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Responses of the Mosses Sphagnum Capillifolium and Polytrichum Strictum to Nitrogen Deposition in a Bog: Growth, Ground Cover, and CO2 Exchange Sari Juutinen, Tim R

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Responses of the Mosses Sphagnum Capillifolium and Polytrichum Strictum to Nitrogen Deposition in a Bog: Growth, Ground Cover, and CO2 Exchange Sari Juutinen, Tim R 127 ARTICLE Responses of the mosses Sphagnum capillifolium and Polytrichum strictum to nitrogen deposition in a bog: growth, ground cover, and CO2 exchange Sari Juutinen, Tim R. Moore, Anna M. Laine, Jill L. Bubier, Eeva-Stiina Tuittila, Allison De Young, and Mandy Chong Abstract: Previous studies have shown that atmospheric nitrogen (N) deposition is detrimental to sphagna, which are a group of mosses that are important for carbon cycling in northern peatlands. Little is known about species interac- tions, such as relative responses of tall moss Polytrichum strictum Menzies ex Brid. and sphagna. We studied the effects of N deposition on growth, abundance, and CO2 exchange of the moss species Sphagnum capillifolium (Ehrh.) Hedw. and Polytrichum strictum in an experiment at a temperate bog. Sphagnum growth and cover decreased significantly with high-dose N treatment (6.4 g N·m−2·year−1) in years 4 and 5 of treatment, whereas the same parameters increased for Polytrichum compared with the control. Net CO2 exchange, gross photosynthesis (Pg), and dark respiration (R)inthe intact moss cores, which were measured in year 5 of treatment, were elevated in the cores that had been treated with the high-dose of N, compared with the control, and this was associated with increased abundance of Polytrichum. The moss cores where Polytrichum was removed, however, had increased mass-based R with the high-dose N treatment. Our results showed that S. capillifolium at Mer Bleue may be close to N saturation, as 5 years of high-dose N loading (6.4 g N·m−2·year−1 + background) was harmful to this species, possibly as a result of increased respiratory cost. Polytrichum strictum had a competitive advantage, at least in the short-term, through allocating excess N to growth. This change in moss layer composition deserves further attention, as a shift to more easily decomposable litter, without correspond- ing increases in plant production, could reduce the carbon sequestration of the bog. Key words: peatland, photosynthesis, chlorophyll fluorescence, respiration, vegetation change, moss. Résumé : Des études précédentes ont montré que le dépôt de N atmosphérique peut être dommageable a` la sphaigne, For personal use only. un genre important au recyclage de C dans les tourbières du nord. On sait peu de choses des interactions entre les espèces, telles les réponses relatives de la mousse Polytricum strictum Menzies ex Brid. et de la sphaigne. Les auteurs ont étudié les effets du dépôt de N sur la croissance en hauteur, l’abondance et l’échange de CO2 des espèces de mousses Sphaghum capillifolium (Ehrh.) Hedw. et Polytricum strictum lors d’une expérience en tourbière tempérée. La croissance en hauteur de Sphaghum et son couvert diminuaient significativement en présence d’une forte concentration de N (6,4 g N·m−2·an−1)a` la quatrième et cinquième années de traitement, alors que ceux de Polytricum augmentaient, relativement au contrôle. L’échange net de CO2, la photosynthèse brute (Pb) et la respiration mitochondriale (R) des mousses intactes, mesurés a` la cinquième année de traitement, étaient accrus par le traitement riche en N relativement aux contrôles, a` cause de l’abondance accrue de Polytricum. Les mousses dont on avait retiré Polytrichum présentaient cependant une R en fonction de la masse accrue par le traitement riche en N. Les résultats obtenus par les auteurs montrent que S. capillifolium de la tourbière de Mer Bleue peut être presqu’a` saturation de N, car 5 ans de charge élevée en N (6,4 g N·m−2·an−1 + niveau de base) ont été dommageables a` cette espèce, possiblement a` cause d’un coût Botany Downloaded from www.nrcresearchpress.com by HELSINKI UNIV on 05/02/16 respiratoire accru. Polytrichum strictum possédait un avantage compétitif, du moins a` court terme, réservant l’excès de N a` la croissance. Ce changement dans la composition du couvert de mousse mérite une plus grande attention car un déplacement vers une litière plus facilement décomposable, sans augmentations correspondantes de la production végétale, peut diminuer la séquestration du C par la tourbière. [Traduit par la Rédaction] Mots-clés : tourbière, photosynthèse, fluorescence de la chlorophylle, respiration, changement de la végétations, mousse. Received 1 October 2015. Accepted 14 November 2015. S. Juutinen* and J.L. Bubier. Environmental Studies Department, Mount Holyoke College, 50 College Street, South Hadley, MA 01075, USA. T.R. Moore, A. De Young, and M. Chong. Department of Geography and Global Environmental & Climate Change Centre, McGill University, 805 Sherbrooke Street West, Montreal, QC H3A 0B9, Canada. A.M. Laine. Department of Forest Sciences, University of Helsinki, P.O. Box 27, FI-00014 Helsinki, Finland. E.S. Tuittila. School of Forest Sciences, University of Eastern Finland, P.O. Box 111, FI-80101 Joensuu, Finland. Corresponding author: Sari Juutinen (email: Sari.juutinen@helsinki.fi). *Present address: Department of Forest Sciences, University of Helsinki, P.O. Box 27, FI-00014 Helsinki, Finland. Botany 94: 127–138 (2016) dx.doi.org/10.1139/cjb-2015-0183 Published at www.nrcresearchpress.com/cjb on 17 November 2015. 128 Botany Vol. 94, 2016 Introduction speaking, excess N from atmospheric deposition only The deposition of atmospheric nitrogen (N) drastically becomes available to vascular plants when the Sphagnum increased N availability in the industrialized world dur- layer becomes saturated with N, after which, vascular ing the Anthropocene (e.g., Galloway et al. 2008). Excess plants allocate N to increased growth and thus gain com- N from deposition threatens many natural ecosystems, petitive advantage over peat mosses (e.g., Malmer et al. by impacting species composition and biogeochemistry, 2003). Polytrichum strictum Menzies ex Brid. is a moss spe- and by increasing the competitive ability of some species cies that often co-occurs with sphagna, but is typically while making conditions unfavorable for others (e.g., more abundant in drier microhabitats, intermediate in Bobbink et al. 2010). Peat mosses (Sphagnum spp.) are key its N requirement, and found to benefit from low-level species in northern nutrient-poor peatlands, but they are addition of N and P compared with sphagna (Vitt 1990; sensitive to the deposition of atmospheric N (Limpens et al. Gunnarsson and Rydin 2000; Berendse et al. 2001; 2011). A decrease in the abundance of Sphagnum could mark- Mitchell et al. 2002; Bubier et al. 2007; Sottocornola et al. edly impact litter quality and decomposability, surface 2007; Bu et al. 2011). Polytrichum strictum is a pioneer spe- structure, and water retention capacity, all of which affect cies that is considered to facilitate the establishment of the important carbon (C) sequestration capacity of peat- sphagna at peatland restoration sites (e.g., Robert et al. lands (e.g., Moore et al. 2007; Straková et al. 2010; Larmola 1999), but can outcompete sphagna through increased et al. 2013). abundance (González et al. 2013). However, studies have Sphagna can increase their growth and production shown that the abundance of P. strictum declines under under increased nutrient availability, but the responses conditions of high N loading (Bubier et al. 2007; Bu et al. depend on species sensitivity, dose, and temporal scale 2011). Therefore, it is important to understand the effects of loading (e.g., Rochefort et al. 1990; Vitt et al. 2003; of N deposition on different peatland plants, and the Gunnarsson et al. 2004). Critical loading of atmospheric possible interactions between the species. N, associated with reduced growth, is considered to be In this study we examined the effects of simulated N ϳ0.5 to 1.5 g N·m−2·year−1 for sphagna (Gunnarsson and deposition on the growth and abundance of Sphagnum Rydin 2000; Vitt et al. 2003; Bragazza et al. 2004; Granath capillifolium (Ehrh.) Hedw. and P. strictum during years 1–5 et al. 2014), and is currently exceeded in parts of Europe, of a fertilization experiment at the temperate ombro- North America, southern China, and south and southeast- trophic peatland, Mer Bleue. We also examined the im- ern Asia (Bobbink et al. 2010). Recognized factors contrib- pact of excess N on the CO2 exchange capacity of moss uting to the negative responses of sphagna to increased after 5 years of fertilization. We hypothesized that N availability are the respiratory cost of storing excess N (i) height growth (hereinafter, growth) of S. capillifolium (Limpens and Berendse 2003; Manninen et al. 2011), com- would decrease, but P. strictum would increase with N For personal use only. petition for light and space with vascular plants (van der addition; (ii)CO2 exchange and chlorophyll fluorescence Heijden et al. 2000b; Berendse et al. 2001), increase in measurements would indicate negative impacts of N depo- parasitic infections (Limpens et al. 2003), decrease in sition on S. capillifolium and positive impacts on P. strictum. other elements, most importantly phosphorus (P) (Aerts Materials and methods et al. 1992; Jauhiainen et al. 1998), and greater sensitivity to drying (van der Heijden et al. 2000a; Manninen et al. Study site The study was conducted at the Mer Bleue peatland, 2011; Fritz et al. 2012). In addition, experiments have in- near Ottawa, Ontario, Canada (46°N, 75.5°W), which has dicated that Sphagnum populations adapted to higher a mean annual temperature of 6.6 °C and an average background N loading are less sensitive to additional N rainfall of 756 mm per year (Canadian Climate Normals input than mosses from areas with low background de- 1981–2010). Nitrogen was applied to randomly assigned Botany Downloaded from www.nrcresearchpress.com by HELSINKI UNIV on 05/02/16 position (Wiedermann et al. 2009a; Granath et al. 2012). A triplicate3m×3mplots per treatment, separated by 1 m- recent meta-analysis on the responses of sphagna to ex- wide buffer zones.
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