Conservation value of forests attacked by bark beetles: Highest number of indicator species is found in early successional stages

Heavy natural disturbance in large protected areas of former commercial forests increasingly evokes European parliaments to call for management intervention because a loss of habitats and species is feared. In contrast, natural early successional habitats have recently been recognised as important for conservation. Current knowledge in this field mostly results from studies dealing only with selected taxa. Here we analyse the success of species across 24 lineages of three kingdoms in the Bavarian Forest National Park (Germany) after 15 years of a European spruce bark beetle (Ips typographus L.) outbreak that led to rapid canopy opening. Using indicator species analysis, we found 257 species with a significant preference for open forests and 149 species with a preference for closed forests, but only 82 species with a preference for the stand conditions transitional between open and closed forests. The large number of species with a preference for open forests across lineages supports the role of this bark beetle as a keystone species for a broad array of species. The slowdown of the outbreak after 15 years in the core zone of the national park resulted in less than half of the area being affected, due to variability in stand ages and tree species mixtures. Our case study is representative of the tree species composition and size of many large protected montane areas in Central European countries and illustrates that (1) natural disturbances increase biodiversity in formerly managed forests and (2) a montane protected area spanning 10,000 ha of low range mountains is likely sufficient to allow natural disturbances without a biased loss of closed-forest species.