, Volume 21, Issue 1, pp 85–97 | Cite as

Dissecting the Effects of Diameter on Wood Decay Emphasizes the Importance of Cross-Stem Conductivity in Fraxinus americana

  • Brad OberleEmail author
  • Kristofer R. Covey
  • Kevin M. Dunham
  • Edgar J. Hernandez
  • Maranda L. Walton
  • Darcy F. Young
  • Amy E. Zanne


Pest outbreaks are driving tree dieback and major influxes of deadwood into forest ecosystems. Understanding how pulses of deadwood impact the climate system requires understanding which factors influence greenhouse gas production during wood decay. Recent analyses identify stem diameter as an important control, but report effects that vary in magnitude and direction. This complexity may reflect interacting effects of soil contact, geometry and variable tissue properties. To dissect these effects, we implemented a three-way factorial experiment in Fraxinus americana, (white ash), an iconic North American species threatened by an invasive beetle. Soil contact accelerated decay rates by an order of magnitude with an effect that varied with stem diameter, not bark presence. After experimentally controlling surface area-to-volume ratio, half-buried wide stems decayed more slowly than half-buried narrow stems but more quickly than the aggregate decay rate of buried and suspended stems. These results closely matched variation in moisture content within and among samples, suggesting that limited vertical conduction of soil moisture through deadwood mediates the effect of stem diameter on wood decay. Soil contact also influenced greenhouse gas concentrations reinforcing recent evidence that deadwood acts as a source for CO2 and CH4 while acting as a sink for N2O. Our results suggest that managing tree species affected by pest outbreaks, including F. americana, for biomass salvage and greenhouse gas mitigation requires understanding traits that mediate wood permeability and diffusivity to soil moisture and greenhouse gases.


carbon dioxide forest carbon emerald ash borer methane nitrous oxide wood decay 



The authors thank the staff of Tyson Research Center and Washington University in St. Louis for access to the site and for logistical support. In particular, we thank T. Mohrman for his help in safely harvesting the large tree. A. Miller helped with initial deployment. S. Hobbie and two anonymous reviewers provided constructive comments that helped improve an earlier draft of this manuscript. Funding was provided by NSF Grant DEB 1302797 to AEZ and NSF Grant DGE 1405135 to KRC.

Supplementary material

10021_2017_136_MOESM1_ESM.jpg (2.2 mb)
FIGURE S1: Experimental set-up, illustrating wide and narrow diameter samples, both with and without bark in both suspended and half-buried vertical positions. (JPG 2296 kb)
10021_2017_136_MOESM2_ESM.pdf (6 kb)
FIGURE S2: Principal components analyses show different patterns of variation in internal gas concentrations in suspended (A) compared to buried (B) Fraxinus stem segments. (PDF 6 kb)
10021_2017_136_MOESM3_ESM.xlsx (9 kb)
TABLE S1: Model selection statistics for estimating initial moisture content. (XLSX 9 kb)
10021_2017_136_MOESM4_ESM.docx (18 kb)
TABLE S2: Summary of hypotheses, predictions and statistical tests. (DOCX 18 kb)


  1. Allen CD, Macalady AK, Chenchouni H, Bachelet D, McDowell N, Vennetier M, Kitzberger T, Rigling A, Breshears DD, Hogg EH. 2010. A Global Overview of Drought and Heat-Induced Tree Mortality Reveals Emerging Climate Change Risks for Forests. Forest Ecology and Management 259:660–84.CrossRefGoogle Scholar
  2. Bates D, Maechler M, Bolker B, Walker S. 2014. lme4: Linear Mixed-Effects Models Using Eigen and S4. R package version 1.1-7.Google Scholar
  3. Bretz F, Hothorn T, Westfall P. 2015. Multiple Comparisons Using R. Boca Raton: CRC Press, Taylor & Francis.Google Scholar
  4. Carll CG, Highley TL. 1999. Decay of Wood and Wood-Based Products Above Ground in Buildings. Jteva 27:150–8.Google Scholar
  5. Cavanaugh JE. 1997. Unifying the Derivations for the Akaike and Corrected Akaike Information Criteria. Statistics & Probability Letters 33:201–8.CrossRefGoogle Scholar
  6. Cornwell WK, Cornelissen JHC, Allison SD, Bauhus J, Eggleton P, Preston CM, Scarff F, Weedon JT, Wirth C, Zanne AE. 2009. Plant Traits and Wood Fates Across the Globe: Rotted, Burned, or Consumed? Global Change Biology 15:2431–49.CrossRefGoogle Scholar
  7. Covey KR, de Mesquita CPB, Oberle B, Maynard DS, Bettigole C, Crowther TW, Duguid MC, Steven B, Zanne AE, Lapin M. 2016. Greenhouse Trace Gases in Deadwood. Biogeochemistry 130:215–26.CrossRefGoogle Scholar
  8. Covey KR, Wood SA, Warren RJ, Lee X, Bradford MA. 2012. Elevated Methane Concentrations in Trees of an Upland Forest. Geophysical Research Letters 39:1–6.CrossRefGoogle Scholar
  9. Dossa GGO, Paudel E, Cao K, Schaefer D, Harrison RD. 2016. Factors Controlling Bark Decomposition and Its Role in Wood Decomposition in Five Tropical Tree Species. Scientific Reports 6:34153.CrossRefPubMedPubMedCentralGoogle Scholar
  10. Dwason H. 2013. Working Group I Contribution to the IPCC Fifth Assessment Report—Summary for Policymakers. Climate Change 2013: The Physical Science Basis vol 53, p 1–36.Google Scholar
  11. Fasth BG, Harmon ME, Sexton J, White P. 2011. Decomposition of Fine Woody Debris in a Deciduous Forest in North Carolina 1. The Journal of the Torrey Botanical Society 138:192–206.CrossRefGoogle Scholar
  12. Frangi JL, Richter LL, Barrera MD, Aloggia M. 1997. Decomposition of Nothofagus Fallen Woody Debris in Forests of Tierra del Fuego, Argentina. Canadian Journal of Forest Research 27:1095–102.CrossRefGoogle Scholar
  13. Freschet GT, Weedon JT, Aerts R, van Hal JR, Cornelissen JHC. 2012. Interspecific Differences in Wood Decay Rates: Insights from a New Short-Term Method to Study Long-Term Wood Decomposition. Journal of Ecology 100:161–70.CrossRefGoogle Scholar
  14. Ganjegunte GK, Condron LM, Clinton PW, Davis MR, Mahieu N. 2004. Decomposition and Nutrient Release from Radiata Pine (Pinus radiata) Coarse Woody Debris. Forest Ecology and Management 187:197–211.CrossRefGoogle Scholar
  15. Garrett LG, Kimberley MO, Oliver GR, Pearce SH, Paul TSH. 2010. Decomposition of Woody Debris in Managed Pinus Radiata Plantations in New Zealand. Forest Ecology and Management 260:1389–98.CrossRefGoogle Scholar
  16. Harmon M, Franklin J, Swanson F, Sollins P, Gregory S, Lattin J, Anderson N, Cline S, Aumen N, Sedell J, Lienkaemper G, Cromack K, Cummins K. 1986. Ecology of Coarse Woody Debris in Temperate Ecosystems. Advances in Ecological Research 15:133–302.CrossRefGoogle Scholar
  17. Klooster WS, Herms DA, Knight KS, Herms CP, McCullough DG, Smith A, Gandhi KJK, Cardina J. 2014. Ash (Fraxinus spp.) Mortality, Regeneration, and Seed Bank Dynamics in Mixed Hardwood Forests Following Invasion by Emerald Ash Borer (Agrilus planipennis). Biological Invasions 16:859–73.CrossRefGoogle Scholar
  18. Krinner G, Viovy N, de Noblet-Ducoudré N, Ogee J, Polcher J, Friedlingstein P, Ciais P, Sitch S, Prentice IC. 2005. A Dynamic Global Vegetation Model for Studies of the Coupled Atmosphere-Biosphere System. Global Biogeochemical Cycles 19:GB1015.CrossRefGoogle Scholar
  19. Kurz WA, Dymond CC, Stinson G, Rampley GJ, Neilson ET, Carroll AL, Ebata T, Safranyik L. 2008. Mountain Pine Beetle and Forest Carbon Feedback to Climate Change. Nature 452:987–90.CrossRefPubMedGoogle Scholar
  20. Laiho R, Prescott CE. 2004. Decay and Nutrient Dynamics of Coarse Woody Debris in Northern Coniferous Forests: A Synthesis. Canadian Journal of Forest Research 34:763–77.CrossRefGoogle Scholar
  21. van der Lee GEM, de Winder B, Bouten W, Tietema A. 1999. Anoxic Microsites in Douglas Fir Litter. Soil Biology and Biochemistry 31:1295–301.CrossRefGoogle Scholar
  22. Moore DJP, Trahan NA, Wilkes P, Quaife T, Stephens BB, Elder K, Desai AR, Negron J, Monson RK. 2013. Persistent Reduced Ecosystem Respiration After Insect Disturbance in High Elevation Forests. Ecology Letters 16:731–7.CrossRefPubMedPubMedCentralGoogle Scholar
  23. Mori S, Itoh A, Nanami S, Tan S, Chong L, Yamakura T. 2014. Effect of Wood Density and Water Permeability on Wood Decomposition Rates of 32 Bornean Rainforest Trees. Journal of Plant Ecology 7:356–63.CrossRefGoogle Scholar
  24. Moser WK, Hansen MH, Brand GJ, Treiman TB. 2007. Missouri’s Forest Resources in 2005. USDA General Technical Report NRS-15. St. Paul, MN.Google Scholar
  25. Næsset E. 1999. Decomposition Rate Constants of Picea abies Logs in Southeastern Norway. Canadian Journal of Forest Research 29:372–81.CrossRefGoogle Scholar
  26. Oberle B, Dunham K, Milo AM, Walton M, Young DF, Zanne AE. 2014. Progressive, Idiosyncratic Changes in Wood Hardness During Decay: Implications for Dead Wood Inventory and Cycling. Forest Ecology and Management 323:1–9.CrossRefGoogle Scholar
  27. Oberle B, Ogle K, Zuluaga JCP, Sweeney J, Zanne AE. 2016. A Bayesian Model for Xylem Vessel Length Accommodates Subsampling and Reveals Skewed Distributions in Species that Dominate Seasonal Habitats. Journal of Plant Hydraulics 3:3.CrossRefGoogle Scholar
  28. Pan Y, Birdsey RA, Fang J, Houghton R, Kauppi PE, Kurz WA, Phillips OL, Shvidenko A, Lewis SL, Canadell JG, Ciais P, Jackson RB, Pacala SW, McGuire AD, Piao S, Rautiainen A, Sitch S, Hayes D. 2011. A Large and Persistent Carbon Sink in the World’s Forests. Science 333:988–93.CrossRefPubMedGoogle Scholar
  29. Parton WJ, Stewart JWB, Cole CV. 1988. Dynamics of C, N, P and S in Grassland Soils: a Model. Biogeochemistry 5:109–31.CrossRefGoogle Scholar
  30. Pyle C, Brown M. 1998. A Rapid System of Decay Classification for Hardwood Logs of the Eastern Deciduous Forest Floor. Journal of the Torrey Botanical Society 125:237–45.CrossRefGoogle Scholar
  31. R Development Core Team. 2010. R: A Language and Environment for Statistical Computing.
  32. Romero LM, Smith TJ, Fourqurean JW. 2005. Changes in Mass and Nutrient Content of Wood During Decomposition in a South Florida Mangrove Forest. Journal of Ecology 93:618–31.CrossRefGoogle Scholar
  33. Rosell JA, Gleason S, Méndez-Alonzo R, Chang Y, Westoby M. 2013. Bark Functional Ecology: Evidence for Tradeoffs, Functional Coordination, and Environment Producing Bark Diversity. The New Phytologist 201:486–97.CrossRefPubMedGoogle Scholar
  34. Song Z, Dunn C, Lü X-T, Qiao L, Pang J-P, Tang J-W. 2017. Coarse Woody Decay Rates Vary by Physical Position in Tropical Seasonal Rainforests of SW China. Forest Ecology and Management 385:206–13.CrossRefGoogle Scholar
  35. Sorz J, Hietz P. 2006. Gas Diffusion Through Wood: Implications for Oxygen Supply. Trees - Structure and Function 20:34–41.CrossRefGoogle Scholar
  36. Spasojevic MJ, Yablon EA, Oberle B, Myers JA. 2014. Ontogenetic Trait Variation Influences Tree Community Assembly Across Environmental Gradients. Ecosphere 5:1–20.CrossRefGoogle Scholar
  37. Taylor AM, Garner BL, Morrell JJ. 2002. Heartwood Formation and Natural Durability—A Review. Wood and Fiber Science 34:587–611.Google Scholar
  38. Thomson AJ, Giannopoulos G, Pretty J, Baggs EM, Richardson DJ. 2012. Biological Sources and Sinks of Nitrous Oxide and Strategies To Mitigate Emissions. Philosophical Transactions of the Royal Society of London Series B, Biological Sciences 367:1157–68.CrossRefPubMedPubMedCentralGoogle Scholar
  39. Whittaker C, Yates NE, Powers SJ, Misselbrook T, Shield I. 2016. Dry Matter Losses and Greenhouse Gas Emissions From Outside Storage of Short Rotation Coppice Willow Chip. Bioenergy Research 9:288–302.CrossRefPubMedGoogle Scholar
  40. Zanne AE, Oberle B, Dunham KM, Milo AM, Walton ML, Young DF. 2015. A Deteriorating State of Affairs: How Endogenous and Exogenous Factors Determine Plant Decay Rates. Journal of Ecology 103:1421–31.CrossRefGoogle Scholar
  41. Zuo J, Berg MP, Klein R, Nusselder J, Neurink G, Decker O, Hefting MM, Sass-Klaassen U, van Logtestijn RSP, Goudzwaard L, van Hal J, Sterck FJ, Poorter L, Cornelissen JHC. 2016. Faunal Community Consequence of Interspecific Bark Trait Dissimilarity in Early-Stage Decomposing Logs. Functional Ecology 30:1957–66.Google Scholar

Copyright information

© Springer Science+Business Media New York 2017

Authors and Affiliations

  1. 1.Division of Natural SciencesNew College of FloridaSarasotaUSA
  2. 2.School of Forestry and Environmental StudiesYale UniversityNew HavenUSA
  3. 3.Department of BiologyUniversity of Missouri-St. LouisSt. LouisUSA
  4. 4.Department of BiologyUniversity of Missouri-St. LouisSt. LouisUSA
  5. 5.Department of BiologyWashington University in St. LouisSt. LouisUSA
  6. 6.Department of Biological SciencesThe George Washington UniversityWashingtonUSA
  7. 7.Center for Conservation and Sustainable DevelopmentMissouri Botanical GardenSt. LouisUSA

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