Communities and attachment networks associated with primary, secondary and alternative foundation species; A case study of stressed and disturbed stands of southern bull kelp

Abstract

Southern bull kelps (Durvillaea spp., Fucales) are ‘primary’ foundation species that control community structures and ecosystem functions on temperate wave-exposed rocky reefs. However, these large foundation species are threatened by disturbances and stressors, including invasive species, sedimentation and heatwaves. It is unknown whether ‘alternative’ foundation species can replace lost southern bull kelps and its associated communities and networks. We compared community structure (by quantifying abundances of different species) and attachment-interaction networks (by quantifying which species were attached to other species) among plots dominated by Durvillaea spp. and plots where Durvillaea spp. were lost either through long-term repeated experimental removals or by recent stress from a marine heatwave. Long-term experimental removal plots were dominated by ‘alternative’ foundation species, the canopy-forming Cystophora spp. (Fucales), whereas the recent heatwave stressed plots were dominated by the invasive kelp Undaria pinnatifida (Laminariales). A network analysis of attachment interactions showed that communities differed among plots dominated by either Durvillaea spp., Cystophora spp. or U. pinnatifida, with different relationships between the primary, or alternative, foundation species and attached epiphytic ‘secondary’ foundation species. For example, native Cystophora spp. were more important as hosts for secondary foundation species compared to Durvillaea spp. and U. pinnatifida. Instead, Durvillaea spp. facilitated encrusting algae, which in turn provided habitat for gastropods. We conclude that (a) repeated disturbances and strong stressors can reveal ecological differences between primary and alternative foundation species, (b) analyses of abundances and attachment-networks are supplementary methods to identify linkages between primary, alternative and secondary foundation species, and (c) interspersed habitats dominated by different types of foundation species increase system-level biodiversity by supporting different species-abundance patterns and species-attachment networks.