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Coarse woody debris

Coarse woody debris (CWD) or coarse woody habitat (CWH) refers to fallen dead trees and the remains of large branches on the ground in forests[1] and in rivers or wetlands.[2] A dead standing tree – known as a snag – provides many of the same functions as coarse woody debris. The minimum size required for woody debris to be defined as "coarse" varies by author, ranging from 2.5–20 cm (1–8 in) in diameter.[3]

Coarse woody debris in Białowieża Forest, Poland

Since the 1970s, forest managers worldwide have been encouraged[by whom?] to allow dead trees and woody debris to remain in woodlands, recycling nutrients trapped in the wood and providing food and habitat for a wide range of organisms, thereby improving biodiversity. The amount of coarse woody debris is considered[by whom?] an important criterion for the evaluation and restoration of temperate deciduous forest.[1] Coarse woody debris is also important in wetlands, particularly in deltas where woody debris accumulates.[2]

Sources

 
Beetle larvae (Prionoplus reticularis) consuming CWD, New Zealand

Coarse woody debris comes from natural tree mortality, disease, and insects, as well as catastrophic events such as fires, storms and floods.[citation needed]

Ancient, or old growth, forest, with its dead trees and woody remains lying where they fell to feed new vegetation, constitutes the ideal woodland in terms of recycling and regeneration. In healthy temperate forests, dead wood comprises up to thirty per cent of all woody biomass. In recent British studies, woods managed for timber had between a third and a seventh less fallen debris than unmanaged woods that had been left undisturbed for many years, while in recently coppiced woods the amount of CWD was almost zero.[citation needed]

In old growth Douglas fir forests of the Pacific Northwest of North America, CWD concentrations were found to be from 72 metric tons/hectare (64,000 pounds/acre) in drier sites to 174 t/ha (155,000 lb/acre) in moister sites.[4] Australian native forests have mean CWD concentrations ranging from 19 t/ha (17,000 lb/acre) to 134 t/ha (120,000 lb/acre), depending on forest type.[5]

Benefits

 
Fungi sprouting from fallen log, Germany

Nutrient cycling

Coarse woody debris and its subsequent decomposition recycles nutrients that are essential for living organisms, such as carbon, nitrogen, potassium, and phosphorus. Saprotrophic fungi and detritivores such as bacteria and insects directly consume dead wood, releasing nutrients by converting them into other forms of organic matter which may then be consumed by other organisms. However CWD itself is extremely scarce in physiologically important nutrients. To fulfill nutritional requirements of consumers, CWD must be first nutritionally enriched by transport of nutrients from outside of CWD.[6][7] Thus CWD is important actor contributing to soil nutrients cycles. CWD, while itself not particularly rich in nitrogen, contributes nitrogen to the ecosystem by acting as a host for nonsymbiotic free-living nitrogen-fixing bacteria.[8]

Scientific studies show that coarse woody debris can be a significant contributor to biological carbon sequestration. Trees store atmospheric carbon in their wood using photosynthesis. Once the trees die, fungi and other saprotrophs transfer some of that carbon from CWD into the soil. This sequestration can continue in old-growth forests for hundreds of years.[9][10]

Habitat

By providing both food and microhabitats for many species, coarse woody debris helps to maintain the biodiversity of forest ecosystems. Up to forty percent of all forest fauna is dependent on CWD. Studies in western North America showed that only five per cent of living trees consisted of living cells by volume, whereas in dead wood it was as high as forty percent by volume, mainly fungi and bacteria.[11] Colonizing organisms that live on the remains of cambium and sapwood of dead trees aid decomposition and attract predators that prey on them and so continue the chain of metabolizing the biomass.[citation needed]

 
Ensatina eschscholtzii, a species of salamander associated with CWD in western North America[12]

The list of organisms dependent on CWD for habitat or as a food source includes bacteria, fungi, lichens, mosses and other plants, and in the animal kingdom, invertebrates such as termites, ants, beetles, and snails,[13] amphibians such as salamanders,[12] reptiles such as the slow-worm, as well as birds and small mammals. One third of all woodland birds live in the cavities of dead tree trunks. Woodpeckers, tits, chickadees, and owls all live in dead trees, and grouse shelter behind woody debris.[citation needed]

Some plants use coarse woody debris as habitat. Mosses and lichens may cover logs, while ferns and trees may regenerate on the top of logs. Large fragments of CWD that provide such habitat for herbs, shrubs, and trees are called nurse logs.[8] CWD can also protect young plants from herbivory damage by acting as barriers to browsing animals. The persistence of coarse woody debris can shelter organisms during a large disturbance to the ecosystem such as wildfire or logging.[citation needed]

Rivers and wetlands

Fallen debris and trees in streams provide shelter for fish, amphibians and mammals by modifying the flow of water and sediment.[14][15] Turtles of many species may also use coarse woody debris for basking.[16] Musk turtles may lay their eggs under logs near wetlands.[17]

Soil

Coarse woody debris, particularly on slopes, stabilizes soils by slowing downslope movement of organic matter and mineral soil. Leaves and other debris collect behind CWD, allowing for decomposition to occur. Infiltration of precipitation is improved as well. During dry weather, CWD slows evaporation of soil moisture and provides damp microhabitats for moisture-sensitive organisms.[8]

Wildfire

 
Coarse woody debris may contribute to the intensity of wildfire.

In fire-prone forests, coarse woody debris can be a significant fuel during a wildfire. High amounts of fuels can lead to increased fire severity and size. CWD may be managed to reduce fuel levels, particularly in forests where fire exclusion has resulted in the buildup of fuels. Reductions in CWD for fire safety should be balanced with the retention of CWD for habitat and other benefits.[18] CWD is classified as 1000-hour fuel by fire managers, referring to the amount of time needed for the moisture level in the wood to react to the surrounding environment.[19]

Regional examples

 
Blera fallax, Belgium

In Glen Affric, Scotland, the Trees for Life group found the black tinder fungus beetle (Bolitothorus reticulatus) is dependent on a particular fungus (Fomes fomentarius), which itself grows only on dead birch. Another insect, the pine hoverfly (Blera fallax), requires rotting Scots Pine in order to reproduce.[11]

In the temperate deciduous forests of Eastern North America, coarse woody debris provides habitat ranging from salamanders to ferns. It is an important indicator for evaluating and restoring this type of forest.[1]

In certain subtropical areas such as Australia where bushfire constitutes a major hazard, the amount of CWD left standing or lying is determined by what may be considered safe in the course of reasonable fire prevention. However, when fires do occur, invertebrates find shelter either within or beneath dead tree logs.[citation needed]

In Canada, bears seek out dead tree logs to tear open and look for and feed on ants and beetles, a fact that has encouraged the authorities to reserve a sufficient amount of coarse woody debris for these purposes. In North America, too, CWD is often used as barriers to prevent browsing deer and elk from damaging young trees.[citation needed]

See also

References

  1. ^ a b c Keddy, P.A.; Drummond, C.G. (1996). "Ecological properties for the evaluation, management, and restoration of temperate deciduous forest ecosystems". Ecological Applications. 6 (3): 748–762. doi:10.2307/2269480. JSTOR 2269480.
  2. ^ a b Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p, p. 225-227.
  3. ^ Lofroth, Eric (1998), , in Voller, J.; Harrison, S. (eds.), Conservation biology principles for forested landscapes, Vancouver, B.C.: UBC Press, pp. 185–214, archived from the original on 2008-05-01, retrieved 2007-12-06
  4. ^ Spies, Thomas A.; Franklin, Jerry F.; Thomas, Ted B. (December 1988). "Coarse Woody Debris in Douglas-Fir Forests of Western Oregon and Washington". Ecology. 69 (6): 1689–1702. doi:10.2307/1941147. JSTOR 1941147.
  5. ^ Woldendorp, G.; Keenan, R. J. (2005). "Coarse woody debris in Australian forest ecosystems: A review". Austral Ecology. 30 (8): 834–843. doi:10.1111/j.1442-9993.2005.01526.x.
  6. ^ Filipiak, Michał; Weiner, January (2014-12-23). "How to Make a Beetle Out of Wood: Multi-Elemental Stoichiometry of Wood Decay, Xylophagy and Fungivory". PLOS ONE. 9 (12): e115104. Bibcode:2014PLoSO...9k5104F. doi:10.1371/journal.pone.0115104. ISSN 1932-6203. PMC 4275229. PMID 25536334.
  7. ^ Filipiak, Michał; Sobczyk, Łukasz; Weiner, January (2016-04-09). "Fungal Transformation of Tree Stumps into a Suitable Resource for Xylophagous Beetles via Changes in Elemental Ratios". Insects. 7 (2): 13. doi:10.3390/insects7020013. PMC 4931425.
  8. ^ a b c Stevens, Victoria (1997), The ecological role of coarse woody debris: an overview of the ecological importance of CWD in B.C. forests (PDF), Working Paper 30/1997, Victoria, B.C.: Research Branch, B.C. Ministry of Forests
  9. ^ Barford, CC; Wofsy, SC; Goulden, ML; Munger, JW; Pyle, EH; Urbanski, SP; Hutyra, L; Saleska, SR; Fitzjarrald, D; Moore, K (23 November 2001). "Factors Controlling Long- and Short-Term Sequestration of Atmospheric CO2 in a Mid-latitude Forest". Science. 294 (5547): 1688–1691. Bibcode:2001Sci...294.1688B. doi:10.1126/science.1062962. PMID 11721047. S2CID 20420952.
  10. ^ Luyssaert, Sebastiaan; -Detlef Schulze, E.; Börner, Annett; Knohl, Alexander; Hessenmöller, Dominik; Law, Beverly E.; Ciais, Philippe; Grace, John (11 September 2008). "Old-growth forests as global carbon sinks" (PDF). Nature. 455 (7210): 213–215. Bibcode:2008Natur.455..213L. doi:10.1038/nature07276. PMID 18784722. S2CID 4424430.
  11. ^ a b Puplett, Dan. . Trees For Life. Archived from the original on 2014-03-01. Retrieved 2011-01-26.
  12. ^ a b Butts, Sally R.; McComb, William C. (January 2000). "Associations of Forest-Floor Vertebrates with Coarse Woody Debris in Managed Forests of Western Oregon". The Journal of Wildlife Management. 64 (1): 95–104. doi:10.2307/3802978. JSTOR 3802978.
  13. ^ Kappes, H. (2005). "Influence Of Coarse Woody Debris On The Gastropod Community Of A Managed Calcareous Beech Forest In Western Europe". Journal of Molluscan Studies. 71 (2): 85–91. doi:10.1093/mollus/eyi011.
  14. ^ Bilby, R. E.; Ward, J. (1991). "Characteristics and function of large woody debris in streams draining old-growth, clear-cut, and 2nd-growth forests in southwestern Washington". Canadian Journal of Fisheries and Aquatic Sciences. 48 (12): 2499–508. doi:10.1139/f91-291. S2CID 67789816.
  15. ^ Crook, D. A.; Robertson, A. I. (1999). "Relationships between riverine fish and woody debris: implications for lowland rivers". Marine and Freshwater Research. 50 (8): 941–53. doi:10.1071/mf99072.
  16. ^ Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p. Figure 8.15.
  17. ^ Ernst, C.H., R.W. Barbour, and J.E. Lovich. 1994. Turtles of the United States and Canada. Smithsonian Institution, Washington. p. 146.
  18. ^ Knapp, E.E.; J.E. Keeley; E. A. Ballenger & T.J. Brennan (2005). "Fuel reduction and coarse woody debris dynamics with early and late season prescribed fire in a Sierra Nevada mixed conifer forest" (PDF). Forest Ecology and Management. 208: 383–397. doi:10.1016/j.foreco.2005.01.016.
  19. ^ "Glossary". Fire Effects Information System. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory. Retrieved March 27, 2012.

Further reading

  • Franklin J. F., Lindenmayer D., MacMahon J. A., McKee A., Magnuson J., Perry D. A., Waide R. & Foster D. (2000). "Threads of Continuity". Conservation Biology in Practice. [Malden, MA] Blackwell Science, Inc. 1(1) pp9–16.
  • Proceedings of the Symposium on the Ecology and Management of Dead Wood in Western Forests. PSW-GTR-181. William F. Laudenslayer, Jr., Patrick J. Shea, Bradley E. Valentine, C. Phillip Weatherspoon, and Thomas E. Lisle Technical Coordinators.
  • Titus, B.D.; Prescott, C.E.; Maynard, D.G.; Mitchell, A.K.; Bradley, R.L.; Feller, M.C.; Koppenaa, R. (2006). "Postharvest nitrogen cycling in clearcut and alternative silvicultural systems in a montane forest in coastal British Columbia" (PDF). The Forestry Chronicle. 82 (6): 844–859. doi:10.5558/tfc82844-6. hdl:10613/2703.

External links

coarse, woody, debris, coarse, woody, habitat, refers, fallen, dead, trees, remains, large, branches, ground, forests, rivers, wetlands, dead, standing, tree, known, snag, provides, many, same, functions, coarse, woody, debris, minimum, size, required, woody, . Coarse woody debris CWD or coarse woody habitat CWH refers to fallen dead trees and the remains of large branches on the ground in forests 1 and in rivers or wetlands 2 A dead standing tree known as a snag provides many of the same functions as coarse woody debris The minimum size required for woody debris to be defined as coarse varies by author ranging from 2 5 20 cm 1 8 in in diameter 3 Coarse woody debris in Bialowieza Forest Poland Since the 1970s forest managers worldwide have been encouraged by whom to allow dead trees and woody debris to remain in woodlands recycling nutrients trapped in the wood and providing food and habitat for a wide range of organisms thereby improving biodiversity The amount of coarse woody debris is considered by whom an important criterion for the evaluation and restoration of temperate deciduous forest 1 Coarse woody debris is also important in wetlands particularly in deltas where woody debris accumulates 2 Contents 1 Sources 2 Benefits 2 1 Nutrient cycling 2 2 Habitat 2 3 Rivers and wetlands 2 4 Soil 3 Wildfire 4 Regional examples 5 See also 6 References 7 Further reading 8 External linksSources Edit Beetle larvae Prionoplus reticularis consuming CWD New Zealand Coarse woody debris comes from natural tree mortality disease and insects as well as catastrophic events such as fires storms and floods citation needed Ancient or old growth forest with its dead trees and woody remains lying where they fell to feed new vegetation constitutes the ideal woodland in terms of recycling and regeneration In healthy temperate forests dead wood comprises up to thirty per cent of all woody biomass In recent British studies woods managed for timber had between a third and a seventh less fallen debris than unmanaged woods that had been left undisturbed for many years while in recently coppiced woods the amount of CWD was almost zero citation needed In old growth Douglas fir forests of the Pacific Northwest of North America CWD concentrations were found to be from 72 metric tons hectare 64 000 pounds acre in drier sites to 174 t ha 155 000 lb acre in moister sites 4 Australian native forests have mean CWD concentrations ranging from 19 t ha 17 000 lb acre to 134 t ha 120 000 lb acre depending on forest type 5 Benefits Edit Fungi sprouting from fallen log Germany Nutrient cycling Edit Coarse woody debris and its subsequent decomposition recycles nutrients that are essential for living organisms such as carbon nitrogen potassium and phosphorus Saprotrophic fungi and detritivores such as bacteria and insects directly consume dead wood releasing nutrients by converting them into other forms of organic matter which may then be consumed by other organisms However CWD itself is extremely scarce in physiologically important nutrients To fulfill nutritional requirements of consumers CWD must be first nutritionally enriched by transport of nutrients from outside of CWD 6 7 Thus CWD is important actor contributing to soil nutrients cycles CWD while itself not particularly rich in nitrogen contributes nitrogen to the ecosystem by acting as a host for nonsymbiotic free living nitrogen fixing bacteria 8 Scientific studies show that coarse woody debris can be a significant contributor to biological carbon sequestration Trees store atmospheric carbon in their wood using photosynthesis Once the trees die fungi and other saprotrophs transfer some of that carbon from CWD into the soil This sequestration can continue in old growth forests for hundreds of years 9 10 Habitat Edit By providing both food and microhabitats for many species coarse woody debris helps to maintain the biodiversity of forest ecosystems Up to forty percent of all forest fauna is dependent on CWD Studies in western North America showed that only five per cent of living trees consisted of living cells by volume whereas in dead wood it was as high as forty percent by volume mainly fungi and bacteria 11 Colonizing organisms that live on the remains of cambium and sapwood of dead trees aid decomposition and attract predators that prey on them and so continue the chain of metabolizing the biomass citation needed Ensatina eschscholtzii a species of salamander associated with CWD in western North America 12 The list of organisms dependent on CWD for habitat or as a food source includes bacteria fungi lichens mosses and other plants and in the animal kingdom invertebrates such as termites ants beetles and snails 13 amphibians such as salamanders 12 reptiles such as the slow worm as well as birds and small mammals One third of all woodland birds live in the cavities of dead tree trunks Woodpeckers tits chickadees and owls all live in dead trees and grouse shelter behind woody debris citation needed Some plants use coarse woody debris as habitat Mosses and lichens may cover logs while ferns and trees may regenerate on the top of logs Large fragments of CWD that provide such habitat for herbs shrubs and trees are called nurse logs 8 CWD can also protect young plants from herbivory damage by acting as barriers to browsing animals The persistence of coarse woody debris can shelter organisms during a large disturbance to the ecosystem such as wildfire or logging citation needed Rivers and wetlands Edit Fallen debris and trees in streams provide shelter for fish amphibians and mammals by modifying the flow of water and sediment 14 15 Turtles of many species may also use coarse woody debris for basking 16 Musk turtles may lay their eggs under logs near wetlands 17 Soil Edit Coarse woody debris particularly on slopes stabilizes soils by slowing downslope movement of organic matter and mineral soil Leaves and other debris collect behind CWD allowing for decomposition to occur Infiltration of precipitation is improved as well During dry weather CWD slows evaporation of soil moisture and provides damp microhabitats for moisture sensitive organisms 8 Wildfire Edit Coarse woody debris may contribute to the intensity of wildfire In fire prone forests coarse woody debris can be a significant fuel during a wildfire High amounts of fuels can lead to increased fire severity and size CWD may be managed to reduce fuel levels particularly in forests where fire exclusion has resulted in the buildup of fuels Reductions in CWD for fire safety should be balanced with the retention of CWD for habitat and other benefits 18 CWD is classified as 1000 hour fuel by fire managers referring to the amount of time needed for the moisture level in the wood to react to the surrounding environment 19 Regional examples Edit Blera fallax Belgium In Glen Affric Scotland the Trees for Life group found the black tinder fungus beetle Bolitothorus reticulatus is dependent on a particular fungus Fomes fomentarius which itself grows only on dead birch Another insect the pine hoverfly Blera fallax requires rotting Scots Pine in order to reproduce 11 In the temperate deciduous forests of Eastern North America coarse woody debris provides habitat ranging from salamanders to ferns It is an important indicator for evaluating and restoring this type of forest 1 In certain subtropical areas such as Australia where bushfire constitutes a major hazard the amount of CWD left standing or lying is determined by what may be considered safe in the course of reasonable fire prevention However when fires do occur invertebrates find shelter either within or beneath dead tree logs citation needed In Canada bears seek out dead tree logs to tear open and look for and feed on ants and beetles a fact that has encouraged the authorities to reserve a sufficient amount of coarse woody debris for these purposes In North America too CWD is often used as barriers to prevent browsing deer and elk from damaging young trees citation needed See also EditLarge woody debris Nurse log Plant litter Snag ecology Soil life Tree hollowReferences Edit a b c Keddy P A Drummond C G 1996 Ecological properties for the evaluation management and restoration of temperate deciduous forest ecosystems Ecological Applications 6 3 748 762 doi 10 2307 2269480 JSTOR 2269480 a b Keddy P A 2010 Wetland Ecology Principles and Conservation 2nd edition Cambridge University Press Cambridge UK 497 p p 225 227 Lofroth Eric 1998 The dead wood cycle in Voller J Harrison S eds Conservation biology principles for forested landscapes Vancouver B C UBC Press pp 185 214 archived from the original on 2008 05 01 retrieved 2007 12 06 Spies Thomas A Franklin Jerry F Thomas Ted B December 1988 Coarse Woody Debris in Douglas Fir Forests of Western Oregon and Washington Ecology 69 6 1689 1702 doi 10 2307 1941147 JSTOR 1941147 Woldendorp G Keenan R J 2005 Coarse woody debris in Australian forest ecosystems A review Austral Ecology 30 8 834 843 doi 10 1111 j 1442 9993 2005 01526 x Filipiak Michal Weiner January 2014 12 23 How to Make a Beetle Out of Wood Multi Elemental Stoichiometry of Wood Decay Xylophagy and Fungivory PLOS ONE 9 12 e115104 Bibcode 2014PLoSO 9k5104F doi 10 1371 journal pone 0115104 ISSN 1932 6203 PMC 4275229 PMID 25536334 Filipiak Michal Sobczyk Lukasz Weiner January 2016 04 09 Fungal Transformation of Tree Stumps into a Suitable Resource for Xylophagous Beetles via Changes in Elemental Ratios Insects 7 2 13 doi 10 3390 insects7020013 PMC 4931425 a b c Stevens Victoria 1997 The ecological role of coarse woody debris an overview of the ecological importance of CWD in B C forests PDF Working Paper 30 1997 Victoria B C Research Branch B C Ministry of Forests Barford CC Wofsy SC Goulden ML Munger JW Pyle EH Urbanski SP Hutyra L Saleska SR Fitzjarrald D Moore K 23 November 2001 Factors Controlling Long and Short Term Sequestration of Atmospheric CO2 in a Mid latitude Forest Science 294 5547 1688 1691 Bibcode 2001Sci 294 1688B doi 10 1126 science 1062962 PMID 11721047 S2CID 20420952 Luyssaert Sebastiaan Detlef Schulze E Borner Annett Knohl Alexander Hessenmoller Dominik Law Beverly E Ciais Philippe Grace John 11 September 2008 Old growth forests as global carbon sinks PDF Nature 455 7210 213 215 Bibcode 2008Natur 455 213L doi 10 1038 nature07276 PMID 18784722 S2CID 4424430 a b Puplett Dan Ecological Features of the Caledonian Forest Dead Wood Trees For Life Archived from the original on 2014 03 01 Retrieved 2011 01 26 a b Butts Sally R McComb William C January 2000 Associations of Forest Floor Vertebrates with Coarse Woody Debris in Managed Forests of Western Oregon The Journal of Wildlife Management 64 1 95 104 doi 10 2307 3802978 JSTOR 3802978 Kappes H 2005 Influence Of Coarse Woody Debris On The Gastropod Community Of A Managed Calcareous Beech Forest In Western Europe Journal of Molluscan Studies 71 2 85 91 doi 10 1093 mollus eyi011 Bilby R E Ward J 1991 Characteristics and function of large woody debris in streams draining old growth clear cut and 2nd growth forests in southwestern Washington Canadian Journal of Fisheries and Aquatic Sciences 48 12 2499 508 doi 10 1139 f91 291 S2CID 67789816 Crook D A Robertson A I 1999 Relationships between riverine fish and woody debris implications for lowland rivers Marine and Freshwater Research 50 8 941 53 doi 10 1071 mf99072 Keddy P A 2010 Wetland Ecology Principles and Conservation 2nd edition Cambridge University Press Cambridge UK 497 p Figure 8 15 Ernst C H R W Barbour and J E Lovich 1994 Turtles of the United States and Canada Smithsonian Institution Washington p 146 Knapp E E J E Keeley E A Ballenger amp T J Brennan 2005 Fuel reduction and coarse woody debris dynamics with early and late season prescribed fire in a Sierra Nevada mixed conifer forest PDF Forest Ecology and Management 208 383 397 doi 10 1016 j foreco 2005 01 016 Glossary Fire Effects Information System U S Department of Agriculture Forest Service Rocky Mountain Research Station Fire Sciences Laboratory Retrieved March 27 2012 Further reading EditFranklin J F Lindenmayer D MacMahon J A McKee A Magnuson J Perry D A Waide R amp Foster D 2000 Threads of Continuity Conservation Biology in Practice Malden MA Blackwell Science Inc 1 1 pp9 16 Proceedings of the Symposium on the Ecology and Management of Dead Wood in Western Forests PSW GTR 181 William F Laudenslayer Jr Patrick J Shea Bradley E Valentine C Phillip Weatherspoon and Thomas E Lisle Technical Coordinators Titus B D Prescott C E Maynard D G Mitchell A K Bradley R L Feller M C Koppenaa R 2006 Postharvest nitrogen cycling in clearcut and alternative silvicultural systems in a montane forest in coastal British Columbia PDF The Forestry Chronicle 82 6 844 859 doi 10 5558 tfc82844 6 hdl 10613 2703 External links Edit Wikimedia Commons has media related to Coarse woody debris Retrieved from https en wikipedia org w index php title Coarse woody debris amp oldid 1134873151, wikipedia, wiki, book, books, library,

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