Urchin Barrens Research Articles

Successful large-scale and long-term kelp forest restoration by culling sea urchins with quicklime and supported by crab predation

In 2013, a large scale (70 hectare) kelp forest restoration experiment was conducted in a sea urchin barren that had been stable for 45 years. We used 200 tons of quicklime to eradicate the sea urchin population. Kelp recovered within a year in the lime-treated sites. Quickliming did not significantly affect the abundance of non-target species. Mobile kelp fauna colonized the restored kelps, but at a slower rate than kelp and other macroalgae, probably due to slower dispersal abilities. The monitoring period (2012–2021) encompassed the expansion of invasive red king crabs, which emerged as a novel predator of sea urchins. This led to a gradual decline in sea urchin populations and facilitated kelp recovery also in the control sites. This delayed recovery does not invalid the conclusions of the short-term efficiency of the quicklime treatment, but it makes it difficult to conclude about the long-term effects. The study shows that quickliming can initiate kelp recovery within large urchin barrens. We postulate that the recovery of kelps outside the limed areas was due to crab predation, and that successful kelp restoration in areas with recruitment of sea urchins depends on a persistent top-down control of the urchins.

Abalone recruitment patterns before and after sea urchin barrens formation in northern California: incorporating climate change

ABSTRACT Understanding the recruitment dynamics of invertebrates in kelp forests is critical to informing climate-ready restoration. Here we examine abalone and sea urchin recruitment (3–20 mm in size) patterns in northern California across a period of drastic change. Annual surveys were conducted before, during and after the MHW (2014–2016), the loss of a major predatory sea star (2012–2016) and the collapse of a bull kelp forest in 2014. Divers surveyed artificial reef recruitment modules (n = 12) over 20 years in an area that once supported dense bull kelp, Nereocystis leutkeana, forests and the world's largest recreational abalone fishery. From 2016 to 2022, we tracked the decline of red abalone, Haliotis rufescens, recruitment and the rise of purple sea urchin, Strongylocentrotus purpuratus, recruitment. Adult densities of purple sea urchins increased as did newly settled sea urchins (<3 mm), while adult and newly settled red abalone declined. Eight years after the kelp forest collapse, red abalone recruitment remained low and sea urchin recruitment continued to increase. Recruitment patterns can inform both abalone restoration targets and sea urchin dynamics as part of a more holistic kelp forest recovery plan that is responsive to climate change drivers.

Phylogeny and genetic diversity of Echinometra sea urchin in Taiwan

ABSTRACT Four closely related Echinometra species exist in the Indo-Pacific region, but little is known about their population genetics in Taiwan, an area rich in marine biodiversity. Echinometra dominates the Douziping coral reef off Liuqiu Island, where urchin barrens are also present. To understand the relationship between Echinometra species and urchin barrens, we investigated the genetic diversity of Echinometra species in Taiwan by using a nondestructive DNA extraction method on the spines of sea urchins. Mitochondrial DNA results showed that E. sp. C is the dominant species in Taiwan, with relatively high interspecific genetic distance. The E. sp. C is the only species in Douziping, and its population is expanding in accordance with the significantly low R2 value, the significantly negative Fu’s Fs values, and mismatch distribution analysis results. Gene flow estimates between the urchin populations in Taiwan and those in the neighbouring waters of Okinawa suggested that the Kuroshio Current could play an important role in Echinometra dispersal. This implies strongly connectivity between these populations. This study reports the first case of E. sp. C-associated urchin barrens in the tropics.

The sunflower sea star reduces grazing rates of purple sea urchins dependent upon urchin starvation state

AbstractEcosystem function is maintained in part by direct species interactions, but indirect interactions and non‐consumptive effects may be of equal ecological importance. Along the west coast of North America, the recent population collapse of the predatory sunflower sea star Pycnopodia helianthoides has been implicated in the proliferation of the purple sea urchin Strongylocentrotus purpuratus, and a concurrent decline in kelp canopy cover in several locales. Recent work began to quantify the predation rates effects (i.e., direct consumptive effects) of Pycnopodia on sea urchins that may lead to density‐mediated indirect effects on kelp. However, the importance of non‐consumptive effects on urchin behavior and the possible trait‐mediated indirect effects of Pycnopodia on kelp are not well understood. This leaves a critical gap in our knowledge about how these predators may be controlling grazer populations and, indirectly, primary production by macroalgae in nearshore habitats. We measured the non‐consumptive behavioral effects of Pycnopodia on S. purpuratus in the laboratory including grazing rates, feeding behavior, and movement of starved versus fed urchins, the latter simulating urchin metabolic conditions within urchin barrens. We found that the presence of a waterborne Pycnopodia cue reduced the grazing rate of fed urchins by 50% over short (~24 h) time scales. In contrast, starved urchins consumed kelp and did not exhibit an escape response in the presence of a Pycnopodia cue. This study highlights a trait‐mediated indirect interaction between Pycnopodia, S. purpuratus, and kelp, showing how the urchin response to a predator cue may differ based on urchin metabolic conditions or ecosystem state, and helps clarify the positive role of Pycnopodia on kelp forest health.

Effects of Rock Type and Food Availability on Bioerosion by the Purple Sea Urchin, Strongylocentrotus purpuratus.

Purple sea urchins (Strongylocentrotus purpuratus) profoundly impact nearshore rocky coasts through their feeding habits. Their intense grazing sculpts substrates through bioerosion using their teeth and spines and controls the alternative stable state dynamic between kelp bed and urchin barrens. These states have contrasting food availability for sea urchins, with abundant food in kelp beds and scarce food in barren grounds. However, the relationship between food availability and bioerosion is unknown. We predicted that when kelp is available, it would ameliorate the action of teeth on the substrate. Our 11-weeks long, 2×2 factorial experiment, crossed community state (kelp present vs absent) and rock type (sandstone vs mudstone). We also quantified the contribution of spine abrasion to bioerosion on the two rock types. The bioerosion rates did not differ between treatments with and without kelp. Although there was no significant difference in net bioerosion between the rock types, there was a large difference between the proportion of bioerosion from teeth vs spine abrasion. Approximately a third of the sandstone bioerosion was from spines whereas less than 2% of mudstone bioerosion could be attributed to spines. As anticipated, growth of sea urchins fed kelp ad-libitum was higher than food-limited sea urchins. Surprisingly, sea urchins on mudstone (which has a higher organic component) grew faster than sea urchins on sandstone. Although bioerosion rates may not differ on a per-urchin basis between community states, the sea urchin population densities between kelp beds and urchin barrens likely causes a difference in net bioerosion between these communities. Our results point to the importance of lithology on the mechanics of sea urchin bioerosion. Differences in texture, grain size, and hardness of rock substrates undoubtedly contribute to bioerosion rates and dynamics.

Sea urchin roe quality within urchin barrens and improvement through kelp restoration

AbstractSea urchin overgrazing is a major cause of kelp loss and consequently sea urchin removal is frequently promoted as a tool for restoring kelp in urchin barrens. While many sea urchin species support high‐end roe fisheries, roe condition is generally lower in barrens than in kelp forests due to reduced food availability. Understanding how roe condition varies between kelp and barren habitats, and in response to kelp restoration, is key to developing effective, culturally and socially acceptable approaches to sea urchin removal and for assessing the potential benefits of kelp restoration. We assessed whether sea urchin roe quality within urchin barrens in northeastern Aotearoa New Zealand can be sufficient for harvest, and whether sea urchin removal can ultimately enhance urchin roe condition (by increasing food quality and quantity). We monitored roe condition (roe weight and gonadosomatic index [GSI]) in kelp forest and urchin barrens monthly for one year at two sites, and quarterly over two years at four sites following large‐scale (1.6–2 ha) sea urchin removal from urchin barrens. While roe condition was generally lower in barrens, the magnitude of this difference varied seasonally and among sites. Sea urchin roe quality within barrens was sufficient for commercial harvest (GSI ≥ 6%) during spring–summer at three sites. Following large‐scale sea urchin removal, kelp and fucoid density increased over two years (~9–58 plants m−2), but remained low in barren control areas. Although variable, sea urchins from removal areas (restored kelp) typically showed increased GSI, roe weight, and improved roe color within nine months following removal. These results suggest that, at some locations, harvest may provide a viable method for removing urchins from urchin barrens for kelp restoration purposes. However, to ensure sea urchin densities are reduced and maintained below the critical levels needed to achieve restoration outcomes, strategic approaches with critical evaluation of methods are required. Furthermore, we demonstrate that effectively removing urchins and promoting kelp recovery can ultimately improve urchin roe quality.

Distribution of sea urchin barrens in shallow algal communities along the eastern Adriatic coast

The sea urchins Paracentrotus lividus and Arbacia lixula are important herbivores in algal communities of the infralittoral rocky bottoms in the Mediterranean Sea. However, grazing by sea urchins that outweighs natural algal recruitment processes might result in the formation of barrens, which are areas dominated by urchins and coralline algae. We present the results of the first large-scale mapping of sea urchin barrens and analyses of sea urchins’ impact on the main algal communities across 1955.5 km of rocky coastline on the eastern side of the Adriatic Sea (Croatia). Since mapping was performed over a geographically wide area and covered a representative quantity of approximately 40% of the coastline of the central part of the Adriatic Sea, the results could be considered to reflect the general situation in that area of the eastern Adriatic coast. Mapping revealed that sea urchin barrens are present along over 35% of the coastline, while the complete absence of algal cover was recorded in 8% of the inspected area. Communities with canopy-forming algae, which are the most valuable shallow-water communities of the Mediterranean Sea, represent the largest proportion of the observed coastline and are affected by sea urchins in over 28% of their extent. Among other factors, the extensive harvesting of date mussels (Lithophaga lithophaga) in the 1970s, 1980s, and early 1990s is likely one of the main activities that resulted in exceptional sea urchin expansion via the removal of micropredators and establishment of shelters for subadult urchins.

Estimating the extent of urchin barrens and kelp forest loss in northeastern Aotearoa, New Zealand

ABSTRACT On shallow rocky reefs in northeastern Aotearoa, New Zealand, urchin barrens are indicators of the ecosystem effects of overfishing reef predators. Yet, information on their extent and variability is lacking. We use aerial imagery to map the urchin barrens and kelp forests on reefs (<30 m depth) across seven locations, including two long-established marine reserves and a marine protected area that allows recreational fishing. Urchin barrens were present in all locations and were restricted to reefs <10–16 m deep. Urchin barrens covered 30% (7–49%) of shallow reefs in fished areas, with variation among and within locations likely related to the relative extent and topography of reefs and wave exposure. Within the marine reserves, barrens covered <2% of shallow reefs. Long-term comparison of historical imagery at two fished locations demonstrates that current areas of urchin barrens were historically dominated by kelp forests, but barrens have persisted since at least the early 2000s. We estimate ∼30 km2 of barrens exist across the region. This demonstrates the widespread nature of barrens, the potential long-term effectiveness of no-take marine protection in restoring urchin barrens, and the need for a multifaceted management approach to restore kelp forests and prevent further expansion of barrens.

Optimizing the biomass balance of macroalgae and sea urchins in kelp beds by removing the urchins

ABSTRACT In this study, we conducted sea urchin removal (SUR) experiments at two sites along Hokkaido’s southwestern and northwestern coasts in the Sea of Japan and one site along the northeast coast in the Sea of Okhotsk, to elucidate the biomass balance between macroalgae and sea urchins at different urchin barren sites. The experiments resulted in the restoration of macroalgal biomass at all sites and an increase in macroalgal diversity, indicating that it was the impact of high grazing pressure reducing the macroalgae, regardless of the particular experimental site. Notably, the recovery of the macroalgal biomass was characterized by increased kelp biomass, including Saccharina japonica var. religiosa and S. japonica var. ochotensis. In addition, negative correlations were observed between macroalgal biomass and sea urchin density. The lowest sea urchin density inducing an increase of macroalgal biomass was approximately the same in the reproductive growth phase (0.1 kg m−2) and in the vegetative growth phase (0.15 kg m−2) of Saccharina spp., the dominant kelps, regardless of the experimental site. These values can be helpful indicators for managing sea urchin density in order to restore macroalgae. In contrast, the maximum sea urchin density inducing the lowest macroalgal biomass varied markedly depending on the study site, indicating that different levels of kelp productivity are influenced by oceanographic conditions. Moreover, a strong positive correlation was observed between macroalgal biomass and sea urchins’ gonadosomatic index (GSI). Based on this correlation, macroalgal biomass supporting sea urchins with commercial value (GSI ≥ 15%) was estimated to be 1.6 kg m−2. This sea urchin density value is also a helpful indicator for managing macroalgal biomass and commercially valuable sea urchins. Moreover, this value will depend on the influence of the oceanographic environment of the site, particularly the nutrient conditions, on macroalgal productivity. In addition, we have also discussed how macroalgae and sea urchins can best coexist at the experimental sites.

Consequences of kelp forest ecosystem shifts and predictors of persistence through multiple stressors.

Ecological communities can be stable over multiple generations, or rapidly shift into structurally and functionally different configurations. In kelp forest ecosystems, overgrazing by sea urchins can abruptly shift forests into alternative states that are void of macroalgae and primarily dominated by actively grazing sea urchins. Beginning in 2014, a sea urchin outbreak along the central coast of California resulted in a patchy mosaic of remnant forests interspersed with sea urchin barrens. In this study, we used a 14-year subtidal monitoring dataset of invertebrates, algae, and fishes to explore changes in community structure associated with the loss of forests. We found that the spatial mosaic of barrens and forests resulted in a region-wide shift in community structure. However, the magnitude of kelp forest loss and taxonomic-level consequences were spatially heterogeneous. Taxonomic diversity declined across the region, but there were no declines in richness for any group, suggesting compositional redistribution. Baseline ecological and environmental conditions, and sea urchin behaviour, explained the persistence of forests through multiple stressors. These results indicate that spatial heterogeneity in preexisting ecological and environmental conditions can explain patterns of community change.

Kelp and sea urchin settlement mediated by biotic interactions with benthic coralline algal species.

Species interactions can influence key ecological processes that support community assembly and composition. For example, coralline algae encompass extensive diversity and may play a major role in regime shifts from kelp forests to urchin-dominated barrens through their role in inducing invertebrate larval metamorphosis and influencing kelp spore settlement. In a series of laboratory experiments, we tested the hypothesis that different coralline communities facilitate the maintenance of either ecosystem state by either promoting or inhibiting early recruitment of kelps or urchins. Coralline algae significantly increased red urchin metamorphosis compared with a control, while they had varying effects on kelp settlement. Urchin metamorphosis and density of juvenile canopy kelps did not differ significantly across coralline species abundant in both kelp forests and urchin barrens, suggesting that recruitment of urchin and canopy kelps does not depend on specific corallines. Non-calcified fleshy red algal crusts promoted the highest mean urchin metamorphosis percentage and showed some of the lowest canopy kelp settlement. In contrast, settlement of one subcanopy kelp species was reduced on crustose corallines, but elevated on articulated corallines, suggesting that articulated corallines, typically absent in urchin barrens, may need to recover before this subcanopy kelp could return. Coralline species differed in surface bacterial microbiome composition; however, urchin metamorphosis was not significantly different when microbiomes were removed with antibiotics. Our results clarify the role played by coralline algal species in kelp forest community assembly and could have important implications for kelp forest recovery.

Large‐scale one‐off sea urchin removal promotes rapid kelp recovery in urchin barrens

Sea urchin overgrazing is a leading cause of kelp forest loss and in such cases their removal is increasingly advocated for kelp forest restoration. However, refining removal approaches is needed to improve the efficiency and success of restoration, as most previous removal studies have been small scale and require ongoing removals to maintain low densities and allow kelp recovery. We investigated the effectiveness of one‐off urchin removal from large, semi‐discrete areas of urchin barrens as a tool to promote kelp recovery. We removed sea urchins (Evechinus chloroticus) from four areas (1.6–2 ha) of urchin barrens in northeastern New Zealand. Exposed urchins were reduced to approximately 7% of initial densities and remained low, yet cryptic urchin densities increased after 2 years. Kelp (Ecklonia radiata) and fucoid (Sargassum sinclairii, Carpophyllum spp.) densities increased rapidly in removal areas, but remained constant or declined in adjacent control urchin barren areas. Macroalgal canopy recovery varied among and within removal areas, but increased on average from approximately 5 to 43% in 2 years. Densities of large sea urchin predators did not increase with kelp recovery, likely due to ongoing fishing within removal areas. Our results demonstrate that a single urchin removal from large, semi‐discrete areas of urchin barrens can effectively and efficiently promote rapid multi‐species macroalgal recovery without additional actions (e.g. repeated urchin removals or macroalgae enhancement) for at least 2 years. However, this approach does not restore whole ecosystems and consequently restoration benefits through kelp recovery will be temporary without longer‐term urchin management and/or rebuilding of predator populations.

Alterations in sea urchin (Mesocentrotus nudus) microbiota and their potential contributions to host according to barren severity

Sea urchins are biotic factors driving the decline of kelp forests in marine ecosystems. However, few studies have analyzed the microbiota of surviving sea urchins in barren regions with scarce diet resources. Here, we analyzed the microbiota in the pharynx and gut of the sea urchin Mesocentrotus nudus located along the coast of an expanding barren region in South Korea. The ecological adaptation of genera in sea urchins was predicted using the neutral assembly model. The pharynx and gut microbiota were different, and microbes in the surrounding habitats dispersed more to the pharynx than to the gut. The gut microbiota in sea urchins is altered by barren severity and plays different roles in host energy metabolism. These findings help to understand the microbiota in sea urchins according to urchin barren and its contribution to the survival of sea urchins in severe barren regions with limited macroalgae.

The role of kelp availability and quality on the energetic state and thermal tolerance of sea urchin and gastropod grazers

Widespread decline in underwater forests, formed by kelps and other macroalgae, is occurring in all oceans. Loss of the vulnerable surface canopy typically leads to alternate rocky reef states dominated by substrate-near, understory vegetation, or reefs without any large fleshy macroalgae such as sea urchin barrens. Such alternate reef states represent a major shift in food availability and quality, and may impact kelp-preferring grazers forced to rely on less nutritious foods. Here we investigate the physiological responses of grazers to changes in food quality and quantity. We quantify the mass-independent oxygen consumption (ṀO2), as a proxy of whole-organism physiology, of four grazing invertebrate species (two sea urchins: Mesocentrotus franciscanus and Strongylocentrotus purpuratus; and two gastropods: Pomaulax gibberosus and Tegula pulligo) provisioned with one of three diet treatments over seven weeks: 1) higher quality, canopy kelp, 2) lower quality, substrate-near kelp, or 3) restricted: no kelp. We further test for differences in the heat resistance of four grazers using an acute, near-lethal heat exposure after completion of the provisioning period. Food restrictions had the strongest influence on oxygen consumption in the two urchins (mass-independent ṀO2 was reduced by 26–78%). By contrast, the two gastropod grazers did not display metabolic depression under food restrictions. Although kelp restrictions had clear physiological consequences in sea urchins, overall, the type of kelp provisioned (Macrocystis and Saccharina) did not relate to oxygen consumption after seven weeks. While heat resistance was also similar across food treatments, unfed individuals showed a slight tendency for lower heat tolerance compared to fed individuals. These results suggest a weak coupling between heat resistance and food quality and quantity, at least for these species and at short time spans. Our work highlights the importance of diet diversity and metabolic depression as strategies to cope with the energetic costs of warming.

Long-term marine protection enhances kelp forest ecosystem stability.

Trophic downgrading destabilizes ecosystems and can drive large-scale shifts in ecosystem state. While restoring predatory interactions in marine reserves can reverse anthropogenic-driven shifts, empirical evidence of increased ecosystem stability and persistence in the presence of predators is scant. We compared temporal variation in rocky reef ecosystem state in New Zealand's oldest marine reserve to nearby fished reefs to examine whether protection of predators led to more persistent and stable reef ecosystem states in the marine reserve. Contrasting ecosystem states were found between reserve and fished sites, and this persisted over the 22-year study period. Fished sites were predominantly urchin barrens but occasionally fluctuated to short-lived turfs and mixed algal forests, while reserve sites displayed unidirectional successional trajectories toward stable kelp forests (Ecklonia radiata) taking up to three decades following protection. This provides empirical evidence that long-term protection of predators facilitates kelp forest recovery, resists shifts to denuded alternate states, and enhances kelp forest stability.

Emergence of the subtropical sea urchin Centrostephanus rodgersii as a threat to kelp forest ecosystems in northern New Zealand

Marine protected areas have long been proposed as a key tool to restore lost food web interactions and increase the resilience of ecosystems to climate change impacts. However, a changing climate can result in the arrival of new species or differentially affect native species, which can alter ecosystem dynamics and make it difficult to predict how ecosystems will respond to protection. The long-spined sea urchin Centrostephanus rodgersii is a well-known range extender with large impacts on kelp forest ecosystems, yet its response to warming and long-term marine protection has not been examined within its native range. We examine long-term trends in C. rodgersii and the endemic sea urchin Evechinus chloroticus following no-take protection within the Poor Knights Islands Marine Reserve, in northeastern Aotearoa New Zealand, from 1999-2022, and compare with population trends at an unprotected island group. Within the marine reserve, E. chloroticus decreased in density, became more cryptic, and urchin barrens associated with this species largely disappeared, whereas in fished areas, populations and extent of barrens remained stable. This differing response between the reserve and fished location is consistent with a top-down effect and greater abundance of predatory fish in the reserve. In contrast, the subtropical sea urchin C. rodgersii increased in abundance by 9.3 times in the Poor Knights Island Marine Reserve and 4.3 times at the fished location, with areas of urchin barrens associated with this species developing at both locations. This increase coincides with substantial warming over the monitoring period (0.25°C decade-1) and low numbers of key predators (rock lobster) at both reserve and fished locations. This highlights the emerging threat of C. rodgersii to rocky reefs in the region and how marine protection alone may not increase resilience to this threat. This suggests multifaceted management approaches are needed to mitigate the impacts of emerging pest species and increase the resilience of temperate reef ecosystems in a warming climate.

Assessing the carbon storage value of kelp forest restoration in the Hauraki Gulf Marine Park, New Zealand: Lessons from no-take Marine Protected Areas

Anthropogenic CO2 emissions are one of the principal driving forces of climate change. Climate change leads to systematic changes of ecosystems that cause significant impacts on species interactions, ecosystem processes, and result in profound socio-economic consequences. While carbon offsetting of terrestrial forests and their monetary value are well accepted, the carbon offsetting potential and value of coastal and marine ecosystems (Blue Carbon, BC) remains excluded from the international carbon accounting framework. In this study, we estimated the carbon offsetting value of kelp forests (Ecklonia radiata) against urchin barrens (dominated by crustose coralline algae) using the standing stock biomass as a metric. We propose that standing stock biomass is a crucial next step in introducing BC to the general carbon offsetting framework. Our approach is informed by long-term ecological research conducted in a marine reserve, which highlights the potential of utilizing trophic cascades to limit the dominance of urchin barrens in shallow coastal habitats. We found that the value of carbon standing stock in current kelp forests with reforestation in the Hauraki Gulf Marine Park (HGMP) is worth up to $NZ 7.9 million, based on 2021 carbon market pricing in New Zealand. This simple step towards realizing the value of marine conservation and its multiple benefits to society highlights the missed opportunities when policy and plans do not align with ecology and economics. Including the ocean's contributions to carbon storage is a critical step in using monetization techniques to overcome climate change challenges. Monetizing the carbon offsetting value of kelp forests contributes to establishing funding mechanisms to attract diverse investments in marine ecosystem restoration while providing ecological benefits in ocean governance.

Effect of Season and Increased Temperature on Survival, Roe Enhancement, and Reproductive Cycle of the Green Sea Urchin (Strongylocentrotus droebachiensis) Collected from Four Relatively Close Sites in Northern Norway

A series of three trials were conducted in the North of Norway at different times of the year. The trials consisted of harvesting sea urchins from four sites, situated within 5 km of each other and then holding them in land-based enhancement facilities. They were held at ambient seawater temperatures as well as ambient plus 3.0°C. Samples were taken from the wild at the time of collection and at the end of the enhancement period as well as from the enhanced sea urchins from each site. Results showed that utilising the sea urchins from a variety of sites, even from areas where there is an abundance of sea urchins (sea urchin barrens) and the Gonad Index (GI) is almost always low, is viable in terms of the final, post roe enhancement GI. The results of the study indicate that differences in initial and final GI and the increase in GI between the sites, despite the relatively close proximity, show site selection is most likely not as important as ease of harvesting and sea urchin availability and the positive environmental impact from removal of sea urchins and these are the factors that harvesting strategies should focus on. The study has shown that it is possible to have a relatively consistent roe enhancement from S. droebachiensis from the North of Norway regardless of seasonality (time of year), site selection, and initial GI. Relatively small changes (up to 3°C) in the seawater temperature appear to have a slightly negative impact on enhancement efforts. However, this may vary when larger differences in temperature are experienced.

Urchin Gonad Response to Kelp Forest Restoration on the Palos Verdes Peninsula, California

Along the Palos Verdes Peninsula in southern California, high densities of Strongylocentrotus purpuratus (purple sea urchin) have consumed almost all macroalgae on large expanses (61 ha) of rocky reef habitat, creating “urchin barrens.” Mesocentrotus franciscanus (red sea urchin) harvesting comprises an important fishery in the region, as their gonads are sold as a high-value sushi product called “uni.” However, with a lack of macroalgal food resources, urchins in barrens are smaller and exist in a starved state, meaning little, if any, gonad product is available to the fishery. To restore local kelp forests and increase gonad biomass available to the M. franciscanus fishery, beginning in October 2013, S. purpuratus were culled in barrens to a target density of 2 per m2 across 5.2 ha of rocky reef on the Palos Verdes Peninsula. Mesocentrotus franciscanus were collected from urchin barren, restoration, and kelp reference sites from April to November 2014 to compare differences in gonad production among the three site types. Culling S. purpuratus resulted in the recovery of normal seasonal M. franciscanus gonad production throughout the 8-month study. Mesocentrotus franciscanus gonad weights at a given test diameter length in restoration sites were equivalent to, and sometimes exceeded, the gonad production of those from the kelp reference sites. The urchin test length distributions of collected M. franciscanus were consistently smaller at urchin barren sites than at kelp reference sites, while those in restoration sites generally fell in between.

Sunflower sea star predation on urchins can facilitate kelp forest recovery.

The recent collapse of predatory sunflower sea stars (Pycnopodia helianthoides) owing to sea star wasting disease (SSWD) is hypothesized to have contributed to proliferation of sea urchin barrens and losses of kelp forests on the North American west coast. We used experiments and a model to test whether restored Pycnopodia populations may help recover kelp forests through their consumption of nutritionally poor purple sea urchins (Strongylocentrotus purpuratus) typical of barrens. Pycnopodia consumed 0.68 S. purpuratus d-1, and our model and sensitivity analysis shows that the magnitude of recent Pycnopodia declines is consistent with urchin proliferation after modest sea urchin recruitment, and even small Pycnopodia recoveries could generally lead to lower densities of sea urchins that are consistent with kelp-urchin coexistence. Pycnopodia seem unable to chemically distinguish starved from fed urchins and indeed have higher predation rates on starved urchins owing to shorter handling times. These results highlight the importance of Pycnopodia in regulating purple sea urchin populations and maintaining healthy kelp forests through top-down control. The recovery of this important predator to densities commonly found prior to SSWD, whether through natural means or human-assisted reintroductions, may therefore be a key step in kelp forest restoration at ecologically significant scales.