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Revista de la Sociedad Entomológica Argentina

versión impresa ISSN 0373-5680versión On-line ISSN 1851-7471

Rev. Soc. Entomol. Argent. v.68 n.1-2 Mendoza ene./jun. 2009


"Jejenes" (Diptera: Simuliidae) of Nahuel Huapi National Park, Patagonia, Argentina: Preliminary results

"Jejenes" (Diptera: Simuliidae) del Parque Nacional Nahuel Huapi, Patagonia, Argentina: Resultados preliminares

Hernández, Luis M.*, Fernanda Montes de Oca**, Malcolm Penn*, Julieta Massaferro***, Analía Garré**** and Stephen J. Brooks*

* The Natural History Museum, London, United Kingdom,
** Universidad Nacional del Comahue, Bariloche, Argentina
*** INIBIOMA, CONICET, Bariloche, Argentina
**** ILPLA, Instituto de Limnología, La Plata, Argentina

ABSTRACT. The Simuliidae is a family of Diptera with approximately 2072 described species worldwide. The females of the majority of the species feed from vertebrates' blood, which makes them a significant plague that affects both men as well as cattle, birds, and other vertebrates. The objective of this paper is to create an inventory of Simuliidae and to reveal certain aspects of the biology and distribution of this family of aquatic insects in the Nahuel Huapi National Park. Moreover, information on the zoogeography of Simuliidae in Patagonia is provided. Five genera, 3 subgenera and 32 species Simuliidae are recorded from Patagonia: Cnesia (three spp.), Cnesiamima (one sp.), Gigantodax (14 spp.), Paraustrosimulium (one sp.), Simulium (Ectemnaspis) (one sp.), S. (Psaroniocompsa) (one sp.) and S. ( Pternaspatha) (11 spp.), At present, we have collected all five genera, one subgenus of Simulium (Pternaspatha), and 19 species of Simuliidae in the park, which amounts to 57% of the Simuliidae fauna in this area. Puerto Blest, a characteristic area of the High-Andean phytogeographical province (humid forest), showed the highest diversity of Simuliidae.

KEY WORDS. Jejenes; Simuliidae; Nahuel Huapi National Park; Patagonia.

RESUMEN. Los simúlidos pertenecen a una familia de Diptera (Simuliidae) con alrededor de 2.072 especies descritas a nivel mundial. Las hembras de la mayoría de las especies se alimentan con sangre de vertebrados, lo cual las convierte en importantes plagas que afectan tanto al hombre como al ganado, aves y otros vertebrados. Los objetivos de este trabajo son llevar a cabo un inventario de Simuliidae y dar a conocer algunos aspectos de la biología y la distribución de esta familia de insectos acuáticos en el Parque Nacional Nahuel Huapi, Argentina. Además, se proporciona información sobre la biogeografía de Simuliidae en la Patagonia. Cinco géneros, un subgénero y 32 especies de simúlidos han sido registrados para Patagonia: Cnesia (3 spp.), Cnesiamima (1 sp.), Gigantodax (14 spp.), Paraustrosimulium (1 sp.), Simulium (Ectemnaspis (1 sp.), S. (Psaroniocompsa) (1 sp.) y Simulium (Pternaspatha) (11 spp.). Hasta el presente, se han colectado los cinco géneros, un subgénero (Pternaspatha) y 19 especies de Simuliidae en el PNNH, lo cual representa el 57% de la fauna de Simuliidae en esta área. La mayor diversidad de Simuliidae se encontró en Puerto Blest, zona característica del bosque de la provincia fitogeográfica Alto-Andina.

PALABRAS CLAVE. Jejenes; Simuliidae; Parque Nacional Nahuel Huapi; Patagonia.

Recibido: 16-02-2009;
Aceptado: 01-04-2009


The Simuliidae ("blackflies" or "jejenes" as they are commonly known) is a family of Diptera containing 2072 formally named species worldwide (Adler & Crosskey, 2008 2009) of which 12 genera and approximately 359 species are found in the Neotropica Region (Coscarón & Coscarón-Arias, 2007) The female in the majority of species requires a blood meal for egg maturation and it is this requirement that makes species in this family important as biting pests and in the transmission of parasites in the blood and skin in both man and warm blooded animals (Crosskey, 1990). The most important parasite transmitted by simuliid blackflies is the nematode Onchocerca volvulus (Leuckart) which is responsible for the human disease known as "onchocerciasis or river blindness" At present, 37 million people are estimated to carry O. volvulus, with 90 million being at risk in Africa (Basánez et al., 2006). They are also responsible for the transmission to humans of the less pathogenic nematode Manzonella ozzardii Manson (Shelley & Coscarón, 2001). Blackflies or jejenes are also well-known vectors of haematozoa parasites (e.g, Leucocytozoon Danilewsky) and arboviruses to a variety of both domesticated and wild animals. Massive blackflies attacks have been known to cause mortality in cattle pigs, and sheep, which is attributed to toxic shock from the salivary injections during their bite (Currie & Adler, 2008). Other economic impacts caused by blackflies are weigh loss, reduced milk production, malnutrition and impotence (Adler et al., 2004; Currie & Adler, 2008).
Apart from their medical importance, the fact that all simuliid species breed in running water makes the family economically important in terms of environmental monitoring of freshwater contamination, particularly because immature stages (larvae and pupae) are highly susceptible to both organic and inorganic pollution (e.g. effluent from sugar mills, slurry from farms, insecticide and fertiliser run off from farms and plantations) (Anthony Shelley, pers. comm.). In addition, blackflies are key organisms in aquatic ecosystems as they are an important food source for many vertebrates (e.g., salmonids) and invertebrates (e.g., Plecoptera) (Currie & Adler, 2008). Furthermore, the filter-feeding of the larva plays an important role in stream ecologies due to their ability to remove particulates and dissolved organic matter from the water column and egested it as nutritious faecal pellets (Currie & Adler, 2008; Malmquivist et al., 2001; 2004).
In Argentina, the jejenes fauna or Paquitas is relatively well known mainly due to the efforts of Coscarón (1987; 1991), Coscarón & Coscarón-Arias (2007), Coscarón & Wygodzinsky (1972) and Wygodzinsky & Coscarón (1967; 1973a, b; 1989). Studies of Simuliidae of Patagonia were first initiated by Edwards (1931) and Wygodzinsky (1953, 1958). In this area the Simuliidae are renowned for the attacks they inflict on people and equines, and the fauna shows a high level of endemicity (Coscarón-Arias, 2002). Nonetheless, the Simuliidae fauna within the National Park Nahuel Huapi (NHNP) is still poorly known. The park comprises a wide variety of habitats and is considered a classical representation of the zoogeographical region of Patagonia. This paper augments the preliminary results obtained for the family Simuliidae, which has been included as a target group in a multidisciplinary project funded by the Darwin Initiative, UK. The projects' basic aim was to undertake baseline research on the wetland invertebrate fauna of northern Patagonia. The distribution and biology of species of Simuliidae in the park is also reviewed.

Previous work on the Simuliidae fauna of Patagonia. The first studies of Simuliidae in Patagonia date back to Blanchard (1852) and Bigot (1888). Philippi (1865) also dealt with the fauna of Chile, which is biogeographically related to Argentina. In these papers the latter authors briefly described four species. The first comprehensive study on Simuliidae in Patagonia is the revision of Edwards (1931). Edwards described nine species new to science, commented on the species previously described by Blanchard, Bigot and Philippi [he also dealt with S. moorei (Silva Figueroa), now a synonym of Paraustrosimulium anthracinum (Bigot), and produced one of the first supraspecific classification systems in Simuliidae. It is worth mentioning a rather neglected paper by Edwards in 1927, in which he gave an account of his field trip from the city of Bariloche (Argentina) to the town of Los Andes (Chile), and provided a detailed insight into the flora and fauna of the region at that time. Enderlein (1925, 1934) also examined material of some species of Simuliidae from South America, and erected several new genera and species of which the genera Cnesia Enderlein (for C. dissimilis (Edwards), as S. gynandrum (Edwards), the type species) and Gigantodax Enderlein (type species G. bolivianus Enderlein) are still valid.
Impetus for the study of Simuliidae in Patagonia was given by Coscarón & Wygodzinsky (1962; 1972), Wygodzinsky & Coscarón (1962; 1967) and Wygodzinsky (1953; 1958). The former paper (Coscarón & Wygodzinsky, 1962) was the first publication in which the Simuliidae were treated in a cladistic framework, although no phylogenetic trees were provided. Subsequently, the latter authors produced a series of publications in which they described numerous new species and/or redescribed other taxa poorly known in Argentina (e.g., Coscarón, 1985; Wygodzinsky & Coscarón, 1967; 1973a-b; 1989). The revision of Wygodzinsky & Coscarón (1973a) still remains the most complete study for the New World "Prosimulinii". In the latter revision, the authors erected the genera Araucnephia Wygodzinsky & Coscarón, Araucnephioides Wygodzinsky & Coscarón, Cnesia and Mayacnephia Wygodzinsky & Coscarón, and also discussed the taxonomic position of the genera Gigantodax, Lutzsimulium D'Andretta & D'Andretta, Paraustrosimulium Wygodzinsky & Coscarón, and Tlalocomyia Wygodzinsky & Díaz Nájera [all these genera now belong to the tribe Simuliini, and Mayacnephia is a synonym of Tlalocomyia; see Adler & Crosskey, 2008]. The reviews of Dumbleton (1963) and Mackerras & Mackerras (1948) on the Australian Simuliidae fauna and the genus Austrosimulium Tonnoir, are pivotal in our understanding of the zoogeographical relationships of the Subantarctic simuliid fauna. The taxonomic position of Paraustrosimulium was discussed in comparison to that of Austrosimulium (Smart, 1945; Stone, 1963).
The control of the Simuliidae in Patagonia has already been reviewed by Coscarón-Arias (1989; 1998a), who also studied the cytotaxonomy of three species of the genus Gigantodax: G. chilensis (Philippi), G. fulvescens (Blanchard), and G. marginalis (Edwards), and of Cnesia dissimilis in the Lanin National Park (Coscarón-Arias, 1998b). Coscarón-Arias (2000) also studied the cytotaxonomy of Simulium limay Wygodzinsky, a species also found in Nahuel Huapi National Park. In 2002, Coscarón-Arias reviewed all species found in Patagonia, and provided a checklist together with an identification key to separate all genera and species based on immature stages. Information on the taxonomy, biology and distribution of all Simuliidae found in Patagonia may be found in Coscarón (1991) and, more recently, in Coscarón & Coscarón-Arias (2007), except for the genus Gigantodax. The latter can be found in Wygodzinsky & Coscarón (1989). Coscarón-Arias (2002) recorded five genera (Cnesia, Cnesiamima Wygodzinsky & Coscarón, Gigantodax, Paraustrosimulium, and Simulium Latreille), three subgenera of Simulium (Ectemnaspis Enderlein, Psaroniocompsa Enderlein, and Pternaspatha Enderlein) and 34 species of Simuliidae from Patagonia. However, this number has now been reduced to 32 due the recent synonymy of Simulium bonaerense Coscarón & Wygodzinsky with Simulium jujuyense (Paterson & Shannon) by Hernández et al. (2007), and Simulium walterwittmeri Wygodzinsky with S. limay by Coscarón & Coscarón Arias (2007).


The NHNP and the National Reserve Nahuel Huapi are located between latitude 40º 08' and 41º 35' south and longitude 71º 02' and 71º 57' west in Argentina. Brooks et al. (2009) provided the characterisation of climate, drainage, geology, topography, and vegetation following the publications of Martin & Mermoz (2005) and Monjeau (2006). Therefore, the reader should consult the latter publications for further details.
Collecting field trips were made in the summer season (October to February) in 2006-2007 and 2007-2008 (Fig. 1). Numerous running water courses were sampled for Simuliidae in different vegetation zones of the NHNP. The collecting, rearing and dissection techniques, and terminology used are those detailed in Shelley et al. (1989; 1997), Hernández & Shelley (2005) and Hernández et al. (2007) [see also "Systematics and Taxonomy of Blackflies, by Hernández, 2007]. Adults collected in Malaise trap and larvae obtained by kick-sampling techniques were also examined for this study. We have followed the classification system of Adler & Crosskey (2008) for the generic and subgeneric arrangements in this paper. All specimens collected during the course of this project are housed at The Natural History Museum, London, UK (BMNH), and Museo La Plata, Argentina (MLP). Further material has been examined from Museum für Naturkunde der Humboldt-Universität, Berlin, Germany (NMHU), National Museum of Natural History, Washington, USA (USNM), and the Canadian National Collection, Ottawa, Canada (CNC).

Fig. 1. Main collecting areas for Simuliidae in Nahuel Huapi National Park, northern Patagonia, Argentina, in the north and central parts of the park.


Coscarón-Arias (2002) and Coscarón & Coscarón-Arias (2007) recorded five genera and 32 species of Simuliidae for Patagonia (see Hernández et al., 2007). At present, we have collected all five genera (Cnesia, Cnesiamima, Gigantodax, Paraustrasimulium, and Simulium), a subgenus of Simulium (Pternaspatha) and 19 species in the NHNP (Table I), which amount to 57% of the Simuliidae fauna in Argentina. We have failed to find Simulium jujuyense (as S. bonaerense) and S. wolffhuegeli (Enderlein) in the park. However, this is hardly surprising as both species are characteristic of hotter climate towards the northern areas of Rio Negro Province, and the phytogeographical provinces of Monte, Espinal and Pampeana (Roig-Juñent & Coscarón, 2001). In addition, Gigantodax trifidus Wygodzinsky & Coscarón, Gigantodax minor Wygodzinsky & Coscarón, and seven species of the subgenus Pternaspatha have also not been collected in the park (see Table I), which highlights the need to continue the survey of Simuliidae in this area. A field trip was made in November 2008 to collect in new water courses to the south of the park and localities near Bariloche, especially Lake Guillelmo, from where Edwards (1931) described several species of Gigantodax.

TABLE I. Checklist of Simuliidae recorded for Patagonia by Coscarón-Arias (2002) and Coscarón & Coscarón-Arias (2007) and those recorded at the Nahuel Huapi National Park, Argentina in this study.

The immature stages (larvae and pupae) of most of the species of Simuliidae were found in cold, clear, fast flowing water streams in NHNP, except for Paraustrosimulium anthracinum that was collected in slow flowing, murky water stream in Isla Victoria. The area of the Valdivian forest (Puerto Blest) showed a higher diversity of Simuliidae, with Gigantodax being the most frequent genus. Cnesia dissimilis and Simulium (Pternaspatha) nemorale Edwards are the most abundant species in the park appearing in more than 50% of the total sites sampled up to March 2008 (184 collecting sites). They have a wider distribution in the park extending from the Nothofagus Blume forests to the Low Steppe and urban areas nearby. The genus Gigantodax is the most diverse in the park with 11 species recorded in this area (see Table I). Two species were collected biting humans in the park, P. anthracinum and S. nemorale. The former species (as S. moorei) was recorded as man-biting in the Taitao Peninsula in Chile by Silva Figueroa (1917), while Edwards (1931) recorded "the painful bite causing swelling of the hand". However, Wygodzinsky & Coscarón (1973a) stated that they had never recorded anthropophilous behaviour in this species during their studies in Patagonia. We only collected a few females of P. anthracinum in the Challhuaco area, although they did not display a marked anthropophily. On the contrary, S. nemorale was very common and highly anthropophilic, biting voraciously in Puerto Blest area, Challhuaco, and the transition forests (ecotone) at Rio Nirihuau.
In general, the Simuliidae fauna of the NHNP and Patagonia shows a high degree of endemism with Cnesia, Cnesiamima, Paraustrosimulium (together with the Chilean taxa Araucnephia, Araucniphiodes) only found in this region (Coscarón & Coscarón-Arias, 1995). The genus Gigantodax is the only taxon that is widely distributed from south USA, along the Mesoamerican mountain range, northern South America (Venezuela) to Tierra del Fuego (Coscarón & Miranda-Esquivel, 1998; Pinto-Sánchez et al., 2005; Wygodzinsky & Coscarón, 1989). The subgenus Pternaspatha is highly specious in Patagonia and the Andes, with a single species (Simulium cotopaxi Wygodzinsky & Coscarón) recorded from Ecuador between 3000-4000m of altitude (Wygodzinsky & Coscarón, 1979; see also Coscarón & Coscarón-Arias, 1996a-b). The biogeography and distribution of the Simuliidae in Patagonia and the Neotropical Region have been reviewed by several authors (e.g., Coscarón & Coscarón-Arias, 1995; Roig-Juñent & Coscarón, 2001), who postulate that the existing fauna suffered a main vicariant event correlated with marine introgression. These authors also stated that the close relationship of Simuliidae with taxa from other continents showed that this family was widely distributed in Gondwana before its fragmentation. Phylogenetical analysis have revealed that the Patagonian Simuliidae fauna is more closely related to the taxa found in the temperate forests of New Zealand, Australia, and areas of southeast Asia (e.g., Gil-Azevedo & Maia-Herzog, 2007; Roig-Juñent & Coscarón, 2001; Wygodzinsky & Coscarón, 1973a-b), which are consistent with the distribution patterns in other insect groups (see detailed review in Morrone, 2006).


The authors would like to thank the Darwin Initiative (Project, Reg. No. 15/025), United Kingdom for providing funds to carry out this study. The first author is grateful to the following colleagues for loans of Simuliidae material: Dr. Sixto Coscarón, Museo La Plata, Argentina; Joachim Ziegler, NMHU, Germany; Wayne Mathis and the late Nancy Adams, USNM; and Scott Brooks and Jeff Cumming, CNC. Special thanks is also given to Susana Seijas and Maria Sage, and the staff at the Headquarter of the Nahual Huapi National Park, Bariloche for their logistical support on this project.


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