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Alien Vs. Native Plants in a Patagonian Wetland: Elemental Ratios and Ecosystem Stoichiometric Impacts

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Alien Vs. Native Plants in a Patagonian Wetland: Elemental Ratios and Ecosystem Stoichiometric Impacts Biol Invasions (2012) 14:179–189 DOI 10.1007/s10530-011-9995-9 ORIGINAL PAPER Alien vs. native plants in a Patagonian wetland: elemental ratios and ecosystem stoichiometric impacts Florencia Cuassolo • Esteban Balseiro • Beatriz Modenutti Received: 19 November 2010 / Accepted: 4 April 2011 / Published online: 19 April 2011 Ó Springer Science+Business Media B.V. 2011 Abstract Wetlands are subject to invasion by exotic of the invasive plant can modify the stoichiometry of plant species, especially during the dry season when the environment. they resemble terrestrial systems; therefore, terrestrial plants could exploit this situation to colonize this Keywords Invasion ecology Á Wetland Á environment. We analyzed P. anserina invading Mycorrhizal fungi Á Stoichiometric strategies Á Patagonian wetlands in terms of elemental ratios that Leachates Á Dissolved organic matter release would modify wetland stoichiometry due to organic matter inputs. We studied the elemental relationship (carbon/nitrogen/phosphorus) of P. anserina in com- parison with native emergent macrophytes (Eleochar- is pachicarpa and Carex aematorrhyncha). These Introduction plant species are common and dominant in the wetland. Additionally, we analyzed the presence of Wetlands are characterized by seasonal fluctuations mycorrhizal fungi in the roots and their proportion of in water level, with alternating wet and dry periods. root infection. Our study reveals that the invasive These ecosystems are especially vulnerable to alien species presented nutrient (especially phosphorus) weed invasions (Zedler and Kercher 2004); therefore, allocation in roots and differences in mycorrhizal many wetland plants fit the definition of invasive infection, with a predominance of arbuscular mycor- plants, which are species or strains that rapidly rhiza, compared with native species. During flooded increase their spatial distribution by expanding into periods with the decay of aerial parts, P. anserina native plant communities (Richardson et al. 2000). stores phosphorus in the roots and releases dissolved The susceptibility of communities to invasion organic matter of high molecular weight molecules, requires more attention (Prieur-Richard and Lavorel high color, and a high C-to-nutrient ratio in compar- 2000) because native plants are now uncommon or ison with native macrophytes. These results show the endangered due to wetland invaders that form strategy of an invasive terrestrial plant in temporary monotypes (Champion et al. 2009), and they can aquatic systems, and how the elemental relationships cause substantial changes in organic matter accumu- lation and carbon sequestration (Zedler and Kercher 2004, 2005). & F. Cuassolo ( ) Á E. Balseiro Á B. Modenutti In aquatic systems, nutrient supply could be an Laboratorio de Limnologı´a, INIBIOMA-CONICET- UNComahue, Quintral 1250, 8400 Bariloche, Argentina important factor in the competitive balance between e-mail: [email protected] plant species, e.g. rapid nutrient uptake allowed a 123 180 F. Cuassolo et al. rapid growth (Zedler and Kercher 2004). In sub- the DOM water content mainly during senescence merged plants, at high fertility, the invasive Hydrilla because of the decomposition of tissue and subse- verticillata was a stronger competitor relative to quent leachates (Bertilsson and Jones 2003). More- Vallisneria americana, whereas under low nutrient over, they are an important source and reservoir of conditions, native species outperform invasive ones essential nutrients, such as phosphorus (Bastidas (Van et al. 1999). Other studies also indicated that Navarro and Modenutti 2010), and contribute nutri- emergent species better adapted to high nutrient ents for heterotrophic production (Bastidas Navarro environment (e.g. Typha dominguensis) can effec- et al. 2009b). These results may imply that variations tively invade with greater competitive capability in the composition of macrophytes could generate under high nutrient conditions (Davis 1991). changes in DOM quality, both in terms of elemental Temporary or semi-temporary wetlands represent composition and in terms of optical features. a particular situation, because during the dry season, Potentilla anserina is an exotic species that is the environment remains without water resembling a widely distributed in wet soils in Patagonia (Correa terrestrial system. The water table fluctuation is a 1984; Ezcurra and Brion 2005). In Patagonia wet- natural disturbance that these systems can tolerate, lands, this hemicriptophyte exotic species may cover which causes changes in both plant diversity and almost the entire bottom and restrict the distribution carbon cycling. However, due to alternation between of native macrophytes (Dı´az Villanueva and Trochine dry and flooded periods, plant species that can 2005; Naiman and Melillo 1984). Recent studies in tolerate a wide range of water levels are more Chile (Hauenstein et al. 2008) have indicated that successful (Raffaele 2004). During dry periods, some P. anserina has intermediate N requirements, which of these species develop their whole growing cycle could create beneficial colonizing environments. We and become an important source of decaying organic have hypothesized that invasive (P. anserina) and matter during the flooded period (Dı´az Villanueva native macrophytes (Eleocharis pachycarpa and and Trochine 2005; Roehm 2005). In this case, the Carex aematorrhyncha) co-occurring in the same elemental composition of the plant tissue would play wetland exhibit different strategies in terms of a crucial role in the resulting organic matter quality nutrient allocation. In that sense, we also propose (Osborne et al. 2007). The differences in macrophyte that the elemental composition of leachates and tissue nutrient concentrations were observed to be tissues of native and invasive macrophytes may species specific with a strict C:N:P (carbon:nitro- distinctively affect the dissolved organic matter gen:phosphorous) stoichiometric ratios (Demars and (DOM) source of the wetland. For that purpose, we Edwards 2007). The elemental composition (N and P) analyzed the elemental relationships (C:N:P) in the in plant tissues is determined by several factors such plant tissues (aerial and radicular parts) and the as (1) the balance of nutrient uptake and carbon leachates of the different species. Additionally, we (C) assimilation (Aerts and Chapin 2000; Sterner and analyzed the presence of mycorrhizal fungi because Elser 2002), (2) losses through leaching, respiration, this type of interaction could be differentially used by exudation, and herbivory (Ventura et al. 2008; Wong exotic and native plants as a strategy for colonizing et al. 2010), and (3) the presence of mycorrhizal fungi temporary environments and obtaining the necessary since mycorrhizal associations favor nutrient uptake nutrients for growth. in wetland plants (Cornwell et al. 2001; Dunham et al. 2003). In the Andean temperate lacustrine district (41°S), there are large and deep lakes, along with small lakes and wetlands (Modenutti Materials and methods et al. 1998). In comparison with deep lakes, shallow lakes and wetlands present a higher concentration of Study area dissolved organic matter (DOM) (Morris et al. 1995). This high content of DOM can be associated with the This study was conducted in Laguna Fantasma, which presence of macrophytes and external watershed is a temporary pond located at 41°070S, 71°270W, contributions (Bastidas Navarro et al. 2009a). Leach- 780 m above sea level in North Patagonia, Argentina. ates of macrophytes can contribute significantly to The pond has an area of 1 ha (170 m length and 80 m 123 Alien vs. native plants in a Patagonian wetland 181 wide approximately), with a maximum depth of 2 m following Valderrama (1981). Sediment samples were during the autumn–winter period. Precipitation during dried at 60°C and were analyzed for particulate carbon autumn and winter (May–August 2009) averaged (C), nitrogen (N), and phosphorous (P) content. C and 208.3 ± 91 mm per month when the flooded area N concentrations were measured with a CN analyzer reached a maximum of 1 ha and was reduced to (Thermo Finingan Flash EA 1112). Phosphorus was 24.2 ± 15 mm per month during late spring and measured after combustion of 2 mg of sediment at summer (September–January 2009). The pond was 450°C for 1 h, followed by the ascorbate-reduced colonized by two native macrophytes, Eleocharis molybdenum method (APHA 2005). pachycarpa and Carex aematorrhyncha, and by the The elemental analysis of vegetal tissue was exotic plant Potentilla anserina. These plant species carried out by carefully washing the plants with tap are common and dominant in the wetland. Riparian water and then with distilled water. The aerial and vegetation was scarce, so other inputs of allochtho- root parts were separated and dried at 60°C for 48 h nous particulate organic matter to the pond were up to constant weight. Afterward, each part was negligible. crushed to powder with a clean mortar, and approx- imately 2 mg of dry weight (DW) was destined for elemental analysis. Carbon (C) and nitrogen (N) con- Field study centrations in the tissues were measured with a CN analyzer (Thermo Finingan Flash EA 1112). Phos- The wetland was sampled on six different occasions phorus was measured after combustion of 2 mg of from April 2009 to January 2010. Water temperature tissue at 450°C for 1 h, followed by the ascorbate- and pH were measured using a terminstor (YSI 85 reduced molybdenum method (APHA 2005). Yellow
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