Lower alpha, higher beta, and similar gamma diversity of saproxylic beetles in unmanaged compared to managed Norway spruce stands

Strong anthropogenic pressures on global forests necessitate that managed forests be evaluated as habitat for biodiversity. The complex pattern of habitat types created in forestry systems is ideal for analyses through the theoretical framework of alpha (local), gamma (total) and beta (compositional) diversity. Here I use saproxylic beetles, a species-rich threatened group, to compare four Norway spruce-dominated habitats representative of the boreal forest landscape of northern Europe: unmanaged semi-natural stands, nature reserves, unthinned middle-aged production stands and commercially thinned production stands. The beetles (in total 38 085 individuals of 312 species), including red-listed ones and three feeding guilds (wood consumers, fungivores and predators) were studied in 53 stands in central-southern Sweden, in two regions with differing amounts of conservation forest. Alpha diversity of saproxylic, but not red-listed, beetles was higher in the thinned stands than in the semi-natural stands, and did not differ for the other forest types. Beta diversity of saproxylic beetles was higher in unmanaged semi-natural stands than in the other forest types, but species composition did not differ noticeably. Furthermore, red-listed saproxylic beetles had higher gamma diversity in unmanaged semi-natural stands in the region with more conservation forest, but not in the one with less such forest. The local factors dead wood volume and dead wood diversity did not influence alpha diversity of beetles, but increasing canopy openness had a minor negative influence on saproxylic and red-listed beetles. While the local scale (alpha diversity) indicates the potential for managed forests to house many saproxylic beetle species associated with spruce forests in this boreal landscape, the larger scales (beta and gamma diversity) indicate the value of unmanaged forests for the conservation of the entire saproxylic beetle fauna. These results show the importance of analyses at multiple levels of diversity (alpha, beta, gamma) for identifying patterns relevant to conservation.