Climate change impacts on oyster aquaculture - Part II: Impact assessment and adaptation measures.

The oyster aquaculture sector plays a major role in food security, providing a sustainable way to obtain food and livelihood for coastal and Island nations. Oysters are one of the preferred choices by aquaculturists because of their resilience to harsh climatic conditions. Nonetheless, climate change will continue to pose threats to its culture. Climate-induced hazards such as floods, storms, disease, and invasive species are some of the key factors limiting oyster production globally. A thriving aquaculture industry needs optimal conditions to maximize exploitation. Here, we continue with the review of the impacts of climate change on oyster aquaculture at the global scale, highlighting climate vulnerability assessment. We also propose a framework for modeling oyster responses to future climate scenarios. Furthermore, we explore the health implications of infected oysters on consumer's health. We also identify knowledge gaps and challenges for sustainable oyster production. Additionally, we document mitigation and adaptation measures and future research directions.

Splitting light pollution: Wavelength effects on the activity of two sandy beach species.

Artificial Light at Night (ALAN) threatens to disrupt most natural habitats and species, including those in coastal settings, where a growing number of studies have identified ALAN impacts. A careful examination of the light properties behind those impacts is important to better understand and manage the effects of this stressor. This study focused on ALAN monochromatic wavelengths and examined which types of light spectra altered the natural activity of two prominent coastal species from the Pacific southeast: the talitroid amphipod Orchestoidea tuberculata and the oniscoid isopod Tylos spinulosus. We compared the natural daylight/night activity of these organisms with the one they exhibit when exposed to five different ALAN wavelengths: lights in the violet, blue, green, amber, and red spectra. Our working hypothesis was that ALAN alters these species' activity at night, but the magnitude of such impact differs depending on light wavelengths. Measurements of activity over 24 h cycles for five consecutive days and in three separate experiments confirmed a natural circadian activity pattern in both species, with strong activity at night (∼90% of probability) and barely any activity during daylight. However, when exposed to ALAN, activity declined significantly in both species under all light wavelengths. Interestingly, amphipods exhibited moderate activity (∼40% of probability) when exposed to red lights at night, whereas isopods shifted some of their activity to daylight hours in two of the experiments when exposed to blue or amber lights, suggesting a possible alteration in this species circadian rhythm. Altogether, our results were consistent with our working hypothesis, and suggest that ALAN reduces night activity, and some wavelengths have differential effects on each species. Differences between amphipods and isopods are likely related to their distinct adaptations to natural low-light habitat conditions, and therefore distinct sensitivity to ALAN.

Artificial light at night alters the feeding activity and two molecular indicators in the plumose sea anemone Metridium senile (L.).

Artificial light at night (ALAN) is becoming a widespread stressor in coastal ecosystems, affecting species that rely on natural day/night cycles. Yet, studies examining ALAN effects remain limited, particularly in the case of sessile species. This study assessed the effects of ALAN upon the feeding activity and two molecular indicators in the widespread plumose sea anemone Metridium senile. Anemones were exposed to either natural day/night or ALAN conditions to monitor feeding activity, and tissue samples were collected to quantify proteins and superoxide dismutase (SOD) enzyme concentrations. In day/night conditions, sea anemones showed a circadian rhythm of activity in which feeding occurs primarily at night. This rhythm was altered by ALAN, which turned it into a reduced and more uniform pattern of feeding. Consistently, proteins and SOD concentrations were significantly lower in anemones exposed to ALAN, suggesting that ALAN can be harmful to sea anemones and potentially other marine sessile species.

Climate change impacts on oyster aquaculture - Part I: Identification of key factors.

Oysters are enriched with high-quality protein and are widely known for their exquisite taste. The production of oysters plays an important role in the local economies of coastal communities in many countries, including Atlantic Canada, because of their high economic value. However, because of the changing climatic conditions in recent years, oyster aquaculture faces potentially negative impacts, such as increasing water acidification, rising water temperatures, high salinity, invasive species, algal blooms, and other environmental factors. Although a few isolated effects of climate change on oyster aquaculture have been reported in recent years, it is not well understood how climate change will affect oyster aquaculture from a systematic perspective. In the first part of this study, we present a systematic review of the impacts of climate change and some key environmental factors affecting oyster production on a global scale. The study also identifies knowledge gaps and challenges. In addition, we present key research directions that will facilitate future investigations.

Predator-prey interactions in a coastal setting: Linking crab feeding rates to small scale distribution of clams.

A coastal predator-prey system, juvenile green crabs (Carcinus maenas) preying upon juvenile hard clams (Mercenaria mercenaria), was used to explore the link between crab predation rates and clam density and small-scale distribution patterns. The channel working area of a racetrack flume was adapted to form a sedimentary arena in a flowing seawater system (5 cm s-1) to assess crab predation rates in relation to clam density and distribution patterns (clams clustered in one patch vs two nearby vs two farther apart). The trials detected significant differences in relation to clam initial density and distribution with strong (∼50%) declines in clam mortality levels among spatial arrangements (one patch > two nearby > two farther apart). Feeding of clams was associated with the time taken by crabs to handle the first clam (first patch), and the frequency of three distinct types of crab behavior (eating, resting, and searching). Altogether these results suggest that small-scale changes in number and distribution of juvenile clams matter and may have unexpectedly strong effects on the outcome of predator-prey interactions.

Cross-examining the influence of upwelling and seaweed quality on herbivores' feeding behavior and growth.

At the regional scale, upwelling conditions are known to influence ecosystems and communities and their primary and secondary productivity. However, the influence of upwelling on local herbivore-algae interactions is less well understood. We address this question by cross-examining herbivores and seaweeds from sites associated with upwelling and downwelling conditions along the Humboldt Current System. Specifically, we quantified the feeding and benefits attained by the black sea urchin (Tetrapygus niger) and the black sea snail (Tegula atra) while consuming a widespread kelp species (Lessonia spicata). We hypothesized that food quality drives herbivores' preference, consumption, and growth rates, regardless of the origin or "prior" conditions of the consumers. Laboratory trials measured algal consumption rates with (preference) and without a choice, and consumer's growth rates, to assess the influence of food quality (algae from upwelling vs downwelling sites) and the site of origin of the consumers. Our results showed that algal quality was a prevailing factor for both herbivores: they chose, consumed more, and grew faster on high quality (upwelling) algae. By comparison, the origin of the consumer was only significant for sea snails: those coming from an upwelling site, consumed significantly more and grew faster than those from downwelling. The bulk of our results provided strong support to our hypothesis and suggest that the high nutritional quality of algae associated with upwelling centers has a strong influence on consumers' preferences, consumption, and performance (growth). The fact that origin was found to be relevant for one of the herbivores suggests that the conditions in which species grow may dictate some of their efficiency as consumers.

Artificial Light at Night (ALAN) causes size-dependent effects on intertidal fish decision-making.

Artificial Light at Night (ALAN) alters cycles of day and night, potentially modifying species' behavior. We assessed whether exposure to ALAN influences decision-making (directional swimming) in an intertidal rockfish (Girella laevisifrons) from the Southeastern Pacific. Using a Y-maze, we examined if exposure to ALAN or natural day/night conditions for one week affected the number of visits and time spent in three Y-maze compartments: dark and lit arms ("safe" and "risky" conditions, respectively) and a neutral "non-decision" area. The results showed that fish maintained in natural day/night conditions visited and spent more time in the dark arm, regardless of size. Instead, fish exposed to ALAN visited and spent more time in the non-decision area and their response was size-dependent. Hence, prior ALAN exposure seemed to disorient or reduce the ability of rock fish to choose dark conditions, deemed the safest for small fish facing predators or other potential threats.

The influence of upwelling on key bivalves from the Humboldt and Iberian current systems.

Eastern Boundary Upwelling Systems (EBUS) deliver cold, nutrient-rich waters, influencing coastal biota from the molecular to the ecosystem level. Although local upwelling (U) and downwelling (DU) conditions are often known, their influence on body attributes of relevant species has not been systematically compared within and between EBUS (i.e., below and above regional scales). Hence, we compared the physical-chemical characteristics of U and DU sites in the Humboldt Current system (Chile) and the Iberian Current system (Portugal). We then assessed the influence of U and DU upon eight body attributes in purple mussels (Perumytilus purpuratus) and Mediterranean mussels (Mytilus galloprovincialis), from the Humboldt and Iberian systems, respectively. We hypothesized that bivalves from U sites display better fitness, as measured by body attributes, regardless of their origin (EBUS). As expected, waters from U sites in both systems showed lower temperatures and pH, and higher nitrite concentrations. We also found that mussels from U sites showed better fitness than those in DU sites in 12 out of 16 direct U vs DU comparisons. Shell length, shell volume, organic content of soft-tissues, and mechanical properties of the shell averaged consistently higher in mussels from U sites in both Current systems. In addition, total weight, soft-tissue weight, shell weight and shell thickness were all higher in the U site at the Humboldt system but had less consistent differences at the Iberian system. Altogether, most results supported our working hypothesis and indicate that U conditions support better fitted mussels. The few attributes that did not exhibit the expected U vs DU differences in the Iberian system suggest that local and species-specific differences also play a role on the attributes of these species. These results may also serve as a reference point for further studies addressing the influence of upwelling in these productive, critically important systems.

Field experimental evidence of sandy beach community changes in response to artificial light at night (ALAN).

Artificial light at night (ALAN) is a pervasive but still under-recognized driver of global change. In coastal settings, a large majority of the studies assessing ALAN impacts has focused on individual species, even though it is unclear whether results gathered from single species can be used to predict community-wide responses. Similarly, these studies often treat species as single life-stage entities, ignoring the variation associated with distinct life stages. This study addresses both limitations by focusing on the effects of ALAN on a sandy beach community consisting of species with distinct early- and late-life stages. Our hypothesis was that ALAN alters community structure and these changes are mediated by individual species and also by their ontogenetic stages. A field experiment was conducted in a sandy beach of north-central Chile using an artificial LED system. Samples were collected at different night hours (8-levels in total) across the intertidal (9-levels) over several days in November and January (austral spring and summer seasons). The abundance of adults of all species was significantly lower in ALAN treatments. Early stages of isopods showed the same pattern, but the opposite was observed for the early stages of the other two species. Clear differences were detected in the zonation of these species during natural darkness versus those exposed to ALAN, with some adult-juvenile differences in this response. These results support our hypothesis and document a series of changes affecting differentially both early and late life stages of these species, and ultimately, the structure of the entire community. Although the effects described correspond to short-term responses, more persistent effects are likely to occur if ALAN sources become established as permanent features in sandy beaches. The worldwide growth of ALAN suggests that the scope of its effect will continue to grow and represents a concern for sandy beach systems.

Hemocyanin as a biological indicator of artificial light at night stress in sandy beach amphipods.

The influence of artificial light at night (ALAN) is becoming evident in marine sandy beaches. These habitats are dominated by species reliant on natural daylight/night regimes, making the identification of biological indicators a priority. We assessed the applicability of hemocyanin, an oxygen-transport protein in the hemolymph of many invertebrates, as an indicator of ALAN-related stress. Unlike total proteins, hemocyanins signal metabolic function and stress, so we expected them to increase in response to ALAN. We adapted spectrophotometry protocols to describe spatial variation in hemocyanins and total proteins in four populations of the talitroid amphipod Americorchestia longicornis. Then, a two-week experiment tested for changes in response to ALAN. Hemocyanin levels increased by 17 % and 40 % with respect to experimental controls after 7 and 14 d, respectively, and were higher than any measurements conducted in the field. These results suggest good prospects for hemocyanin as an indicator of ALAN effects.

Upwelling enhances seaweed nutrient quality, altering feeding behavior and growth rates in an intertidal sea urchin, Loxechinus albus.

Upwelling systems deliver nutrient-rich water into coastal ecosystems, influencing primary productivity and potentially altering seaweed-herbivore interactions. Upwelling bottom-up effects on distinct trophic levels are well-known. However, their influence on seaweed biomolecules and on algae-herbivore interactions and growth are less known. The aim of this study was threefold: i) to compare physical-chemical characteristics and nutrient levels in the water of upwelling (U) and downwelling (DU) zones, ii) to quantify their influence on the content of protein and carbohydrates in seaweed tissues of representative U and DU locations, and iii) to experimentally assess their effect on the feeding behavior and growth of a prominent intertidal herbivore, the sea urchin Loxechinus albus. Waters from U zones showed lower temperatures and pH, and higher phosphate concentrations than those from downwelling zones. Similarly, the tissue of seaweeds from a U location had significantly more proteins and carbohydrates than those from a DU location. The origin location of the sea urchins had a significant influence on consumption and growth rates: in general, those coming from a site with U conditions consumed and grew more than those coming from DU conditions. The quality of the algae was a significant factor on consumption rates, although in the case of preference trials, this factor interacted with sea urchin origin location. Our results show that the availability and quality of the food in upwelling zones has an influence on herbivore-seaweed direct interactions. However, these interactions and the growth of the sea urchins were also related to the coastal site and conditions from which the sea urchins came from. These results are relevant considering the expected impact of climate change on the world's oceans, and the importance of U zones as thermal (cold water) refuges for marine ectotherms.

Artificial light at night (ALAN) causes variable dose-responses in a sandy beach isopod.

Artificial Light at Night (ALAN) is expanding worldwide, and the study of its influence remains limited mainly to documenting impacts, overlooking the variation in key characteristics of the artificial light such as its intensity. The potential dose-response of fitness-related traits to different light intensities has not been assessed in sandy beach organisms. Hence, this study explored dose-responses to ALAN by exposing the intertidal sandy beach isopod Tylos spinulosus to a range of light intensities at night: 0 (control), 20, 40, 60, 80 and 100 lx. We quantified the response of this species at the molecular (RNA:DNA ratios), physiological (absorption efficiency) and organismal (growth rate) levels. Linear and non-linear regressions were used to explore the relationship between light intensity and the isopod response. The regressions showed that increasing light intensity caused an overall ~ threefold decline in RNA:DNA ratios and a ~ threefold increase in absorption efficiency, with strong dose-dependent effects. For both response variables, non-linear regressions also identified likely thresholds at 80 lx (RNA:DNA) and 40 lx (absorption efficiency). By contrast, isopod growth rates were unrelated (unaltered) by the increase in light intensity at night. We suggest that ALAN is detrimental for the condition of the isopods, likely by reducing the activity and feeding of these nocturnal organisms, and that the isopods compensate this by absorbing nutrients more efficiently in order to maintain growth levels.

Morphological, physiological and behavioral responses of an intertidal snail, Acanthina monodon (Pallas), to projected ocean acidification and cooling water conditions in upwelling ecosystems.

Ocean acidification (OA) is expected to rise towards the end of the 21st century altering the life history traits in marine organisms. Upwelling systems will not escape OA, but unlike other areas of the ocean, cooling effects are expected to intensify in these systems. Regardless, studies evaluating the combined effects of OA and cooling remain scarce. We addressed this gap using a mesocosm system, where we exposed juveniles of the intertidal muricid snail Acanthina monodon to current and projected pCO2 (500 vs. 1500 ppm) and temperature (15 vs. 10 °C) from the southeast Pacific upwelling system. After 9 weeks of experimental exposure to those conditions, we conducted three estimations of growth (wet weight, shell length and shell peristomal length), in addition to measuring calcification, metabolic and feeding rates and the ability of these organisms to return to the normal upright position after being overturned (self-righting). Growth, feeding and calcification rates increased in projected cooling conditions (10 °C) but were unaffected by pCO2 or the interaction between pCO2 and temperature. Instead, metabolic rates were driven by pCO2, but a significant interaction with temperature suggests that in cooler conditions, metabolic rates will increase when associated with high pCO2 levels. Snail self-righting times were not affected across treatments. These results suggest that colder temperatures projected for this area would drive this species growth, feeding and calcification, and consequently, some of its population biology and productivity. However, the snails may need to compensate for the increase in metabolic rates under the effects of ocean acidification. Although A. monodon ability to adjust to individual or combined stressors will likely account for some of the changes described here, our results point to a complex dynamic to take place in intertidal habitats associated with upwelling systems.

Ocean acidification alters anti-predator responses in a competitive dominant intertidal mussel.

Widespread intertidal mussels are exposed to a variety of natural and anthropogenic stressors. Even so, our understanding of the combined influence of stressors such as predation risk and ocean acidification (OA) on these species remains limited. This study examined the response of the purple mussel (Perumytilus purpuratus), a species distributed along Pacific southeastern rocky shores, to the effects of predation risk and OA. Using a laboratory 2 × 2 cross design, purple mussels were either devoid or exposed to predator cues from the muricid snail Acanthina monodon, while simultaneously exposing them to current (500 ppm) or projected OA conditions (1500 ppm). The response of purple mussels to these factors was assessed using growth, calcification, clearance, and metabolic rates, in addition to byssus production. After 60 d, the presence of predator cues reduced mussel growth in width and length, and in the latter case, OA enhanced this response making the effects of predator cues more severe. Calcification rates were driven by the interaction between the two stressors, whereas clearance rates increased only in response to OA, likely explaining some of the growth results. Mussel byssus production also increased with pCO2 but interacted with predation risk: in the absence of predator cues, byssus production increased with OA. These results suggest that projected levels of OA may alter and in some cases prevail over the natural response of purple mussels to predation risk. Considering the role played by this mussel as a dominant competitor and ecosystem engineer in rocky shores, these results have community-wide implications.

Artificial Light at Night (ALAN) negatively affects the settlement success of two prominent intertidal barnacles in the southeast Pacific.

Many coastal processes are regulated by day/night cycles and are expected to be altered by Artificial Light at Night (ALAN). The goal of this study was to assess the influence of ALAN on the settlement rates of intertidal barnacles. A newly designed settlement plate equipped with a small central LED light source was used to quantify settlement rates in presence/absence of ALAN conditions. "ALAN plates" as well as regular settlement plates were deployed in the mid rocky intertidal zone. Both ALAN and control plates collected early and late settlers of the barnacles Notochthamalus scabrosus and Jehlius cirratus. Early settlers (pre-metamorphosis cyprids) were not affected by ALAN. By contrast, the density of late settlers (post-metamorphosis spats) was significantly lower in ALAN than in control plates for both species, suggesting detrimental ALAN impacts on the settlement process. The new ALAN plates represent an attractive and alternative methodology to study ALAN effects.

Artificial light at night alters the activity and feeding behaviour of sandy beach amphipods and pose a threat to their ecological role in Atlantic Canada.

Artificial light at night (ALAN) is a growing source of stress for organisms and communities worldwide. These include species associated with sandy beaches, which consume and process stranded seaweeds (wrack) in these ecosystems. This study assessed the influence of ALAN on the activity and feeding behaviour of Americorchestia longicornis, a prominent talitrid amphipod living in sandy beaches of Prince Edward Island, Atlantic Canada. First, two parallel field surveys were conducted to document the natural daily cycle of activity of this species. Then, three related hypotheses were used to assess whether ALAN disrupts its locomotor activity, whether that disruption lasts over time, and whether it affects the feeding behaviour and growth of the amphipods. Tanks equipped with actographs recorded amphipod locomotor activity for ~7 days and then its potential recovery (after ALAN removal) for additional ~3 days. Separate tanks were used to compare amphipod food consumptions rates, absorption efficiency and growth rates under natural daylight / night (control) and altered conditions (ALAN). The results of these manipulations provide support to two of the three hypotheses proposed and indicate that ALAN was temporarily detrimental for (i.e. significantly reduced) the surface activity, consumption rates and absorption efficiency of the amphipods, whereas growth rates remained unaffected. The results also rejected the remaining hypothesis and suggest that the plasticity exhibited by these amphipods confer them the capacity to recover their natural rhythm of activity shortly after ALAN was removed from the system. Combined, these results suggest that ALAN has a strong, albeit temporary, influence upon the abundant populations of A. longicornis. Such influence has implications for the ecosystem role played by these amphipods as consumers and processors of the subsidy of stranded seaweeds entering these ecosystems.

Effects of artificial light at night and predator cues on foraging and predator avoidance in the keystone inshore mollusc Concholepas concholepas.

The growth of Artificial Light At Night (ALAN) is potentially having widespread effects on terrestrial and coastal habitats. In this study we addressed both the individual effects of ALAN, as well as its combined effect with predation risk on the behaviour of Concholepas concholepas, a fishery resource and a keystone species in the southeastern Pacific coast. We measured the influence of ALAN and predation risk on this mollusc's feeding rate, use of refuge for light and crawling out of water behaviour. These behavioural responses were studied using light intensities that mimicked levels that had been recorded in coastal habitat exposed to ALAN. Cues were from two species known to prey on C. concholepas during its early ontogeny: the crab Acanthocyclus hassleri and the seastar Heliaster helianthus. The feeding rates of C. concholepas were 3-4 times higher in darkness and in the absence of predator cues. In contrast, ALAN-exposed C. concholepas showed lower feeding activity and were more likely to be in a refuge than those exposed to control conditions. In the presence of olfactory predator cues, and regardless of light treatment, C. concholepas tended to crawl-out of the waterline. We provide evidence to support the hypothesis that exposure to either ALAN or predation risk can alter the feeding behaviour of C. concholepas. However, predator cue recognition in C. concholepas was not affected by ALAN in situations where ALAN and predator cues were both present: C. concholepas continued to forage when predation risk was low, i.e., in darkness and away from predator cues. Whilst this response means that ALAN may not lead to increased predation mortality in C. concholepas, it will reduce feeding activity in this naturally nocturnal species in the absence of dark refugia. Such results may have implications for the long-term health, productivity and sustainability of this keystone species.

Artificial light at night alters the settlement of acorn barnacles on a man-made habitat in Atlantic Canada.

Human growth has caused an unprecedented increase in artificial light at night (ALAN). In coastal habitats, many species rely on day/night cycles to regulate various aspects of their life history and these cycles can be altered by this stressor. This study assessed the influence of ALAN on the early (cyprid) and late (spat) settlement stages of the acorn barnacle Semibalanus balanoides, a species widely distributed in natural and man-made coastal habitats of the North Atlantic. A newly designed settlement plate, originally for studies in rocky intertidal habitats in the southeast Pacific, was adapted to measure settlement rates on man-made habitats -wharf seawalls- located in Atlantic Canada. Plates equipped with a small LED diode powered by an internal battery (ALAN plates) were used to quantify settlement rates in comparison to plates lacking a light source (controls). These plates were deployed for 6 d in the mid-intertidal levels, where adult barnacles were readily visible. ALAN and control plates collected large number of settlers and showed to be suitable for this type of man-made habitats. The number of early settlers (cyprids) did not differ between plates but the number of late settlers (spat) was significantly lower in ALAN plates than in controls. These results suggest that light pollution has little influence on the early stages of the acorn barnacle settlement but is clearly detrimental to its late stages. As barnacles dominate in many natural and man-made hard substrates, it is likely that ALAN also has indirect effects on community structure.

Community disruption in small biogenic habitats: A coastal invader overcomes habitat complexity to alter community structure.

Non-indigenous species are often identified as threats to native species and communities. Yet, the mechanisms that enable many of these invaders to thrive and alter their newly invaded habitats are still not fully understood. This applies to habitats such as widespread sedimentary shorelines characterized by the presence of scattered biogenic clumps of blue mussels (Mytilus edulis) structurally more complex than bare sediments. In Atlantic Canada, some of these shorelines are numerically dominated by native mud crabs (Dyspanopeus sayi) but have been gradually invaded by the European green crab (Carcinus maenas). This study describes between-habitat (mussel clump vs. bare sediment) differences in density and diversity of invertebrates. It also tests the impact of juvenile green crabs in comparison to native mud crabs using two approaches: First, measuring habitat-related differences in these crabs' feeding rates on a common prey (soft-shell clams, Mya arenaria). Second, measuring their influence on invertebrate communities associated with mussel clumps. The results show that mussel clumps hold higher invertebrate density and diversity than surrounding sedimentary bottoms. In the laboratory, the feeding rates of native mud crabs were dependent on the type of habitat (sand flat > mussel clump), whereas those of green crabs were significantly higher and unrelated to the habitat in which predation occurred. In field experiments, juvenile green crabs were also the only predators that changed community structure in the mussel clump habitat. These results indicate that green crabs can cause a significant impact on native species and communities. Moreover, they suggest that the ability of this species to overcome the refuge provided by complex biogenic habitats for prey may represent an unexplored mechanism to explain this invader's expansion here and elsewhere.

The positive effect of coexisting ecosystem engineers: a unique seaweed-mussel association provides refuge for native mud crabs against a non-indigenous predator.

In marine sedimentary bottoms, mussels and macroalgae have long been recognized as important autogenic engineers that create habitat and modify abiotic conditions. The structural complexity added by bivalves and macroalgae may also mediate intraguild predation amongst marine decapod crustaceans. While spatial distributions of these ecosystem engineers frequently overlap, there is limited understanding of compounded effects when more than one engineer is present. Here we demonstrate that the coexistence of two ecosystem engineers may create habitat valuable for the survival of a small native species, the Atlantic mud crab (Panopeus herbstii), in the presence of the invasive green crab (Carcinus maenas). Using laboratory and field habitat mimics, we measured mud crab survival rates as a proxy for refuge quality. We compared the refuge provided by a unique association between shells of blue mussels (Mytilus edulis) and the giant strain of Irish moss (Chondrus crispus) to that provided by bare substrate, and by each engineer alone. These experiments revealed that the association of giant Irish moss with blue mussel shells positively and non-additively increased mud crab survival compared to the other less complex habitat mimics. In contrast, parallel experiments revealed that high habitat complexity was less important for young green crabs to survive predation from large conspecifics. These results suggest that the impact of ecosystem engineers on trophic dynamics should be considered in a broader, whole-community context encompassing multiple habitat-forming species present.