Massive barnacle recruitment on the Gulf of St. Lawrence coast of Nova Scotia (Canada) in 2024 linked to increased sea surface temperature.

With the ongoing climate and oceanographic change, an increasing number of studies are reporting dramatic population losses caused by thermal extremes in intertidal habitats. Under moderate warming, however, populations can fare better in places where species normally experienced suboptimal temperatures. This article reports the massive recruitment of the barnacle Semibalanus balanoides on the Gulf of St. Lawrence coast of Nova Scotia (Canada) in 2024. As recruits appear mostly during May in this region, coastal sea surface temperature (SST) in April is critical for the ecological performance of larvae, as they are pelagic and live in the water column for weeks before intertidal settlement. Thus, a study that spanned 12 years (2005 to 2016) on this coast found that annual barnacle recruitment was positively correlated to April SST. In April 2024, coastal SST was 116% higher than for the same month averaged over those 12 years (4.1 vs. 1.9 °C). This SST spike was followed by an elevated recruitment that was 111% higher than the average for those 12 years (1,278 vs. 607 recruits dm-2). Overall for the studied years, the amount of variation in annual barnacle recruitment statistically explained by April SST was 51%. While the southern distribution limit of S. balanoides has moved northwards in recent decades due to lethal warming, our results support the notion of improving reproductive success with seawater warming on colder northern shores.

Does foil-like debris impair barnacles by covering them?

On 24 June 2024, we detected foil that tightly adhered to an intertidal wall in Vigo harbor (Spain) during low tide. It covered multiple barnacles, potentially threatening their survival. We present photos of this novel debris-animal interaction and discuss possible effects that such cover could have on barnacles.

Musseling through: Mytilus byssal thread production is unaffected by continuous noise.

Anthropogenic low-frequency noise (ALFN) is a rising pollutant in the world oceans. Despite the ubiquity of ALFN, its effect on marine invertebrates is still poorly understood. Here, we tested how continuous low-frequency noise (CLFN), a substantial component of ALFN, affects the byssal thread production of Mytilus, a cosmopolitan genus of mussels with high ecological and economic importance. The effects of acute CLFN exposure and predator cues on byssogenesis by Mytilus spp. were explored in both the presence and absence of predator cues. While predator effluents increased thread production, CLFN had seemingly no effect on thread counts. Further, trends suggested a synergistic effect of CLFN and predator cues. The behavioral indifference of Mytilus spp. toward CLFN could contribute to the observed prevalence of these animals in inherently disturbed habitats. This would partly explain their success in colonizing and persisting on artificial substrata rife with disturbances.

Changes in the composition of invertebrate assemblages from wave-exposed intertidal mussel stands along the Nova Scotia coast, Canada.

Rocky intertidal habitats occur worldwide and are mainly characterized by primary space holders such as seaweeds and sessile invertebrates. Some of these organisms are foundation species, as they can form structurally complex stands that host many small invertebrates. The abundance of primary space holders is known to vary along coastlines driven directly or indirectly by environmental variation. However, it is less clear if the invertebrate assemblages associated to a foundation species may remain relatively unchanged along coastlines, as similar stands of a foundation species can generate similar microclimates. We examined this question using abundance data for invertebrate species found in mussel stands of a similar structure in wave-exposed rocky habitats at mid-intertidal elevations along the Atlantic coast of Nova Scotia (Canada). While the most abundant invertebrate species were found at three locations spanning 315 km of coastline, species composition (a combined measure of species identity and their relative abundance) differed significantly among the locations. One of the species explaining the highest amount of variation among locations (a barnacle) exhibited potential signs of bottom-up regulation involving pelagic food supply, suggesting benthic-pelagic coupling. The abundance of the species that explained the highest amount of variation (an oligochaete) was positively related to the abundance of their predators (mites), further suggesting bottom-up forcing in these communities. Overall, we conclude that species assemblages associated to structurally similar stands of a foundation species can show clear changes in species composition at a regional scale.

Characterization of three plastic forms: Plasticoncrete, plastimetal and plastisessiles.

Plastic forms, including plastiglomerate, pyroplastic, plasticrusts, anthropoquinas, plastistone and plastitar, were recorded worldwide. These plastic forms derive from geochemical or geophysical interactions such as heat-induced plastic fusion with rock in campfires, incomplete plastic combustion, water motion-driven plastic abrasion in the rocky intertidal zone, plastic deposition in hardened sediments and plastic bonding with tar. Thereby, these interactions can profoundly influence the fate of plastics in the environment. This study characterized three novel plastic forms (plasticoncrete, plastimetal and plastisessiles) discovered on Helgoland island (North Sea). Plasticoncrete consisted of common polyethylene (PE) and polypropylene (PP) fibers hardened in concrete. Plastimetal included PE fibers rusted with metal. Plastisessiles consisted of PE fibers attached to benthic substrates by sessile invertebrates (oysters and polychaetes). Plasticoncrete and plastimetal are the first plastic forms composed of two man-made materials. Plastisessiles show that plastic forms not only result from human- or environment-mediated interactions but also from biological interactions between invertebrates and plastic. All plastic forms (bulk density ≥ 1.4 g/cm3) sunk during floating tests and hardly changed their positions during a 13-day field experiment and 153- to 306-day field monitorings, indicating their local formation, limited mobility and longevity. Still, experimentally detached plastic fibers floated, confirming that the formation of these plastic forms influences the fate of plastic fibers in the environment. Furthermore, the experiment showed that plasticoncrete got deposited in beach sand under wavy and windy conditions, indicating that coastal waves and onshore winds drive plasticoncrete deposition in coastal sediments. We also provide first records of plasticoncrete on Mallorca island (Mediterranean Sea) and plastimetal on Hikoshima island (Sea of Japan), respectively, which show that these plastic forms are no local phenomena. Thereby, our study contributes to the growing fundamental knowledge of plastic forms that is essential to understand the role and fate of these pollutants in coastal habitats worldwide.

Plasticrust generation and degeneration in rocky intertidal habitats contribute to microplastic pollution.

Plasticrusts are a plastic form that consists of plastic encrusting intertidal rocks. To date, plasticrusts have been reported on Madeira island (Atlantic Ocean), Giglio island (Mediterranean Sea) and in Peru (Pacific Ocean) but information on plasticrust sources, generation, degeneration and fate is largely missing. To address these knowledge gaps, we combined plasticrust field surveys, experiments and monitorings along the Yamaguchi Prefecture (Honshu, Japan) coastline (Sea of Japan) with macro-, micro- and spectroscopic analyses in Koblenz, Germany. Our surveys detected polyethylene (PE) plasticrusts that derived from very common PE containers and polyester (PEST) plasticrusts that resulted from PEST-based paint. We also confirmed that plasticrust abundance, cover and distribution were positively related to wave exposure and tidal amplitude. Our experiments showed that plasticrusts are generated by cobbles scratching across plastic containers, plastic containers being dragged across cobbles during beach clean-ups, and waves abrading plastic containers on intertidal rocks. Our monitorings found that plasticrust abundance and cover decreased over time and the macro- and microscopic examinations indicated that detached plasticrusts contribute to microplastic pollution. The monitorings also suggested that hydrodynamics (wave occurrence, tidal height) and precipitation drive plasticrust degeneration. Finally, floating tests revealed that low density (PE) plasticrusts float whereas high density (PEST) plasticrusts sink suggesting that polymer type floatability influences the fate of plasticrusts. By tracking the entire lifespan of plasticrusts for the first time, our study contributes fundamental knowledge of plasticrust generation and degeneration in the rocky intertidal zone and identified plasticrusts as novel microplastic sources.

Frequent observations of novel plastic forms in the Ariho River estuary, Honshu, Japan.

Pyroplastic and plastiglomerate are novel plastic forms that are currently being reported from coastal beaches worldwide. Pyroplastic is burned plastic with a rock-like appearance. Plastiglomerate is a solid bond consisting of either melted plastic attached to rock (in-situ plastiglomerate) or a melted plastic matrix containing (in)organic material (clastic plastiglomerate). Both plastic forms have been related to the (un)intentional burning of plastic. Yet, information on pyroplastic and plastiglomerate from estuarine habitats is limited to a pilot study (for this study) and knowledge of pyroplastic and plastiglomerate dynamics as well as the underlying drivers is missing. To address these knowledge gaps, we frequently surveyed stranded pyroplastics and plastiglomerates in the Ariho River estuary (Honshu, Japan) over seven months and studied the collected samples at the lab. In total, 37 pyroplastics (consisting of polyethylene, polypropylene, polystyrene, alkyd resin, polyacrylate styrene and polyvinyl chloride) and seven plastiglomerates (consisting of polyethylene and polypropylene) occurred. While pyroplastics occurred frequently, plastiglomerates occurred occasionally which indicates that both forms are common. Pyroplastic (but not plastiglomerate) occurrence and density (items/m2) were related to intertidal elevation. Strandline pyroplastic density, that contributed heavily to the pyroplastic and plastiglomerate entirety, increased under prevailing onshore winds which shows that such winds are environmental drivers of pyroplastic density. Floating tests revealed that clastic plastiglomerate can float. Macro-, micro- and spectroscopic examinations indicated only slight pyroplastic and plastiglomerate weathering which suggests the regional and/or recent formation of both plastic forms. Additionally, we detected the first plastiglomerate with clastic and in-situ features (a plastic matrix containing (in)organic material firmly melted to a rock) which constituted a novel plastiglomerate subtype that we termed "clastic/in-situ plastiglomerate". Overall, our study initiates the development of the fundamental understandings of pyroplastic and plastiglomerate dynamics and the underlying drivers in estuaries.

Microplastic load and polymer type composition in European rocky intertidal snails: Consistency across locations, wave exposure and years.

Microplastics (<5 mm) are emerging pollutants in oceans worldwide. As such small particles are easily ingested, microplastics are found in numerous pelagic and benthic organisms. However, information on microplastics in rocky intertidal organisms and habitats is relatively scant. Therefore, we examined snails and water samples from wave-sheltered and wave-exposed rocky intertidal habitats in Helgoland (North Sea), Cap Ferrat and Giglio (Mediterranean) and Madeira (Atlantic Ocean) in 2019-2020 for microplastics. Furthermore, we examined snails from the same habitats in Helgoland, Cap Ferrat and Giglio in 2007-2009. In total, we performed 362 individual micro-Fourier-transform infrared spectroscopy (µFTIR) measurements on the snails and water samples. While the snails contained 50 microplastics (composed of nine polymer types), the water samples contained 24 microplastics (comprising six polymer types). Microplastic load and polymer type composition in the snails were rather similar across locations, wave exposure and years. Also, microplastic load and polymer composition in the water samples were similar across locations and wave exposure. Moreover, snail and water microplastic loads were significantly correlated which indicates that snails are useful bioindicators for microplastic loads in rocky intertidal habitats. Interestingly, the majority of the microplastics consisted of paint chips that likely derived from ships. Overall, our study provides the first comprehensive microplastic record in rocky intertidal organisms across locations, wave exposure and years that can serve as a baseline to examine historic and future microplastic dynamics in rocky intertidal systems.

Field observations in pebble beach habitats link plastiglomerate to pyroplastic via pebble clasts.

Plastiglomerate and pyroplastic are two novel plastic debris forms that were originally discovered on sandy beaches in Hawaii and the UK, respectively. While plastiglomerate consists of plastic melted together with rocks or pebbles, pyroplastic is melted plastic. Although both plastic debris forms were related to campfires, it is unclear whether they are related to each other. Also, plastiglomerate and pyroplastic records from other shore types are missing. Therefore, we surveyed pebble beach habitats in Madeira Island (Atlantic Ocean) for plastiglomerate and pyroplastic. We detected one plastiglomerate (PG1, including a pebble) and four pyroplastics (PP1-4). While PP2-4 consisted of polypropylene, PG1 and PP1 consisted of polyethylene and polypropylene. Furthermore, PG1 and PP1 included previously undescribed pebble shaped clasts that unequivocally linked plastiglomerate to pyroplastic. Thereby, our findings provide the first record of plastiglomerate and pyroplastic from pebble beach habitats worldwide and establish the link between these two novel plastic debris forms.

Plasticrusts derive from maritime ropes scouring across raspy rocks.

Plasticrusts are a novel form of plastic debris which has only recently been discovered in Madeira Island, NE Atlantic Ocean. Plasticrusts consist of plastic encrusting wave-exposed rocky intertidal habitats and are presumably generated by waves smashing plastic debris against intertidal rocks. However, direct observations of this process are lacking and it is unknown which type of plastic debris the plasticrusts derive from. Therefore, we examined the Madeira rocky intertidal for signs of plasticrust formation and collected plasticrust and co-occurring plastic debris pieces of matching colors. We examined all collected materials using digital microscopy and Fourier-transform infrared spectroscopy. We found that plasticrusts can result from maritime ropes being scoured across raspy intertidal rocks and that the plasticrusts and the corresponding ropes consisted of polypropylene (PP) and high-density polyethylene (HDPE). Furthermore, we show that high temperatures contribute to plasticrust formation. Thereby, our study provides first insights into the complex plasticrust formation process.

Limpet disturbance effects on barnacle recruitment are related to recruitment intensity but not recruit size.

Intertidal limpets are important grazers along rocky coastlines worldwide that not only control algae but also influence invertebrates such as common barnacles. For instance, grazing limpets ingest settling barnacle cyprid larvae (hereafter cyprids) and push cyprids and barnacle recruits off the substrate. Such limpet disturbance effects (LDEs) can limit barnacle recruitment, a key demographic variable affecting barnacle population establishment and persistence. In this study, we examined limpet (Lottia cassis) disturbance to barnacle (Chthamalus dalli, Balanus glandula) recruitment on the Pacific coast of Hokkaido, Japan, as information on limpet-barnacle interactions from this region is missing. We investigated, for the first time, whether barnacle size and recruitment intensity influence LDEs on barnacle recruitment. Small barnacles may be less susceptible to LDEs than larger barnacles, because small size may reduce the propbability of limpet disturbance. Moreover, recruitment intensity can influence LDEs, as high recruitment can compensate for LDEs on barnacle recruitment density. In Hokkaido, C. dalli cyprids are smaller than B. glandula cyprids, and C. dalli recruitment is higher than B. glandula recruitment. Thus, we hypothesized that LDEs on C. dalli recruitment would be weaker than those on B. glandula recruitment. To test our hypothesis, we conducted a field experiment during which we manipulated limpet presence/absence on the interior surfaces of ring-shaped cages. After four weeks, we measured barnacle recruitment and recruit size on the interior surfaces of the cages and found negative LDEs on C. dalli and B. glandula recruitment and recruit size. As hypothesized, the LDEs on C. dalli recruitment were weaker than the LDEs on B. glandula recruitment. Additionally, C. dalli recruits were smaller than B. glandula recruits. However, the LDEs on C. dalli recruit size were as strong as the LDEs on B. glandula recruit size, indicating that the smaller C. dalli recruits are not less susceptible to LDEs than B. glandula recruits. Since C. dalli recruitment was higher than B. glandula recruitment, we propose that the higher C. dalli recruitment compensated for the LDEs on C. dalli recruitment. Our findings indicate that the detected differences in LDEs on barnacle recruitment are related to barnacle recruitment intensity but not recruit size.

First record of 'plasticrusts' and 'pyroplastic' from the Mediterranean Sea.

We report the presence of 'plasticrusts' and 'pyroplastic' from coastal habitats in Giglio island, Tyrrhenian Sea, Italy. These novel plastic debris types have only recently been described for the first time from Madeira island (NE Atlantic Ocean) and the United Kingdom, respectively. While 'plasticrusts' are generated by sea waves smashing plastic debris against intertidal rocks, 'pyroplastic' derives from (un)deliberately burnt plastic waste. Using Fourier-transform infrared (FTIR) spectroscopy, we identified the 'plasticrust' material as polyethylene (PE) and the 'pyroplastic' material as polyethylene terephthalate (PET). These polymers are widely used in everyday products and, therefore, contribute heavily to plastic pollution in aquatic and terrestrial environments worldwide. Furthermore, our field surveys suggest that 'plasticrust' abundance is related to wave-exposure and that the 'pyroplastic' derived from beverage bottles which we frequently found along the Giglio coast. Overall, our findings corroborate the notion that 'plasticrusts' and 'pyroplastic' are common debris types in marine coastal habitats.

A 5-year study (2014-2018) of the relationship between coastal phytoplankton abundance and intertidal barnacle size along the Atlantic Canadian coast.

Benthic-pelagic coupling refers to the ecological relationships between benthic and pelagic environments. Studying such links is particularly useful to understand biological variation in intertidal organisms along marine coasts. Filter-feeding invertebrates are ecologically important on marine rocky shores, so they have often been used to investigate benthic-pelagic coupling. Most studies, however, have been conducted on eastern ocean boundaries. To evaluate benthic-pelagic coupling on a western ocean boundary, we conducted a 5-year study spanning 415 km of the Atlantic coast of Nova Scotia (Canada). We hypothesized that the summer size of intertidal barnacles (Semibalanus balanoides) recruited in the preceding spring would be positively related to the nearshore abundance (biomass) of phytoplankton, as phytoplankton constitutes food for the nauplius larvae and benthic stages of barnacles. Every year between 2014 and 2018, we measured summer barnacle size in clearings created before spring recruitment on the rocky substrate at eight wave-exposed locations along this coast. We then examined the annual relationships between barnacle size and chlorophyll-a concentration (Chl-a), a proxy for phytoplankton biomass. For every year and location, we used satellite data to calculate Chl-a averages for a period ranging from the early spring (when most barnacle larvae were in the water) to the summer (when barnacle size was measured after weeks of growth following spring benthic recruitment). The relationships were always positive, Chl-a explaining nearly half, or more, of the variation in barnacle size in four of the five studied years. These are remarkable results because they were based on a relatively limited number of locations (which often curtails statistical power) and point to the relevance of pelagic food supply to explain variation in intertidal barnacle size along this western ocean boundary coast.

Unimodal relationship between small-scale barnacle recruitment and the density of pre-existing barnacle adults.

Recruitment is a key demographic process for population persistence. This paper focuses on barnacle (Semibalanus balanoides) recruitment. In rocky intertidal habitats from the Gulf of St. Lawrence coast of Nova Scotia (Canada), ice scour is common during the winter. At the onset of intertidal barnacle recruitment in early May (after sea ice has fully melted), mostly only adult barnacles and bare substrate are visible at high elevations in wave-exposed habitats. We conducted a multiannual study to investigate if small-scale barnacle recruitment could be predicted from the density of pre-existing adult barnacles. In a year that exhibited a wide adult density range (ca. 0-130 individuals dm-2), the relationship between adult density and recruit density (scaled to the available area for recruitment, which excluded adult barnacles) was unimodal. In years that exhibited a lower adult density range (ca. 0-40/50 individuals dm-2), the relationship between adult and recruit density was positive and resembled the lower half of the unimodal relationship. Overall, adult barnacle density was able to explain 26-40% of the observed variation in recruit density. The unimodal adult-recruit relationship is consistent with previously documented intraspecific interactions. Between low and intermediate adult densities, the positive nature of the relationship relates to the previously documented fact that settlement-seeking larvae are chemically and visually attracted to adults, which might be important for local population persistence. Between intermediate and high adult densities, where population persistence may be less compromised and the abundant adults may limit recruit growth and survival, the negative nature of the relationship suggests that adult barnacles at increasingly high densities stimulate larvae to settle elsewhere. The unimodal pattern may be especially common on shores with moderate rates of larval supply to the shore, because high rates of larval supply may swamp the coast with settlers, decoupling recruit density from local adult abundance.

A 12-year record of intertidal barnacle recruitment in Atlantic Canada (2005-2016): relationships with sea surface temperature and phytoplankton abundance.

On the Gulf of St. Lawrence coast of Nova Scotia (Canada), recruitment of the barnacle Semibalanus balanoides occurs in May and June. Every year in June between 2005 and 2016, we recorded recruit density for this barnacle at the same wave-exposed rocky intertidal location on this coast. During these 12 years, mean recruit density was lowest in 2015 (198 recruits dm-2) and highest in 2007 (969 recruits dm-2). The highest recruit density observed in a single quadrat was 1,457 recruits dm-2 (in 2011) and the lowest was 34 recruits dm-2 (in 2015). Most barnacle recruits appear during May, which suggests that most pelagic larvae (which develop over 5-6 weeks before benthic settlement) are in the water column in April. An AICc-based model selection approach identified sea surface temperature (SST) in April and the abundance of phytoplankton (food for barnacle larvae, measured as chlorophyll-a concentration -Chl-a-) in April as good explanatory variables. Together, April SST and April Chl-a explained 51% of the observed interannual variation in recruit density, with an overall positive influence. April SST was positively related to March-April air temperature (AT). April Chl-a was negatively related to the April ratio between the number of days with onshore winds (which blow from phytoplankton-limited offshore waters) and the number of days with alongshore winds (phytoplankton is more abundant on coastal waters). Therefore, this study suggests that climatic processes affecting April SST and April Chl-a indirectly influence intertidal barnacle recruitment by influencing larval performance.

Adult Prey Neutralizes Predator Nonconsumptive Limitation of Prey Recruitment.

Recent studies have shown that predator chemical cues can limit prey demographic rates such as recruitment. For instance, barnacle pelagic larvae reduce settlement where predatory dogwhelk cues are detected, thereby limiting benthic recruitment. However, adult barnacles attract conspecific larvae through chemical and visual cues, aiding larvae to find suitable habitat for development. Thus, we tested the hypothesis that the presence of adult barnacles (Semibalanus balanoides) can neutralize dogwhelk (Nucella lapillus) nonconsumptive effects on barnacle recruitment. We did a field experiment in Atlantic Canada during the 2012 and 2013 barnacle recruitment seasons (May-June). We manipulated the presence of dogwhelks (without allowing them to physically contact barnacles) and adult barnacles in cages established in rocky intertidal habitats. At the end of both recruitment seasons, we measured barnacle recruit density on tiles kept inside the cages. Without adult barnacles, the nearby presence of dogwhelks limited barnacle recruitment by 51%. However, the presence of adult barnacles increased barnacle recruitment by 44% and neutralized dogwhelk nonconsumptive effects on barnacle recruitment, as recruit density was unaffected by dogwhelk presence. For species from several invertebrate phyla, benthic adult organisms attract conspecific pelagic larvae. Thus, adult prey might commonly constitute a key factor preventing negative predator nonconsumptive effects on prey recruitment.

First visual record of rare purple-colored dogwhelks ( Nucella lapillus) on the Atlantic coast of Nova Scotia, Canada.

The dogwhelk Nucella lapillus is a rocky intertidal gastropod of the North Atlantic coast. Individual shell color varies. Common colors range between white and brown, with darker dogwhelks being more affected by heat stress than lighter-colored conspecifics. Other reported shell colors are purple, black, mauve, pink, yellow, and orange from UK coasts, red and gray from the Bay of Fundy coast of New Brunswick and Nova Scotia (Canada), and purple, black, gray, yellow, and orange from the coasts of Maine and Massachusetts (USA), with purple being considered as a rare color. On the Atlantic coast of Nova Scotia, dogwhelks are active from April until November, but information on dogwhelk shell color is missing for this coast. On 16 June 2016, we found two purple-colored dogwhelks in the mid-to-high intertidal zone of a moderately wave-exposed rocky shore near Duncans Cove, on the Atlantic coast of Nova Scotia while collecting dogwhelks (n= 1000) during low tide for manipulative field experiments. All other dogwhelks collected on that day were of common white and brown colors. During earlier dogwhelk collections in Atlantic Nova Scotia (between 2011-2013) and field surveys in Duncans Cove (between 2014-2016), we did not find any purple-colored dogwhelks, indicating the rareness of this color in that region. Apparently, our observations provide the first visual record of rare purple-colored dogwhelks on the Atlantic coast of Nova Scotia, Canada.

Predator nonconsumptive effects on prey recruitment weaken with recruit density.

We investigated the nonconsumptive effects (NCEs) of predatory dogwhelks (Nucella lapillus) on intertidal barnacle (Semibalanus balanoides) recruitment through field experiments on the Gulf of St. Lawrence coast and the Atlantic coast of Nova Scotia, Canada. We studied the recruitment seasons (May-June) of 2011 and 2013. In 2011, the Gulf coast had five times more nearshore phytoplankton (food for barnacle larvae and recruits) during the recruitment season and yielded a 58% higher barnacle recruit density than the Atlantic coast at the end of the recruitment season. In 2013, phytoplankton levels and barnacle recruit density were similar on both coasts and also lower than for the Gulf coast in 2011. Using the comparative-experimental method, the manipulation of dogwhelk presence (without allowing physical contact with prey) revealed that dogwhelk cues limited barnacle recruitment under moderate recruit densities (Atlantic 2011/2013 and Gulf 2013) but had no effect under a high recruit density (Gulf 2011). Barnacle recruits attract settling larvae through chemical cues. Thus, the highest recruit density appears to have neutralized dogwhelk effects. This study suggests that the predation risk perceived by settling larvae may decrease with increasing recruit density and that prey food supply may indirectly influence predator NCEs on prey recruitment.