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Broadleaf Litter Controls Feather Moss Growth in Black Spruce and Birch Forests of Interior Alaska

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Abstract

Plant–soil feedbacks can maintain or reinforce alternative states within ecological systems. In Alaskan boreal forests, changes in fire characteristics have stimulated the replacement of needle-leaf black spruce (Picea mariana) by broadleaf deciduous trees. Feather mosses have strong associations with forest type: They dominate black spruce forest understories and are uncommon in broadleaf stands, with consequences for nutrient cycling and carbon storage. Here we test a long-standing hypothesis that broadleaf litter directly excludes mosses with a field experiment in broadleaf paper birch (Betula neoalaskana) and black spruce stands. We established 30 plots (15 each in birch and spruce dominated areas) with three Hylocomium splendens transplants treated with one of three treatments in each plot (ambient leaf litter deposition, birch leaf litter exclusion or addition), and 30 natural H. splendens areas. We measured moss growth and reproductive potential over 3 years. A 1-year experiment assessed leaf leachate and physical structure impacts on moss growth. Moss shoot growth in natural patches was larger in spruce than in birch stands (24.8 vs. 17.3 mg) and H. splendens made large contributions to ecosystem productivity in spruce stands. In both stand types, we observed a 40% reduction in moss biomass between litter addition and exclusion treatments and litter additions decreased sporophyte production. We found no difference in growth for mosses treated for 1 year with leaf leachates or physical litter structures. Leaf litter effects appear strong enough to exclude mosses from broadleaf forests, providing experimental support for hypothesized plant–soil interactions that may stabilize alternate forest types.

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References

  • Apps MJ, Kurz WA, Luxmoore RJ, Nilsson LO, Sedjo RA, Schmidt R, Simpson LG, Vinson TS. 1993. Boreal forests and tundra. Water Air Soil Pollut 70:39–53.

    CAS  Google Scholar 

  • Bates D, Maechler M, Bolker B, Walker S. 2014. lme4: Linear mixed-effects models using Eigen and S4. R Packag version 1:1–23.

    Google Scholar 

  • Bolker BM, Brooks ME, Clark CJ, Geange SW, Poulsen JR, Stevens MHH, White J-SS. 2009. Generalized linear mixed models: a practical guide for ecology and evolution. Trends Ecol Evol 24:127–35.

    PubMed  Google Scholar 

  • Busby JR, Bliss LC, Hamilton CD. 1978. Microclimate control of growth-rates and habitats of boreal forest mosses, Tomenthypnum nitens and Hylocomium splendens. Ecol Monogr 48:95–110.

    Google Scholar 

  • Calef MP, McGuire AD, Epstein HE, Rupp TS, Shugart HH. 2005. Analysis of vegetation distribution in Interior Alaska and sensitivity to climate change using a logistic regression approach. J Biogeogr 32:863–78.

    Google Scholar 

  • Callaghan TV, Collins NJ, Callaghan CH. 1978. Photosynthesis, growth and reproduction of Hylocomium splendens and Polytrichum commune in Swedish Lapland. Strategies of growth and population dynamics of tundra plants 4. Oikos 31:73–88.

    Google Scholar 

  • Chapin FS, Hollingsworth TN, Murray DF, Viereck LA, Walker MD. 2006. Floristic diversity and vegetation distribution in the Alaskan boreal forest. In: Oswood MW, Van Cleve K, Viereck LA, Verbyla DL, Chapin FS, Eds. Alaska’s changing boreal forest. New York: Oxford University Press. p 81–99.

    Google Scholar 

  • Crawley MJ. 2007. Statistical modelling. In: Crawley MJ, Ed. The R Book. Chichester: Wiley. p 338–448.

    Google Scholar 

  • Cronberg N. 2002. Colonization dynamics of the clonal moss Hylocomium splendens on islands in a Baltic land uplift area: Reproduction, genet distribution and genetic variation. J Ecol 90:925–35.

    Google Scholar 

  • Davey ML, Currah RS. 2006. Interactions between mosses (Bryophyta) and fungi. Can J Bot 84:1509–19.

    Google Scholar 

  • De Grandpré L, Gagnon D, Bergeron Y. 1993. Changes in the understory of Canadian southern boreal forest after fire. J Veg Sci 4(6):803–10.

    Google Scholar 

  • DeLuca TH, Zackrisson O, Gentili F, Sellstedt A, Nilsson MC. 2007. Ecosystem controls on nitrogen fixation in boreal feather moss communities. Oecologia 152:121–30.

    PubMed  Google Scholar 

  • DeLuca TH, Zackrisson O, Nilsson M-C, Sellstedt A. 2002. Quantifying nitrogen-fixation in feather moss carpets of boreal forests. Lett Nat 419:917–20.

    CAS  Google Scholar 

  • Ehrenfeld JG, Ravit B, Elgersma K. 2005. Feedback in the Plant–Soil System. Annu Rev Environ Resour 30:75–115.

    Google Scholar 

  • Flanagan PW, Van Cleve K. 1983. Nurient cycling in relation to decomposition and organic-matter quality in taiga ecosystems. Can J For Res 13:795–816.

    CAS  Google Scholar 

  • Frazer GW, Canham CD, Lertzman KP. 1999. Gap Light Analyzer (GLA).

  • Frego KA. 2007. Bryophytes as potential indicators of forest integrity. For Ecol Manag 242:65–75.

    Google Scholar 

  • Greene DF, Macdonald SE, Haeussler S, Domenicano S, Noel J, Jayen K, Charron I, Gauthier S, Hunt S, Gielau ET, Bergeron Y, Swift L. 2007. The reduction of organic-layer depth by wildfire in the North American boreal forest and its effect on tree recruitment by seed. Can J For Res 37:1012–23.

    Google Scholar 

  • Gunderson LH. 2000. Ecological Resilience-in theory and application. Annu Rev Ecol Syst 31:425–39.

    Google Scholar 

  • Hart SA, Chen HYH. 2006. Understory vegetation dynamics of North American boreal forests. CRC Crit Rev Plant Sci 25:381–97.

    Google Scholar 

  • Hinzman LD, Bolton RW, Petrone KC, Jones JB, Adams PC. 2006. Watershed Hydrology and Chemistry in the Alaskan Boreal Forest The Central Role of Permafrost. Alaska’s Chang boreal For:269.

  • Jaeger B. 2016. r2glmm: Computes R squared for mixed (multilevel) models (LMMs and GLMMs). R package version 0.1.0.

  • Jean M. 2017. Effects of leaf litter and environment on bryophytes in boreal forests of Alaska. Ph.D. dissertation. University of Saskatchewan, Saskatoon, SK, Canada.

  • Jean M, Alexander HD, Mack MC, Johnstone JF. 2017a. Patterns of bryophyte succession in a 160-year chronosequence in deciduous and coniferous forests of boreal Alaska. Can J For Res 1032:1021–32.

    Google Scholar 

  • Jean M, Mack MC, Johnstone JF. 2017b. Murphy Dome: Covariates measured on the sampling units of moss (Hylocomium splendens) from a moss transplant and leaf litter manipulation experiment in three pairs of adjacent black spruce and birch stands. Bonanza Creek LTER—University of Alaska Fairbanks (BNZ:673), https://doi.org/10.6073/pasta/56123c8ae622186e9a91b6b4553bcc87.

  • Jean M, Mack MC, Johnstone JF. 2017c. Murphy Dome: Hourly temperature of air and soil, PAR, relative humidity and soil moisture in three pairs of adjacent black spruce and birch stands. Bonanza Creek LTER—University of Alaska Fairbanks (BNZ:670), https://doi.org/10.6073/pasta/7e2e261a7cb914d5e4b7e7a74a0e6606.

  • Jean M, Mack MC, Johnstone JF. 2017d. Murphy Dome: Moss (Hylocomium splendens and Pleurozium schreberi) biomass results from a birch leaf litter and leachates manipulation experiment in a black spruce stand. Bonanza Creek LTER—University of Alaska Fairbanks (BNZ:672), https://doi.org/10.6073/pasta/84237e605d4be5185dbeffa72330c342.

  • Jean M, Mack MC, Johnstone JF. 2017e. Murphy Dome: Moss (Hylocomium splendens) biomass results from a moss transplant and leaf litter manipulation experiment in three pairs of adjacent black spruce and birch stands. Bonanza Creek LTER—University of Alaska Fairbanks (BNZ:671), https://doi.org/10.6073/pasta/7df8bb07dbbd38f8eede86cfff05be04.

  • Johnson PCD. 2014. Extension of Nakagawa and Schielzeth’s R2GLMM to random slopes models. Methods Ecol Evol 5:944–6.

    PubMed  PubMed Central  Google Scholar 

  • Johnstone JF, Chapin FS. 2006. Effects of soil burn severity on post-fire tree recruitment in boreal forest. Ecosystems 9:14–31.

    Google Scholar 

  • Johnstone JF, Chapin FS, Hollingsworth TN, Mack MC, Romanovsky V, Turetsky M. 2010a. Fire, climate change, and forest resilience in interior Alaska. Can J For Res 40:1302–12.

    Google Scholar 

  • Johnstone JF, Hollingsworth TN, Chapin FS, Mack MC. 2010b. Changes in fire regime break the legacy lock on successional trajectories in Alaskan boreal forest. Glob Chang Biol 16:1281–95.

    Google Scholar 

  • Kardol P, Spitzer CM, Gundale MJ, Nilsson MC, Wardle DA. 2016. Trophic cascades in the bryosphere: the impact of global change factors on top-down control of cyanobacterial N2-fixation. Ecol Lett 19:967–76.

    PubMed  Google Scholar 

  • Kolari P, Pumpanen J, Kulmala L, Ilvesniemi H, Nikinmaa E, Grönholm T, Hari P. 2006. Forest floor vegetation plays an important role in photosynthetic production of boreal forests. For Ecol Manag 221:241–8.

    Google Scholar 

  • Kuznetsova A, Brockhoff PB, Christensen RHB. 2015. lmerTest: tests in linear mixed effects models. R package version 2.0-20. https//cran rproject org/web/packages/lmerTest. Accessed 15:2016.

  • Landhäusser SM, Lieffers VJ. 2003. Seasonal changes in carbohydrate reserves in mature northern Populus tremuloides clones. Trees 17:471–6.

    Google Scholar 

  • Lindo Z, Gonzalez A. 2010. The bryosphere: an integral and influential component of the Earth’s biosphere. Ecosystems 13:612–27.

    Google Scholar 

  • Lindo Z, Nilsson MC, Gundale MJ. 2013. Bryophyte-cyanobacteria associations as regulators of the northern latitude carbon balance in response to global change. Glob Chang Biol 19:2022–35.

    PubMed  Google Scholar 

  • Mann DH, Scott Rupp T, Olson MA, Duffy PA. 2012. Is Alaska’s boreal forest now crossing a major ecological threshold? Arctic, Antarct Alp Res 44:319–31.

    Google Scholar 

  • Melvin AM, Mack MC, Johnstone JF, McGuire AD, Genet H, Schuur EAG. 2015. Differences in ecosystem carbon distribution and nutrient cycling linked to forest tree species composition in a mid-successional boreal forest. Ecosystems 18:1472–88.

    CAS  Google Scholar 

  • Miyanishi K, Johnson EA. 2002. Process and patterns of duff consumption in the mixedwood boreal forest. Can J For Res 32:1285–95.

    Google Scholar 

  • Nilsson M-C, Zackrisson O. 1992. Inhibition of Scots pine seedling establishment byEmpetrum hermaphroditum. J Chem Ecol 18:1857–70.

    CAS  PubMed  Google Scholar 

  • Oechel WC, Van Cleve K. 1986. The role of bryophytes in nutrient cycling in the taiga. For Ecosyst Alaskan Taiga Synth Struct Funct 57:121–37.

    Google Scholar 

  • Økland RH. 1995. Population biology of the clonal moss Hylocomium splendens in Norwegian boreal spruce forests. 1. Demography. J Ecol 83:697–712.

    Google Scholar 

  • Payette S. 1992. Fire as a controlling process in the North American boreal forest. A Syst Anal Glob Boreal For:144–69.

  • R Development Core Team. 2016. R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing; 2014.

  • Rochefort L. 2000. Sphagnum—a keystone genus in habitat restoration. Bryologist 103:503–8.

    Google Scholar 

  • Rydgren K, Økland RH. 2001. Sporophyte production in the clonal moss Hylocomium splendens: the importance of shoot density. J Bryol 23:91–6.

    Google Scholar 

  • Rydgren K, Økland RH. 2002a. Sex distribution and sporophyte frequency in a population of the clonal moss Hylocomium splendens. J Bryol 24:207–14.

    Google Scholar 

  • Rydgren K, Økland RH. 2002b. Ultimate costs of sporophyte production in the clonal moss Hylocomium splendens. Ecology 83:1573–9.

    Google Scholar 

  • Rydgren K, Økland RH, Økland T. 1998. Population biology of the clonal moss Hylocomium splendens in Norwegian boreal spruce forests. 4. Effects of experimental fine-scale disturbance. Oikos:5–19.

    Google Scholar 

  • Skaug H, Fournier D, Nielsen A, Magnusson A, Bolker B. 2011. glmmADMB: generalized linear mixed models using AD Model Builder. R package version 0.8.0.

  • Sorensen PL, Michelsen A. 2011. Long-term warming and litter addition affects nitrogen fixation in a subarctic heath. Glob Chang Biol 17:528–37.

    Google Scholar 

  • Startsev N, Lieffers VJ, Landhäusser SM. 2008. Effects of leaf litter on the growth of boreal feather mosses: implication for forest floor development. J Veg Sci 19:253–60.

    Google Scholar 

  • Sulyma R, Coxson DS. 2001. Microsite displacement of terrestrial lichens by feather moss mats in late seral pine-lichen woodlands of North-central British Columbia. Bryologist 104:505–16.

    Google Scholar 

  • Sveinbjornsson B, Oechel WC. 1992. Controls on growth and productivity of bryophytes: environmental limitations under current and anticipated conditions. Bryophyt lichens a Chang Environ/Ed by Jeffrey W Bates Andrew M Farmer:77–102.

  • Tamm CO. 1953. Growth, yield and nutrition in carpets of a feather moss (Hylocomium splendens). Menddelanded Statense Skogsforsknings Inst 43:1–140.

    Google Scholar 

  • Trugman AT, Fenton NJ, Bergeron Y, Xu X, Welp LR, Medvigy D. 2016. Climate, soil organic layer, and nitrogen jointly drive forest development after fire in the North American boreal zone. J Adv Model Earth Syst 8:1180–209.

    Google Scholar 

  • Turetsky MR. 2003. The role of bryophytes in carbon and nitrogen cycling. Bryologist 106:395–409.

    Google Scholar 

  • Turetsky MR, Bond-Lamberty BP, Euskirchen ES, Talbot J, Frolking S, McGuire AD, Tuittila E-S. 2012. The resilience and functional role of moss in boreal and arctic ecosystems. New Phytol 196:49–67.

    CAS  PubMed  Google Scholar 

  • Turetsky MR, Mack MC, Hollingsworth TN, Harden JW. 2010. The role of mosses in ecosystem succession and function in Alaska’s boreal forestT. Can J For Res 40:1237–64.

    Google Scholar 

  • Van Cleve K, Dyrness CT, Viereck LA, Fox J, Chapin FS, Oechel W. 1983a. Taiga ecosystems in interior Alaska. Bioscience 33:39–44.

    Google Scholar 

  • Van Cleve K, Oliver L, Schlentner RE, Viereck LA, Dyrness CT. 1983b. Productivity and nutrient cycling in taiga forest ecosystems. Can J For Res 13:747–66.

    Google Scholar 

  • Yarie J, Billings S. 2002. Carbon balance of the taiga forest within Alaska: present and future. Can J For Res 32:757–67.

    Google Scholar 

  • Zeileis A, Kleiber C, Jackman S. 2008. Regression models for count data in R. J Stat Softw 27:1–25.

    Google Scholar 

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Acknowledgements

Funding came from the Department of Defense’s Strategic Environmental Research and Development Program (project RC-2109), the Natural Science and Engineering Research Council of Canada, Canada’s Northern Scientific Training Program, and the Bonanza Creek Long Term Ecological Research program supported by the National Science Foundation (NSF DEB- 0620579) and USDA Forest Service, Pacific Northwest Research Station (PNW01-JV11261952-23). We thank Alix Conway, Samantha Miller, Patricia Tomchuk, Dominic Olver, Alexandre Truchon-Savard, Xanthe Walker, Nicolas Boldt, and students from the University of Saskatchewan and University of Florida for field and laboratory work.

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  1. April M. Melvin

    Present address: , Washington, DC, USA

Authors and Affiliations

  1. Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan, S7N 5E2, Canada

    Mélanie Jean & Jill F. Johnstone

  2. Center for Ecosystem Science and Society, Northern Arizona University, Flagstaff, Arizona, USA

    Mélanie Jean & Michelle C. Mack

  3. Department of Biology, University of Florida, Gainesville, Florida, 32611, USA

    April M. Melvin

  4. Institute of Arctic Biology, University of Alaska Fairbanks, Fairbanks, Alaska, USA

    Jill F. Johnstone

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  1. Mélanie Jean
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Correspondence to Mélanie Jean.

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MJ led the field, laboratory, and data analyses and drafted the manuscript with guidance from JFJ. AMM led the field and laboratory work in 2012 when the experiment was established. JFJ, MCM, and AMM conceived the ideas, study approach, and designed the methodology. All authors contributed to manuscript drafts and gave final approval for publication.

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Jean, M., Melvin, A.M., Mack, M.C. et al. Broadleaf Litter Controls Feather Moss Growth in Black Spruce and Birch Forests of Interior Alaska. Ecosystems 23, 18–33 (2020). https://doi.org/10.1007/s10021-019-00384-8

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