Occurrence of Arbuscular Mycorrhizal Fungi in High Altitude Sites of the Patagonian Altoandina Region in Nahuel Huapi National Park (Argentina)
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Acta Botanica Brasilica - 30(4): 521-531. October-December 2016. doi: 10.1590/0102-33062016abb0223 Occurrence of arbuscular mycorrhizal fungi in high altitude sites of the Patagonian Altoandina region in Nahuel Huapi National Park (Argentina) María Silvana Velázquez 1*, Sidney Luiz Stürmer 2, Clara Bruzone 3, Sonia Fontenla 3, Marcelo Barrera 4 and Marta Cabello 1 Received: June 22, 2016 Accepted: September 3, 2016 . ABSTRACT Knowledge of the occurrence and diversity of arbuscular mycorrhizal fungi (AMF) in National Parks is essential for the establishment of policies for conservation. Th e aim of this study was to characterize the AMF communities in the Patagonian Altoandina region in Nahuel Huapi National Park, Argentina. We surveyed AMF spores associated with the rhizospheres of 9 plant species in the Patagonian Steppe (PS), Challhuaco Hill (ChH), Catedral Hill (CH), and Tronador Hill (TH) regions and detected a total of 27 Glomeromycota species. Acaulospora laevis was dominant at all sites. Th e AMF community was dominated by Acaulosporaceae, as regards the number of species and contribution of each one to the total number of spores. Th ree Glomeromycota families were detected at PS, the site with the lowest elevation; whereas fi ve to six families were detected at ChH, CH, and TH. Cluster analysis indicated that the AMF communities were grouped according to habitat. We concluded that certain patterns of the AMFcommunity structure detected were equivalent to those of high-altitude environments from other studies, while others were unique to the Patagonian region; thus suggesting that historical infl uences like dispersion and speciation played a critical role in shaping AMF community composition in such high-altitude environments. Keywords: Acaulosporaceae, altitudinal gradient, Glomeromycota, species diversity, spore numbers, steppe wide biologic diversity because of substantial environmental Introduction variability; and are generally not modifi ed by anthropic interference (Ferreyra et al. 2006). Th is last feature feature High-mountain ecosystems extend above the upper limit gives those elevated landscapes a sacrosanct status in this of the timberline and exhibit many features of great ecologic day and age. Owing to the harsh living conditions in high value worldwide: Constituting one-fi fth of the land area of mountain environments, the resident biota of such regions the planet; such elevated regions represent one of the main display adaptive strategies that are of special interest from sources of fresh water, serve as refuge for various animals the biologic, ecologic, and biogeographic points of view and plants that are not found at lower altitudes, exhibit a (Körner 1999). High-mountain ecosystems – typically 1 Instituto de Botánica Spegazzini, Universidade Nacional de La Plata, 53 # 477 (1900), La Plata, Buenos Aires, Argentina 2 Departamento de Ciências Naturais, Universidade Regional de Blumenau, R. Antonio da Veiga, 140, 89030-903, Blumenau, SC, Brazil 3 Laboratorio de Microbiología Aplicada y Biotecnología, Centro Regional Universitario Bariloche, Universidad Nacional del Comahue Quintral 1250, San Carlos de Bariloche (8400), Río Negro, Argentina 4 Laboratorio de Investigación en Sistemas Ecológicos y Ambientales, Facultad de Ciencias Agrarias y Forestales-Facultad de Ciencias Naturales y Museo, UNLP. Diagonal 113, #469 (1900), La Plata, Buenos Aires, Argentina * Corresponding author: [email protected] Diagramação e XML SciELO Publishing Schema: www.editoraletra1.com.br María Silvana Velázquez, Sidney Luiz Stürmer, Clara Bruzone, Sonia Fontenla, Marcelo Barrera and Marta Cabello unstable and immature – are characterized by shallow surface soils; low temperatures but with a wide temperature Materials and methods range; rainfall in all four seasons; snow accumulation during at least some part of the year (i. e., between May Study site and November in Argentina); low relative humidity; high radiation levels; strong winds; and low partial pressures The study area is located in the Northwest Patagonia of oxygen, and carbon dioxide (Ferreyra 1995). These within the Nahuel Huapi National Park (705,000 ha) in the ecosystems are home to different habitats, distributed in Province of Río Negro, Argentina. That region – part of the the form of islands (such as screes, meadows, and high- Patagonian Phytogeographic Province (Cabrera & Willink altitude steppes) that allow the development of diverse plant 1980; Fig. 1) – is characterized by a semiarid to subhumid communities adapted to those environmental conditions. climate (mean annual isotherm below 10 °C; Soriano et The high-Andean environments harbor more than 300 plant al. 1983), with rain and snow being concentrated in the species (Ferreyra 1995; Ferreyra et al. 2006), of which flora autumn and winter seasons and a high frequency of strong little is known regarding its association with mycorrhizal westerly winds. The spring and summer, however, are cool fungi or other fungal endophytes. and dry, though also windy (Ayesa et al. 1995). Arbuscular mycorrhizal fungi (AMF; phylum Glomeromycota) are common soil denizens that establish Sampling design an arbuscular mycorrhizal association with a great majority of land plants (Wang & Qiu 2006). Those fungi provide The sites selected for this study at the national park several ecosystem services including promoting plant were the Patagonian Steppe (PS), Challhuaco Hill (ChC), growth and nutrient uptake, increasing plant resistance to drought along with protection against biotic stresses, and Catedral Hill (CH), and Tronador Hill (TH; Tab. 1). The sites enhancing soil stability and water retention (Gianinazzi et are distributed over a precipitation and altitude gradient al. 2010). Therefore, that symbiotic association constitutes ranging from 3,500 mm and 3,484 m, respectively, in the an key strategy for assisting plants to cope with extreme westernmost site (TH) to 500 mm and 830 m, respectively, environmental conditions (Smith & Read 2008), such as in the easternmost site (PS). Owing to differences in altitude occur in high-altitude ecosystems. Previous investigations within each site, all samples in these high-mountain have indicated that plants occurring in the high Andes of environments were collected between 1.500 – 1,900 m (Tab. Peru at altitudes of up to 5,391 m (Schmidt et al. 2008) 1). Within each site, two to four different environments were and in the Bolivian Andes at heights between 3,700 and selected (i. e., steppe, high steppe, forest, scree, timberline, 4,000 m (Urcelay et al. 2011) are associated with AMF and and meadow). In each environment, rhizosphere-soil dark septate endophytes. The species richness of AMF in samples (500 g each) from three individuals of each plant high-altitude environments has also been reported (Lugo species were collected. The plant individuals were selected et al. 2008; Oehl et al. 2011a; Oehl & Koerner 2014) and by means of a random-walk method and were at least five new resident species described (Oehl et al. 2012; Palenzuela meters away from each other. We selected nine plant species et al. 2014). with different life forms plus a wide altitude distribution Despite the ubiquitous presence of AMF in high-altitude (830 – 3500 m) and precipitation regimen (500 – 3,500 environments, only scarce information is available on the mm/year; Table 1). occurrence of those fungi in the high-Andean Patagonia in Argentina. The Northwest region of the Patagonia is Soil analysis characterized by marked climatic and vegetation gradients. The precipitation levels range from more than 3,500 mm per All the individual soil samples from each environment year in the mountains of the border between Argentina and were pooled to obtain a composite specimen to be used Chile, in the west, to less than 500 mm per year in the east (Pereyra et al. 2005). Therefore, the aim of this work was: for physical and chemical analysis. The proportions of clay, i) to assess the occurrence and diversity of AMF associated silt, and sand were determined by the hydrometer method with the rhizospheres of different plant species growing in (Bouyocus 1962); soil pH measured in a 1:2.5 (w/v) soil-to- diverse environments over an altitude gradient and ii) to water ratio; organic carbon assessed by the wet-oxidation investigate the effect of host-plant identity and altitude on method of Walkley & Black (1934); total N estimated by AMF community composition. The results reported here will the micro-Kjedahl method (Jackson 1967); and phosphorus increase our knowledge regarding the distribution of this concentration assayed by mineralization of the sample and group of fungi and in so doing facilitate the development dry digestion. All the analyses were carried out by the Soil of strategies directed at the management and protection of Group from the Centro Regional Universitario Bariloche, these biota in the different high-mountain environments. Universidad Nacional del Comahue, Río Negro, Argentina. 522 Acta Botanica Brasilica - 30(4): 521-531. October-December 2016 Diagramação e XML SciELO Publishing Schema: www.editoraletra1.com.br Occurrence of arbuscular mycorrhizal fungi in high altitude sites of the Patagonian Altoandina region in Nahuel Huapi National Park (Argentina) Nahuel Huapi National Park Argentina Steppes Patagonica Hill Tronador Hill Catedral Hill Challhuaco Figure 1. Map of Argentina showing the Nahuel Huapi National Park and in the detailed inset indicating the location of the sampling sites: the Patagonian