QUESTION: What is the impact of simulated historical tree litter removal on understorey plants and soil properties in a temperate deciduous forest? What is the role of seasonal timing of tree litter removal on understorey plants? LOCATION: Podyjí National Park, Czech Republic. METHODS: We conducted an experiment in a randomized complete block design of 45 plots (5 × 5 m). Each block (N = 15) consisted of one plot for each of the three treatments. Treatments consisted of (i) tree litter removal during spring, (ii) tree litter removal during autumn, or (iii) no litter removal as control treatment. These treatments were repeated for a duration of four years. In each plot we recorded the understorey plant species composition and collected soil samples prior to treatment (year 0) and in each subsequent year (years 1-4). Temporal trends in species richness were analysed using repeated measures ANOVAs. The impact of the treatments on vegetation composition over time was analysed using Principal Response Curves. RESULTS: Total species richness per plot significantly changed over time, but this was not related to treatment. Annual species richness increased significantly, but only for the autumn treatment. Annual species also showed the highest inter-annual variation. Endangered species were not affected. When compared to the control treatment, the effect of autumn raking on species composition was stronger than the effect of spring raking. Although the amount of removed nutrients substantially exceeded ambient nitrogen input, no changes in soil conditions were detected. CONCLUSIONS: The season in which tree litter removal took place had a small but significant impact on the understorey vegetation, in particular affecting the germination and establishment of annual species. The large inter-annual variation in species richness calls for a long-term field experiment. The removal of nutrients via litter raking greatly exceeds atmospheric nutrient deposition, warranting a further investigation of litter raking as a potential tool for forest conservation.

Department of Botany Faculty of Science Palacký University Šlechtitelů 11 783 71 Olomouc Czech Republic

DST NRF Centre of Excellence for Invasion Biology Department of Zoology University of Johannesburg Auckland Park 2006 South Africa

Institute of Botany Department of Vegetation Ecology The Czech Academy of Sciences Lidická 25 27 Brno 602 00 Czech Republic

Masaryk University Department of Botany and Zoology Kotlářská 2 Brno 611 37 Czech Republic

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Baskin CC, Baskin JM. Seeds: ecology, biogeography, and evolution of dormancy and germination. Academic Press; San Diego, CA: 2014.

Beatty SW, Sholes ODV. Leaf litter effect on plant species composition of deciduous forest treefall pits. Canadian Journal of Forest Research. 1988;18:553–559.

Benkobi L, Trlica MJ, Smith JL. Soil loss as affected by different combinations of surface litter and rock. Journal of Environmental Quality. 1993;22:657–661.

Bergmeier E, Petermann J, Schröder E. Geobotanical survey of wood-pasture habitats in Europe: diversity, threats and conservation. Biodiversity and Conservation. 2010;19:2995–3014.

Bobbink R, Hicks K, Galloway JN, Spranger T, Alkemade R, Ashmore M, Bustamante M, Cinderby S, Davidson E, Dentener F, Emmett B, et al. Global assessment of nitrogen deposition effects on terrestrial plant diversity: a synthesis. Ecological Applications. 2010;20:30–59. PubMed

Bobbink R, Hornung M, Roelofs JGM. The effects of air-borne nitrogen pollutants on species diversity in natural and semi-natural European vegetation. Journal of Ecology. 1998;86:717–738.

Bocock K. Changes in the amounts of dry matter, nitrogen, carbon and energy in decomposing woodland leaf litter in relation to the activities of the soil fauna. Journal of Ecology. 1964;52:273–284.

Bürgi M. A case study of forest change in the Swiss lowlands. Landscape Ecology. 1999;14:567–575.

Bürgi M, Gimmi U. Three objectives of historical ecology: the case of litter collecting in Central European forests. Landscape Ecology. 2007;22:77–87.

Calado J, Basch G, de Carvalho M. Appearance of spontaneous plants from disturbed and undisturbed soil under mediterranean conditions. Revista de Ciências Agrárias. 2008;31:68–78.

Carlisle A, Brown A, White E. The nutrient content of tree stem flow and ground flora litter and leachates in a sessile oak (Quercus petraea) woodland. Journal of Ecology. 1967;55:615–627.

CHMI. Odhad celkové roční depozice uvedených složek na plochu České republiky (78 841 km2) v tunách, 2010. [accessed 29 December 2014];2010 URL: http://portal.chmi.cz/

CHMI. ČHMÚ Historická data – meteorologie a klimatologie. [accessed 14 October 2014];2014 URL: http://www.chmi.cz/

Chytrý M, Tichý L. Diagnostic, constant and dominant species of vegetation classes and alliances of the Czech Republic: a statistical revision. Folia facultatis scientiarum naturalium universitatis Masarykianae Brunensis. 2003;108:1–231.

Čarni A, Košir P, Marinšek A, Šilc U, Zelnik I. Changes in structure, floristic composition and chemical soil properties in a succession of birch forests. Periodicum Biologorum. 2007;109:13–20.

Dengler J, Chytrý M, Ewald J. Phytosociology. In: Jørgensen SE, Fath BD, editors. Encyclopedia of Ecology. Elsevier; Oxford, UK: 2008. pp. 2767–2779.

Dickens ED, Moorhead DJ, Morris LA. Pine straw – an economically important forest product in Georgia. University of Georgia; Athens, GA USA: 2012.

Dzwonko Z, Gawroński S. Effect of litter removal on species richness and acidification of a mixed oak-pine woodland. Biological Conservation. 2002a;106:389–398.

Dzwonko Z, Gawroński S. Influence of litter and weather on seedling recruitment in a mixed oak-pine woodland. Annals of Botany. 2002b;90:245–251. PubMed PMC

Ebermayer E. Die gesammte Lehre der Waldstreu mit Rücksicht auf die chemische Statik des Waldbaues. Springer; Berlin, Germany: 1876.

Eriksson O. Seedling recruitment in deciduous forest herbs: the effects of litter, soil chemistry and seed bank. Flora. 1995;190:65–70.

Facelli JM, Pickett S. Plant litter: its dynamics and effects on plant community structure. Botanical Review. 1991;57:1–32.

Gabrielová J, Münzbergová Z, Tackenberg O, Chrtek J. Can we distinguish plant species that are rare and endangered from other plants using their biological traits? Folia Geobotanica. 2013;48:449–466.

Gimmi U, Bürgi M. Using oral history and forest management plans to reconstruct traditional non-timber forest uses in the Swiss Rhone Valley (Valais) since the late nineteenth century. Environment and History. 2007;13:211–246.

Gimmi U, Bürgi M, Stuber M. Reconstructing anthropogenic disturbance regimes in forest ecosystems: a case study from the Swiss Rhone valley. Ecosystems. 2007;11:113–124.

Gimmi U, Poulter B, Wolf A, Portner H, Weber P, Bürgi M. Soil carbon pools in Swiss forests show legacy effects from historic forest litter raking. Landscape Ecology. 2013;28:835–846.

Gimmi U, Wohlgemuth T. Land-use and climate change effects in forest compositional trajectories in a dry Central-Alpine valley. Annals of Forest Science. 2010;67:701.

Glatzel G. Causes and consequences of forest growth trends in Europe: Results of the recognition project. In: Karjalainen T, Spieker H, Laroussinie O, editors. Causes and consequences of accelerated tree growth in Europe. European Forest Institute; Joensuu, Finland: 1999. pp. 65–74.

Glatzel G. The impact of historic land use and modern forestry on nutrient relations of Central European forest ecosystems. Fertilizer Research. 1991;27:1–8.

Grime JP. Plant strategies, vegetation processes, and ecosystem properties. John Wiley and Sons; Chichester, UK: 2001.

Grulich V. Red List of vascular plants of the Czech Republic: 3rd edition. Preslia. 2012;84:631–645.

Hellström K, Huhta A-P, Rautio P, Tuomi J. Seed introduction and gap creation facilitate restoration of meadow species richness. Journal for Nature Conservation. 2009;17:236–244.

Hofmeister J, Oulehle F, Krám P, Hruška J. Loss of nutrients due to litter raking compared to the effect of acidic deposition in two spruce stands, Czech Republic. Biogeochemistry. 2008;88:139–151.

Hüttl RF, Schaaf W. Nutrient supply of forest soils in relation to management and site history. Plant and Soil. 1995;168-169:31–41.

Ito E, Toriyama J, Araki M, Kiyono Y, Kanzaki M, Tith B, Keth S, Chandararity L, Chann S. Physicochemical surface-soil properties after litter-removal manipulation in a Cambodian lowland dry evergreen forest. Japan Agricultural Research Quarterly. 2014;48:195–211.

Klotz S, Kühn I. BIOLFLOR – Eine Datenbank zu biologisch-ökologischen Merkmalen der Gefäßpflanzen in Deutschland. Bundesamt für Naturschutz; Bonn, Germany: 2002.

Koorem K, Price JN, Moora M. Species-specific effects of woody litter on seedling emergence and growth of herbaceous plants. PLoS ONE. 2011;6:e26505. PubMed PMC

Kopáček J, Veselý J, Stuchlík E. Sulphur and nitrogen fluxes and budgets in the Bohemian Forest and Tatra Mountains during the Industrial Revolution (1850–2000) Hydrology and Earth System Sciences. 2001;5:391–405.

Kubát K, Hrouda L, Chrtek J Jr, Kaplan Z, Kirschner J, Štěpánek J, editors. Klíč ke květeně České republiky. Academia, Praha, Czech Republic; 2002.

Leff JW, Wieder WR, Taylor PG, Townsend AR, Nemergut DR, Grandy A, Cleveland CC. Experimental litterfall manipulation drives large and rapid changes in soil carbon cycling in a wet tropical forest. Global Change Biology. 2012;18:2969–79. PubMed

Li X, Niu J, Xie B. The effect of leaf litter cover on surface runoff and soil erosion in Northern China. PLoS ONE. 2014;9:e107789. PubMed PMC

Loqué D, Eudes A, Yang F. Biomass availability and sustainability for biofuels. In: Simmons B, editor. Chemical and biochemical catalysis for next generation biofuels. Royal Society of Chemistry; Cambridge, UK: 2011.

Monk CD, Gabrielson F. Effects of shade, litter and root competition on old-field vegetation in South Carolina. Bulletin of the Torrey Botanical Club. 1985;112:383–392.

Olsen RS. Phosphorus. In: Page AL, editor. Methods in Soil Analysis. Madison, Wisconsin, USA: 1982.

Oksanen J, Blanchet GF, Kindt R, Legendre P, Minchin PR, O’Hara RB, Simpson G, Solymos P, Stevens MHH, Wagner HH. Vegan: Community Ecology Package. R package version 2.0-10. 2014.

Osono T, Takeda H. Organic chemical and nutrient dynamics in decomposing beech leaf litter in relation to fungal ingrowth and succession during 3-year decomposition processes in a cool temperate deciduous forest in Japan. Ecological Research. 2001;16:649–670.

Piepho H, Büchse A, Richter C. A mixed modelling approach for randomized experiments with repeated measures. Journal of Agronomy & Crop Science. 2004;190:230–247.

Prietzel J, Kaiser KO. De-eutrophication of a nitrogen-saturated Scots pine forest by prescribed litter-raking. Journal of Plant Nutrition and Soil Science. 2005;168:461–471.

Quinn G, Keough M. Experimental design and data analysis for biologists. Cambridge University Press; Cambridge, UK: 2002.

Sayer EJ. Using experimental manipulation to assess the roles of leaf litter in the functioning of forest ecosystems. Biological Reviews. 2005;81:1–31. PubMed

Šilc U, Čarni A, Košir P, Marinšek A, Zelnik I. Litter-raking forests in SE Slovenia and in Croatia. Hacquetia. 2008;7:71–88.

Tamm CO. Nitrogen in terrestrial ecosystems: Questions of productivity, vegetational changes, and ecosystem stability. Springer; Berlin, Germany: 1991.

Tolasz R, Míková T, Valeriánová T, Voženílek V. Climate atlas of Czechia. Czech Hydrometeorological Institute and Palacký University; Olomouc, Czech Republic: 2007.

Van den Brink P, Ter Braak C. Principal response curves: analysis of time-dependent multivariate responses of biological community to stress. Environmental Toxicology and Chemistry. 1999;18:138–148.

Vincent EM, Roberts EH. The interaction of light, nitrate and alternating temperature in promoting the germination of dormant seeds of common weed species. Seed Science and Technology. 1977;5:659–670.

Wilson S, Tilman D. Quadratic variation in old-field species richness along gradients of disturbance and nitrogen. Ecology. 2002;83:492–504.

Xiong S, Nilsson C. The effects of plant litter on vegetation: a meta-analysis. Journal of Ecology. 1999;87:984–994.

Xu S, Liu LL, Sayer EJ. Variability of above-ground litter inputs alters soil physicochemical and biological processes: a meta-analysis of litterfall-manipulation experiments. Biogeosciences. 2013;10:7423–7433.