Global kelp forest restoration: past lessons, present status, and future directions.

Kelp forest ecosystems and their associated ecosystem services are declining around the world. In response, marine managers are working to restore and counteract these declines. Kelp restoration first started in the 1700s in Japan and since then has spread across the globe. Restoration efforts, however, have been largely disconnected, with varying methodologies trialled by different actors in different countries. Moreover, a small subset of these efforts are 'afforestation', which focuses on creating new kelp habitat, as opposed to restoring kelp where it previously existed. To distil lessons learned over the last 300 years of kelp restoration, we review the history of kelp restoration (including afforestation) around the world and synthesise the results of 259 documented restoration attempts spanning from 1957 to 2020, across 16 countries, five languages, and multiple user groups. Our results show that kelp restoration projects have increased in frequency, have employed 10 different methodologies and targeted 17 different kelp genera. Of these projects, the majority have been led by academics (62%), have been conducted at sizes of less than 1 ha (80%) and took place over time spans of less than 2 years. We show that projects are most successful when they are located near existing kelp forests. Further, disturbance events such as sea-urchin grazing are identified as regular causes of project failure. Costs for restoration are historically high, averaging hundreds of thousands of dollars per hectare, therefore we explore avenues to reduce these costs and suggest financial and legal pathways for scaling up future restoration efforts. One key suggestion is the creation of a living database which serves as a platform for recording restoration projects, showcasing and/or re-analysing existing data, and providing updated information. Our work establishes the groundwork to provide adaptive and relevant recommendations on best practices for kelp restoration projects today and into the future.

Density-dependence and seasonal variation in reproductive output and sporophyte production in the kelp, Ecklonia radiata.

The kelp, Ecklonia radiata, is an abundant subtidal ecosystem engineer in southern Australia. Density-dependent changes in the abiotic environment engineered by Ecklonia may feedback to affect reproduction and subsequent recruitment. Here, we examined: 1) how the reproductive capacity of Ecklonia individuals in the field (zoospores released · mm-2 reproductive tissue) varied with adult density and time, and 2) how the recruitment of microscopic gametophytes and sporophytes was influenced by zoospore density at two times. Zoospore production did not vary with adult density, with only one month out of ten sampled over a 2-y period showing a significant effect of density. However, zoospore production varied hugely over time, being generally highest in mid-autumn and lowest in mid-late summer. There were strong effects of initial zoospore density on gametophyte and sporophyte recruitment with both a minimum and an optimum zoospore density for sporophyte recruitment, but these varied in time. Almost no sporophytes developed when initial zoospore density was <6.5 · mm-2 in spring or <0.5 · mm-2 in winter with optimum densities of 90-355 · mm-2 in spring and 21-261 · mm-2 in winter, which resulted in relatively high recruitment of 4-7 sporophytes · mm-2 . Sporophyte recruitment declined at initial zoospore densities >335 · mm-2 in spring and >261 · mm-2 in winter and was zero at very high zoospore densities. These findings suggest that although adult Ecklonia density does not affect per-capita zoospore production, because there is a minimum zoospore density for sporophyte production, a decline in population-level output could feedback to impact recruitment.

Interactive effects of canopy-driven changes in light, scour and water flow on microscopic recruits in kelp.

Ecosystem engineering kelp forms habitat and influences associated communities by altering abiotic conditions. These conditions can also affect the engineer's own demographic rates but the mechanisms underpinning these feedbacks are not well known. Here, we tested the interactive effects of three abiotic factors engineered by the Australasian kelp Ecklonia radiata (light, water flow and scour) on the early survivorship and growth of its outplanted microscopic recruits. After six weeks, recruit survivorship was high in the absence of scour and low light (2-3 times higher than when scour was present) and under low water flow-ambient light conditions. Growth of sporophytes was strongly related to light, with recruits under ambient light approximately four times larger after six weeks. Overall, reduced scour (for survivorship) and ambient light (for growth) appear crucial for maximising E. radiata recruitment suggesting a healthy forest can provide microenvironments to enhance survivorship while gaps in the canopy enhance growth.

High kelp density attracts fishes except for recruiting cryptobenthic species.

As foundation species, kelp support productive and species rich communities; however, the effects of kelp structure on mobile species within these complex natural systems are often difficult to assess. We used artificial reefs with transplanted kelp to quantify the influence of kelp patch size and density on fish assemblages including the arrival of recruiting cryptobenthic species. Large patches with dense kelp supported the highest abundance, species richness, and diversity of fishes, with the addition of dense kelp tripling biomass and doubling richness. The abundance of recruits in artificial collectors declined with patch size and was halved on reefs with sparse kelp compared to reefs with dense kelp or no kelp. These results highlight the importance of dense kelp cover in facilitating biodiversity and indicate that kelp addition could support the recovery of degraded coastal ecosystems. Kelp also apparently drives complex interactions affecting the recruitment/behaviour of some cryptobenthic species.

Resilience and stability of kelp forests: The importance of patch dynamics and environment-engineer feedbacks.

Habitat forming 'ecosystem engineers' such as kelp species create complex habitats that support biodiverse and productive communities. Studies of the resilience and stability of ecosystem engineers have typically focussed on the role of external factors such as disturbance. However, their population dynamics are also likely to be influenced by internal processes, such that the environmental modifications caused by engineer species feedback to affect their own demography (e.g. recruitment, survivorship). In numerous regions globally, kelp forests are declining and experiencing reductions in patch size and kelp density. To explore how resilience and stability of kelp habitats is influenced by this habitat degradation, we created an array of patch reefs of various sizes and supporting adult Ecklonia radiata kelp transplanted at different densities. This enabled testing of how sub-canopy abiotic conditions change with reductions in patch size and adult kelp density, and how this influenced demographic processes of microscopic and macroscopic juvenile kelp. We found that ecosystem engineering by adult E. radiata modified the environment to reduce sub-canopy water flow, sedimentation, and irradiance. However, the capacity of adult kelp canopy to engineer abiotic change was dependent on patch size, and to a lesser extent, kelp density. Reductions in patch size and kelp density also impaired the recruitment, growth and survivorship of microscopic and macroscopic juvenile E. radiata, and even after the provisioning of established juveniles, demographic processes were impaired in the absence of sufficient adult kelp. These results are consistent with the hypothesis that ecosystem engineering by adult E. radiata facilitates development of juvenile conspecifics. Habitat degradation seems to impair the ability of E. radiata to engineer abiotic change, causing breakdown of positive intraspecific feedback and collapse of demographic functions, and overall, leading to reductions in ecosystem stability and resilience well before local extirpation.