Temperate Rocky Shores Research Articles

Sea anemone microhabitats enhance the diversity and biomass of mobile invertebrates on temperate rocky shores

Sea anemones of the genus Anthopleura are abundant in rocky intertidal communities of the eastern Pacific, but their role as habitat and facilitators of biodiversity in this system has been overlooked. We provide evidence for a previously undescribed mechanism of facilitation associated with these cnidarians on temperate rocky shores. Sea anemones in the genus Anthopleura slowly release stored water at low tide, which helps them maintain ideal temperature conditions via evaporative cooling. This behavior results in leakage of water to surrounding intertidal habitats, producing unique microhabitats in the upper intertidal zone. We hypothesized that mobile invertebrate communities would use this habitat during low tides to ameliorate abiotic stress. In an in situ experiment spanning 3 sites along the coast of California, USA, it was found that maximum temperatures and relative desiccation rates were lower in sea anemone-associated microhabitats as compared to adjacent rock habitats. Observational surveys revealed higher richness and biomass of mobile invertebrates in sea anemone microhabitats, and these differences were supported by manipulative experiments that included both adding and removing sea anemones. Our results support a novel community-wide facilitation driven by an adaptation of intertidal sea anemones. The effects of sea anemone microhabitat on upper shore biodiversity were associated with site and sea anemone species and may suggest a facilitative role of a temperate cnidarian-algal mutualism in mitigating stress and enhancing biodiversity.

Could the annual Saccorhiza polyschides replace a sympatric perennial kelp (Laminaria ochroleuca) when it comes to supporting the holdfast-associated fauna?

Kelp forests dominate temperate rocky shores worldwide but are declining globally with consequences for organisms that depend on them. In NW Iberia, the golden kelp (Laminaria ochroleuca) commonly occurs alongside a fast-growing annual that, unlike the golden kelp, does not seem to have receded in recent times (Saccorhiza polyschides). Here, we assessed whether the bulbous holdfast of S. polyschides could replace the intricate holdfast of L. ochroleuca as epifaunal habitat provider. Richness, diversity and total abundance of epifauna was similar in both seaweeds, while colonial/encrusting fauna was more abundant in L. ochroleuca. More importantly, each host supported a distinctive assemblage structure, indicating that S. polyschides seems an unsuitable replacement for L. ochroleuca as habitat provider for holdfast epifauna. Therefore, while S. polyschides may contribute substantially to the kelp forest canopy in some seasons, a regional decline of L. ochroleuca will likely alter the patterns of biodiversity within kelp stands.

Seasonal ecophysiology of Fucus vesiculosus (Phaeophyceae) in the Northern Baltic Sea

ABSTRACT The brown macroalga Fucus vesiculosus is a foundation species in temperate rocky shores, subjected to seasonally fluctuating environmental conditions. To obtain a more complete picture of the seasonality of F. vesiculosus ecophysiology in the northern Baltic Sea, in situ photochemistry, elemental ratios and chlorophyll a and c content of the alga were investigated in field campaigns conducted in different months throughout the year during 2017. Carbon, nitrogen, carbon to nitrogen ratio and chlorophyll a and c content of the alga varied substantially throughout the year, with highest carbon content observed in summer, and highest nitrogen content in winter. C:N ratio in F. vesiculosus apical tissue ranged from 8.6 in February to 48.3 in July. Chlorophyll a and c content followed inversely the seasonal patterns of ambient irradiance. High chlorophyll a and c content in winter was associated with higher maximum photosynthetic efficiency of energy conversion (Fv/Fm), but not with efficiency of photosynthetic energy conversion under light limitation (α). Electron transport rate correlated strongly with seawater temperature, and the highest electron transport rates were observed in summer and correlated with highest internal carbon content of the alga. Redundancy analysis conducted on measured environmental variables against physiological responses identified day of year, temperature and macronutrients in seawater as the most important variables driving the observed seasonal patterns in F. vesiculosus ecophysiology. The results suggest elevated temperatures may increase Fucus growth and photosynthesis rates in the study area. Highlights Two distinct physiological states of Fucus vesiculosus were identified. The most important variables driving responses were season, temperature and macronutrients. Fucus ecophysiology shows substantial seasonality in the northern Baltic Sea.

The surface bacterial community of an Australian kelp shows cross-continental variation and relative stability within regions.

Epiphytic microbial communities often have a close relationship with their eukaryotic host, assisting with defence, health, disease prevention and nutrient transfer. Shifts in the structure of microbial communities could therefore have negative effects on the individual host and indirectly impact the surrounding ecosystem, particularly for major habitat-forming hosts, such as kelps in temperate rocky shores. Thus, an understanding of the structure and dynamics of host-associated microbial communities is essential for monitoring and assessing ecosystem changes. Here, samples were taken from the ecologically important kelp, Ecklonia radiata, over a 17-month period, from six different sites in two distinct geographic regions (East and West coasts of Australia), separated by ∼3,300 kms, to understand variation in the kelp bacterial community and its potential environmental drivers. Differences were observed between kelp bacterial communities between the largely disconnected geographical regions. In contrast, within each region and over time the bacterial communities were considerably more stable, despite substantial seasonal changes in environmental conditions.

Modelling the annual primary production of an intertidal brown algal community based on in situ measurements

Intertidal communities dominated by canopy-forming brown algae (Phaeophyceae) usually prove to be highly productive systems, based on short-term measurements. However, long-term metabolism (primary production and respiration) is sensitive to several factors acting on different time scales (e.g. tidal cycle, seasonality), making its assessment challenging. Here, we used mathematical modelling to investigate the metabolism of aFucus serratus-dominated community on daily and annual time scales. This widespread community, which is usually characteristic of the low mid-intertidal level of temperate rocky shores, is submerged for approximately 83% of the time at our study site (Brittany, France). The model incorporated a large spectrum of physiological (e.g. estimates of primary production versus irradiance parameters) and environmental (e.g. temperature, incident irradiance, depth of the water column) parameters measuredin situ. The model simulation predicted that net community primary production (NCP) peaks at 8.0 gC m-2d-1in late spring, when environmental conditions are most favorable. In contrast, during fall and winter, respiration frequently overcomes primary production, making the system heterotrophic on a daily basis. For the year as a whole, simulation predicted that the community acts as an autotrophic system, with its annual gross primary production amounting to ca. 1301 gC m-2and annual respiration to ca. 899 gC m-2. According to this simulation, the annual NCP of our intertidalF. serratuscommunity therefore reached 402 gC m-2, which is comparable to subtidal communities dominated by canopy-forming brown algae. Although theF. serratuscommunity lives mostly underwater, it was particularly autotrophic during aerial exposures.

First comparative assessment of the reproductive cycle of three species of Chiton on a temperate rocky shore of the southeastern Pacific

AbstractThe present study provides the first assessment of the reproductive cycle of three common and widespread species of the genus Chiton, sympatric in rocky shores of Coquimbo, northern Chile (~30°S): C. granosus, C. cumingsii, and C. magnificus. From specimens seasonally sampled during a 2‐year period, a histological study was performed in order to describe the gonad structure and stages of gonadal development for each species. The morphological structure of the gonad was similar in the three species, with sexual cells developing from the gonad wall towards its center. The three species were gonochoric and iteroparous, with asynchronous gamete maturation at population level; in all seasons, sexual maturity was evident in small to large individuals of each species (≥25 mm in total length for C. cumingsii and C. granosus, and ≥47 mm for C. magnificus).

Summer and Winter Marine Heatwaves Favor an Invasive Over Native Seaweeds.

Marine heatwaves (MHWs) are emerging as forceful agents of ecosystem change and are increasing in frequency, duration, and intensity with climate change. During MHWs, physiological thresholds of native species may be exceeded while the performance of invasive species with warm affinities may be enhanced. As a consequence, MHWs could significantly alter an ecosystem's invasive dynamics, but such interactions are poorly understood. Following a 10-d acclimation period, we investigated the physiological resistance and resilience of an intertidal rock pool assemblage invaded by the seaweed Sargassum muticum to realistic 14-d marine heatwave scenarios (+1.5°C, +2.0°C, +3.5°C) followed by a 14-d recovery period. We conducted mesocosm experiments in both summer and winter to investigate temporal variability of MHWs. MHW treatments had clear negative impacts on native seaweeds (Fucus serratus and Chondrus crispus) while enhancing the performance of S.muticum. This pattern was consistent across season indicating that acclimation to cooler ambient temperatures results in winter MHWs having significant impacts on native species. As climate warming advances, this may ultimately lead to changes in competitive interactions and potentially exclusion of native species, while invasive species may proliferate and become more conspicuous within temperate rocky shore environments.

Disappearing Blue Mussels – Can Mesopredators Be Blamed?

Abstract Despite many theories, the recent evident decreases in blue mussel (Mytilus edulis) abundance in southern Norway and western Sweden (eastern North Sea) have not yet been explained. To test the possible role of increased predation, an ongoing mesocosm experiment exploring general effects of two mesopredators on the structure of littoral macroalgal and macrofaunal communities was used. These mesopredators were the green crab (Carcinus maenas) and the goldsinny wrasse (Ctenolabrus rupestris) which were distributed in a crossed manner to 12 large mesocosms containing diverse rocky shore communities. For the purposes of this study, boulders covered with recently recruited blue mussels and barnacles (Balanus improvisus) from a natural shore were brought in to the mesocosms during two seasons (August and October), and the coverage of the animals (just blue mussels in summer, both mussels and barnacles in autumn) was registered repeatedly over 24 hours. The mussels were rapidly consumed by crabs and wrasses, whereas high survival was recorded on boulders in the controls without predators. The barnacles were only eaten by the crabs and not before most of the mussels had been consumed. As both the green crab and the goldsinny wrasse have been reported to increase in abundance, probably related to overfishing of top predators, the resulting higher predation pressure on especially small blue mussels (recruits) may contribute to the mussel decline along these temperate rocky shores.

Characterizing biogeochemical fluctuations in a world of extremes: A synthesis for temperate intertidal habitats in the face of global change.

Coastal and intertidal habitats are at the forefront of anthropogenic influence and environmental change. The species occupying these habitats are adapted to a world of extremes, which may render them robust to the changing climate or more vulnerable if they are at their physiological limits. We characterized the diurnal, seasonal and interannual patterns of flux in biogeochemistry across an intertidal gradient on a temperate sandstone platform in eastern Australia over 6years (2009-2015) and present a synthesis of our current understanding of this habitat in context with global change. We used rock pools as natural mesocosms to determine biogeochemistry dynamics and patterns of eco-stress experienced by resident biota. In situ measurements and discrete water samples were collected night and day during neap low tide events to capture diurnal biogeochemistry cycles. Calculation of pHT using total alkalinity (TA) and dissolved inorganic carbon (DIC) revealed that the mid-intertidal habitat exhibited the greatest flux over the years (pHT 7.52-8.87), and over a single tidal cycle (1.11 pHT units), while the low-intertidal (pHT 7.82-8.30) and subtidal (pHT 7.87-8.30) were less variable. Temperature flux was also greatest in the mid-intertidal (8.0-34.5°C) and over a single tidal event (14°C range), as typical of temperate rocky shores. Mean TA and DIC increased at night and decreased during the day, with the most extreme conditions measured in the mid-intertidal owing to prolonged emersion periods. Temporal sampling revealed that net ecosystem calcification and production were highest during the day and lowest at night, particularly in the mid-intertidal. Characterization of biogeochemical fluctuations in a world of extremes demonstrates the variable conditions that intertidal biota routinely experience and highlight potential microhabitat-specific vulnerabilities and climate change refugia.

Top-down release of mesopredatory fish is a weaker structuring driver of temperate rocky shore communities than bottom-up nutrient enrichment

Ecological impacts of top-down trophic cascades in combination with bottom-up nutrient enrichment are increasingly being reported. Such effects may be triggered by decline in the abundance and size of piscivore fish leading to a release of smaller-sized mesopredatory fish that are capable of reducing mesograzers and their buffering herbivorous effects, thus intensifying eutrophication symptoms. Hitherto, such mesopredator release has not been studied in controlled manner in macroalgal-dominated rocky shore communities. This study utilised twelve littoral mesocosms in southeastern Norway to investigate the impact of increased nutrient levels and increased abundance of the mesopredatory goldsinny wrasse (Ctenolabrus rupestris) over 6 months, from spring (April) to autumn (October). The study mainly revealed typical eutrophication responses in both July and in October with nutrient enrichment leading to more ephemeral algae and less perennial Fucacean and red macroalgae. Significant responses to the addition of mesopredatory fish alone or with nutrients in combination were clearly fewer in July and almost non-existent in October. Mesopredatory fish reduced the number of some mesograzers, but not others, and some herbivores even benefitted. Few joint effects occurred between fish and nutrients and the anticipated negative consequences for perennial macrophytes were largely absent; the possible reasons for these observations are discussed in depth. Curiously, the presence of mesopredatory fish seems to favour some canopy-forming macrophytes. This last finding warrants further investigation as the labrid fishery in coastal waters of southern Norway and western Sweden is increasing and a too heavy reduction of mesopredators may have its own unforeseen ecosystem implications.

Gene regulatory response to hyposalinity in the brown seaweed Fucus vesiculosus

BackgroundRockweeds are among the most important foundation species of temperate rocky littoral shores. In the Baltic Sea, the rockweed Fucus vesiculosus is distributed along a decreasing salinity gradient from the North Atlantic entrance to the low-salinity regions in the north-eastern margins, thus, demonstrating a remarkable tolerance to hyposalinity. The underlying mechanisms for this tolerance are still poorly understood. Here, we exposed F. vesiculosus from two range-margin populations to the hyposaline (2.5 PSU - practical salinity unit) conditions that are projected to occur in the region by the end of this century as a result of climate change. We used transcriptome analysis (RNA-seq) to determine the gene expression patterns associated with hyposalinity acclimation, and examined the variation in these patterns between the sampled populations.ResultsHyposalinity induced different responses in the two populations: in one, only 26 genes were differentially expressed between salinity treatments, while the other population demonstrated up- or downregulation in 3072 genes. In the latter population, the projected future hyposalinity induced an acute response in terms of antioxidant production. Genes associated with membrane composition and structure were also heavily involved, with the upregulation of fatty acid and actin production, and the downregulation of ion channels and alginate pathways. Changes in gene expression patterns clearly indicated an inhibition of the photosynthetic machinery, with a consequent downregulation of carbohydrate production. Simultaneously, energy consumption increased, as revealed by the upregulation of genes associated with respiration and ATP synthesis. Overall, the genes that demonstrated the largest increase in expression were ribosomal proteins involved in translation pathways. The fixation rate of SNP:s was higher within genes responding to hyposalinity than elsewhere in the transcriptome.ConclusionsThe high fixation rate in the genes coding for salinity acclimation mechanisms implies strong selection for them. The among-population differentiation that we observed in the transcriptomic response to hyposalinity stress suggests that populations of F. vesiculosus may differ in their tolerance to future desalination, possibly as a result of local adaptation to salinity conditions within the Baltic Sea. These results emphasise the importance of considering interspecific genetic variation when evaluating the consequences of environmental change.

Feeding behaviour of Patella caerulea L. and P. rustica L. under spring and neap simulated tides. An innovative approach for quick quantification of grazing activity

Limpets are one of the most important herbivorous grazers of temperate rocky shores, promoting diversity and influencing the structure and functioning of intertidal communities. In the current context of global change, harvesting pressures on accessible intertidal and habitat modifications caused by coastal urbanization are threatening these organisms in many coastal areas. In this regard, the improvement of ex situ maintenance techniques, combined with the understanding of limpets’ behavioural traits, would help to improve the effectiveness of conservation strategies and mitigation efforts. In this study, Patella caerulea and Patella rustica grazing activity was quantified under a scenario of simulated tides. After leaving limpets to graze 10 days under two tidal amplitude regimes: Spring and Neap, radula mark coverage was obtained from the surface of wax discs with a quick image-processing methodology, using the dry-brush technique and Adobe© Photoshop and ImageJ software. Feeding activity, quantified as radula mark coverage, was higher for P. caerulea than P. rustica and, within P. caerulea, for spring over neap tides. Therefore, tidal range has to be taken into consideration when limpet specimens are maintained in captive conditions.

Quantification of the foraging activity of Patella caerulea and P. rustica under simulated tides using photographic analysis of radula scratches

Quantification of the foraging activity of Patella caerulea and P. rustica under simulated tides using photographic analysis of radula scratches

The multiple roles of β-diversity help untangle community assembly processes affecting recovery of temperate rocky shores.

Metacommunity theory highlights the potential of β-diversity as a useful link to empirical research, especially in diverse systems where species exhibit a range of stage-dependent dispersal characteristics. To investigate the importance of different components and scales of β-diversity in community assembly, we conducted a large-scale disturbance experiment and compared relative recovery across multiple sites and among plots within sites on the rocky shore. Six sites were spread along 80 km of coastline and, at each site, five plots were established, matching disturbed and undisturbed quadrats. Recovery was not complete at any of the sites after 1 year for either epibenthos (mostly composed of macroalgae and, locally, mussels) or infauna. Significant differences in recovery among sites were observed for epibenthos but not for infauna, suggesting that different community assembly processes were operating. This was supported by epibenthos in the recovering plots having higher species turnover than in undisturbed sediment, and recovery well predicted by local diversity, while infaunal recovery was strongly influenced by the epibenthic community's habitat complexity. However, infaunal community recovery did not simply track formation of habitat by recovering epibenthos, but appeared to be overlain by within-site and among-site aspects of infaunal β-diversity. These results suggest that documenting changes in the large plants and animals alone will be a poor surrogate for rocky shore community assembly processes. No role for ecological connectivity (negative effect of among-site β-diversity) in driving recovery was observed, suggesting a low risk of effects from multiple disturbances propagating along the coast, but a limited resilience at the site scale to large-scale disturbances such as landslides or oil spills.

First Detailed Record of Symbiosis Between a Sea Anemone and Homoscleromorph Sponge, With a Description of Tempuractis rinkai gen. et sp. nov. (Cnidaria: Anthozoa: Actiniaria: Edwardsiidae).

A new species in a new genus of sea anemone, Tempuractis rinkai gen. et sp. nov., was discovered at several localities along the temperate rocky shores of Japan. The new species is approximately 4 mm in length and has been assigned to family Edwardsiidae, because it has eight macrocnemes, lacks sphincter and basal muscles, and possesses rounded aboral end. The sea anemone, however, also has a peculiar body shape unlike that of any other known taxa. This new species resembles some genera, especially Drillactis and Nematostella, in smooth column surface without nemathybomes or tenaculi, but is distinguishable from them by several morphological features: the presence of holotrichs and absence of nematosomes. Furthermore, this edwardsiid species exhibits a peculiar symbiotic ecology with sponges. Therefore, a new genus, Tempuractis, is proposed for this species. In the field, T. rinkai sp. nov. was always found living inside homosclerophorid sponge of the genus Oscarella, which suggests a possible obligate symbiosis between Porifera and Actiniaria. The benefit of this symbiosis is discussed on the basis of observations of live specimens, both in the aquarium and field. This is the first report of symbiosis between a sea anemone and a homoscleromorph sponge.

Can secondary species maintain a primary role? Consistent inter-regional effects of understory algae on diversity

Loss of habitat is a global threat to biodiversity. Habitat-forming species in particular are shifting their distribution at local and regional scales, changing habitat aspect globally. In temperate intertidal rocky shores, this poleward shift of canopy-forming seaweeds is leading to a shift to mat- or turf-dominated communities. These changes and their consequences are poorly understood and can vary globally. Most studies, however, have focused on local and regional processes. Using rocky low intertidal algal-dominated systems, this study aims to understand the independent role of a rich understory vegetation layer formed by cespitose algae as community drivers. Moreover, the study aimed to understand whether the observed patterns can be consistent over spatial regions, thus three distinct regional zones of the North Atlantic (Eastern shores of Nova Scotia, Canada; the Poitou-Charentes coast, France; the Eastern shores of Northern Ireland, UK) were sampled as part of this study. From surveys, results showed that in the intertidal zone with low desiccation potential, canopies generally do not drive understory cespitose species richness or distribution. Cespitose algae on the other hand positively influenced richness in all regions. Community composition was also influenced by cespitose algal species, which increased the number of mobile epifauna species, while decreasing the number of encrusting epifauna species. This has wide implications: (i) understory species that are often overlooked may play a primary role as habitat formers; (ii) understory species may help maintain biodiversity in a changing world where primary species distribution is shifting.

Novel ssDNA Viruses Detected in the Virome of Bleached, Habitat-Forming Kelp Ecklonia radiata

Kelp forests provide essential habitats for organisms in temperate rocky shores. Loss of kelp forests has occurred over large areas in a number of temperate regions, including in Australia, where the dominant kelp Ecklonia radiata has been lost from substantial areas of the shoreline. Loss of E. radiata has been associated with environmental stressors, including increased temperature and anthropogenic contaminants, as well as biological factors such as herbivory. Disease may also play a role, but there is little information on the role of disease in the loss of kelp from coastal ecosystems or on the potential role of pathogenic microorganisms, such as viruses. E. radiata across much of its distribution in Australia can develop a “bleached” phenotype, which may be a disease. To investigate whether the phenotype was associated with a potential viral agent, we shotgun sequenced viral particles that were isolated from kelp with normal (healthy) and bleached phenotypes. Each virome consisted of ~380,000 reads, of which ~25 % were similar to known viruses. All samples were dominated by bacteriophages, but novel ssDNA virus sequences were detected that were almost exclusively in viromes from the bleached kelp phenotype. These ssDNA viruses are covered by eleven contigs that contained complete capsids and characteristic rep genes that were 30 – 60 % similar to those of circular, Rep-encoding ssDNA viruses (CRESS-DNA viruses). CRESS-DNA viruses have not previously been described from macroalgae, and the rep genes were similar to CRESS-DNA viruses from marine water samples, snails, crabs, anemones, but also dragonflies. This raises the interesting possibility that the kelp could be a vector of the CRESS-DNA viruses to other organisms that are associated with the bleached state.

Environmental factors influencing the settlement of Diplodus spp. juveniles in temperate marine rocky shores

Environmental factors influencing the settlement of Diplodus spp. juveniles in temperate marine rocky shores

Artificial light at night alters trophic interactions of intertidal invertebrates.

Despite being globally widespread in coastal regions, the impacts of light pollution on intertidal ecosystems has received little attention. Intertidal species exhibit many night-time-dependent ecological strategies, including feeding, reproduction, orientation and predator avoidance, which are likely negatively affected by shifting light regimes, as has been observed in terrestrial and aquatic taxa. Coastal lighting may shape intertidal communities through its influence on the nocturnal foraging activity of dogwhelks (Nucella lapillus), a widespread predatory mollusc that structures biodiversity in temperate rocky shores. In the laboratory, we investigated whether the basal and foraging activity of this predator was affected by exposure to night-time lighting both in the presence and absence of olfactory predator cues (Carcinus maenas, common shore crab). Assessments of dogwhelks' behavioural responses to night-time white LED lighting were performed on individuals that had been acclimated to night-time white LED lighting conditions for 16days and individuals that had not previously been exposed to artificial light at night. Dogwhelks acclimated to night-time lighting exhibited natural refuge-seeking behaviour less often compared to control animals, but were more likely to respond to and handle prey irrespective of whether olfactory predator cues were present. These responses suggest night-time lighting likely increased the energetic demand of dogwhelks through stress, encouraging foraging whenever food was available, regardless of potential danger. Contrastingly, whelks not acclimated under night-time lighting were more likely to respond to the presence of prey under artificial light at night when olfactory predator cues were present, indicating an opportunistic shift towards the use of visual instead of olfactory cues in risk evaluation. These results demonstrate that artificial night-time lighting influences the behaviour of intertidal fauna such that the balance of interspecific interactions involved in community structuring may be affected.

Long-term exposure to elevated pCO2 more than warming modifies early-life shell growth in a temperate gastropod

Co-occurring global change drivers, such as ocean warming and acidification, can have large impacts on the behaviour, physiology, and health of marine organisms. However, whilst early-life stages are thought to be most sensitive to these impacts, little is known about the individual level processes by which such impacts take place. Here, using mesocosm experiments simulating ocean warming (OW) and ocean acidification (OA) conditions expected for the NE Atlantic region by 2100 using a variety of treatments of elevated pCO2 and temperature. We investigated their impacts on bio-mineralization, microstructure, and ontogeny of Nucella lapillus (L.) juveniles, a common gastropod predator that exerts important top-down controls on biodiversity patterns in temperate rocky shores. The shell of juveniles hatched in mesocosms during a 14 month long experiment were analysed using micro-CT scanning, 3D geometric morphometrics, and scanning-electron microscopy. Elevated temperature and age determined shell density, length, width, thickness, elemental chemistry, shape, and shell surface damages. However, co-occurring elevated pCO2 modified the impacts of elevated temperature, in line with expected changes in carbonate chemistry driven by temperature. Young N. lapillus from acidified treatments had weaker shells and were therefore expected to be more vulnerable to predation and environmental pressures such as wave action. However, in some instances, the effects of both higher CO2 content and elevated temperature appeared to have reversed as the individuals aged. This study suggests that compensatory development may therefore occur, and that expected increases in juvenile mortality under OA and OW may be counteracted, to some degree, by high plasticity in shell formation in this species. This feature may prove advantageous for N. lapillus community dynamics in near-future conditions.