Annals of Forest Science

, Volume 72, Issue 7, pp 955–965 | Cite as

European beech, silver fir, and Norway spruce differ in establishment, height growth, and mortality rates on coarse woody debris and forest floor—a study from a mixed beech forest in the Western Carpathians

  • Olga OrmanEmail author
  • Janusz Szewczyk
Original Paper


Key message

In mixed forests, coarse woody debris promotes the successful establishment and growth of conifers and beech. In contrast to beech and fir, older spruce seedlings were only present on coarse woody debris and not on the forest floor.


Coarse woody debris (CWD) is considered a suitable seedbed for small-seeded and light-demanding species. Its role in enhancing tree regeneration is well reported in boreal or subalpine spruce forests. Less is known about its role in the establishment, growth, and survival of other species, particularly in mixed forests.


We analyzed the role of CWD in seedling establishment, growth, and survival for European beech, silver fir, and Norway spruce.


We tracked the growth and survival of all germinants and seedlings over 5 years.


Conifers were relatively more successful than beech at colonizing on CWD. The density of seedlings was variable in all CWD decay classes but was the highest on well-decomposed CWD. CWD supported the growth of all species. Beech cohorts and older seedlings had similar mortality rates on both microsites. Spruce germinants did not survive on the forest floor for more than a year, and older seedlings were only observed on CWD. Fir cohorts had similar mortality rates on both microsites, but older seedlings survived better on the forest floor.


Although the three species differed in their preferred microsite for establishment, CWD can be considered a suitable regeneration microsite for all three species by enhancing their growth and, in the case of spruce, both short- and long-term survival.


Regeneration Colonization pattern Decaying wood Microsite 



The assistance of Wojciech Cieślik, Irena Kulikowska, Maciej Pawlaczek, and Tomasz Pachowicz in field data collection is appreciated. We thank Jerzy Szwagrzyk and Maki Suzuki, and the anonymous reviewers for their comments on the manuscript. We thank Gregory J. Sproull for revising the English.


This study was supported by the research grants: N N304 325436 (2009-12) and N N309 716440 (2011-14) of the Polish Ministry of Science and Higher Education.


  1. Bače R, Svoboda M, Janda P (2011) Density and height structure of seedlings in subalpine spruce forests of Central Europe: Logs vs. Stumps as a favourable substrate. Silva Fennica 45:1065–1078CrossRefGoogle Scholar
  2. Bače R, Svoboda M, Pouska V, Janda P, Červenka J (2012) Natural regeneration in Central-European subalpine spruce forests: Which logs are suitable for seedling recruitment? For Ecol Manag 266:254–262. doi: 10.1016/j.foreco.2011.11.025 CrossRefGoogle Scholar
  3. Baier R, Ettl R, Hahn C, Göttlein A (2006) Early development and nutrition of Norway spruce (Picea abies (L.) Karst.) seedlings on different seedbeds in the Bavarian Limestone Alps – a bioassay. Ann For Sci 63:339–348. doi: 10.1051/forest:2006014 CrossRefGoogle Scholar
  4. Baier R, Meyer J, Göttlein A (2007) Regeneration niches of Norway spruce (Picea abies [L.] Karst.) saplings in small canopy gaps in mixed mountain forests of the Bavarian Limestone Alps. Eur J Forest Res 126:11–22CrossRefGoogle Scholar
  5. Bellingham PJ, Richardson SJ (2006) Tree seedling growth and survival over 6 years across different microsites in a temperate rain forest. Can J For Res 36:910–918CrossRefGoogle Scholar
  6. Brunner A, Kimmins JP (2003) Nitrogen fixation in coarse woody debris of Thuja plicata and Tsuga heterophylla forests on northern Vancouver Island. Can J For Res 33:1670–1682. doi: 10.1139/x03-085 CrossRefGoogle Scholar
  7. Carlton GC, Bazzaz FA (1998) Regeneration of three sympatric birch species on experimental hurricane blowdown microsites. Ecol Monograph 68:99–120CrossRefGoogle Scholar
  8. Christie DA, Armesto JJ (2003) Regeneration microsites and tree species coexistence in temperate rain forests of Chiloe Island, Chile. J Ecol 91:776–784. doi: 10.1046/j.1365-2745.2003.00813.x CrossRefGoogle Scholar
  9. Christy EJ, Mack RN (1984) Variation in demography of juvenile Tsuga heterophylla across the substratum mosaic. J Ecol 72:75–91CrossRefGoogle Scholar
  10. Grodziński W, Sawicka-Kapusta K (1970) Energy values of tree-seeds eaten by small mammals. Oikos 21:52–58CrossRefGoogle Scholar
  11. Harmon ME (1989a) Effects of bark fragmentation on plant succession on conifer logs in the Picea-Tsuga forests of Olympic National Park, Washington. Am Midl Nat 121:112–124CrossRefGoogle Scholar
  12. Harmon ME (1989b) Retention of needles and seed on logs in Picea sitchensisTsuga heterophylla forests of coastal Oregon and Washington. Can J Bot 67:1833–1837CrossRefGoogle Scholar
  13. Harmon ME, Franklin JF, Swanson FJ, Sollins P, Gregory SV, Lattin JD, Anderson NH, Cline SP, Aumen NG, Sedell JR, Lienkaemper GW, Cromack K Jr, Cummins KW (2004) Ecology of woody debris in temperate ecosystems. Adv Ecol Res Classic Papers 34:59–234CrossRefGoogle Scholar
  14. Holeksa J (2003) Relationship among field-layer vegetation and canopy openings in a Carpathian subalpine spruce forest. Plant Ecol 168:57–67. doi: 10.1023/A:1024457303815 CrossRefGoogle Scholar
  15. Hunziker U, Brang P (2005) Microsite patterns of conifer seedling establishment and growth in a mixed stand in the southern Alps. For Ecol Manag 210:67–79. doi: 10.1016/j.foreco.2005.02.019 CrossRefGoogle Scholar
  16. Iijima H, Shibuya M (2010) Evaluation of suitable conditions for natural regeneration of Picea jezoensis on fallen logs. J For Res 15:46–54. doi: 10.1007/s10310-009-0133-9 CrossRefGoogle Scholar
  17. Iijima H, Shibuya M, Saito H, Takahashi K (2006) The water relation of seedlings of Picea jezoensis on fallen logs. Can J For Res 36:664–670. doi: 10.1139/x05-285 CrossRefGoogle Scholar
  18. Iijima H, Shibuya M, Saito H (2007) Effects of surface and light conditions of fallen logs on the emergence and survival of coniferous seedlings and saplings. J For Res 12:262–269. doi: 10.1007/s10310-007-0012-1 CrossRefGoogle Scholar
  19. Ježek K (2004) Contribution of regeneration on dead wood to spontaneous regeneration of a mountain forest. J For Sci 50:405–414CrossRefGoogle Scholar
  20. Knapp AK, Smith WK (1982) Factors influencing understory seedling establishment of Engelman spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa) in southeast Wyoming. Can J Bot 60:2753–2761CrossRefGoogle Scholar
  21. Laiho R, Prescott CE (2004) Decay and nutrient dynamics of coarse woody debris in northern coniferous forests: a synthesis. Can J For Res 34:763–777. doi: 10.1139/X03-241 CrossRefGoogle Scholar
  22. Lusk CH, Kelly CK (2003) Interspecific variation in seed size and safe sites in a temperate rain forest. New Phytol 158:535–541. doi: 10.1046/j.1469-8137.2003.00760.x CrossRefGoogle Scholar
  23. Maser C, Tarrant RF, Trappe JM, Franklin JF (1988) From the Forest to the Sea: A Story of Fallen Trees – General Technical Report PNW-GTR-229. USDA Forest Service, Portland, OregonCrossRefGoogle Scholar
  24. McKenny HJA, Kirkpatrick JB (1999) The role of fallen logs in the regeneration of tree species in Tasmanian mixed forest. Austral J Bot 47:745–753. doi: 10.1071/BT98001 CrossRefGoogle Scholar
  25. Moles AT, Westoby M (2004) Seedling survival and seed size: a synthesis of the literature. J Ecol 92:372–383. doi: 10.1111/j.0022-0477.2004.00884.x CrossRefGoogle Scholar
  26. Mori A, Mizumachi E (2005) Season and substrate effects on the first-year establishment of current-year seedlings of major conifer species in an old-growth subalpine forest in central Japan. For Ecol Manag 210:461–467. doi: 10.1016/j.foreco.2005.02.027 CrossRefGoogle Scholar
  27. Mori A, Mizumachi E, Osono T, Doi Y (2004) Substrate-associated seedling recruitment and establishment of major conifer species in an old-growth subalpine forest in central Japan. For Ecol Manag 196:287–297. doi: 10.1016/j.foreco.2004.03.027 CrossRefGoogle Scholar
  28. Narukawa Y, Yamamoto S (2002) Effects of dwarf bamboo (Sasa sp.) and forest floor microsites on conifer seedling recruitment in a subalpine forest, Japan. For Ecol Manag 163:61–70. doi: 10.1016/S0378-1127(01)00527-8 CrossRefGoogle Scholar
  29. Narukawa Y, Iida S, Tanouchi H, Abe S, Yamamoto S (2003) State of fallen logs and the occurrence of conifer seedlings and saplings in boreal and subalpine old-growth forests in Japan. Ecol Res 18:267–277. doi: 10.1046/j.1440-1703.2003.00553.x CrossRefGoogle Scholar
  30. Noguchi M, Yoshida T (2004) Tree regeneration in partially cut conifer-hardwood mixed forests in northern Japan: roles of establishment substrate and dwarf bamboo. For Ecol Manag 190:335–344. doi: 10.1016/j.foreco.2003.10.024 CrossRefGoogle Scholar
  31. Ódor P, Heilmann-Clausen J, Christensen M, Aude E, van Dort KW, Piltaver A, Siller I, Veerkamp MT, Walleyn R, Standovár T, van Hees AFM, Kosec J, Matočec N, Kraigher H, Grebenc T (2006) Diversity of dead wood inhabiting fungi and bryophytes in semi-natural beech forests in Europe. Biol Conserv 131:58–71. doi: 10.1016/j.biocon.2006.02.004 CrossRefGoogle Scholar
  32. Ota T, Masaki T, Sugita H, Kanazashi T, Abe H (2012) Properties of stumps that promote the growth and survival of Japanese cedar saplings in a natural old-growth forest. Can J For Res 42:1976–1982. doi: 10.1139/x2012-141 CrossRefGoogle Scholar
  33. Parent S, Walsh D, Simard MJ, Morin H, Gagnon R (2006) Emergence of balsam fir seedlings under increasing broadleaf litter thickness. Can J For Res 36:1976–1981. doi: 10.1139/x06-091 CrossRefGoogle Scholar
  34. Pawlaczek M (2010) Skład gatunkowy i struktura drzewostanu w rezerwacie Dolina Łopusznej w Gorczańskim Parku Narodowym. BSc thesis, University of Agriculture in Cracow [in Polish]Google Scholar
  35. Pouska V, Svoboda M, Lepšová A (2010) The diversity of wood-decaying fungi in relation to changing site conditions in old-growth mountain spruce forest, Central Europe. Eur J For Res 129:219–231. doi: 10.1007/s10342-009-0324-0 CrossRefGoogle Scholar
  36. Różański W, Czarnota P, Stefanik M, Tomasiewicz J (2006) Gorczański Park Narodowy – 25 lat ochrony dziedzictwa przyrodniczego i kulturowego Gorców. Gorczański Park Narodowy, Poręba Wielka [in Polish]Google Scholar
  37. Saniga M, Kucbel S, Anić I, Mikac S, Prebeg M (2011) Structure, growing stock, coarse woody debris and regeneration process in virgin forest Dobroč (Slovakia) and Čorkova Uvala (Croatia). Beskydy 4:39–50Google Scholar
  38. Simon A, Gratzer G, Sieghardt M (2011) The influence of windthrow microsites on tree regeneration and establishment in an old growth mountain forest. For Ecol Manag 262:1289–1297. doi: 10.1016/j.foreco.2011.06.028 CrossRefGoogle Scholar
  39. Sollins P (1982) Input and decay of coarse woody debris in coniferous stands in western Oregon and Washington. Can J For Res 12:18–28CrossRefGoogle Scholar
  40. Stancioiu PT, O’Hara KL (2006) Regeneration growth in different light environments of mixed species, multiaged, mountainous forests of Romania. Eur J Forest Res 125:151–162CrossRefGoogle Scholar
  41. Stokland JN, Siitonen J, Jonsson BG (2012) Biodiversity in dead wood. Cambridge University Press, CambridgeCrossRefGoogle Scholar
  42. Svoboda M, Fraver S, Janda P, Bače R, Zenáhlíková J (2010) Natural development and regeneration of a Central European montane spruce forest. For Ecol Manag 260:707–714. doi: 10.1016/j.foreco.2010.05.027 CrossRefGoogle Scholar
  43. Szewczyk J, Szwagrzyk J (1996) Tree regeneration on rotten wood and on soil in old-growth stand. Vegetatio 122:37–46CrossRefGoogle Scholar
  44. Szwagrzyk J, Szewczyk J, Bodziarczyk J (2001) Dynamics of seedling banks in beech forest: result of 10-years of study on germination, growth and survival. For Ecol Manag 141:237–250. doi: 10.1016/S0378-1127(00)00332-7 CrossRefGoogle Scholar
  45. Takahashi M, Sakai Y, Ootomo R, Shiozaki M (2000) Establishment of tree seedlings and water-soluble nutrients in coarse woody debris in an old-growth Picea-Abies forest in Hokkaido, northern Japan. Can J For Res 30:1148–1155. doi: 10.1139/x00-042 CrossRefGoogle Scholar
  46. Ulanova NG (2000) The effects of windthrow on forests at different spatial scales: a review. For Ecol Manag 135:155–167. doi: 10.1016/S0378-1127(00)00307-8 CrossRefGoogle Scholar
  47. Vrška T, Adam D, Hort L, Kolár T, Janík D (2009) European beech (Fagus sylvatica L.) and silver fir (Abies alba Mill.) rotation in the Carpathians – A development cycle or a linear trend induced by man? For Ecol Manag 258:347–356. doi: 10.1016/j.foreco.2009.03.007 CrossRefGoogle Scholar
  48. Zielonka T (2006) When does dead wood turn into a substrate for spruce replacement? J Veg Sci 17:739–746. doi: 10.1111/j.1654-1103.2006.tb02497.x CrossRefGoogle Scholar
  49. Zielonka T, Niklasson M (2001) Decomposition rate of dead wood and regeneration pattern in natural spruce forest in the Tatra Mountains. Ecol Bulletins 49:159–163Google Scholar
  50. Zielonka T, Piątek G (2004) The herb and dwarf shrubs colonization of decaying logs in subalpine forest in the Polish Tatra Mountains. Plant Ecol 172:63–72. doi: 10.1023/B:VEGE.0000026037.03716.fc CrossRefGoogle Scholar

Copyright information

© The Author(s) 2015

Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (, which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made.

Authors and Affiliations

  1. 1.Department of Forest Biodiversity, Forest Ecology and Silviculture InstituteUniversity of AgricultureKrakówPoland
  2. 2.Department of Natural Environmental Studies, Graduate School of Frontier SciencesThe University of TokyoKashiwaJapan

Personalised recommendations