Forest biodiversity and ecosystem services from spruce-birch mixtures: The potential importance of tree spatial arrangement

Abstract There is increasing empirical support for the biodiversity and ecosystem service (ES) benefits of mixed-species production forests. However, few studies control for the spatial arrangement of the trees within mixtures to determine the influence that clustering the tree species (patch scale mixtures), versus evenly dispersing them (intimate scale mixtures), may have for biodiversity and ES outcomes. To highlight the potential implications of altering tree spatial arrangement in mixtures, and the need to fill related knowledge gaps, here we provide a qualitative multi-disciplinary overview of ecological and socio-economic drivers with the potential to alter biodiversity, ecosystem services, and management-related outcomes from patch versus intimate scale mixtures. We focused our overview on even-aged mixtures of Norway spruce (Picea abies) and birch (Betula pendula or B. pubescens) in Sweden, which enabled us to contrast findings within a biogeographical and silvicultural setting. Specifically, we targeted implications for biodiversity (understory vascular plants, epiphytic lichens, saproxylic beetles, birds), biomass production, harvesting costs, management ease, recreation and aesthetics, cervid game, as well as abiotic and biotic risks (wind, fire, pathogens, pests, browsing damage). In the absence of direct empirical evidence, we primarily relied on expert inference from theory and relevant empirical studies sourced from the Fennoscandian region, and further afield if needed. Collectively these efforts allowed us to develop a number of informed hypotheses indicating that for spruce-birch mixtures in this region, patch scale mixtures may have the potential to favour the diversity of several forest dependent taxonomic groups, cervid game and reduce harvesting costs, whereas intimate mixtures may have the potential to reduce pathogen and pest damage, and likewise, potentially benefit production outcomes. Current knowledge was too limited, inconsistent or context dependent to even tentatively infer outcomes for fire risk, wind damage, browsing damage, management ease, recreational and aesthetic outcomes. We emphasize that our hypotheses require testing, but are sufficient to i) highlight the likely importance of spatial-scale to biodiversity and ecosystem services outcomes in mixed-species production forests, ii) caution against generalization from mixture studies that lack scale considerations, and iii) motivate the targeted consideration of spatial grain in future mixture studies.