Dieback of managed spruce stands in western Germany promotes beetle diversity

Abstract

Biodiversity loss is a significant problem at a global scale and may be amplified by climate change. In recent years, coniferous forests have had substantial dieback across Europe due to drought and subsequent bark-beetle outbreaks. As many studies on the consequences of disturbance and subsequent management have focused on natural stands, management implications for managed spruce stands are not well understood, even though such stands are widespread throughout Europe. In this study, beetle taxonomy, conservation value, and community composition are compared among spruce plantations and four post-disturbance management approaches: standing deadwood, lying deadwood, clear cuts, and long-term succession. Diversity and community composition differed significantly among management categories, while different beetle families responded similarly. Intact spruce stands harbored the lowest beetle diversity while the highest taxonomic diversity and conservation value was on clear cuts and stands with lying or standing deadwood. The proportion of forest specialists was highest in successional forests. In summary, different forest management categories harbored distinct beetle communities at the family-, species-, and ecological guild levels. Therefore, post-disturbance management should consider the landscape scale and include different management types. This enhances landscape heterogeneity and thus overall biodiversity but could also mitigate negative impacts of natural disturbances on ecosystem services.

Keywords

Bark-beetle outbreak / Forest dieback / Spruce / Deadwood / Nature conservation / Beetle biodiversity / Coleoptera / Coniferous forests

References

[1]	Barnosky AD, Matzke N, Tomiya S, Wogan GO, Swartz B, Quental TB, Marshall C, McGuire JL, Lindsey EL, Maguire KC, Mersey B, Ferrer EA (2011) Has the earth’s sixth mass extinction already arrived? Nature 471(7336):51–57. https://doi.org/10.1038/nature09678
[2]	Bellard C, Bertelsmeier C, Leadley P, Thullier W, Courchamp F (2012) Impacts of climate change on the future of biodiversity. Ecol Lett 15(4):365–377. https://doi.org/10.1111/j.1461-0248.2011.01736.x
[3]	Beudert B, B?ssler C, Thorn S, Noss R, Schr?der B, Dieffenbach-Fries H, Foullois N, Müller J (2015) Bark beetles increase biodiversity while maintaining drinking water quality. Conserv Lett 8(4):272–281. https://doi.org/10.1111/conl.12153
[4]	Bivand RS, Wong DWS (2018) Comparing implementations of global and local indicators of spatial association. TEST 27(3):716–748. https://doi.org/10.1007/s11749-018-0599-x
[5]	BMEL (2021) [Bundesministerium für Ern?hrung und Landwirtschaft] Ergebnisse der Waldzustandserhebung 2021, 1–76 (in German)
[6]	Brodribb TJ, Powers J, Cochard H, Choat B (2020) Hanging by a thread? For Drought Sci 368(6488):261–266. https://doi.org/10.1126/science.aat7631
[7]	Bruun HH, Heilmann-Clausen J (2021) What is unmanaged forest and how does it sustain biodiversity in landscapes with a long history of intensive forestry? J Appl Ecol 58(9):1813–1816. https://doi.org/10.1111/1365-2664.13754
[8]	Cardinale BJ, Duffy JE, Gonzalez A, Hooper D, Perrings C, Venail P, Narwani A, MacE G, Timan D, Wardle D, Kinzig A, Daily G, Loreau M, Grace J, Larigauderie A, Srivastava D, Naeem S (2012) Biodiversity loss and its impact on humanity. Nature 486(7401):59–67. https://doi.org/10.1038/nature11148
[9]	Cobb TP, Morissette JL, Jacobs JM, Koivula MJ, Spence JR, Langor DW (2011) Effects of postfire salvage logging on deadwood-associated beetles. Conserv Biol 25(1):94–104. https://doi.org/10.1111/j.l523-17392010.01566.x
[10]	Conner RN, Rudolph DC, Kulhavy D, Snow A (1991) Causes of mortality of red-cockaded woodpecker cavity trees. J Wildl Manage 55(1):531–537
[11]	De Warnaffe GDB, Lebrun P (2004) Effects of forest management on carabid beetles in Belgium: implications for biodiversity conservation. Biol Conserv 118(2):219–234. https://doi.org/10.1016/j.biocon.2003.08.015
[12]	Dirzo R, Young HS, Galetti M, Ceballos G, Isaac NJB, Collen B (2014) Defaunation in the anthropocene. Science 345(6195):401–406. https://doi.org/10.1126/science.1251817
[13]	Doerfler I, Gossner MM, Müller J, Seibold S, Weiser WW (2018) Deadwood enrichment combining integrative and segregative conservation elements enhances biodiversity of multiple taxa in managed forests. Biol Conserv 228:70–78. https://doi.org/10.1016/j.biocon.2018.10.013
[14]	Dormann CF, McPherson JM, Araújo MB, Bivand R, Bolliger J, Carl G, Davies RG, Hirzel A, Jetz W, Kissling D, Kühn I, Ohlemüller R, Peres-Neto PR, Reineking B, Schr?der B, Schurr FM, Wilson R (2007) Methods to account for spatial autocorrelation in the analysis of species distributional data: a review. Ecography 30:609–628. https://doi.org/10.1111/j.2007.0906-7590.05171.x
[15]	Fahrig L (2003) Effects of habitat fragmentation on biodiversity. Annu Rev Ecol Evol Syst 34:487–515. https://doi.org/10.1146/annurev.ecolsys.34.011802.132419
[16]	FAO (2020) [Food and Agriculture Organization of the United Nations] Global forest resources assessment. https://fra-data.fao.org/WO/fra2020/home/ Accessed 15 Jun 2022
[17]	Freude H, Harde KW, Lohse GA (1983) Die K?fer Mitteleuropas. Vol. 1–11. Goecke & Evers Verlag, Krefeld, 3761 pages (in German)
[18]	García-Tejero S, Spence JR, O’Halloran J, Bourassa S, Oxbrough A (2018) Natural succession and clearcutting as drivers of environmental heterogeneity and beta diversity in North American boreal forests. PLoS ONE 13(11):e0206931. https://doi.org/10.1371/journal.pone.0206931
[19]	Geiser R (1998) Rote Liste der K?fer (Coleoptera). In: Bundesamt für Naturschutz (Hrsg): Rote Liste gef?hrdeter Tiere Deutschlands. Schriftenr Landschaftspflege Naturschutz 55: 178–179 (in German).
[20]	Georgiev KB, B?ssler C, Feldhaar H, Heibl C, Karasch P, Müller J, Perlik M, Weiss I, Thorn S (2022) Windthrow and salvage logging alter β-diversity of multiple species groups in a mountain spruce forest. For Ecol Manage 520:120401. https://doi.org/10.1016/j.foreco.2022.120401
[21]	Gerlach J, Samways M, Pryke J (2013) Terrestrial invertebrates as bioindicators: an overview of available taxonomic groups. J Insect Conserv 17:831–850. https://doi.org/10.1007/s10841-013-9565-9
[22]	G?rn S, Fischer K (2011) Niedermoore Nordostdeutschland bewerten. Nat Landsch 43(7):211–217 ((in German))
[23]	Grove SJ (2002) Saproxylic insect ecology and the sustainable management of forests. Annu Rev Ecol Syst 33:1–23. https://doi.org/10.1146/annurev.ecolsys.33.010802.150507
[24]	Hagge J, Leibl F, Müller J, Plechinger M, Soutinho JG, Thorn S (2019a) Reconciling pest control, nature conservation, and recreation in coniferous forests. Conserv Lett. https://doi.org/10.1111/conl.12615
[25]	Hagge J, Müller J, B?ssler C, Biebl SS, Brandl R, Drexler M, Gruppe A, Hotes S, Hothorn T, Langhammer P, Stark H, Wirtz R, Zimmerer V, Mysterud A (2019b) Deadwood retention in forests lowers short-term browsing pressure on silver fir saplings by overabundant deer. For Ecol Manag. https://doi.org/10.1016/j.foreco.2019.117531
[26]	Hagge J, Müller J, Birkemoe T, Buse J, Bojesen Christensen RH, Gossner MM, Gruppe A, Heibl C, Jarzabek-Müller A, Seibold S, Siitonen J, Soutinho JG, Sverdrup-Thygeson A, Thorn S, Drag L (2021) What does a threatened saproxylic beetle look like? Modelling extinction risk using a new morphological trait database. J Anim Ecol 90(8):1934–1947. https://doi.org/10.1111/1365-2656.13512
[27]	Hallmann CA, Sorg M, Jongejans E, Siepel H, Hofland N, Schwan H, Stenmans W, Müller A, Sumser H, H?rren T, Goulson D, de Kroon H (2017) More than 75 percent decline over 27 years in total flying insect biomass in protected areas. PLoS ONE. https://doi.org/10.1371/journal.pone.0185809
[28]	Hlásny T, Krokene P, Liebhold A, Montagné-Huck C, Müller J, Qin H, Raffa KJ, Schelhaas MJ, Seidl R, Svoboda M, Viiri H (2017) Living with bark beetles: impacts, outlook and management options. From Sci Policy 8:1–52. https://doi.org/10.36333/fs08
[29]	Hlásny T, K?nig L, Krokene P, Lindner M, Montagné-Huck C, Müller J, Qin H, Raffa KF, Schelhaas M-J, Svoboda M, Viiri H, Seidl R (2021) Bark beetle outbreaks in Europe: state of knowledge and ways forward for management. Curr for Rep 7(3):138–165. https://doi.org/10.1007/s40725-021-00142-x
[30]	Homburg K, Drees C, Boutaud E, Nolte D, Schuett W, Zumstein P, von Ruschkowski E, Assmann T (2019) Where have all the beetles gone? Long-term study reveals carabid species decline in a nature reserve in Northern Germany. Insect Conserv Divers 12:268–277. https://doi.org/10.1111/icad.12348
[31]	Horák J, Brestovanská T, Mladenovi? S, Kout J, Bogusch P, Halda JP, Zasadil P (2019) Green desert?: Biodiversity patterns in forest plantations. For Ecol Manag 433:343–348. https://doi.org/10.1016/j.foreco.2018.11.019
[32]	Hothorn T, Bretz F, Westfall P (2008) Simultaneous inference in general parametric models. Biom J 50(3):346–363. (package = "multcomp")
[33]	Hsieh TC, Ma KH, Chao A (2016) iNEXT: An R package vor interpolation and extrapolation of species diversity (Hill numbers). Methods Ecol Evol 7:1451–1456
[34]	IPCC (2018) [Intergovernmental Panel on Climate Change]. Summary for policymakers. In: Global warming of 1.5 °C. An IPCC special report on the impacts of global warming of 1.5 °C above pre-industrial levels and related global greenhouse gas emission pathways, in the context of strengthening the global response to the threat of climate change, sustainable development, and efforts to eradicate poverty.
[35]	IPCC (2022) [Intergovernmental Panel on Climate Change]. Climate change 2022: impacts, adaptation, and vulnerability. Contribution of working group II to the sixth assessment report of the intergovernmental panel on climate change.
[36]	Jost L (2006) Entropy and diversity. Oikos 113(2):363–375. https://doi.org/10.1111/j.2006.0030-1299.14714.x
[37]	Kausrud K, ?kland B, Skarpaas O, Grégoire J-C, Erbilgin N, Stenseth NC (2012) Population dynamics in changing environments: the case of an eruptive forest pest species. Biol Rev 87(1):34–51. https://doi.org/10.1111/j.1469-185X.2011.00183.x
[38]	Koch K (1996) Die K?fer Mitteleuropas - ?kologie, Vol. 1–3. Goecke & Evers Verlag, Krefeld, 1211 pages (in German)
[39]	Koivula M, Spence JR (2006) Effects of post-fire salvage logging on boreal mixed-wood ground beetle assemblages (Coleoptera, Carabidae). For Ecol Manag 236(1):102–112. https://doi.org/10.1016/j.foreco.2006.09.004
[40]	Koivula M, Kukkonen J, Niemela J (2002) Boreal carabid-beetle (Coleoptera, Carabidae) assemblages along the clear-cut originated succession gradient. Biodivers Conserv 11:1269–1288
[41]	Koprowski JL, Alanen MI, Lynch AM (2005) Nowhere to run and nowhere to hide: response of endemic Mt. Graham red squirrels to catastrophic forest damage. Biol Conserv 126(4):491–498. https://doi.org/10.1016/j.biocon.2005.06.028
[42]	Kortmann M, Müller JC, Baier R, B?ssler C, Buse J, Cholewińska O, F?rschler MI, Georgiev KB, Hilszczański J, Jaroszewicz B, Jaworski T, Kaufmann S, Kuijper D, Lorz J, Lotz A, ?ubek A, Mayer M, Mayerhofer S, Meyer S, Morinière J, Popa F, Reith H, Roth N, Seibold S, Seidl R, Stengel E, Wolski GJ, Thorn S (2021) Ecology versus society: impacts of bark beetle infestations on biodiversity and restorativeness in protected areas of Central Europe. Biol Conserv. https://doi.org/10.1016/j.biocon.2020.108931
[43]	Kortmann M, Roth N, Buse J, Hilszczański J, Jaworski T, Morinière J, Seidl R, Thorn S, Müller J (2022) Arthropod dark taxa provide new insights into diversity responses to bark beetle infestations. Ecol Appl. https://doi.org/10.1002/eap.2516
[44]	Lacasella F, Gratton C, De Felici S, Isaia M, Zapparoli M, Marta S, Sbordoni V (2015) Asymmetrical responses of forest and “beyond edge” arthropod communities across a forest–grassland ecotone. Biodivers Conserv 24:447–465. https://doi.org/10.1007/s10531-014-0825-0
[45]	Landesamt für Geologie und Bergbau Rheinland-Pfalz (2013a) Kartenviewer - Fachanwendungen und Fachthemen: Boden - BFD 50 - Bodenformengesellschaft. https://mapclient.lgb-rlp.de/?app=lgb&view_id=19. Accessed 28 Jun 2022 (in German)
[46]	Landesamt für Geologie und Bergbau Rheinland-Pfalz (2013b) Kartenviewer - Fachanwendungen und Fachthemen: Geologie: Geologische übersichtskarte 1:300,000. https://mapclient.lgb-rlp.de/?app=lgb&view_id=19. Accessed 28 Jun 2022] (in German)
[47]	Lange M, Türke M, Pa?ali? E, Boch S, Hessenm?ller D, Müller J, Prati D, Socher SA, Fischer M, Weisser WW, Gossner MM (2014) Effects of forest management on ground-dwelling beetles (Coleoptera; Carabidae, Staphylinidae) in Central Europe are mainly mediated by changes in forest structure. For Ecol Manage 329:166–176. https://doi.org/10.1016/j.foreco.2014.06.012
[48]	Leather SR (2005) Insect sampling in forest ecosystems. Blackwell Publishing, Oxford, p 303
[49]	Lehnert LW, B?ssler C, Brandl R, Burton PJ, Müller J (2013) Conservation value of forests attacked by bark beetles: highest number of indicator species is found in early successional stages. J Nat Conserv 21(2):97–104. https://doi.org/10.1016/j.jnc.2012.11.003
[50]	Leverkus AB, Lindenmayer DB, Thorn S, Gustafsson L (2018) Salvage logging in the world’s forests: interactions between natural disturbance and logging need recognition. Global Ecol Biogeogr 27(10):1140–1154. https://doi.org/10.1111/geb.12772
[51]	Lompe A (2002) Die K?fer Europas - ein Bestimmungswerk im Internet. http://coleonet.de/coleo/index.htm. Accessed 12 Jan 2022 (in German)
[52]	López-Bedoya PA, Magura T, Edwards FA, Edwards DP, Rey-Benayas JM, L?vei GL, Noriega JA (2021) What level of native beetle diversity can be supported by forestry plantations? Glob Synth Insect Conserv Divers 14(16):736–747. https://doi.org/10.1111/icad.12518
[53]	LVermGeo (2021) [Landesamt für Vermessung und Geobasisinformation Rheinland-Pfalz]. Luftbilder Rheinland-Pfalz DOP40. In: Open Data - Digitale Orthophotos (DOP), Bodenaufl?sung 40cm. https://www.geoportal.rlp.de/mapbender/php/mod_inspireDownloadFeed.php?id=2b009ae4-aa3e-ff21-870b-49846d9561b2&type=SERVICE&generateFrom=wmslayer&layerid=61676. Accessed 27 Nov 2022] (in German)
[54]	Magura T (2002) Carabids and forest edge: spatial pattern and edge effect. For Ecol Manag 157:23–37
[55]	Magura T, Tóthmérész B, Bordán Z (2000) Effects of nature management practice on carabid assemblages (Coleoptera: Carabidae) in a non-native plantation. Biol Conserv 93:95–102
[56]	Magura T, Tóthmérész B, Elek Z (2006) Changes in carabid beetle assemblages as Norway spruce plantations age. Community Ecol 7(1):1–12. https://doi.org/10.1556/ComEc.7.2006.1.1
[57]	Ministerium für Umwelt Energie Ern?hrung und Forsten Rheinland-Pfalz (2020) WALDZUSTANDS-BERICHT 2020 (in German)
[58]	Ministerium für Umwelt Energie Ern?hrung und Forsten Rheinland-Pfalz (2021) WALDZUSTANDS-BERICHT 2021 (in German)
[59]	Moretti M, Obrist MK, Duelli P (2004) Arthropod biodiversity after forest fires: winners and losers in the winter fire regime of the southern Alps. Ecography 27:173–186
[60]	Morris JL, Cottrell S, Fettig CJ, Hansen WD, Sherriff RL, Carter VA, Clear JL, Clement J, DeRose RJ, Hicke JA, Higuera PE, Mattor KM, Seddon AWR, Seppa HT, Stednick JD, Seybold SJ (2016) Managing bark beetle impacts on ecosystems and society: priority questions to motivate future research. J Appl Ecol 54(3):750–760. https://doi.org/10.1111/1365-2664.12782
[61]	Mullen K, O’Halloran J, Breen J, Giller P, Pithon J, Kelly T (2008) Distribution and composition of carabid beetle (Coleoptera, Carabidae) communities across the plantation forest cycle—implications for management. For Ecol Manage 256(4):624–632. https://doi.org/10.1016/j.foreco.2008.05.005
[62]	Müller J, Bu?ler H, Go?ner M, Rettelbach T, Duelli P (2008) The European spruce bark beetle Ips typographus in a national park: from pest to keystone species. Biodivers Conserv 17(12):2979–3001. https://doi.org/10.1007/s10531-008-9409-1
[63]	Müller J, Noss RF, Thorn S, B?ssler C, Leverkus AB, Lindenmayer D (2019) Increasing disturbance demands new policies to conserve intact forest. Conserv Lett. https://doi.org/10.1111/conl.12449
[64]	Muys B, Angelstam P, Bauhus J, Bouriaud L, Jactel H, Kraigher H, Müller J, Pettorelli N, P?tzelsberger E, Primmer E, Svoboda M, Thorsen BJ, Van Meerbeek K (2022) Forest biodiversity in Europe. From Sci Policy 13:1–80
[65]	Newbold T, Hudson LN, Hill SLL, Contu S, Lysenko I, Senior RA, B?rger L, Bennett DJ, Choimes A, Collen B, Day J, De Palma A, Díaz S, Echeverria-Londo?o S, Edgar MJ, Feldman A, Garon M, Harrison MLK, Alhusseini T, Ingram DJ, Itescu Y, Kattge J, Kemp V, Kirkpatrick L, Kleyer M, Laginha Pinto Correia D, Martin CD, Meiri S, Novosolov M, Pan Y, Phillips HRP, Purves DW, Robinson A, Simpson J, Tuck SL, Weiher E, White HJ, Ewers RM, Mace GM, Scharlemann JPW, Purvis A (2015) Global effects of land use on local terrestrial biodiversity. Nature 520(7545):45–50. https://doi.org/10.1038/nature14324
[66]	Oksanen J, Blanchet FG, Friendly M, Kindt R, Legendre P, McGlinn D, Minchin P, O’Hara R, Simpson G, Solymos P, Stevens M, Szoecs E, Wagner H (2020) Vegan: community ecology package. R-package. https://CRAN.R-project.org/package=vegan
[67]	Overbeck M, Schmidt M (2012) Modelling infestation risk of Norway spruce by Ips typographus (L.) in the Lower Saxon Harz Mountains (Germany). For Ecol Manag 266:115–125. https://doi.org/10.1016/j.foreco.2011.11.011
[68]	Paradis E, Schliep K (2019) ape 5.0: an environment for modern phylogenetics and evolutionary analyses in R. Bioinformatics 35(3):526–528. https://doi.org/10.1093/bioinformatics/bty633
[69]	Pearce JL, Venier LA (2006) The use of ground beetles (Coleoptera: Carabidae) and spiders (Araneae) as bioindicators of sustainable forest management: a review. Ecol Indic 6(4):780–793. https://doi.org/10.1016/j.ecolind.2005.03.005
[70]	Pinheiro J, Bates D (2020) Linear and nonlinear mixed effect models (nlme-package). R-package. https://CRAN.R-project.org/package=nlme
[71]	Prestemon JP, Wear DN, Stewart FJ, Holmes TP (2006) Wildfire, timber salvage, and the economics of expediency. For Policy Econ 8(3):312–322. https://doi.org/10.1016/j.forpol.2004.07.003
[72]	Pukkala T (2018) Effect of species composition on ecosystem services in European boreal forest. J for Res 29(2):261–272. https://doi.org/10.1007/s11676-017-0576-3
[73]	Purchart L, Tuf IH, Hula V, Suchomel J (2013) Arthropod assemblages in Norway spruce monocultures during a forest cycle—a multi-taxa approach. For Ecol Manag 306:42–51. https://doi.org/10.1016/j.foreco.2013.06.012
[74]	QGIS (2002) QGIS Development Team: QGIS Geographic Information System. Version 3.4.7—Madeira. Open Source Geospatial Foundation Project.
[75]	R Core Team (2020) R: a language and environment for statistical computing. R Foundation for statistical computing, Vienna
[76]	Agrarmeteorologie Rheinland-Pfalz (2022a) Langj?hrige Mittelwerte Wetterstation Isert. https://www.dlr.rlp.de/Internet/AM/NotesAM.nsf/cc619a07ed6c4634c1257800003dca56/07804c9e9e2baf08c1257171002e8a40?OpenDocument&TableRow=2.1.1,2.6#2.1. Accessed 1 Feb 2022 (in German)
[77]	Agrarmeteorologie Rheinland-Pfalz (2022b) Langj?hrige Mittelwerte Hydrometeorologische Station Hachenburg. https://www.dlr.rlp.de/Internet/AM/NotesAM.nsf/cc619a07ed6c4634c1257800003dca56/ce8964b69eb243f7c12573ef002f58b1?OpenDocument&TableRow=2.4#2. Accessed 1 Feb 2022 (in German)
[78]	Rothacher J, Hagge J, B?ssler C, Brandl R, Gruppe A, Müller J (2023) Logging operations creating snags, logs, and stumps under open and closed canopies promote stand-scale beetle diversity. For Ecol Manag. https://doi.org/10.1016/j.foreco.2023.121022
[79]	Sánchez-Bayo F, Wyckhuys KAG (2019) Worldwide decline of the entomofauna: a review of its drivers. Biol Conserv 232:8–27
[80]	Schall P, Gossner MM, Heinrichs S, Fischer M, Boch S, Prati D, Jung K, Baumgartner V, Blaser S, B?hm S, Buscot F, Daniel R, Goldmann K, Kaiser K, Kahl T, Lange M, Müller J, Overmann J, Renner SC, Schulze E-D, Sikorski J, Tschapka M, Türke M, Weisser WW, Wemheuer B, Wubet T, Ammer C (2018) The impact of even-aged and uneven-aged forest management on regional biodiversity of multiple taxa in European beech forests. J Appl Ecol 55(1):267–278. https://doi.org/10.1111/1365-2664.12950
[81]	Schall P, Heinrichs S, Ammer C, Ayasse M, Boch S, Buscot F, Fischer M, Goldmann K, Overmann J, Schulze E-D, Sikorski J, Weisser WW, Wubet T, Gossner MM (2021) Among stand heterogeneity is key for biodiversity in managed beech forests but does not question the value of unmanaged forests: response to Bruun and Heilmann-Clausen (2021). J Appl Ecol 58(9):1817–1826. https://doi.org/10.1111/1365-2664.13959
[82]	Schmidl J (2005) Entomologiebedarf-Eklektoren / FIT’s Rahn/Mini. In: Bioform-entomology & equipment. https://www.bioform.de/shop.php?wg=1&action=wgstart. Accessed 12 Apr 2023 (in German)
[83]	Seibold S, Brandl R, Buse J, Hothorn T, Schmidl J, Thorn S, Müller J (2015) Association of extinction risk of saproxylic beetles with ecological degradation of forests in Europe. Conserv Biol 29(2):382–390. https://doi.org/10.1111/cobi.12427
[84]	Seibold S, B?ssler C, Baldrian P, Reinhard L, Thorn S, Ulyshen MD, Wei? I, Müller J (2016a) Dead-wood addition promotes non-saproxylic epigeal arthropods but effects are mediated by canopy openness. Biol Conserv 204:181–188. https://doi.org/10.1016/j.biocon.2016.09.031
[85]	Seibold S, B?ssler C, Brandl R, Büche B, Szallies A, Thorn S, Ulyshen MD, Müller J (2016b) Microclimate and habitat heterogeneity as the major drivers of beetle diversity in dead wood. J Appl Ecol 53(3):934–943. https://doi.org/10.1111/1365-2664.12607
[86]	Seibold S, Gossner MM, Simons NK, Blüthgen N, Müller J, Ambarli D, Ammer C, Bauhus J, Fischer M, Habel JC, Linsenmair KE, Nauss T, Penone C, Prati D, Schall P, Schulze E-D, Vogt J, W?llauer S, Weisser WW (2019) Arthropod decline in grasslands and forests is associated with landscape-level drivers. Nature 574(7780):671–674. https://doi.org/10.1038/s41586-019-1684-3
[87]	Seibold S (2015) Biodiversity of dead-wood dependent insects–effects of forest management and prospects of conservation. Dissertation, Wissenschaftszentrum Weihenstephan für Ern?hrung, Landnutzung und Umwelt, Technical University of Munich, 1–55
[88]	Seidl R, Schelhaas MJ, Rammer W, Verkerk PJ (2014) Increasing forest disturbances in Europe and their impact on carbon storage. Nat Clim Chang 4(9):806–810. https://doi.org/10.1038/nclimate2318
[89]	Seidl R, Thom D, Kautz M, Martin-Benito D, Peltoniemi M, Vacchiano G, Wild J, Ascoli D, Petr M, Honkaniemi J, Lexer MJ, Trotsiuk V, Mairota P, Svoboda M, Fabrika M, Nagel TA, Reyer CPO (2017) Forest disturbances under climate change. Nat Clim Chang 7(6):395–402. https://doi.org/10.1038/nclimate3303
[90]	Senf C, Mori AS, Müller J, Seidl R (2020) The response of canopy height diversity to natural disturbances in two temperate forest landscapes. Landsc Ecol 35(9):2101–2112. https://doi.org/10.1007/s10980-020-01085-7
[91]	Sessions J, Bettinger P, Buckman R, Newton M, Hamann J (2004) Hastening the return of complex forests following fire: the consequences of delay. J for 102(3):38–45
[92]	Sommerfeld A, Rammer W, Heurich M, Hilmers T, Müller J, Seidl R (2021) Do bark beetle outbreaks amplify or dampen future bark beetle disturbances in Central Europe? J Ecol 109(2):737–749. https://doi.org/10.1111/1365-2745.13502
[93]	Statistisches Landesamt Rheinland-Pfalz (2020) Fl?chennutzung Verbandsgemeinde Hachenburg. http://infothek.statistik.rlp.de/MeineHeimat/content.aspx?id=103&l=2&g=0714302&tp=1025. Accessed 1 Feb 2022 (in German)
[94]	Stephens SS, Wagner MR (2007) Forest plantations and biodiversity: a fresh perspective. J for 105(6):307–313. https://doi.org/10.1093/jof/105.6.307
[95]	Swanson ME, Franklin JF, Beschta RL, Crisafulli CM, DellaSala DA, Hutto RL, Lindenmayer DB, Swanson FJ (2011) The forgotten stage of forest succession: early-successional ecosystems on forest sites. Front Ecol Environ 9(2):117–125. https://doi.org/10.1890/090157
[96]	Thom D, Seidl R (2016) Natural disturbance impacts on ecosystem services and biodiversity in temperate and boreal forests. Biol Rev 91(3):760–781. https://doi.org/10.1111/brv.12193
[97]	Thom D, Rammer W, Seidl R (2017) The impact of future forest dynamics on climate: interactive effects of changing vegetation and disturbance regimes. Ecol Monogr 87(4):665–684. https://doi.org/10.1002/ecm.1272
[98]	Thorn S, B?ssler C, Gottschalk T, Hothorn T, Bussler H, Raffa K, Müller J (2014) New insights into the consequences of post-windthrow salvage logging revealed by functional structure of saproxylic beetles assemblages. PLoS ONE. https://doi.org/10.1371/journal.pone.0101757
[99]	Thorn S, B?ssler C, Bernhardt-R?mermann M, Cadotte M, Heibl C, Sch?fer H, Seibold S, Müller J (2015) Changes in the dominant assembly mechanism drive species loss caused by declining resources. Ecol Lett 19(2):163–170. https://doi.org/10.1111/ele.12548
[100]	Thorn S, B?ssler C, Svoboda M, Müller J (2016) Effects of natural disturbances and salvage logging on biodiversity—lessons from the Bohemian Forest. For Ecol Manag 388:113–119. https://doi.org/10.1016/j.foreco.2016.06.006
[101]	Thorn S, B?ssler C, Brandl R, Burton PJ, Cahall R, Campbell JL, Castro J, Choi C-Y, Cobb T, Donato DC, Durska E, Fontaine JB, Gauthier S, Hebert C, Hothorn T, Hutto RL, Lee E-J, Leverkus AB, Lindenmayer DB, Obrist MK, Rost J, Seibold S, Seidl R, Thom D, Waldron K, Wermelinger B, Winter M-B, Zmihorski M, Müller J (2018) Impacts of salvage logging on biodiversity: a meta-analysis. J Appl Ecol 55(1):279–289. https://doi.org/10.1111/1365-2664.12945
[102]	Thorn S, Seibold S, Leverkus AB, Michler T, Müller J, Noss RF, Stork N, Vogel S, Lindenmayer DB (2020) The living dead: acknowledging life after tree death to stop forest degradation. Front Ecol Environ 18(9):505–512. https://doi.org/10.1002/fee.2252
[103]	Thorn S (2016) Effects of salvage logging on biodiversity—ecological evidence for conservation-oriented management of natural disturbances. Dissertation, Wissenschaftszentrum Weihenstephan für Ern?hrung, Landnutzung und Umwelt, Technical University of Munich, 1–62
[104]	Uhl B, Krah FS, Baldrian P, Brandl R, Hagge J, Müller J, Thorn S, Vojtech T, B?ssler C (2022) Snags, logs, stumps, and microclimate as tools optimizing deadwood enrichment for forest biodiversity. Biol Conserv. https://doi.org/10.1016/j.biocon.2022.109569
[105]	Uhl B, Pouska V, ?ervenka J, Karasch P, B?ssler C (2023) Response of moth communities (Lepidoptera) to forest management strategies after disturbance. Eur J Entomol 120:35–41. https://doi.org/10.14411/eje.2023.005
[106]	Vele A, Holusa J, Frouz J, Konvicka O (2011) Local and landscape drivers of ant and carabid beetle communities during spruce forest succession. Eur J Soil Biol 47(6):349–356. https://doi.org/10.1016/j.ejsobi.2011.09.003
[107]	Vogel S, Gossner MM, Mergner U, Müller J, Thorn S (2020) Optimizing enrichment of deadwood for biodiversity by varying sun exposure and tree species: an experimental approach. J Appl Ecol 57(10):2075–2085. https://doi.org/10.1111/1365-2664.13648
[108]	Wermelinger B, Flückiger PF, Obrist MK, Duelli P (2007) Horizontal and vertical distribution of saproxylic beetles (Col., Buprestidae, Cerambycidae, Scolytinae) across sections of forest edges. J Appl Entomol 131(2):104–114. https://doi.org/10.1111/j.1439-0418.2006.01128.x
[109]	White SP, Pickett ST (1985) The ecology of natural disturbance and patch dynamics. Academic Press, New York
[110]	Winter MB, Ammer C, Baier R, Donato DC, Seibold S, Müller J (2015) Multi-taxon alpha diversity following bark beetle disturbance: evaluating multi-decade persistence of a diverse early-seral phase. For Ecol Manag 338:32–45. https://doi.org/10.1016/j.foreco.2014.11.019
[111]	Zumr V, Reme? J, Pulkrab K (2021) How to increase biodiversity of saproxylic beetles in commercial stands through integrated forest management in Central Europe. Forests. https://doi.org/10.3390/f12060814

Funding

Universit?t Koblenz (6377)