Facilitation of animals is stronger during summer marine heatwaves and around morphologically complex foundation species.

Foundation species create biogenic habitats, modify environmental conditions, augment biodiversity, and control animal community structures. In recent decades, marine heatwaves (MHWs) have affected the ecology of foundation species worldwide, and perhaps also their associated animal communities. However, no realistic field experiment has tested how MHWs affect animals that live in and around these foundation species. We therefore tested, in a four-factorial field experiment, if colonisation by small mobile marine animals (epifauna) onto plates with attached single versus co-occurring foundation species of different morphological complexities, were affected by 3-5°C heating (that mirrored a recent extreme MHW in the study area) and if the heating effect on the epifauna varied within and between seasons. For this experiment mimics of turf seaweed represented the single foundation species and holdfasts of seven common canopy-forming seaweed represented the co-occurring foundation species with different morphological complexities. We found that the taxonomic richness and total abundance of epifauna, dominated by copepods, generally were higher on heated plates with complex seaweed holdfasts in warmer summer trials. Furthermore, several interactions between test-factors were significant, e.g., epifaunal abundances, were, across taxonomic groups, generally higher in warmer than colder summer trials. These results suggest that, in temperate ecosystems, small, mobile, short-lived, and fast-growing marine epifauna can be facilitated by warmer oceans and morphologically complex foundation species, implying that future MHWs may increase secondary production and trophic transfers between primary producers and fish. Future studies should test whether these results can be scaled to other ecological species-interactions, across latitudes and biogeographical regions, and if similar results are found after longer MHWs or within live foundation species under real MHW conditions.

Long-term community shifts driven by local extinction of an iconic foundation species following an extreme marine heatwave.

Gradual ocean warming combined with stronger marine heatwaves (MHWs) can reduce abundances of foundation species that control community structures, biodiversity, and ecosystem functioning. However, few studies have documented long-term succession trajectories following the more extreme events that cause localized extinctions of foundation species. Here, we documented long-term successional changes to marine benthic communities in Pile Bay, New Zealand, following the Tasman 2017/18 MHW, which caused localized extinctions of dominant southern bull kelp (Durvillaea sp.). Six years on, multiscale annual and seasonal surveys show no sign of Durvillaea recolonization. Instead, the invasive annual kelp (Undaria pinnatifida), rapidly colonized areas previously dominated by Durvillaea, followed by large changes to the understory community, as Durvillaea holdfasts and encrusting coralline algae were replaced by coralline turf. Between 3 and 6 years after the total loss of Durvillaea, smaller native fucoids colonized in high densities. Although Undaria initially colonized plots throughout Durvillaea's tidal range, later in the succession Undaria only retained dominance in the lower intertidal zone and only in spring. Ultimately, the tidal zone was slowly replaced by alternative foundation species, composed of different canopy-forming brown seaweeds that dominated different intertidal elevations, resulting in a net increase in canopy and understory diversity. This study is a rare example of long-term effects following an extreme MHW that caused extinctions of a locally dominant canopy-former, but these events and their associated dramatic changes to community structures and biodiversity are expected to become increasingly common as MHWs continue to increase in strength, frequency, and duration.

Spatiotemporal stressors, not secondary structures or small temperature increases, control rapid facilitation of intertidal epifauna.

Small animals (epifauna) are ubiquitous in marine systems. Epifauna have high secondary production and provide trophic linkages between primary producers and higher-order consumers, like fish. Despite their importance, little is known about how these animals respond to warming or how their communities vary across spatiotemporal gradients. Here we use mimics of turf seaweed and invasive kelp holdfast to test, in a 5-factorial field experiment, whether intertidal epifauna are facilitated by different habitat structures, temperature conditions, and along cooccurring spatiotemporal gradients. We found that facilitation of epifauna by intertidal turf seaweed peaked in summer, at low elevation, in older habitats and at a less wave-exposed site. However, epifauna were not affected by the presence of a secondary structure like kelp holdfast mimics or small temperature increases from passive solar heating of black and white mimics. There were many significant two-way, but few higher order interactions, showing stronger facilitation under specific environmental conditions, like at low elevation in summer, or low elevation in old habitats. These results highlight that turf-associated epifauna are controlled by vertical elevation, season, hydrodynamics, and habitat age, and appear to be resilient to small temperature increases. Findings are important to better understand linkages between primary producers and higher order consumers and system-wide productivity, and because fast growing turf, facilitated by global warming and eutrophication, are increasingly outcompeting slower growing large perennial canopy forming seaweeds, like kelp and rockweeds.

Unravelling seasonal trends in coastal marine heatwave metrics across global biogeographical realms.

Marine heatwaves (MHWs) can cause dramatic changes to ecologically, culturally, and economically important coastal ecosystems. To date, MHW studies have focused on geographically isolated regions or broad-scale global oceanic analyses, without considering coastal biogeographical regions and seasons. However, to understand impacts from MHWs on diverse coastal communities, a combined biogeographical-seasonal approach is necessary, because (1) bioregions reflect community-wide temperature tolerances and (2) summer or winter heatwaves likely affect communities differently. We therefore carried out season-specific Theil-Sen robust linear regressions and Pettitt change point analyses from 1982 to 2021 on the number of events, number of MHW days, mean intensity, maximum intensity, and cumulative intensity of MHWs, for each of the world's 12 major coastal biogeographical realms. We found that 70% of 240 trend analyses increased significantly, 5% decreased and 25% were unaffected. There were clear differences between trends in metrics within biogeographical regions, and among seasons. For the significant increases, most change points occurred between 1998 and 2006. Regression slopes were generally positive across MHW metrics, seasons, and biogeographical realms as well as being highest after change point detection. Trends were highest for the Arctic, Northern Pacific, and Northern Atlantic realms in summer, and lowest for the Southern Ocean and several equatorial realms in other seasons. Our analysis highlights that future case studies should incorporate break point changes and seasonality in MHW analysis, to increase our understanding of how future, more frequent, and stronger MHWs will affect coastal ecosystems.