Abstract
Woody detritus (WD), created by mortality of trees and their associated parts, is an important component of forested ecosystems with roles in energy flow, hydrologic and geomorphologic processes as well as in carbon and nutrient cycling. Although likely to be increasingly important as forest systems respond to climatic and other human induced changes, WD-related science is just beginning relative to other aspects of forested ecosystems. WD differs from other litter forms and soil in key ways (i.e., size range, rigidity, and heterogeneity) that limit the application of many paradigms currently used in studying and modeling decomposition. Thus, while temperature and concentrations of lignin and nitrogen are important controls, others factors related to moisture and its interaction with canopy openness, WD size, position relative to the soil surface, and decomposers need to be better understood. Moreover, the unique attributes of WD decomposers need to be acknowledged as they have evolved over hundreds of millions of years to efficiently process this high lignin, low nutrient substrate. Given the heterogeneity within and among WD entities, WD behavior can be extremely non-linear, often resulting in cascades of activity rather than all or nothing behavior. Substantial improvements in understanding and modeling the respiration, fragmentation, leaching, and burial involved during WD decomposition are required to accurately assess the impact of global increases in tree mortality. Additionally understanding how the perception of WD-related processes changes with scale and organizational level is required to fully understand WD’s role in past, present, and future biogeochemical cycling.











Similar content being viewed by others
Data availability
The models used to generate the figures and conduct analyzes are available upon request to the author.
References
Achat DL, Deleuze C, Landmann G, Pousse N, Ranger J, Augusto L (2015) Quantifying consequences of removing harvesting residues on forest soils and tree growth—a meta-analysis. For Ecol Manag 348:124–141. https://doi.org/10.1016/j.foreco.2015.03.042
Albini FA, Reinhardt ED (1995) Modeling ignition and burning rate of large woody natural fuels. Int J Wildland Fire 5:81–91. https://doi.org/10.1071/WF9950081
Albini FA, Reinhardt ED (1997) Improved calibration of a large fuel burnout model. Int J Wildland Fire 7:21–28. https://doi.org/10.1071/WF9970021
Allen CD, Macalady AK, Chenchouni H, Bachelet D, McDowell N, Vennetier M, Kitzberger T, Rigling A, Breshears DD, Hogg ET, Gonzalez P (2010) A global overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests. For Ecol Manag 259:660–684. https://doi.org/10.1016/j.foreco.2009.09.001
Ambrose AR, Sillett SC, Koch GW, Van Pelt R, Antoine ME, Dawson TE (2010) Effects of height on treetop transpiration and stomatal conductance in coast redwood (Sequoia sempervirens). Tree Physiol 30:1260–1272. https://doi.org/10.1093/treephys/tpq064
Anderegg LD, Anderegg WR, Berry JA (2013) Not all droughts are created equal, translating meteorological drought into woody plant mortality. Tree Physiol 33:701–712. https://doi.org/10.1093/treephys/tpt044
Anderson NH, Sedell JR (1979) Detritus processing by macroinvertebrates in stream ecosystems. Annu Rev Entomol 24:351–377. https://doi.org/10.1146/annurev.en.24.010179.002031
Brauman A, Kane MD, Labat M, Breznak JA (1992) Genesis of acetate and methane by gut bacteria of nutritionally diverse termites. Science 257:1384–1387. https://doi.org/10.1126/science.257.5075.1384
Berner RA (2003) The long-term carbon cycle, fossil fuels and atmospheric composition. Nature 426:323–326. https://doi.org/10.1038/nature02131
Bond-Lamberty B, Gower ST (2008) Decomposition and fragmentation of coarse woody debris: re-visiting a boreal black spruce chronosequence. Ecosystems 11:831–840. https://doi.org/10.1007/s10021-008-9163-y
Boussu MF (1954) Relationship between trout populations and cover on a small stream. J Wildl Manag 18:229–239. https://doi.org/10.2307/3797719
Brown SL, Schroeder PE (1999) Spatial patterns of aboveground production and mortality of woody biomass for eastern US forests. Ecol Appl 9:968–980. https://doi.org/10.1890/1051-0761(1999)009[0968:SPOAPA]2.0.CO;2
Brown JK, Marsden MA, Ryan KC, Reinhardt ED (1985) Predicting duff and woody fuel consumed by prescribed fire in the northern Rocky Mountains. Res. Pap. INT-RP-337. USDA Forest Service. Intermountain Research Station, Ogden, UT
Brown JK, Reinhardt ED, Fischer WC (1991) Predicting duff and woody fuel consumption in northern Idaho prescribed fires. For Sci 37:1550–1566. https://doi.org/10.1093/forestscience/37.6.1550
Campbell JL, Green MB, Yanai RD, Woodall CW, Fraver S, Harmon ME, Hatfield MA, Barnett CJ, See CR, Domke GM (2019) Estimating uncertainty in the volume and carbon storage of downed coarse woody debris. Ecol Appl 29:e01844. https://doi.org/10.1002/eap.1844
Chen H, Harmon ME, Griffiths RP (2001) Decomposition and nitrogen release from decomposing woody roots in coniferous forests of the Pacific Northwest: a chronosequence approach. Can J For Res 31:246–260. https://doi.org/10.1139/x00-167
Covey KR, Wood SA, Warren RJ, Lee X, Bradford MA (2012) Elevated methane concentrations in trees of an upland forest. Geophys Res Lett 39(15). https://doi.org/10.1029/2012GL052361
Cowling EB, Merrill W (1966) Nitrogen in wood and its role in wood deterioration. Can J Bot 44:1539–1554. https://doi.org/10.1139/x06-070
Domke GM, Woodall CW, Walters BF, Smith JE (2013) From models to measurements: comparing downed dead wood carbon stock estimates in the U.S. Forest Inventory. PLoS One 8(3):e59949. https://doi.org/10.1371/journal.pone.0059949
Dukes JS (2003) Burning buried sunshine: human consumption of ancient solar energy. Clim Change 61:31–44. https://doi.org/10.1023/A:1026391317686
Elton CS (1966) The pattern of animal communities. Springer, Dordrecht. https://doi.org/10.1007/978-94-009-5872-2
Esau K (1965) Plant anatomy, 2nd edn. Wiley, New York
Feulner G (2017) Formation of most of our coal brought Earth close to global glaciation. Proc Natl Acad Sci 114:11333–11337. https://doi.org/10.1073/pnas.1712062114
Floudas D, Binder M, Riley R, Barry K, Blanchette RA, Henrissat B, Martínez AT, Otillar R, Spatafora JW, Yadav JS, Aerts A (2012) The Paleozoic origin of enzymatic lignin decomposition reconstructed from 31 fungal genomes. Science 336:1715–1719. https://doi.org/10.1126/science.1221748
Frandsen WH (1987) The influence of moisture and mineral soil on the combustion limits of smoldering forest duff. Can J For Res 17:1540–1544. https://doi.org/10.1139/x87-236
Franklin J, Bell D, Shaw D (2020) Long-term growth, mortality and regeneration of trees in permanent vegetation plots in the Pacific Northwest, 1910 to present. Long-Term Ecological Research. Forest Science Data Bank, Corvallis, OR. [Database]. http://andlter.forestry.oregonstate.edu/data/abstract.aspx?dbcode=TV010. https://doi.org/10.6073/pasta/2b389d702b0813701d07f52fd6df43e0. Accessed 25 Aug 2020
Fridman J, Walheim M (2000) Amount, structure, and dynamics of dead wood on managed forestland in Sweden. For Ecol Manag 131:23–36. https://doi.org/10.1016/S0378-1127(99)00208-X
Gibling MR, Davies NS (2012) Palaeozoic landscapes shaped by plant evolution. Nat Geosci 5:99–105. https://doi.org/10.1038/ngeo1376
Gosz JR, Likens GE, Bormann FH (1973) Nutrient release from decomposing leaf and branch litter in the Hubbard Brook Forest, New Hampshire. Ecol Monogr 43:173–191. https://doi.org/10.2307/1942193
Griffiths RP, Harmon ME, Caldwell BA, Carpenter SE (1993) Acetylene reduction in conifer logs during early stages of decomposition. Plant Soil 148:53–61. https://doi.org/10.1007/BF02185384
Hafner SD, Groffman PM, Mitchell MJ (2005) Leaching of dissolved organic carbon, dissolved organic nitrogen, and other solutes from coarse woody debris and litter in a mixed forest in New York State. Biogeochemistry 74:257–282. https://doi.org/10.1007/s10533-004-4722-6
Harmon ME, Sexton J (1995) Water balance of conifer logs in early stages of decomposition. Plant Soil 172:141–152. https://doi.org/10.1007/BF00020868
Harmon ME, Bell DM (2020) Mortality in forested ecosystems: suggested conceptual advances. Forests 11:572. https://doi.org/10.3390/f11050572
Harmon ME, Franklin JF, Swanson FJ, Sollins P, Lattin JD, Anderson NH, Gregory SV, Cline SP, Aumen NG, Sedell JR, Lienkaemper GW, Cromack K Jr, Cummins KW (1986) The ecology of coarse woody debris in temperate ecosystems. Adv Ecol Res 15:133–302. https://doi.org/10.1016/S0065-2504(08)60121-X
Harmon ME, Sexton J, Caldwell BA, Carpenter SE (1994) Fungal sporocarp mediated losses of Ca, Fe, K, Mg, Mn, N, P, and Zn from conifer logs in the early stages of decomposition. Can J For Res 24:1883–1893. https://doi.org/10.1139/x94-243
Harmon ME, Krankina ON, Yatskov M, Mathews E (2001) Predicting broad-scale carbon stores of woody detritus from plot-level data. In: Lai R, Kimble J, Stewart BA (eds) Assessment methods for soil carbon. CRC Press, New York, pp 533–552
Harmon ME, Bible K, Ryan MJ, Shaw D, Chen H, Klopatek J, Li X (2004) Production, respiration, and overall carbon balance in an old-growth Pseudotsuga/Tsuga forest ecosystem. Ecosystems 7:498–512. https://doi.org/10.1007/s10021-004-0140-9
Harmon ME, Bond‐Lamberty B, Tang J, Vargas R (2011) Heterotrophic respiration in disturbed forests: a review with examples from North America. J Geophys Res Biogeosci 116(G4). https://doi.org/10.1029/2010JG001495
Harmon ME, Fasth BG, Yatskov M, Kastendick D, Rock J, Woodall CW (2020) Release of coarse woody detritus-related carbon: a synthesis across forest biomes. Carbon Balance Manag 15:1–21. https://doi.org/10.1186/s13021-019-0136-6
Harris WF, Henderson GS, Todd DE (1972) Measurement of turnover of biomass and nutrient elements from the woody component of forest litter on walker branch watershed. Eastern Deciduous Forest Biome Memo Report #72-146 Oak Ridge National Laboratory, Oak Ridge, TN, USA
Hart SC (1999) Nitrogen transformations in fallen tree boles and mineral soil of an old-growth forest. Ecology 80:1385–1394. https://doi.org/10.1890/0012-9658(1999)080[1385:NTIFTB]2.0.CO;2
Heede BH (1972) Influences of a forest on the hydraulic geometry of two mountain streams 1. JAWRA J Am Water Resour Assoc 8:523–530. https://doi.org/10.1111/j.1752-1688.1972.tb05174.x
Hicks WT, Harmon ME (2002) Diffusion and seasonal dynamics of O2 in woody debris from the Pacific Northwest, USA. Plant Soil 243:67–79. https://doi.org/10.1023/A:1019906101359
Hollis JJ, Anderson WR, McCaw WL, Cruz MG, Burrows ND, Ward B, Tolhurst KG, Gould JS (2011) The effect of fireline intensity on woody fuel consumption in southern Australian eucalypt forest fires. Aust For 74:81–96. https://doi.org/10.1080/00049158.2011.10676350
Kahl T, Mun M, Bauhus J, Schulze ED (2012) Dissolved organic carbon from European beech logs: patterns of input to and retention by surface soil. Ecoscience 19:364–373. https://doi.org/10.2980/19-4-3501
Keller EA, Swanson FJ (1979) Effects of large organic material on channel form and fluvial processes. Earth Surf Process 4:361–380. https://doi.org/10.1002/esp.3290040406
Kimetu JM, Lehmann J, Ngoze SO, Mugendi DN, Kinyangi JM, Riha S, Verchot L, Recha JW, Pell AN (2008) Reversibility of soil productivity decline with organic matter of differing quality along a degradation gradient. Ecosystems 11:726. https://doi.org/10.1007/s10021-008-9154-z
Knorr W, Lehsten V, Arneth A (2012) Determinants and predictability of global wildfire emissions. Atmos Chem Phys 12(15):6845–6861. https://doi.org/10.5194/acp-12-6845-2012
Knorr W, Jiang L, Arneth A (2016) Climate, CO 2 and human population impacts on global wildfire emissions. Biogeosciences 13(1):267–282. https://doi.org/10.5194/bg-13-267-2016
Laiho R, Prescott CE (1999) The contribution of coarse woody debris to carbon, nitrogen, and phosphorus cycles in three Rocky Mountain coniferous forests. Can J For Res 29:1592–1603. https://doi.org/10.1139/x99-132
Lajtha K, Bowden RD, Crow S, Fekete I, Kotroczó Z, Plante A, Simpson MJ, Nadelhoffer KJ (2018) The detrital input and removal treatment (DIRT) network: insights into soil carbon stabilization. Sci Total Environ 640:1112–1120. https://doi.org/10.1016/j.scitotenv.2018.05.388
Lambert RL, Lang GE, Reiners WA (1980) Loss of mass and chemical change in decaying boles of a subalpine balsam fir forest. Ecology 61:1460–1473. https://doi.org/10.2307/1939054
Lodge DJ (1993) Nutrient cycling by fungi in wet tropical forests. In: Isaac S, Frankland JC, Walting R, Whalley AJS (eds) Aspects of tropical mycology. British Mycological Society, Cambridge, pp 37–57
Mann LK, Johnson DW, West DC, Cole DW, Hornbeck JW, Martin CW, Riekerk H, Smith CT, Swank WT, Tritton LM, Van Lear DH (1988) Effects of whole-tree and stem-only clearcutting on postharvest hydrologic losses, nutrient capital, and regrowth. For Sci 34:412–428. https://doi.org/10.1093/forestscience/34.2.412
McFee WW, Stone EL (1966) The persistence of decaying wood in the humus layers of northern forests. Soil Sci Soc Am J 30:513–516. https://doi.org/10.2136/sssaj1966.03615995003000040032x
Morgan RA, Litton CD, Salisbury CR (1987) Trackways and tree trunks—dating Neolithic oaks in the British Isles. Tree Ring Bull 47:61–69
Moroni MT, Morris DM, Shaw C, Stokland JN, Harmon ME, Fenton NJ, Merganičová K, Merganič J, Okabe K, Hagemann U (2015) Buried wood: a common yet poorly documented form of deadwood. Ecosystems 18:605–628. https://doi.org/10.1007/s10021-015-9850-4
Nelsen MP, DiMichele WA, Peters SE, Boyce CK (2016) Delayed fungal evolution did not cause the Paleozoic peak in coal production. Proc Natl Acad Sci 113:2442–2447. https://doi.org/10.1073/pnas.1517943113
Niinemets Ü (2001) Global-scale climatic controls of leaf dry mass per area, density, and thickness in trees and shrubs. Ecology 82:453–469. https://doi.org/10.1890/0012-9658(2001)082[0453:GSCCOL]2.0.CO;2
Nordén B, Götmark F, Tönnberg M, Ryberg M (2004) Dead wood in semi-natural temperate broadleaved woodland: contribution of coarse and fine dead wood, attached dead wood and stumps. For Ecol Manag 194:235–248. https://doi.org/10.1016/j.foreco.2004.02.043
Odum HT (1970) Chapter I-10 Summary: an emerging view of the ecological system at El Verde. In: Odum HT, Pigeon HF (eds) A tropical rain forest: a study of irradiation and ecology at El Verde, Puerto. US Atomic Energy Commission Technical Information Center, Oak Ridge, pp I-191–I-281
Ohlson M, Dahlberg B, Økland T, Brown KJ, Halvorsen R (2009) The charcoal carbon pool in boreal forest soils. Nat Geosci 2:692–695. https://doi.org/10.1038/ngeo617
Olsson BA, Bengtsson J, Lundkvist H (1996) Effects of different forest harvest intensities on the pools of exchangeable cations in coniferous forest soils. For Ecol Manag 84:135–147. https://doi.org/10.1016/0378-1127(96)03730-9
Pan Y, Birdsey RA, Fang J, Houghton R, Kauppi PE, Kurz WA, Phillips OL, Shvidenko A, Lewis SL, Canadell JG, Ciais P (2011) A large and persistent carbon sink in the world’s forests. Science 333:988–993. https://doi.org/10.1126/science.1201609
Parker GG (1983) Throughfall and stemflow in the forest nutrient cycle. Adv Ecol Res 13:57–133. https://doi.org/10.1016/S0065-2504(08)60108-7
Pausas JG (2015) Bark thickness and fire regime. Funct Ecol 29:315–327. https://doi.org/10.1111/1365-2435.12372
Pettersen RC (1984) The chemical composition of wood. In: Rowell RM (ed) The chemistry of solid wood. American Chemical Society, Washington, DC, pp 57–126
Prichard SJ, Wright CS, Vihnanek RE, Ottmar RD (2006) Predicting forest floor and woody fuel consumption from prescribed burns in ponderosa pine forests. In: Proceedings of the Third International Fire Ecology and Management Conference: Fire as a Global Process, San Diego, CA
Pugh TA, Rademacher T, Shafer SL, Steinkamp J, Barichivich J, Beckage B, Haverd V, Harper A, Heinke J, Nishina K, Rammig A (2020) Understanding the uncertainty in global forest carbon turnover. Biogeosciences 17:3961–3989. https://doi.org/10.5194/bg-17-3961-2020
Reich PB, Luo Y, Bradford JB, Poorter H, Perry CH, Oleksyn J (2014) Temperature drives global patterns in forest biomass distribution in leaves, stems, and roots. Proc Natl Acad Sci 111:13721–13726. https://doi.org/10.1073/pnas.1216053111
Reichle DE (ed) (1981) Dynamic properties of forest ecosystems, vol 23. Cambridge University Press, Cambridge
Rhoades CC, Battaglia MA, Rocca ME, Ryan MG (2012) Short-and medium-term effects of fuel reduction mulch treatments on soil nitrogen availability in Colorado conifer forests. For Ecol Manag 276:231–238. https://doi.org/10.1016/j.foreco.2012.03.028
Robinson JM (1990) Lignin, land plants, and fungi: biological evolution affecting Phanerozoic oxygen balance. Geology 18:607–610. https://doi.org/10.1130/0091-7613(1990)018%3c0607:LLPAFB%3e2.3.CO;2
Roskoski JP (1980) Nitrogen fixation in hardwood forests of the northeastern United States. Plant Soil 54:33–44. https://doi.org/10.1007/BF02181997
Rothermel RC (1972) A mathematical model for predicting fire spread in wildland fuels. Res. Pap. INT-115 US Department of Agriculture, Intermountain Forest and Range Experiment Station, Ogden UT
Sanderson MG (1996) Biomass of termites and their emissions of methane and carbon dioxide: a global database. Glob Biogeochem Cycles 10:543–557. https://doi.org/10.1029/96GB01893
Scheffer TC (1966) Natural resistance of wood to microbial deterioration. Annu Rev Phytopathol 4:147–168. https://doi.org/10.1146/annurev.py.04.090166.001051
Schlesinger WH (1978) Community structure, dynamics and nutrient cycling in the Okefenokee cypress swamp-forest. Ecol Monogr 48:43–65. https://doi.org/10.2307/2937359
Spears JDH, Lajtha K (2004) The imprint of coarse woody debris on soil chemistry in the western Oregon Cascades. Biogeochemistry 71:163–175. https://doi.org/10.1007/s10533-004-6395-6
Spears JD, Holub SM, Harmon ME, Lajtha K (2003) The influence of decomposing logs on soil biology and nutrient cycling in an old-growth mixed coniferous forest in Oregon, USA. Can J For Res 33:2193–2201. https://doi.org/10.1139/x03-148
Steudler PA, Bowden RD, Melillo JM, Aber JD (1989) Influence of nitrogen fertilization on methane uptake in temperate forest soils. Nature 341:314–316
Svensson M, Dahlberg A, Ranius T, Thor G (2014) Dead branches on living trees constitute a large part of the dead wood in managed boreal forests, but are not important for wood-dependent lichens. J Veg Sci 25:819–828. https://doi.org/10.1111/jvs.12131
Swift MJ, Heal OW, Anderson JM (1979) Decomposition in terrestrial ecosystems, vol 5. University of California Press, Berkeley
Taylor BR, Parkinson D, Parsons WF (1989) Nitrogen and lignin content as predictors of litter decay rates: a microcosm test. Ecology 70:97–104. https://doi.org/10.2307/1938416
Thiffault E, Hannam KD, Paré D, Titus BD, Hazlett PW, Maynard DG, Brais S (2011) Effects of forest biomass harvesting on soil productivity in boreal and temperate forests—a review. Environ Rev 19:278–309. https://doi.org/10.1139/a11-009
Tinker DB, Knight DH (2000) Coarse woody debris following fire and logging in Wyoming lodgepole pine forests. Ecosystems 3:472–483. https://doi.org/10.1007/s100210000041
Uhl D, Jasper A, Hamad AMA, Montenari M (2008) Permian and Triassic wildfires and atmospheric oxygen levels. 1st WSEAS International Conference on Environmental and Geological Science and Engineering (EG’08), pp 179–187
Walker GM, White NA (2017) Introduction to fungal physiology. In: Kavanagh K (ed) Fungi: biology and applications, 3rd edn. Wiley-Blackwell, Hoboken, pp 1–35
Walmsley JD, Jones DL, Reynolds B, Price MH, Healey JR (2009) Whole tree harvesting can reduce second rotation forest productivity. For Ecol Manag 257:1104–1111. https://doi.org/10.1016/j.foreco.2008.11.015
Wei X, Waterhouse MJ, Qi G, Wu J (2020) Long-term logging residue loadings affect tree growth but not soil nutrients in Pinus contorta Doug. ex Loud. forests. Ann For Sci 77:1–10. https://doi.org/10.1007/s13595-020-00968-8
Weng JK, Chapple C (2010) The origin and evolution of lignin biosynthesis. New Phytol 187:273–285. https://doi.org/10.1111/j.1469-8137.2010.03327.x
Williams AP, Allen CD, Macalady AK, Griffin D, Woodhouse CA, Meko DM, Swetnam TW, Rauscher SA, Seager R, Grissino-Mayer HD, Dean JS (2013) Temperature as a potent driver of regional forest drought stress and tree mortality. Nat Clim Change 3:292–297. https://doi.org/10.1038/nclimate1693
Worthington EB (1965) The international biological programme. Nature 208:223–226
Zechmeister-Boltenstern S, Keiblinger KM, Mooshammer M, Peñuelas J, Richter A, Sardans J, Wanek W (2015) The application of ecological stoichiometry to plant–microbial–soil organic matter transformations. Ecol Monogr 85:133–155. https://doi.org/10.1890/14-0777.1
Acknowledgements
Data were provided by the HJ Andrews Experimental Forest research program, funded by the National Science Foundation’s Long-Term Ecological Research Program (DEB 1440409), US Forest Service Pacific Northwest Research Station, and Oregon State University. This work was also supported by the National Science Foundation’s OPUS (DEB-1353159). I am grateful for the helpful suggestions and corrections provided by two anonymous reviewers.
Funding
Funding for the research related to this manuscript came from the National Science Foundation, the U.S. Forest Service Pacific Northwest Research Station, and Oregon State University.
Author information
Authors and Affiliations
Contributions
MEH is the sole contributor to this manuscript.
Corresponding author
Ethics declarations
Conflict of interest
There are no conflicts of interest or competing interests related to this manuscript.
Additional information
Responsible Editor: Kate Lajtha.
Publisher's Note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
This paper is an invited contribution to the 35th Anniversary Special Issue, edited by Sujay Kaushal, Robert Howarth, and Kate Lajtha.
Supplementary Information
Below is the link to the electronic supplementary material.
Rights and permissions
About this article
Cite this article
Harmon, M.E. The role of woody detritus in biogeochemical cycles: past, present, and future. Biogeochemistry 154, 349–369 (2021). https://doi.org/10.1007/s10533-020-00751-x
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s10533-020-00751-x


