H. pampeana, H. pernix, H. pseudoazteca, and H. warmingii); and (5) Subantarctic (H. patagonica H. costera, H. curvispina, H. chiloensis, H. falklandensis, H. franciscae, H. neonoma, H. patagonica, and H. simplex). One node was found at lake Titicaca (intersection of generalized tracks 1, 2, and 3).

Keywords: Chile, Hyalella South America, biogeography


Se estudió el patrón de distribución de especies Sudamericanas del género de anfípodo de agua dulce Hyalella mediante una aproximación panbiogeográfica. Se encontraron cinco trazos (1) Norte de los Andes al Lago Titicaca (H. dielaii, H. meinerti, H. dybowskii, H.jelskii, H. lubominsky, y H. pauperocavae; (2) Lago Titicaca (H. armata, H. cuprea, H. latinamus, H. lucifugax, H. montforti, H. neveulemairei, H. robusta, H. tiwanaku, H. simplex simplex, y H. solida); (3) Andes centrales (H. fossamancinii y H. kochi); (4) Pampas (H. bonariensis, H. caeca, H. castroi, H. longispina, H. montenegrinae, H. pampeana, H. pernix, H. pseudoazteca, y H. warmingii); y b (5) Subantártico (H. araucana, H. costera, H. curvispina, H. chiloensis, H. falklandensis, H. franciscae, H. neonoma, H. patagonica, y H. simplex). Se encontró un nodo en el lago Titicaca que correspondió a la intersección de los trazos 1; 2 y 3.

Palabras clave: Chile, Hyalella Sudamérica, biogeografía



Freshwater amphipods inhabit superficial and underground waters. They have different trophic roles: herbivores, carnivores, detritivores or omnivores (Witt and Hebert 2000; Vainola et al. 2008). In South America, amphipods are represented by species inhabiting subterranean (Bogidellidae) and superficial (Hyalellidae) waters (Vainola et al. 2008). Hyalellidae are represented in the Americas by the genus Hyalella, which has been recently studied from a taxonomic perspective (Witt and Hebert 2000; Witt et al. 2003; González and Wattling 2002a-c, 2003a-c; González 2003; González et al. 2006), and is possibly found in New Zealand (Pugh et al. 2002).

Recent studies on Chilean species of Hyalella indicate that they inhabit subsaline inland waters, such as streams and shallow ponds in northern Chile (González 2003; De los Ríos et al. 2010), coastal inland waters (González 2003), and oligotrophic inland waters of Patagonia between 38-51° S (González 2003; De los Ríos et al. 2007; De los Ríos and Roa 2010). Similar studies have investigated Central Argentina and Patagonia (Lopretto 1982, 1983a, b; Miserendino 2001; Jergentz et al. 2004; Miserendino et al. 2008; Zilli et al. 2008).

Integrative biogeographic studies of the species of Hyalella have not yet been conducted. Here we undertake a preliminary track analysis of the South American species of this genus to improve current understanding of their patterns of geographical distribution.


Distributional data for this study were obtained from the literature (Lopretto 1982, 1983a, b; Pereira, 1985, 1989, 2004 Pereira et al. 1985, 1989; Stock and Plavoet 1991; Bond-Buckup and Araujo 1998; Casset et al. 2001; González and Coleman 2002; González and Wattling 2002a-c, 2003a-d; Carrera and Günkel 2003; González 2003; Jergentz et al. 2004; Pereira 2004; Coleman & González, 2006; Galan & Herrera, 2006; González et al. 2006; Da Silva and Bond-Buckup 2008; Dos Santos et al. 2008; Dutra et al. 2008; Zilli et al. 2008; Rauque and Semenas 2009; Spaccesi & Rodríguez. 2009; De los Ríos et al. 2007, 2010; De los Ríos and Roa 2010; Cardoso et al., 2011; Bastos-Pereira & De Padua-Bueno, 2012). It was not cosidered the species names H. curvispina cangallensis, H. aff. curvispina, H. thomseni, H. sapropelica and H. lalage (Brehm-Lunz, 1928, 1935; Gonzalez & Wattling, 2003c; González et al., 2006)

Coordinates of localities were obtained from the literature or calculated on maps and rounded to minutes. Within each country, localities are ordered roughly in a south-north direction. For details on the localities of each species see Appendix.

The panbiogeographic approach involves plotting distributions of different taxa on maps and joining their separate localities with lines called individual tracks. These tracks represent the geographical coordinates of species or higher taxa. Operationally, they consist of lines drawn on a map connecting the localities at which the taxa occur according to their geographical proximity. When different individual tracks are superimposed, the resulting summary lines are considered generalized tracks, which are interpreted as indicating the pre-existence of ancestral biotas that subsequently became fragmented by tectonic and/or climatic change. If two or more generalized tracks intersect in a given area, they determine a node, which indicates that different ancestral biotic and geological fragments interrelate in space and time as a consequence of terrain collision, docking or suturing. This node thus constitutes a composite area. For further details about panbiogeographic methods, see Morrone (2004, 2006, 2009).


Individual tracks and localities of the species analyzed are represented in Figures 1-9. In northern South America, H. anophthalma, H. azteca and H. inermis has been reported from Venezuela (Fig. 1a) and H. paramoensis from northern Colombia (Fig. 1b). Hyalella quindioensis has been reported in inland mountain waters of the Colombian Andes (Fig. 2a). In the northern Andean inland waters, H. simplex cangallensis (Fig. 1c), H. pauperocavae (Fig. 2b), H. lubomirskii (Fig. 3a), H. dybowskii (Fig. 2c), and H. meinerti (Fig. 3b) have been reported. In coastal zones of Lake Titicaca (Peru and Bolivia), 17 species have been reported: H. echinus, H. crawfordi, H. gauthieri, H. longipalma, H. neveulemairei, H. cuprea, H. nefrens, H. armata, H. lucifugax, H. montforti, H. tiwanaku, H. longipes, H. latimanus, H. robusta, H. longispina, H. solida, and H. jelskii (Fig. 4a). Hyalella fossamancinii has been reported from inland waters of the Andes Argentina and northern Chile (Fig. 4b). Hyalella kochi occurs in Argentinean and Chilean mountain inland waters (Fig. 5a). Several species have been reported near the Atlantic coast: H. gracilicornis (Fig. 4c), H. warmingi (Fig. 5b), H. pernix (Fig. 5c), H. spelaea, H. longistila (Fig. 6a), H. caeca (Fig. 6b), and H. dielaiii (Fig. 4d). The following species have been reported from southern Brazil: H. carstica (Fig. 2e), H. longistila (Fig. 2f), H. pseudoazteca (Fig. 5d), H. montenegrinae (Fig. 6c), H. castroi (Fig. 7a), and H. pleoacuta (Fig. 7c). Two species are restricted to streams from Buenos Aires province, Argentina: H. bonariensis (Fig. 6d) and H. pampeana (Fig. 7b). The species H. costera has been reported from coastal wetlands, streams, and shallow ponds from northern Chilean Patagonia to northern Chile, in the Antofagasta region (Fig. 7d). Hyalella chiloensis has been reported from Central Patagonia, coastal lakes in Chiloé Island, and Concepción in central Chile (Fig. 6e). Hyalella falklandensis (Fig. 7e) and H. neonoma (Fig. 5e) have been reported from the Falkland Islands (Islas Malvinas). In Patagonia, including nearby islands, H. patagonica has been reported (Fig. 1e). In Argentina, H. curvispina has been reported from southern, central and northern Argentinean Patagonia in large and deep lakes and in Buenos Aires province; it has also been reported from the Falkland Islands, Uruguay, southern Brazil specifically in coastal inland waters, and from Peru (Fig. 8a). Hyalella rionegrina has been reported from one locality in Chubut province, in Argentinean Patagonia (Fig. 2d). A species restricted to Chile is H. simplex, is known from southern Chilean Patagonia, specifically from coastal inland waters of the Strait of Magellan strait and from a high-mountain shallow lake in northern Chilean Patagonia

(Fig. 3d). Hyalella franciscae has been reported from lakes of Torres del Paine National Park, in the Magellan region

(Fig. 7f).

Based on the overlap of the different individual tracks, we identified five generalized tracks (Fig. 9):

(1) Northern Andes. Inland waters of Venezuela and Peru between 10° N-15° S. The species assigned to this track are

H. anophtalma, H. inermis, H. quindioensis, H. paramoensis, H. meinerti, H. dybowskii, H.jelskii, H. lubominsky, and H. pauperocavae.

(2) Lake Titicaca. The species assigned to this track are H.

armata, H. crawfordi, H. cuprea, H. echinus, H. gauthieri H. latinamus, H. lucifugax, H. montforti, H. neveulemailei, H. robusta, H. tiwanaku, H. simplex simplex and H. solida.

(3) Central Andes. Inland waters from central Argentina (31° S) to northern Chile (18-26° S). The species assigned to this track are H. fossamancinii and H. kochi.

(4) Pampas. Inland waters close to the Atlantic coast from Buenos Aires province (35° S) in central Argentina to Uruguay and southern Brazil (19° S). The species assigned to this track are H. bonaeriensis, H. caeca, H. carstica, H. castroi, H. dielaii, H. longistila, H. montenegrinae, H. pampeana, H. pernix, H. pseudoazteca, and H. warmingii.

(5) Subantarctic. Andean inland waters of Argentina and Chile between 39-51° S, southern Patagonian plains between 51-53° S, Tierra del Fuego, and the Falkland Islands. Species assigned to this track are H. curvispina, H. chiloensis, H. falklandensis, H. franciscae, H. neonoma, H. patagonica, and H. simplex.

One node (Fig. 9) was found where generalized tracks 1, 2, and 3 intersect, at lake Titicaca.


The generalized tracks identified are similar to those described by Menu-Marque et al. (2000), who applied this procedure to the copepod genus Boeckella. That study identified a generalized track in southern Patagonia, the Atlantic coast of Argentina, the Andean mountains of northern Argentina and Chile, and the Andean regions of Bolivia, Colombia and Ecuador. Nevertheless, the nodes reported in the present study differ from those identified by Menu-Marque et al. (2000). Species of Hyalella are also known from Mexico and Central America, Bousfield (1996) supported it, and suggested the presence of Austrohyalella and Mesohyalella genus in South America. Nevertheless, the recent literature mentioned the presence of Hyalella genus for all American continent (Vainola et al., 2008; Witt & Hebert 2000; Wit et al., 2006)

The absence of nodes in southern Patagonia agrees with analyses for Subantarctic islands (Pugh et al. 2002) and southern South America (Dos Santos et al. 2008), about similarities between continental Subantarctic species. Our results contrast with those from calanoid copepods (Menu-Marque et al. 2000) and fishes of the genus Galaxias (Cussac et al. 2004), that are known from southern Patagonia, New Zealand and adjacent islands, a different situation was reported for species of Hyalella that are widespread in South North America (Witt & Hebert, 2000; Witt et al. 2003). The results of geographical distribution of Hyalella genus are similar in comparison to freshwater crabs of Aegla genus that is located in central and southern Argentina and Chile, and east South America, specifically Brazil and Uruguay (Pérez-Lozada et al., 2002, 2004, 2009). Nevertheless, species reports for practically all South American inland waters are rather scarce, so future analyses may modify these results, specifically between 18-8° N, the Titicaca basin, and eastern South America.


The present study was funded by projects DGI-CDA-2007-01 and MECESUP Project UCT 0804. The authors recognize the contributions of Luciano Parra to the preparation of the present manuscript.


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H. anophthalma (Ruffo, 1957). Venezuela: Cuevas de Hueque (Galán & Herrera, 2006).

H. armata (Faxon 1876). BOLIVIA: Achacache (16º 00' S; 68º 42' W). PERU: Coata bay (15º 30' S; 69º 54' W), Taquili/Amantane (15º 42' S; 69º 42' W), Chuquito (15º 53' S; 69º 52' W), Juli (16º 12' S; 69º 29' W), Isla Suana (16º 20' S; 68º 51' W), Golfo Desaguadero (16º 33' S; 69º 02' W) (González & Coleman 2002).

H. bonariensis Bond-Buckup et al. 2008. ARGENTINA: Salto, Buenos Aires province (34º 18' S; 60º 20' W), Pergamino river (35º 56' S; 60º 28' W), Cañada Honda, San Antonio de Areco (34º 04' S; 59º 30' W), Arroyo Manantiales (33º 42' S; 60º 20' W) (Dos Santos et al. 2008).

H. brasiliensis Bousfi eld 1996. BRAZIL: stream close to Prudentopolis (25º 12' S; 50º 57' W).

H. caeca Pereira & Goulart 1989. BRAZIL: Gruta Tobias de Baixo, Município de Ipiranga (around 23º 26' S; 47º 25' W) (Pereira 1989). Hyalella carstica (Bastos-Pereira & Bueno, 2012). BRAZIL: High Sao Francisco River (20°19'59,6' S; 45°36'25.3''W) (Bastos-Pereira & Bueno, 2012).

H. castroi González et al. 2006. BRAZIL: Vale das Trutas (28º 47' S; 49º 51' W) (González et al. 2006).

H. chiloensis González & Watling 2001. CHILE: Notuco, Chiloé (42º 39' S; 73º 49' W), Laguna Redonda, Concepción (36º 48' S; 73º 04'W), río Ñirepan, Coihaique (around 45º 34' S; 72º 03' W) (González 2003).

H. costera González & Watling 2001. CHILE: Quebrada Paposo, Antofagasta (25° 01' S, 70° 28' W), Limache (32º 59' S; 71º 15' W), El Molle (29º 58' S; 70º 56' W), isla Teja, Valdivia (39º 48' S; 73º 12' W) (González 2003).

H. crawfordi (Coleman & González, 2006). Perú: Molinopampa lake Titicaca (6º11' S; 77º37'W):,Catachaca, Bay of Puno (15º50' S; 71º08' W) (Coleman & González, 2006).

H. cuprea (Faxon 1876). BOLIVIA: Santa Rosa, lake Titicaca (16º 18' S; 60º 00' W), Lake Titicaca, unknown site (15º 50' S; 69º 25' W) (González & Watling 2003d).

H. curvispina Shoemaker 1942. ARGENTINA: Falkland Islands (51º 49' S; 59º 22' W), Argentinean lake (50º 15' S; 72º 33' W), Puerto Bandera (50º 18' S; 72º 47' W); laguna larga, Futalaufquen (42º 50' S; 71º 41' W); Lacar lake (40º 10' S; 71º 22' W); Horqueta stream, city of Arrefi ces, Buenos Aires province (34º 04' S; 60º 06' W) (Jergentz et al. 2004); Lujan river (34° 28' S; 59° 00' W); Samborombón river, Brandsen (35º 07' S; 60º 30' W), Samborombón river, Chascomús (35º 34' S; 58º 02' W), Samborombón Bay (35º 50' S; 57º 23' W). CHILE: Punta Arenas (53º 11' S; 70º 56' W). PERU: Pampa of Cangallo (13º 25' S; 74º 20' W). URUGUAY: Carrasco creek, Montevideo (around 34º 53' S; 56º 03' W) (Grosso & Peralta 1999; Casset et al. 2001; Dos Santos et al. 2008; Dutra et al. 2008; Spaccesi & Rodrígues 2009).

H. dielaiii Pereira 2004. BRAZIL: Alto da Serra, São Paulo (around 27º 01' S; 52º 47' W) (Pereira 2004).

H. dybowskii (Wrzesniowski 1879). PERU: Paucal, Montaña Nacho (07º 00' S; 79º 08' W) (González & Watling 2002b).

H. echinus (Coleman & González, 2006). PERU: Siripata, Uruñi bay, lake Titicaca (15º50' S; 71º08' W) (Coleman & González, 2006).

H. falklandensis Bousfi eld 1996. ARGENTINA: West Falkland, 4 sites (around 51º 39' S; 59º 58' W); East Falkland, 4 sites (around 51º 49' S; 58º 52' W) (Stock & Plavoet 1991; Dos Santos et al. 2008).

H. fossamanchini Cavalieri 1959. ARGENTINA: San Juan province (31º 30' S; 68º 32' W). CHILE: Santa Bárbara (21º 58' S; 68º 36' W), Conchi (22º 00' S; 68º 36' W), Salado (20º 20' S; 68º 39' W), Salvador (22º 23' S; 69º 31' W); Toconao (23º 11' S; 68º 00' W), Miscanti (23º 44' S; 67º 47' W), Miniques (23º 45' S; 67º 47' W); Río La Ola, Copiapó (around 27º 21' S; 70º 18' W), Surire, Arica (18º 50' S; 69º 02' W), Putana river (22º 34' S; 67º 52' W), El Tatio (22º 21' S; 68º 00' W), Salar de Punta Negra (24º 35' S; 68º 58' W) (González 2003; Dos Santos et al. 2008; De los Ríos et al. 2010; De los Ríos-Escalante 2011).

H. franciscae González & Watling 2003. CHILE: Laguna El Paso, Torres del Paine (50º 29' S; 72º 58' W), Lago Risopatrón (44º 15' S; 72º 31' W), Laguna Melliza (51º 00' S; 73º 00' W), Laguna Redonda (51º 00' S; 73º 00' W), Laguna Larga (53º 20' S; 69º 00' W), Lago Sarmiento (51º 04' S; 72º 45' W); Laguna Melliza (51º 03' S; 72º 55' W); Tres Puentes wetland (53° 06' S; 70° 52' W), Porvenir pond (53° 17' S; 70° 19' W) (González 2003; González & Watling 2003a). Hyalella gauthieri (Coleman & González, 2006). Perú: Oruñi Bay, Taquili island (15º46' S; 69º41' S) (Coleman & González, 2006).

H. gracilicornis (Faxon 1876). BRAZIL: Campos, Rio de Janeiro (21º 28' S; 41º 13' W), Represa Belvedere, Campus da UFV, Minas Gerais State (18º 55' S; 48º 01' W) (González & Watling 2003c).

H. inermis Smith 1875. ECUADOR: Río Itambi (07º 11' S; 78º 12' W). VENEZUELA: Lake Valencia (10º 11' N; 68º 00' W) (Carrera & Gunkel 2003; González & Watling 2003b).

H. jelskii (Wrzesniowski 1879). PERU: Cordillera de Punamarca (14º 02' S; 76º 16' W) (González & Watling 2002b).

H. kochi González & Watling 2001. ARGENTINA: Jujuy province, Yavi Chico River (22º 05' S; 65º 25' W), reservoir near Escuela Agrotécnica in Huamahuaca (23º 11' S; 65º 20' W), Tilcara, lake north of Tilcara, near Río Grande (23º 33' S; 65º 23' W), Tucumán province, streamlet tributary of Río de Los Sojas, route 307 in the direction of Tucumán with Tafi del Valle (26º 56' S; 65º 39' W). CHILE: Guallatire, Arica (18º 29' S; 70º 16' W), Bofedal de Arabilla, Arica (19º 14' S; 68º 51' W), El Tatio (22º 21' S; 68º 00' W), Salar de Punta Negra (24º 35' S; 68º 58' W), Loa river, Antofagasta (22º 14' S; 68º 36' W); Santa Bárbara (21º 58' S; 68º 36' W), Conchi (22º 00' S; 68º 36' W), Salado (20º 20' S; 68º 39' W). PERU (12º 03' S; 75º 07' W) (González 2003; González & Watling 2002b; Dos Santos et al. 2008; De los Ríos et al. 2010).

H. latimanus (Faxon 1876). BOLIVIA: Llampopata, lake Titicaca (16º 00' S; 69º 15' W), Isla Coati, lake Titicaca (16º 15' S; 68º 40' W), Huatajata, lake Titicaca (16º 15' S; 68º 45' W) (González & Watling 2003d).

H. longipalma (Faxon 1876). BOLIVIA: lake Titicaca (16º 25' S; 68º 45' W) (González & Watling 2003d). Gayana 76(2), 2012 160

H. longipes (Faxon 1876). BOLIVIA: Achacache, lake Titicacaca (16º 06' S; 68º 26' W) (González & Watling 2003d).

H. longispina González & Coleman 2002. BOLIVIA: Paton (15º 38' S; 69º 15' W), Ancoraimes (15º 54' S; 68º 42' W), Siripata Bay (16º 05' S; 69º 03' W). PERU: Molinopampa (around 16º 13' S; 69º 22' W), lake Titicaca, Choccocoyo Bay (15º 24' S; 69º 30' W), Capachica (15º 39' S; 69º 47' W), Tamar (15º 37' S; 69º 52' W) (González & Coleman 2002).

H. longistila (Faxon 1876). BRAZIL: swamp 3 miles from Campos, Rio de Janeiro (21º 28' S; 41º 13' W) (González & Watling 2003c). Ijaci, Minas Gerais State (21°10'24' S; 44°56'24.2''W), Lagoa Feia, Rio do Janeiro State (21°05' 53.8'' S; 41°20' 22.4'' W)(Bastos-Pereira & Bueno, 2012).

H. lubomirskii (Wrzesniowski 1879). PERU: Pacasmayo (07º 25' S; 79º 34' W) (González & Watling 2002b).

H. lucifugax (Faxon 1876). BOLIVIA: Juli, lake Titicaca (16º 10' S; 69º 30' W) (González & Watling 2003d).

H. meinerti Stebbing 1899. BRAZIL: Sao Paulo (23º 20' S; 46º 22' W). COLOMBIA: Río Meléndez (08º 16' N; 73º 36' W), Rio Cali (07º 10' N; 73º 28' W), Río Piendamo (03º 00' N; 76º 00' W), Laguna de San Rafael (02º 50' N; 76º 15' W). PERU: Apurimac river, NE Ayacucho (15º 00' S; 72º 00' W). VENEZUELA: Laguna de Espino (08º 23'S; 66º 06' W. ECUADOR: Camar province (03º 20' S; 46º 22' W) (González & Watling 2002b, 2003b).

H. montenegrinae Bond-Buckup & Araujo 1998. BRAZIL: Rio Grande do Sul, municipio de Sao José dos Ausentes (29º 29' S; 50º 15' W) (Bond-Buckup & Araujo 1998).

H. montforti Chevreux 1907. BOLIVIA: Bay of Challa (16°30' S; 68°50' W; Isla del Sol (16°00' S; 69° 15' W); Achacache Bay (15° 58' S; 68° 50' W), Río Suchez, lake Titicaca (15º 37' S; 69º 05' W). PERU: small lake in Isle of Ampura (15°40' S; 69° 52' W) (González & Watling 2003d).

H. nefrens González & Watling 2003. BOLIVIA: Santa Rosa, lake Titicaca (16º 18' S; 60º 00' W) (González & Watling 2003d).

H. neonoma Stock & Platvoet 1991. ARGENTINA: West Falkland: Pebble Island (51º 19' S; 59º 34' W); New Island (51º 42' S; 61º 16' W); Big Pond, Pebble Island (51º 19' S; 59º 34' W); East Falkland: small pond at Bertha's Beach (51º 53'S; 58º 22' W); unnamed creek on Wireless Ridge near Stanley (around 51º 40' S; 57º 55' W); Murrell River, around Stanley (around 51º 42' S;57º 50' W); Mullet Creek stream, Fitzroy river, unnamed stream at the crossing with Mount Pleasant (all around 51º 49' S; 58º 26' W) (Stock & Plavoet 1991; Dos Santos et al. 2008).

H. neveulemairei Chevreux 1904. BOLIVIA: Bay of Challa (16º 30' S; 68º 50' W), Isla del Sol (16º 00' S; 69º 15' W), Bay of Achacache (15º 58' S; 68º 50' W); near archipiélago de Campanario, lago Grande (15º 40' S; 69º 20' W), Escoma, lake Titicaca (15º 35' S; 69º 10' W). PERU: Small lake in Isle of Ampura (Yapura) (15º 40' S; 69º 52' W) (González & Watling 2003d).

H. pampeana Cavalieri 1968. ARGENTINA: Arroyo Vitel, Chascomús lagoon (35º 33' S; 58º 03' W); Laguna de Cardiel (Lopretto 1983a; Dos Santos et al. 2008).

H. patagonica (Cunningham 1871) (= H. araucana Grosso & Peralta 1999, see Morrone 2001). ARGENTINA: Peninsula near Ushuaia, streamlet Gente Grande (54º 43' S; 68º 01' W), Fagnano lake (54° 34' S, 68° 00' W), Santa Cruz, Fossiles river (50º 20' 21'' S; 72º 15' 22'' W), Puelo lake (42° 10' S, 71° 39' W), Lacar lake (40º 10' S; 71º 22' W); Verde lagoon, Chapelco (40º 07' S; 71º 13' W); Blanca lagoon, Neuquén (39º 02' S; 70º 21' W); Santa Cruz province, arroyo Calafate (50º 20' S; 72º 15' W), lake Escondido, Ushuaia (54º 39' S; 67º 49' W), stream near Lake Escondido (54º 43' S; 68º 01 'W); Olnie river, Santa Cruz (47º 45' S; 71º 30' W) (Santos et al. 2008); lake Mascardi (41° 17' S; 71° 38' W). CHILE: river of los Dervos (Ciervos), Magallanes (around 53º 10' S; 70º 56' W); Península Brunswick (53º 30' S; 71º 25' W); Deseado lake (54º 22' S;68º 40' W), Azopardo river (54º 28'S; 68º 58'W); De las Minas river, Pta. Arenas (53º 09' S; 70º 54'W), Puerto Natales (51º 55' S; 72º 27' W); Pólux lake (45° 43' S; 71° 53' W), Chorrillo de la Piedra (52º 12'S; 74º 46'W); Seco river (53º 05' S; 70º 52' W); Lago de los Ciervos, Punta Arenas (53º 11' S; 70º 56' W); Lago Sarmiento (51º 04' S; 72º 45' W); Negra lagoon (39° 15' S; 71° 42' W); Tinquilco lake (39º 10' S; 71º 43' W), Toro lake (39º 08' S; 71º 42' W), Chico lake (39º 08' S; 71º 42' W), Verde II lake (39º 08' S; 71º 42' W), Los Patos lagoon (39º 10' S; 71º 42' W); Conguillío lagoon (38° 38'; 71° 37' W), Verde I lagoon (38° 41' S; 71° 46' W), Arcoiris lagoon (38° 40' S; 71° 37' W), Captren lagoon (38° 38' S; 71° 42' W); Tres Puentes wetland (53º 07' S; 70º 52' W); Isla Picton (55º 03' S; 66º 55' W) (Schallenberg 1931; González 2003; De los Ríos et al. 2007; Santos et al. 2008; De los Ríos & Romero- Mieres 2009; Rauque & Semenas 2009; De los Ríos & Roa, 2010).

H. paramoensis Andres 1988. COLOMBIA: Laguna Negra, Paramo de Chisaca (04º 20' N; 74º 12' W) (González & Watling 2003b).

H. pauperocavae González & Watling 2002. PERU: trout culture station “El Ingenio”, Huancayo (12º 03' S; 75º 07' W) (González & Watling 2002b).

H. pernix (Moreira 1903). BRAZIL: Lagoa Esgotada, Itatiaia, Rio de Janeiro (around 22º 30' S; 44º 37' W) (Pereira 1985).

H. pleoacuta González et al. 2006. BRAZIL: Rio Grande Do Sul, Sao José Dos Ausentes, Vale das Trutas (28º 47' S; 49º 50' W) (González et al. 2006; Da Silva & Bond-Buckup 2008).

H. pseudoazteca González & Watling 2003. BRAZIL: Reserva Ecológica de Taim, Rio Grande do Sul State (32º 27' S; 52º 38' W) (González & Watling 2003c).

H. quindioensis González & Watling 2003. COLOMBIA: Quindio (06º 14' N; 73º 26' W) (González & Watling 2003b).

H. rionegrina Grosso & Peralta 1999. ARGENTINA: Prov. Río Negro: El Bolsón (around 41º 58' S; 71º 30' W) (Dos Santos et al. 2008).

H. robusta Chevreux 1907. BOLIVIA: Bay of Achacache (16º 06' S; 68º 26' W) (González & Watling 2003d).

H. simplex Schellenberg 1943. CHILE: Punta Arenas (53º 10' S; 70º 54' W), Punta Delgada, Magallanes (52º 15' S; 69º 45' W), Lago Quillehue (39º 34' S; 71º 32' W); Redonda lagoon (51° 01' S; 72' 52' W); Larga lagoon (51° 01' S ; 72' 52' W); Monserrat lagoon (51°07' S; 72° 57' W); Puerto Natales ponds (51° 42' S; 72° 47' W); Río Seco ponds (53° 06' S; 70° 53' W); Tres Puentes wetland (53° 06' S; 70° 52' W); Porvenir pond (53° 17' S; 70° 19' W) (González 2003; González & Watling 2003a).

H. simplex cangallensis Schellenberg 1943. PERU: Pampa de Cangallo (13º 25' S; 74º 20' W), Laguna Urcos, Cuzco (13º 41' S; 71º 37' W) (González & Watling 2002b). 161 Hyalella en Sudamerica: DE LOS RÍOS-ESCALANTE, P. ET AL.

H. solida (Chevreux 1907). PERU: Small lake in isle of Ampura (Yapura) (15º 40' S; 69º 52' W) (González & Watling 2003d).

H. tiwanaku González & Watling 2003d. BOLIVIA: Tiquina, lake Titicaca (16º 10' S; 68º 45' W), Moro, lake Titicaca (16º 00' S; 69º 15' W), Majal, lake Titicaca (16º 00' S; 69º 15' W), Puerto Pérez, lake Titicaca (16º 20' S; 68º 35' W), Chua, lake Titicaca (16º 10' S; 68º 45' W), Lago Mullini (17º 04' S; 66º 37' W) (González & Watling 2003d).

H. warmingi (Stebbing 1899). BRAZIL: Lagoa Santa, Minas Gerais State (19º 38' S; 43º 53' W), Gruta Mirasol, Sao Paulo State (20º 00' S; 49º 00' W) (González & Watling 2003c).

Recibido: 15.12.11 Aceptado: 13.08.12

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