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Revista de la Sociedad Entomológica Argentina

versión impresa ISSN 0373-5680

Rev. Soc. Entomol. Argent. vol.71 no.1-2 Mendoza ene./jun. 2012

 

TRABAJOS CIENTÍFICOS

Aquatic Coleoptera from Mburucuyá National Park (Corrientes Province, Argentina)

Los coleópteros acuáticos del Parque Nacional Mburucuyá (Provincia de Corrientes, Argentina)

 

Torres, Patricia L. M.*, Mariano C. Michat, María L. Libonatti*, Liliana A. Fernández**, Adriana Oliva*** and Axel O. Bachmann

*CONICET - Laboratorio de Entomología, Departamento de Biodiversidad y Biología Experimental, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Argentina; e-mail: patricialmtorres@yahoo.com.ar
**CONICET - División Entomología, Museo de La Plata.
***CONICET - Museo Argentino de Ciencias Naturales (Bernardino Rivadavia).

Recibido: 23-XI-2011
Aceptado: 23-II-2012

 


ABSTRACT. A list of the species of aquatic Coleoptera collected in Mburucuyá National Park is presented. One hundred and twenty-eight species included in 44 genera and seven families were identified. Ten species are new for Argentina: Agaporomorphus mecolobus Miller and Bidessonotus obtusatus Régimbart (Dytiscidae); Mesonoterus laevicollis Sharp, Suphisellus hyeroglyphicus Zimmermann, S. rufulus Zimmermann, S. sexnotatus (Régimbart), and Notomicrus traili Sharp (Noteridae); Berosus pluripunctatus Mouchamps, Enochrus guarani Fernández and Helobata corumbaensis Fernández & Bachmann (Hydrophilidae). Two families (Scirtidae and Spercheidae), 12 genera and 69 species are first cited for Corrientes Province.

KEY WORDS. Aquatic Coleoptera; Biodiversity Inventory; Iberá marshes; Neotropical Region

RESUMEN. Se presenta una lista de las especies de coleópteros acuáticos colectadas en el Parque Nacional Mburucuyá. Se identificaron 128 especies, incluidas en 44 géneros y siete familias. Diez especies se citan por primera vez para la Argentina: Agaporomorphus mecolobus Miller y Bidessonotus obtusatus Régimbart (Dytiscidae); Mesonoterus laevicollis Sharp, Suphisellus hyeroglyphicus Zimmermann, S. rufulus Zimmermann, S. sexnotatus (Régimbart) y Notomicrus traili Sharp (Noteridae); Berosus pluripunctatus Mouchamps, Enochrus guarani Fernández y Helobata corumbaensis Fernández & Bachmann (Hydrophilidae). Dos familias (Scirtidae y Spercheidae), 12 géneros y 69 especies son citados por primera vez para la provincia de Corrientes.

PALABRAS CLAVE. Coleópteros acuáticos; Biodiversidad; Inventario; Esteros del Iberá; Región Neotropical.


 

INTRODUCTION

Mburucuyá National Park is a protected area that covers 17,660 ha. It is located in the north-western part of Corrientes Province, Argentina, between 27°58´ to 26°05´ S and 57°59´ to 58°08´ W (Fig. 1). The park was created in 1991, and according to Ringuelet (1961) it belongs to the Mesopotamic District, which is within the Subtropical Dominion of the Neotropical Region. The area has a wide range of habitats: marshes, ponds, streams, flood-prone areas, forests and grasslands, and the landscape presents single-layered semixerophilous vegetation, herbaceous grasses, halophilic sedges, rushes, reeds, grasses and trees.


Fig. 1. Mburucuyá National Park (Corrientes Province) and sampling sites. (A) Marsh 1 at Santa Teresa Ranch; (B) Pond 1; (C) Marsh 2; (D) Pond 2 "facing pond"; (E) First stream heading to Portillo Gulley; (F) Portillo Stream; (G) Pond 3; (H) Second stream heading to Portillo Gulley; (I) Pond 4 "facing pond"; (J) Pond 5.

Several authors postulated the use of aquatic macroinvertebrates as indicators of quality conditions in freshwater ecosystems (e.g., Usseglio-Polatera et al., 2000), and others found a correlation between the composition of aquatic invertebrate communities and environmental conditions (e.g., Baldigo et al., 2009; Beyene et al., 2009). More specifically, aquatic Coleoptera have proven to be very useful as biological indicators and suitable for developing conservation criteria (Eyre & Foster, 1989; Eyre & Rushton, 1989; Foster et al., 1990; Ribera & Foster, 1993). Therefore, in order to evaluate possible future changes in water ecosystems it is important to know with a reasonable detail the current structure of the Coleoptera assemblages of a particular region.
This paper is part of a series of recent studies on the aquatic Coleoptera from different protected areas of Argentina (Torres et al., 2007, 2008; von Ellenrieder, 2007; Fernández et al., 2008, 2010). The aim of the present paper is to provide the first inventory of aquatic Coleoptera from the Iberá marshes eco-region, based on material collected in Mburucuyá National Park.

MATERIAL AND METHODS

Study area. Mburucuyá N. P. (Fig. 1) belongs to the Iberá marshes eco-region and has a vast variety of habitats, which represent important refuges for the native flora and fauna. Altitude within the park ranges between 60 and 98 MASL, average annual precipitation is 1200 mm, and mean annual temperature is 22°C. Prevailing winds are from the east. The landscape is mainly represented by rolling hills, marshes and extensive flatlands with a slight SW slope. Water accumulates throughout the valleys, originating wetlands such as marshes, swamps, gulleys, ponds and temporary pools.

Lentic habitats sampled

Marsh 1 (Fig. 1-A). Small marsh at Santa Teresa Ranch. Location: 28°00' 59.18'' S, 58°01' 50.71'' W. Dimensions: 185 x 200 m. Depth at sampling site: 40 cm. Insolation: very high. Vegetation: Eichhornia sp. (water hyacinth), Nymphoides indica (water snowflake), Salvinia sp. (water fern), Ludwigia sp. (primrose-willow). Soil: sandy. Turbidity: medium. Organic matter: high.
Pond 1 (Fig. 1-B). Round lagoon 200 m apart from the main road of the park. Location: 28° 02' 2.76'' S, 58°04' 48.93'' W. Dimensions: 200 m x 230 m. Depth at sampling site: 40 cm. Insolation: very high. Vegetation: large marginal patches of Eichhornia crassipes (common water hyacinth), Salvinia sp., Myriophyllum sp. (water milfoil), Ricciocarpus sp. (fringed heartwort). Soil: muddy. Turbidity: very low. Organic matter: very high.
Marsh 2 (Fig. 1-C). Large, irregularly shaped marsh about 500 m apart from the main road of the park. Sampling site location: 28°01' 10.01'' S, 58°04' 10.69'' W. Depth at sampling site: 5-10 cm. Insolation: very high. Vegetation: unidentified Gramineae, Schoenoplectus californicus (California bulrush), Salvinia sp. covering most of surface, Eichhornia azurea (anchored water hyacinth), some Pontederia cordata (pickerelweed). Soil: muddy. Turbidity: high. Organic matter: high.
Pond 2 (Fig. 1-D). Small rounded lagoon besides the main road, known as one of
the facing ponds. Location: 28°01' 17.30'' S, 58°03' 40.89'' W. Dimensions: 150 x 225 m. Depth at sampling site: 30-50 cm. Insolation: very high. Vegetation: Nymphoides indica, Ludwigia grandiflora (large-flower primrose-willow), Salvinia sp., Eichhornia sp., unidentified Gramineae. Soil: muddy. Turbidity: medium. Organic matter: medium.
Pond 3 (Fig. 1-G). Small rounded lagoon at the side of the main road of the park.
Location: 28°01' 05.89'' S, 58°03' 01.50'' W. Dimensions: 140 x 140 m. Depth at sampling site: 90 cm. Insolation: very high. Vegetation: covering 2/3 of the surface. Eichhornia sp., Salvinia sp., some Nymphoides indica. Soil: muddy. Turbidity: medium. Organic matter: medium.
Pond 4 (Fig. 1-I). The other facing pond, in front of pond 2. Location: 28°01' 29.95'' S, 58°03' 32.07'' W. Dimensions: 235 x
280 m. Depth at sampling site: 30-50 cm. Insolation: very high. Vegetation: Nymphoides indica, Ludwigia grandiflora, Eichhornia sp., unidentified Gramineae. Soil: muddy. Turbidity: medium. Organic matter: medium.
Pond 5 (Fig. 1-J). Rounded lagoon about 50 m aside from the main road. Location: 28° 01' 54.42'' S, 58°04' 40.48'' W. Dimensions: 220 x 250 m. Depth at sampling site: 30-50 cm. Insolation: very high. Vegetation: Eichhornia azurea, Schoenoplectus californicus (California bulrush), Nymphoides indica, Salvinia sp., Ludwigia sp. Soil: sandy Turbidity: medium. Organic matter: high.

Lotic habitats sampled

First stream heading to Portillo Gulley (Fig. 1-E). Several shallow ditches belonging to the first stream in the road to Portillo Gulley. Location: 28°01' 41.89'' S, 58°05' 49.13'' W. Depth at sampling site: 50 cm. Insolation: high. Vegetation: Nymphoides indica, Ludwigia grandiflora, Eichhornia crassipes, Cortaderia selloana (Pampas grass), Schoenoplectus californicus, unidentified Gramineae. Soil: muddy. Turbidity: medium. Organic matter: medium. Current: very slow.
Portillo Stream (Fig. 1-F). Main stream of the park, close to its entrance, running with NS direction between Fragosa Gulley and Santa Lucía Marsh. Sampling site location: 28° 02' 11.08'' S, 58°06' 33.27'' W. Insolation: very high. Depth at sampling site: 20-30 cm. Vegetation: Nymphoides indica, Eichhornia sp., Sagittaria montevidensis (giant arrowhead), Pontederia cordata, unidentified Gramineae. Soil: muddy. Turbidity: low. Organic matter: low. Current: slow.
Second stream heading to Portillo Gulley (Fig. 1-H). Location: 28°0' 9.10'' S, 58°05' 43.94'' W. Depth at sampling site: 20 cm. Insolation: very high. Vegetation: unidentified Gramineae, Salvinia sp., Pontederia cordata. Soil: muddy. Turbidity: medium. Organic matter: high. Current: very slow.

Insect sampling and taxonomic identification

Samples came from a field trip to Mburu-cuyá N. P. from 14 to 18 January 2008. Techniques employed to collect specimens included the use of aquatic nets (0,5-1,0 mm mesh), hand-picking, and mercury light traps (250 watts). When collecting in aquatic habitats, the net was passed from the bottom to the surface of water and from the shoreline to the reachable deepest edge of the water body. The vegetation was swept with the net for several minutes, and the content of the net was then transferred to a tray with clear water. Visible insects were sorted from debris and hand-picked with the aid of a pipette and fixed in small vials containing 96% ethyl alcohol. The remaining net contents were also fixed with 96% ethyl alcohol and examined later in the laboratory. The light traps were set in four occasions from sunset to past midnight in the surroundings of Santa Teresa Ranch. One particular light trap (17.i.2008) provided several thousand specimens. All insects collected at light were fixed with 96% ethyl alcohol.
Taxonomic identifications were done to the lowest possible taxonomic level, using available keys and literature (Young, 1974; Grosso, 1979; Trémouilles, 1989; Young, 1990; Grosso, 1993; Miller, 2000, 2001b, 2002; Oliva et al., 2002; Trémouilles et al., 2004, 2005; Miller, 2005; Archangelsky et al., 2009; Libonatti et al., 2011). The material is held in the collection of the Laboratory of Entomology, Buenos Aires University, Argentina.

RESULTS

In total, 128 species of Coleoptera included in 44 genera and seven families were identified (Table I). Dytiscidae was the richest family with regards to genera and species (20, 41), followed by Hydrophilidae (12, 38); Noteridae (6, 27); Scirtidae (2, 13), Gyrinidae (2, 2); Hydrochidae (1, 5), and Spercheidae (1, 1). Ten species are new for Argentina: Agaporomorphus mecolobus Miller and Bidessonotus obtusatus Régimbart (Dytiscidae); Mesonoterus laevicollis Sharp, Suphisellus hyeroglyphicus Zimmermann, S. rufulus Zimmermann, S. sexnotatus (Régimbart), and Notomicrus traili Sharp (Noteridae); Berosus pluripunctatus Mouchamps, Enochrus guarani Fernández and Helobata corumbaensis Fernández & Bachmann (Hydrophilidae).

Table I. List of species of aquatic Coleoptera collected at Mburucuyá National Park. Letters A-J refer to the habitats detailed in Fig. 1 and in Material and Methods.

Two families (Scirtidae and Spercheidae) and 12 genera (Dytiscidae: Copelatus Erichson, Laccophilus Leach; Noteridae: Mesonoterus Sharp, Pronoterus Sharp, Notomicrus Sharp; Hydrophilidae: Anacaena Thomson, Chaetarthria Stephens, Chasmogenus Sharp, Paracymus Thomson; Spercheidae: Spercheus Kugelann; Scirtidae: Ora Clark, Scirtes Illiger) are herein first recorded for Corrientes Province. Sixty-nine species (Table I) are first cited for Corrientes, rising to 165 the number of species of aquatic Coleoptera recorded for the province. Eighty-three taxa were collected exclusively with light traps; of these, 40 species are new records for the province.

DISCUSSION Suborder Adephaga

Family Dytiscidae

The Dytiscidae ae currently represented in the Argentinean fauna by 119 species in 31 genera (Libonatti et al., 2011). Dytiscids are abundant and widespread in Mburucuyá N. P., with 20 genera and 41 species (Table I) in four of the eight subfamilies present in Argentina. The subfamily Copelatinae was represented by two genera: Copelatus and Agaporomorphus Zimmermann. The collection of four species of Copelatus (Table I) is the first mention of the genus for Corrientes Province. Copelatus had been previously cited from Buenos Aires, Entre Ríos, Misiones, Salta, Jujuy and Chaco Provinces (Trémouilles, 1998; Torres et al., 2008). On the other hand, Agaporomorphus was recently cited as new for Argentina (Libonatti et al., 2011) based on material from Mburucuyá N. P. and from Misiones Province; here we report A. mecolobus Miller as new for the Argentinean fauna, which was only known from near São Paulo, Brazil (Miller, 2001b).
The subfamily Dytiscinae was represented by three genera: Thermonectus Dejean, Notaticus Zimmermann, and Megadytes Sharp. Thermonectus has five species in Argentina. Both T. nobilis Zimmermann and T. succinctus (Aubé) (Table I) had been previously recorded from Corrientes (Trémouilles, 1989). However, this is the first concrete locality record for T. nobilis. The single species of Notaticus, N. fasciatus Zimmermann, has been cited from northern Argentina, and in particular from two localities in Corrientes Province, Santo Tomé (Trémouilles & Bachmann, 1981) and Mburucuyá N. P. (Michat & Alarie, 2009). The genus Megadytes was only collected as larvae in this study. Although several species of Megadytes were expected in the area, only M. fallax (Aubé) has been cited from Mburucuyá N. P. (Michat, 2010).
Hydrodytinae was erected by Miller (2001a, 2002) to include several species previously placed in Agaporomorphus, and was recently reported as new for Argentina by Libonatti et al. (2011). Only Hydrodytes opalinus (Zimmermann), a widespread species distributed from northern South America to Honduras (Miller, 2002), is present in Corrientes and Misiones Provinces.
The subfamily Hydroporinae accounted for 12 genera and 23 species in this study (Table I). Libonatti et al. (2011) mentioned
Bidessodes Régimbart and Queda Sharp as first records for Argentina, and Anodocheilus Babington, Bidessonotus Régimbart, Derovatellus Sharp, Desmopachria Babington, Hemibidessus Zimmermann and Pachydrus Sharp as first records for Corrientes Province, without giving reference to a specific locality. However, all these findings are based on material coming from this study. Three specimens of Queda were captured in Portillo Stream (Fig. 1, F), which represents the first note about the habitat of the genus. We also enlarge the distributional range of Bidessonotus obtusatus Régimbart, which was known only from Brazil and Paraguay (Nilsson, 2001). Other novelties for Corrientes Province are Hydrovatus crassulus Sharp (known from Chaco Province) and four species of Desmopachria (Table I). Desmopachria concolor Sharp was known from Buenos Aires, Entre Ríos, Santa Fe and Chaco Provinces (Trémouilles, 1998).
Laccophilinae is a speciose subfamily represented in Argentina by Laccophilus and an unpublished genus (Toledo & Michat, in prep.) that occurs in Chaco, Corrientes and Entre Ríos Provinces (Libonatti et al., 2011). Both genera were collected in this study (Table I). Laccophilus is frequent and widespread in Argentina; however, this is the first record for Corrientes Province. The genus is in need of a taxonomic revision for the Neotropical Region; seven unidentified species were captured in this survey (Table I).

Family Gyrinidae

The family is represented in Argentina by three genera (Andogyrus Ochs, Gyretes Brullé and Gyrinus Müller) and about 26 species (Michat & Archangelsky, in press). We found two species in this study (Table I). Gyrinus violaceus Régimbart is reported as new for Corrientes Province. This species, known from Argentina, Brazil, and Uruguay, had been previously cited from Salta and Misiones Provinces (Bruch, 1927; Fernández et al., 2008; Michat & Archangelsky, in press).

Family Noteridae

Noterids include six genera in Argentina, all of which were captured in Mburucuyá N. P. (Table I). Twenty-seven species were recognized, five of which are new records for the country: Mesonoterus laevicollis, Suphisellus hyeroglyphicus, S. rufulus, S. sexnotatus, and Notomicrus traili. Also, three genera are regarded as new to Corrientes Province: Mesonoterus, Pronoterus, and Notomicrus. The genus Mesonoterus has four species in total and two in the Argentinean fauna, M. crassicornis (Régimbart) cited by Zimmermann (1920) but without giving any concrete locality, and M. laevicollis, distributed in Bolivia, Brazil, Cuba, Guatemala, Mexico, Paraguay, and new to the Argentinean fauna. Our record constitutes the first locality for the genus in Argentina. On the other hand, the genus Pronoterus has three species, of which only P. punctipennis Sharp is distributed in South America (Argentina and Brazil); this species was previously cited from Buenos Aires and Chaco Provinces (Benetti et al., 2003) and our record is the first for Corrientes Province. Within Notomicrus, nine species are recognized (Nilsson & van Vondel, 2005). Only N. brevicornis Sharp had been previously cited from Argentina (Santa Fe Province) (Bruch, 1927); this is the first record for Corrientes. Notomicrus traili is added to the Argentinean fauna; it was known from Brazil, Bolivia and Suriname (Nilsson & van Vondel, 2005). Nineteen species of Suphisellus are known to inhabit Argentina (Bruch, 1915, 1927; Benetti et al., 2003). Suphisellus flavopictus (Régimbart) and S. grossus (Sharp) were mentioned for the Paraná River floodplain (Poi de Neiff & Neiff, 2006), but only S. remator (Sharp) has been precisely cited for Corrientes (Bruch, 1915, 1927). Here we report 11 Suphisellus species new to the fauna of the province (Table I) as well as three species new to Argentina: S. hyeroglyphicus and S. sexnotatus, known from Brazil, and S. rufulus, known from Bolivia and Brazil. For S. curtus (Sharp) this is the first locality record in Argentina. Hydrocanthus Say is represented by six species in Argentina (Nilsson & van Vondel, 2005). H. sharpi Zimmermann and H. socius Sahlberg in Mburucuyá N. P. constitute the first records for Corrientes Province; both species had been cited only from Chaco Province (Benetti et al., 2003), though Bruch (1915, 1927) mentioned H. sharpi (under the names H. atripennis Say and H. iricolor Say) for Buenos Aires and Misiones Provinces. The genus Suphis Aubé is represented by six species in Argentina, of which only S. cimicoides Aubé and S. notaticollis Zimmermann were previously recorded from Corrientes Province (Grosso, 1993). We report the occurrence of S. freudei Mouchamps as new to Corrientes; this species was known from Buenos Aires and Chaco Provinces (Grosso, 1993).

Suborder Polyphaga

Family Hydrochidae

A monogeneric family with about 180 worldwide species and represented by 17 species in Argentina, most of them distributed in the northern part of the country (Oliva et al., 2002). Several species have been cited from Corrientes Province (Fernández & Bachmann, 1998). In this paper we report the presence of H. ducalis Knisch as new to Corrientes; it is also known from Buenos Aires, Santa Fe, Chaco and Formosa Provinces. Other species occurring in Mburucuyá N. P. are H. purpureus Knisch and H. pumilio Knisch (Oliva, pers. obs, 1997).

Family Hydrophilidae

The argentinean hydrophilids are represented by approximately 150 species in 22 genera and two subfamilies. Thirty-eight species in 12 genera where recorded in this study, all belonging to the subfamily Hydrophilinae (Table I). The tribe Acidocerini was represented in our samplings by the four genera occurring in Argentina: Chasmogenus Sharp, Enochrus Thomson, Helobata Bergroth and Helochares Mulsant. For Chasmogenus, this is the first record for Corrientes Province. The sole species of Chasmogenus present in Argentina, C. sapucay Fernández, was previously known from Formosa and Santiago del Estero Provinces (Oliva et al., 2002). Within Enochrus, E. guarani is recorded as new to Argentina. This species was known from Paraguay (Fernández, 1988). Also, Enochrus circumcinctus (Bruch), widespread in the northern half of Argentina, and E. sublongus (Fall), only known to occur in Chaco and Formosa Provinces, are herein listed as new for Corrientes. With regards to Helobata, H. corumbaensis is new to Argentina; this species was known from Brazil (Fernández & Bachmann, 1987). In addition, H. confusa Fernández & Bachmann, cited only from Entre Ríos Province, is reported here as first record for Corrientes. Eight species of Helochares were collected in the park (Table I), of which two are new to Corrientes Province: H. chaquensis Fernández, known to occur in Chaco and Formosa Provinces, and H. pallipes (Brullé), previously recorded from Buenos Aires, Entre Ríos, Santa Fe and Santiago del Estero Provinces (Oliva et al., 2002).
The tribe Anacaenini has two genera in Argentina, Anacaena and Paracymus, both of which are first records for Corrientes Province. Komarek (2005) recently revised the Neotropical Anacaena and listed A. suturalis (Leconte) from Buenos Aires and Santiago del Estero Provinces, and A. coruscalis Orchymont from Buenos Aires; Torres et al. (2008) cited an unidentified species from Jujuy. On the other hand, Paracymus is in need of a revision; according to Oliva et al. (2002) and Torres et al. (2007) four species are known from Argentina but many others are expected to occur. The known localities for Paracymus are in Buenos Aires, Entre Ríos, Misiones, Tucumán and San Luis Provinces, and our record is the first for Corrientes.
Within the tribe Berosini, two genera were collected: Berosus Leach and Derallus Sharp (Table 1). Berosus is a speciose genus, accounting for 39 species in Argentina (Oliva et al., 2002). Sixteen species were already known from Corrientes (Oliva et al., 2002) and here we cite one species new to Argentina (B. pluripunctatus) and one new to Corrientes (B. pedregalensis Jensen-Haarup). Berosus pluripunctatus was known from Venezuela and Brazil.
The genus Chaetarthria is the only
representative of the tribe Chaetarthriini in Argentina with six species. The occurrence of two unidentified species in Mburucuyá N. P. enlarges the distribution of the genus to Corrientes Province, being the first record for the Mesopotamia.
The tribe Hydrophilini was represented in this study by three genera and nine species (Table I), most of them widespread and frequent. Tropisternus collaris (Fabricius), a common species in South America and the northern part of Argentina, is first cited for Corrientes Province.

Family Spercheidae

This monogeneric family has 18 species and two in the Neotropical Region, Spercheus fimbriicollis Bruch and S. halophilus Archangelsky, both present in Argentina. The occurrence of S. fimbriicollis in Mburucuyá N. P. represents the first record of the family for Corrientes Province. Spercheus fimbriicollis distributes in Buenos Aires, Santa Fe, Chaco and Formosa Provinces, whereas S. halophilus is only known from Córdoba Province.

Family Scirtidae

In spite of being cosmopolitan and including 1300 species in almost 50 genera all over the world, the current state of knowledge of this family is extremely deficient in the Neotropical Region (Klausnitzer, 2009). Only 11 species and four genera (Cyphon Paykull, Microcara Thomson, Ora Clark and Scirtes Illiger) have been cited so far in Argentina (Trémouilles et al., 1995). In this study, we report the finding of seven unidentified species of Scirtes and six unidentified species of Ora, representing the first records of the family and those genera for Corrientes Province.

ACKNOWLEDGEMENTS

We thank the Administration of National Parks and the park guard Hector Bal, for allowing and helping us with our activities in Mburucuyá National Park. We also thank two anonymous referees for their comments on the manuscript. This work was funded by projects PIP 112-200801-02759 (CONICET), PICT-2007-01438 and PICT-2010-0526 (ANPCyT) and UBACyT-20020090300135 (UBA).

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