Detailed research has documented gradual changes to biological communities attributed to increases in global average temperatures. However, localized and abrupt temperature anomalies associated with heatwaves may cause more rapid biological changes. We analyzed temperature data from the South Island of New Zealand and investigated whether the hot summer of 2017/18 affected species of bull kelp, Durvillaea antarctica, D. poha, and D. willana. Durvillaea spp. are large iconic seaweeds that inhabit the low intertidal zone of exposed coastlines, where they underpin biodiversity and ecosystem functioning. Sea surface temperatures (SST) during the summer of 2017/18 included the strongest marine heatwaves recorded in 38 years of existing oceanic satellite data for this region. Air temperatures were also high, and, coupled with small wave heights, resulted in strong desiccation stress during daytime low tides. Before-After analysis of drone images of four reef platforms (42, 42, 44, and 45 • S) was used to evaluate changes to bull kelp over the hot summer. Bull kelp loss varied among species and reefs, with the greatest (100%) loss of D. poha at Pile Bay in Lyttelton Harbor (44 • S). In Pile Bay, SST exceeded 23 • C and air temperatures exceeded 30 • C, while Durvillaea was exposed for up to 3 h per day during low tide. Follow-up surveys showed that all bull kelps were eliminated from Pile Bay, and from all reefs within and immediately outside of Lyttelton Harbor. Following the localized extinction of bull kelp in Pile Bay, the invasive kelp Undaria pinnatifida recruited in high densities (average of 120 m −2 ). We conclude that bull kelps are likely to experience additional mortalities in the future because heatwaves are predicted to increase in magnitude and durations. Losses of the endemic D. poha are particularly concerning due to its narrow distributional range.

It has long been recognized that primary foundation species (FS), such as trees and seagrasses, enhance biodiversity. Among the species facilitated are secondary FS, including mistletoes and epiphytes. Case studies have demonstrated that secondary FS can further modify habitat-associated organisms ('inhabitants'), but their net effects remain unknown. Here we assess how inhabitants, globally, are affected by secondary FS. We extracted and calculated 2,187 abundance and 397 richness Hedges' g effect sizes from 91 and 50 publications, respectively. A weighted meta-analysis revealed that secondary FS significantly enhanced the abundance and richness of inhabitants compared to the primary FS alone. This indirect facilitation arising through sequential habitat formation was consistent across environmental and experimental conditions. Complementary unweighted analyses on log response ratios revealed that the magnitude of these effects was similar to the global average strength of direct facilitation from primary foundation species and greater than the average strength of trophic cascades, a widely recognized type of indirect facilitation arising through sequential consumption. The finding that secondary FS enhance the abundance and richness of inhabitants has important implications for understanding the mechanisms that regulate biodiversity. Integrating secondary FS into conservation practice will improve our ability to protect biodiversity and ecosystem function.

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