Introduction
Coenogonium Ehrenb. is characterized by the formation of biatorine apothecia with a yellow to orange or brown disc, a paraplectenchymatous excipulum, unitunicate asci with entirely thin walls, uniseptate (rarely simple), colourless ascospores, and a trentepohlioid photobiont (Rivas Plata et al. Reference Rivas Plata, Lücking, Aptroot, Sipman, Chavez, Umaña and Lizano2006; Lücking Reference Lücking2008). According to its current circumscription, it comprises c. 130 species of mainly tropical to subtropical lichens, with a filamentous or crustose thallus, previously divided into the genera Coenogonium s. str. and Dimerella (Lücking & Kalb Reference Lücking and Kalb2000; Rivas Plata et al. Reference Rivas Plata, Lücking, Aptroot, Sipman, Chavez, Umaña and Lizano2006; Lücking Reference Lücking2008). This transfer has not been accepted by all workers (Vězda Reference Vězda2004; Lackovičoca & Guttová Reference Lackovičoca and Guttová2005). Molecular studies, however, based on DNA sequences from the nuclear small and large subunit ribosomal RNA genes, support the synonymy of Coenogonium and Dimerella (Kauff & Lutzoni Reference Kauff and Lutzoni2002). Recent evidence based on ontogenetic, anatomical and morphological studies also shows that the genus belongs in its own family Coenogoniaceae (Kauff & Büdel Reference Kauff and Büdel2005; Baloch et al. Reference Baloch, Lücking, Lumbsch and Wedin2010).
In South America, c. 55 species of Coenogonium have been recorded so far (Lücking Reference Lücking2008; Feuerer Reference Feuerer2011), but there are some countries where the genus is still poorly known. In Argentina, 14 species of this genus have been recorded, and in Paraguay, only eight (Ferraro Reference Ferraro1978; Calvelo & Liberatore Reference Calvelo and Liberatore2002; Lücking Reference Lücking2008; Feuerer Reference Feuerer2011; Lumbsch et al. Reference Lumbsch, Ahti, Altermann, Amo De Paz, Aptroot, Arup, Bárcenas Peña, Bawingan, Benatti and Betancourt2011).
As a result of the study of collections from Argentina and Paraguay 27 species of Coenogonoium (six filamentous and 21 crustose) have been studied; four new species are described and illustrated, five species are recorded for the first time for South America, and the distribution of the existing 24 species is extended, substantially increasing the knowledge of this genus in southern South America. A revision of the species of this genus from both countries is also presented.
Material and Methods
Most of the specimens studied were collected by the authors in Argentina and Paraguay. They are deposited in the herbaria CTES, FCQ, LG, PY, and Czeczuga-Bialistok. The morphological characters were examined using stereoscopic and light microscopes (Leica MZ6 and Leica CME respectively). Apothecia and pycnidia, when present, were sectioned by hand with a razor blade and then mounted in 5% KOH and Lugol's solution for examination.
The New Species
Coenogonium albomarginatum Michlig & L. I. Ferraro sp. nov.
MycoBank No.: MB563945
Differing from Coenogonium pyrophthalmum by its smaller apothecia.
Type: Argentina, Prov. Corrientes, Dept. San Cosme, 3 km from crossing to Paso de la Patria, on the way to Puerto González, 21 May 1979, L. I. Ferraro et al. 1928 (CTES—holotype).
(Fig. 1A)
Fig. 1. New species of Coenogonium from southern South America. A & B, C. verrucosum (type); A, thallus with apothecia; B, wart-shaped pycnidia. C, C. albomarginatum (type), thallus with apothecia. D, C. flavovirens (type), thallus with apothecia. Scale=0·5 mm.
Thallus crustose, corticolous, verrucose, yellowish greyish green, 25–45 mm diam.; prothallus white, shiny, conspicuous.
Apothecia sessile, rounded, rarely slightly irregular in outline, 0·2–0·6 mm diam., 200–215 µm high; disc pale orange, flat to slightly concave first, then flat to rarely convex; margin thick, prominent, cream-coloured, paler than the disc. Excipulum paraplectenchymatous with radiating cell rows, I−. Hypothecium 26 µm high, colourless, I+ yellow. Hymenium 65 µm high, colourless, I+ blue turning greenish. Ascospores uniseriate, oblong-elongated, 1-septate, 8–11×2·5–3·0 µm.
Pycnidia absent.
Notes
The specific epithet of this new species refers to the whitish and shiny prothallus. It is close to Coenogonium pyrophthalmum (Mont.) Lücking et al., but differs in having smaller apothecia (0·2–0·6 mm diameter).
Coenogonium brasiliense L. I. Ferraro & Michlig sp. nov.
MycoBank No.: MB803411
Differing from Coenogonium dilucidum by the paler thallus and the larger and oblong ascospores.
Type: Argentina, Prov. Corrientes, Dept. Santo Tomé, Ea. Timbó, coast of Uruguay River, 26 km SE of Garabí colony, in lowland, on Styrax leprosum leaves, 18 November 1980, A. Schinini 20984 (CTES—holotype).
Thallus crustose, foliicolous, smooth, greyish to whitish green, extended, 3–13 mm, distributed into confluent patches, generally with a sinuous margin, somewhat raised above the substratum in some regions.
Apothecia with a constricted base, rounded, 1·0–1·5 mm diam.; disc pale brown to reddish yellow, flat, transparent; margin inconspicuous. Excipulum paraplectenchymatous, irregular cells, colourless, c. 3–5 µm diam. Hypothecium 24 µm high. Hymenium 42–50 µm high, I+ yellow, paraphysis with widened apices. Asci clavate, 45–50 µm long, 3–4 µm wide. Ascospores uniseriate, oblong, slightly wider in the central region, 1-septate, 9–11×2–3 µm.
Notes
Coenogonium brasiliense is based on Dimerella brasiliensis Vězda & Kalb, nomen nudum. This name, used for a long time, was never formerly described. It was mentioned in a checklist of lichens from Paraguay (Ferraro Reference Ferraro1990). This species is close to Coenogonium dilucidum (Kremp.) Kalb & Lücking in the colour and transparent aspect of the apothecia, especially the younger ones which are also shiny. Both species differ in the shape and size of the ascospores and the colour of the thallus. The ascospores in C. brasiliense are larger, oblong, and widened in the central region.
Additional material examined. Argentina: Corrientes: Ituzaingó, mouth of Garapé stream on Paraná River, 45 km E of Ituzaingó, on orange leaves, on a grown wild orange, inside forest, 1975, Ferraro et al. 710 (CTES).—Paraguay: Amambay: Cerro Guazú, 26°5′S; 56°W, over Thelypteris sp. leaves (Pteridophyta), 1980, Schinini et al. 20730 (CTES). Guairá: Ybituruzú mountain range, over ferns, 1979, Schinini & Bordas 19798 (CTES).
Coenogonium flavovirens L. I. Ferraro & Michlig sp. nov.
MycoBank No.: MB563946
Differing from Coenogonium aciculatum by the yellowish green thallus and smaller ascospores.
Type: Argentina, Prov. Salta, Dept. Santa Victoria, Los Toldos, El Nogalar de Los Toldos Reserve, 24 April 2009, L. I. Ferraro et al. 9079 (CTES—holotype).
(Fig. 1B)
Thallus crustose, corticolous, continuous, thin, smooth, yellowish green, 30 mm diam; prothallus absent. Photobiont Trentepohlia, cells angular to rounded, densely grouped in some areas of the thallus.
Apothecia sessile, rounded, 1·5–3·0 mm diam., 100–220 µm high; disc plane, intense yellow to yellow-orange; margin thin, not prominent, cream-coloured, paler than the disc. Excipulum paraplectenchymatous with radiating cell rows. Hypothecium 20–30 µm high, pale yellow, I+ yellow. Hymenium 50–60 µm high, colourless, I+ pale blue then red. Ascospores biseriate, oblong, 1-septate, 15–17×2·0–2·5 µm.
Pycnidia absent.
Notes
The specific epithet of this new species refers to the yellow greenish colour of its thallus. Most species of Coenogonium are characterized by the presence of a green to greyish green or brownish thallus. The species is otherwise characterized by its large, bright yellow apothecia and long, narrow ascospores, which resemble C. aciculatum Lücking & Aptroot. The latter differs in its greenish grey thallus and narrower ascospores (1·5–2·0 µm wide). Most similar are C. subsquamosum (Aptroot & Seaward) Lücking et al. from the Seychelles and C. pertenue (Stirt.) Kalb & Lücking from Australia, but both have broader ascospores (c. 3–4 µm broad).
Coenogonium verrucosum Michlig & L. I. Ferraro sp. nov.
MycoBank No.: MB563947
Differing from other species of Coenogonium by the verrucose thallus, yellow prothallus, medium sized apothecia, wart-shaped pycnidia, and non-septate, ellipsoid-bacillar conidia.
Type: Argentina, Prov. Corrientes, Dept. Ituzaingó, Apipé Grande island, Punta Arazá, in marginal forest, on bark of Ocotea sp., 26 November 1988, Ferraro et al. 3606 (CTES—holotype).
Thallus crustose, corticolous, verrucose, thin, brownish green; prothallus yellow.
Apothecia sessile, rounded to slightly irregular in outline, (0·2)0·3–0·8(1·0) mm diam., 40–50 µm high; disc slightly concave first then flat, orange; margin thick and prominent first, then thin and not prominent, cream-coloured to pale orange. Excipulum paraplectenchymatous with radiating cell rows, 40–50 µm high, colourless, I+ pale yellow. Hypothecium 25 µm high, colourless, I+ pale yellow. Hymenium 40–50 µm high, colourless, I+ blue then quickly sordid green. Ascospores irregularly biseriate, ellipsoid, 1-septate, 8–9×2·5–3·0 µm.
Pycnidia present, wart-shaped; conidia broadly ellipsoid-bacillar, non-septate, 3–5×1–2 µm.
Notes
This new species is characterized by the presence of a yellow prothallus, medium-sized apothecia, wart-shaped pycnidia, non-septate ellipsoid-bacillar conidia, and a verrucose thallus. These verrucae are formed by calcium oxalate crystals. Other species with a verrucose thallus are Coenogonium degeneri (Kalb & Vězda) Kalb & Lücking and C. tavaresianum (Vězda) Lücking et al. & Sipman, but these differ in the lack of a prothallus and larger ascospores. A yellow prothallus is also found in C. atroluteum (Vain.) Lücking et al., but it differs in its pale yellow-brown apothecia with dark margin and a smooth thallus. Other similar species are C. zonatum (Müll. Arg.) Kalb & Lücking and C. subluteum (Rehm) Kalb & Lücking, but both differ in the presence of uniseptate conidia. The former is also distinguished by its white prothallus. Coenogonium roumeguerianum (Müll. Arg.) Kalb also has apothecia with a concave orange disc but differs in the presence of a white prothallus and broader ascospores (4–6 µm broad).
New records for Argentina and Paraguay
The species listed below represent 14 species recorded here for the first time from Argentina, and five from Paraguay. The new records for Argentina and Paraguay are indicated with a circle (•) and a square (▪), respectively. The known species of Coenogonium in both countries are shown in Table 1.
Table 1. List of reported Coenogonium species from Argentina (ARG) and Paraguay (PAR). The contributions of this paper are in bold.
Coenogonium atroluteum (Vain.) Lücking et al.
Fungal Diversity 23: 312 (2006).
This species has been previously recorded from Argentina (Calvelo & Liberatore 2002) and Paraguay (Feuerer Reference Feuerer2011).
Selected specimes examined. Paraguay: Misiones: West of Yacyretá island, in forest, 1988, Ferraro 3576 (CTES).
Coenogonium confervoides Nyl. (•)
Flora 41: 380 (1858).
In South America, this species has previously been reported only from Colombia (Sipman et al. Reference Sipman, Hekking and Aguirre2008).
Selected specimen examined. Argentina: Misiones: San Pedro, Yaboty Biosphere Reserve, Moconá Provincial Park, Chachi trail, over a fallen branch on the side of the trail, 2009, Michlig & Niveiro 1886 (CTES).
Coenogonium eximium (Nyl.) Kalb & Lücking (•)
Fungal Diversity 23: 297 (2006).
In South America, this species has previously been reported only from Colombia (Sipman et al. Reference Sipman, Hekking and Aguirre2008).
Selected specimen examined. Argentina: Corrientes: San Cosme, Paso de la Patria, San Juan stream and Paraná River, in marginal forest, 1979, Ferraro 1688 (CTES).
Coenogonium implexum Nyl. (•)
Ann. Sci. Nat. Bot. Ser. 4, 16: 92 (1862).
In South America, this species has been reported only from Colombia (Sipman et al. Reference Sipman, Hekking and Aguirre2008).
Selected specimen examined. Argentina: Corrientes: Mburucuyá, Potrero 1 grande, over a fallen tree on the edge of the swamp, 2006, Ferraro et al. 8126 (CTES).
Coenogonium interplexum Nyl. (▪)
Ann. Sci. Nat. Bot. Ser. 4, 16: 92 (1862).
This is the first record of this species for north-western Argentina; it was previously recorded from Buenos Aires, Corrientes, and Misiones Provinces (Calvelo & Liberatore Reference Calvelo and Liberatore2002).
Selected specimens examined. Argentina: Salta: Santa Victoria, Los Toldos, El Nogalar de Los Toldos Reserve, 2009, Ferraro 9076 (CTES).—Paraguay: Paraguarí: Ybicuí National Park, 1990, Pérez 841 (CTES, PY).
Coenogonium interpositum Nyl.
Ann. Sci. Nat. Bot. Ser. 4, 16: 91 (1862).
This is the first record of this species for northern Argentina; it was previously recorded from Buenos Aires province (Calvelo & Liberatore Reference Calvelo and Liberatore2002). In Paraguay, it has been recorded by Malme (Reference Malme1937).
Selected specimens examined. Argentina: Misiones: San Pedro, Yaboty Biosphere Reserve, Moconá Provincial Park, Piedra Bugre wharf, 2008, Michlig et al. 897 (CTES).—Paraguay: Misiones: West of Yacyretá island, in scrubland, near a sandy area, 1988, Ferraro & Popoff 3575 (CTES, FCQ, LG).
Coenogonium isidiatum (G. Thor & Vězda) Lücking et al. (•)
Fungal Diversity 23: 297 (2006).
This species is recorded here for the first time from South America.
Selected specimen examined. Argentina: Corrientes: Empedrado, 12 road and Pehuajó stream, in an islet of scrubland degraded by grazing, 2007, Michlig et al. 265 (CTES).
Coenogonium isidiigerum (Vězda & Osorio) Lücking et al.
Fungal Diversity 23: 297 (2006).
This is the first record of this species for northern Argentina; it has been previously reported from Buenos Aires province (Vězda Reference Vězda1989).
Selected specimen examined. Argentina: Misiones: San Ignacio, 1 km from Quiroga's home, on way to the river, 1981, Ferraro et al. 2240 (CTES).
Coenogonium linkii Ehrenb. (▪)
Horae Phys. Berol.: 120 (1820).
This is the first record of this species for north-western Argentina; it was previously recorded from Misiones province (Calvelo & Liberatore Reference Calvelo and Liberatore2002).
Selected specimens examined. Argentina: Salta: entrance to the El Rey National Park, over branches with moss, 2007, Ferraro 8317 (CTES).—Paraguay: Guairá: Independencia Colony, alt. 250 m, on bark of Sebastiania sp., 1986, Schinini 25356 (CTES, LG). Caazapá: Yute District, 15 km S of Capitindy, clearing in the forest, edge of the stream, flooded grassland, 1987, Arbo 2920 (CTES, LG). Alto Paraná: Ytabó Biological Reserve, Tangará trail, Krapovickas 43411 (CTES, hb. Czeczuga-Bialistok, LG). Amambay: 7 km N of 5 road, western boundary of Cerro Corá National Park, 1988, Ferraro et al. 3480 (CTES, LG).
Coenogonium luteocitrinum Rivas Plata et al. (•)
Fungal Diversity 23: 283 (2006).
In South America, this species has previously been recorded only from Bolivia (Flakus Reference Flakus2008).
Selected specimens examined. Argentina: Corrientes: Mburucuyá, Mburucuyá National Park, Maizal paddock, in the palm grove of Butia yatay, 2006, Ferraro & Popoff 8183 (CTES). Salta: El Rey National Park, at the entrance of the trail to the cascade ‘Los Lobitos’, 2005, Ferraro et al. 7934 (CTES).
Coenogonium luteum (Dicks.) Kalb & Lücking (▪)
Bot. Jahrb. Syst. 122: 32 (2000).
This is the first record of this species for north-western Argentina; it was previously recorded from the Corrientes, Misiones, and Río Negro provinces (Calvelo & Liberatore Reference Calvelo and Liberatore2002).
Selected specimens examined. Argentina: Jujuy: Calilegua National Park, Aguas negras Camping, over bark, 2005, Ferraro et al. 7584 (CTES). Salta: Santa Victoria, Los Toldos, El Nogalar Reserve, 2009, Ferraro 9074 (CTES). Neuquén: Nahuel Huapí National Park, trail from de Blest to Los Cántaros, 2001, Ferraro & Giaiotti 6510 (CTES).—Paraguay: Guairá: Independencia Colony, 250 m, over leaves of Trichilia catigua, in forest, 1986, Schinini 25324 (CTES). Itapúa: Yacyretá Island, over leaves of Chrisophyllum gonocarpum, 1988, Ferraro 3654 (CTES).
Coenogonium magdalenae Rivas Plata et al. (•)
Fungal Diversity 23: 286 (2006).
This species is recorded here for the first time from South America.
Selected specimens examined. Argentina: Corrientes: Santo Tomé, Santo Tomé, first section of chakras, in shrubland, 1978, Ferraro 1368 (CTES).
Coenogonium moniliforme Tuck. (•)
Proc. Amer. Acad. Sci. 5: 416 (1862).
This species has been recently recorded from Paraguay (Lücking Reference Lücking2008).
Selected specimens examined. Argentina: Misiones: General Manuel Belgrano, San Antonio Strict Nature Reserve, trail that marks the beginning of the protected area, over a tree in the forest, 2009, Michlig & Niveiro 1955 (CTES).
Coenogonium nepalense (G. Thor & Vězda) Lücking et al. (•)
Fungal Diversity 23: 297 (2006).
In South America, this species has previously been recorded only from Brazil (Cáceres Reference Cáceres2007).
Selected specimens examined. Argentina: Corrientes: Mburucuyá, Mburucuyá National Park, Tung paddock, 2006, Ferraro 8166 (CTES). Misiones: San Pedro, Yaboty Biosphere Reserve, Esmeralda Provincial Park, on the way to pine forest, over a tree on the side of the trail, 2008, Michlig et al. 757 (CTES).
Coenogonium persistens (Malme) Lücking et al. (•)
Fungal Diversity 23: 297 (2006).
This species is here recorded for the first time from South America.
Selected specimens examined. Argentina: Corrientes: Mburucuyá, Mburucuyá National Park, Yatay trail, 2006, Ferraro 8042 (CTES); Concepción, 118 route, 13 km from Florida, Paraje PUCE, 2008, Ferraro 8492 (CTES). Misiones: Iguazú, Iguazú National Park, Macuco trail from the CIES to the Arrechea Falls, over bark, 2003, Ferraro & Popoff 6901 (CTES).
Coenogonium pusillum (Mont.) Lücking et al. (▪)
Fungal Diversity 23: 298 (2006).
This species is recorded here for the first time from South America. It has been previously reported from Costa Rica (Rivas Plata et al. Reference Rivas Plata, Lücking, Aptroot, Sipman, Chavez, Umaña and Lizano2006) and Cuba (Montagne Reference Montagne and de la Sagra1842).
Selected specimen examined. Paraguay: Misiones: W of the Yacyretá island, in the forest, 1988, Ferraro 3567 (CTES).
Coenogonium pyrophthalmum (Mont.) Lücking et al.
Fungal Diversity 23: 314 (2006).
This species has been previously recorded in Argentina from the Corrientes and Buenos Aires provinces (Calvelo & Liberatore Reference Calvelo and Liberatore2002).
Selected specimen examined. Argentina: Misiones: Iguazú, Apepú Reserve, 15 km from the entrance, 1982, Ferraro 2497 (CTES).
Coenogonium roumeguerianum (Müll. Arg.) Kalb
Sched. Lich. Neotrop. 13: 3 (2001).
This species has been previously recorded in Argentina from Misiones province (Calvelo & Liberatore Reference Calvelo and Liberatore2002), and Paraguay (Feuerer Reference Feuerer2011).
Selected specimens examined. Argentina: Corrientes: Empedrado, Las Tres Marías Ranch, in marginal forest of the Paraná River, 1979, Ferraro 1742b (CTES); San Martín, Drews rice paddy, Cambá Trapo swamp, in scrubland over Scutia sp. (Mirtaceae), 1976, Ferraro & Tressens 947 (CTES).
Coenogonium siquirrense (Lücking) Lücking (•)
Flora Neotropica 103: 580 (2008).
In South America, this species has been recorded from Ecuador, Peru (Lücking Reference Lücking2008), and Venezuela (Feuerer Reference Feuerer2011).
Selected specimens examined. Argentina: Misiones: Iguazú, Iguazú National Park, Ñandú Camping, over mosses, 2004, Ferraro & Popoff 7401 (CTES).
Coenogonium strigosum Rivas Plata et al. (•) (▪)
Fungal Diversity 23: 290 (2006).
In South America, this species has previously been recorded from Brazil (Cáceres Reference Cáceres2007).
Selected specimens examined. Argentina: Corrientes: General Paz, 26 km W of Caá Catí, over Celtis sp., 1978, Ahumada 2483 (CTES); Capital, Molina Punta, in scrubland on bank of the river, 1978, Ferraro 1327 (CTES); Santo Tomé, Bertrán (Infrán Cué) Ranch, 23 km SW of Virasoro, in an islet of forest, 1992, Tressens et al. 4155 (CTES). Misiones: Iguazú, Iguazú National Park, 101 road, near Garganta del Diablo, over bark, 2003, Ferraro & Popoff 7083 (CTES); San Ignacio, Teyú Cuaré, 1981, Ferraro 2362 (CTES). Salta: Santa Victoria, Los Toldos, El Nogalar Reserve, 2009, Ferraro et al. 9075 (CTES).—Paraguay: Misiones: W of Yacyretá island, in open path in forest, 1988, Ferraro 3568 (CTES).
Coenogonium subdentatum (Vězda & G. Thor) Rivas Plata et al. (•)
Fungal Diversity 23: 298 (2006).
Selected specimen examined. Argentina: Misiones: General Manuel Belgrano, San Antonio Strict Nature Reserve, trail that marks the beginning of the protected area, over a tree in the forest, 2009, Michlig & Niveiro 1947 (CTES).
Coenogonium subluteum (Rehm) Kalb & Lücking
Bot. Jahrb. Syst. 122: 34 (2000).
This is the first record of this species for north-western Argentina; it was previously recorded from the Corrientes and Misiones provinces (Calvelo & Liberatore Reference Calvelo and Liberatore2002). In Paraguay, it has been previously recorded by Ferraro (Reference Ferraro1997) and Lücking (Reference Lücking2008).
Selected specimens examined. Argentina: Jujuy: Ledesma, Calilegua National Park, La Lagunita Trail, 2005, Ferraro et al. 7820 (CTES).—Paraguay: Cordillera: Valenzuela, Yhacá stream, in marginal forest, foliicolous, over Myrtaceae, 1989, Ferraro et al. 3917 (CTES).
Coenogonium weberi (Vězda) Lücking et al. (•)
Fungal Diversity 23: 298 (2006).
This species is recorded here for the first time from South America.
Selected specimens examined. Argentina: Corrientes: Mburucuyá, Mburucuyá National Park, Santa Teresa Ranch, 2006, Ferraro 8093 (CTES). Jujuy: Ledesma, Calilegua National Park, Tataupa trail, over bark in forest, 2005, Ferraro et al. 7747 (CTES).
Coenogonium zonatum (Müll. Arg.) Kalb & Lücking (•)
Bot. Jahrb. Syst. 122: 34 (2000).
In South America, this species has been recorded from Paraguay (Lücking Reference Lücking2008; Feuerer Reference Feuerer2011) and Colombia (Sipman et al. Reference Sipman, Hekking and Aguirre2008).
Selected specimens examined. Argentina: Corrientes: Mburucuyá, Mburucuyá National Park, Potrero 17, at the edge of Santa Lucia swamp, 2006, Ferraro 8215 (CTES).—Paraguay: Itapúa: Tirol Hotel, foliicolous, over leaves of Monstera, 1985, Krapovickas & Cristóbal 40084a (CTES).
The authors specially wish to thank Dr Robert Lücking for critically reading this manuscript, and for discussing the identification of some species during the GLAL IX. We are also grateful to the Myndel Botanica Foundation, the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), the Secretaría General de Ciencia y Técnica (SGCyT-UNNE), and the Área de Manejo Integral Reserva de Biosfera Yaboty (AMIRBY) for financial support of our fieldwork. We also wish to thank the authorities of the National Parks of Argentina for providing collection permits.
