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Balanomorph (acorn) barnacles are found throughout the world’s coastal oceans, and their success is dependent on a hard, mineralized, outer shell. Although macro-scale morphology of barnacle shells has been studied extensively, relatively little is known about shell properties at the micron-scale and if such properties vary among species. We assessed shell structure, mechanics, and composition in seven species of balanomorph barnacles from five genera. Three species, Amphibalanus amphitrite, Amphibalanus improvisus, and Austrominius modestus, were laboratory-reared, enabling direct comparison of shell properties of barnacles grown under the same conditions for the same duration. Four other species, Semibalanus balanoides, Amphibalanus eburneus, Chthamalus stellatus, and Tetraclita rubescens, were field-collected. At the macro- and meso-scales, shell properties varied markedly among species, with differences in the number of shell plates, the presence of canals within the plates, mineralization of the base, and shell plate thickness. At the micron-scale, however, structure was remarkably similar among species. Plates of all species were constructed of irregular micron-scale crystallites, with a broad range of crystallite dimensions observed within the same shell. Similarly, micromechanical properties did not vary among species, regardless of testing orientation. Calcium carbonate was identified as calcite in all species assessed with no other mineral phases present, and calcium content did not vary among species. Hence, despite variation in the overall macro- and meso-scale morphology of barnacles, all appear to be built using the same, evolutionarily conserved, mineralization pathway. 

Barnacles are ancient arthropods that, as adults, are surrounded by a hard, mineralized, outer shell that the organism produces for protection. While extensive research has been conducted on the glue-like cement that barnacles use to adhere to surfaces, less is known about the barnacle exoskeleton, especially the process by which the barnacle exoskeleton is formed. Here, we present data exploring the changes that occur as the barnacle cyprid undergoes metamorphosis to become a sessile juvenile with a mineralized exoskeleton. Scanning electron microscope data show dramatic morphological changes in the barnacle exoskeleton following metamorphosis. Energy-dispersive X-ray spectroscopy indicates a small amount of calcium (8%) 1 h post-metamorphosis that steadily increases to 28% by 2 days following metamorphosis. Raman spectroscopy indicates calcite in the exoskeleton of a barnacle 2 days following metamorphosis and no detectable calcium carbonate in exoskeletons up to 3 h post-metamorphosis. Confocal microscopy indicates during this 2 day period, barnacle base plate area and height increases rapidly (0.001 mm 2 h −1 and 0.30 µm h −1 , respectively). These results provide critical information into the early life stages of the barnacle, which will be important for developing an understanding of how ocean acidification might impact the calcification process of the barnacle exoskeleton. 

ABSTRACT Ocean acidification can affect the ability of calcifying organisms to build and maintain mineralized tissue. In decapod crustaceans, the exoskeleton is a multilayered structure composed of chitin, protein and mineral, predominately magnesian calcite or amorphous calcium carbonate (ACC). We investigated the effects of acidification on the exoskeleton of mature (post-terminal-molt) female southern Tanner crabs, Chionoecetes bairdi. Crabs were exposed to one of three pH levels – 8.1, 7.8 or 7.5 – for 2 years. Reduced pH led to a suite of body region-specific effects on the exoskeleton. Microhardness of the claw was 38% lower in crabs at pH 7.5 compared with those at pH 8.1, but carapace microhardness was unaffected by pH. In contrast, reduced pH altered elemental content in the carapace (reduced calcium, increased magnesium), but not the claw. Diminished structural integrity and thinning of the exoskeleton were observed at reduced pH in both body regions; internal erosion of the carapace was present in most crabs at pH 7.5, and the claws of these crabs showed substantial external erosion, with tooth-like denticles nearly or completely worn away. Using infrared spectroscopy, we observed a shift in the phase of calcium carbonate present in the carapace of pH 7.5 crabs: a mix of ACC and calcite was found in the carapace of crabs at pH 8.1, whereas the bulk of calcium carbonate had transformed to calcite in pH 7.5 crabs. With limited capacity for repair, the exoskeleton of long-lived crabs that undergo a terminal molt, such as C. bairdi, may be especially susceptible to ocean acidification. 

Abstract Crustaceans comprise an ecologically and morphologically diverse taxonomic group. They are typically considered resilient to many environmental perturbations found in marine and coastal environments, due to effective physiological regulation of ions and hemolymph pH, and a robust exoskeleton. Ocean acidification can affect the ability of marine calcifying organisms to build and maintain mineralized tissue and poses a threat for all marine calcifying taxa. Currently, there is no consensus on how ocean acidification will alter the ecologically relevant exoskeletal properties of crustaceans. Here, we present a systematic review and meta‐analysis on the effects of ocean acidification on the crustacean exoskeleton, assessing both exoskeletal ion content (calcium and magnesium) and functional properties (biomechanical resistance and cuticle thickness). Our results suggest that the effect of ocean acidification on crustacean exoskeletal properties varies based upon seawaterpCO₂and species identity, with significant levels of heterogeneity for all analyses. Calcium and magnesium content was significantly lower in animals held atpCO₂ levels of 1500–1999 µatm as compared with those under ambientpCO₂. At lowerpCO₂ levels, however, statistically significant relationships between changes in calcium and magnesium content within the same experiment were observed as follows: a negative relationship between calcium and magnesium content atpCO₂of 500–999 µatm and a positive relationship at 1000–1499 µatm. Exoskeleton biomechanics, such as resistance to deformation (microhardness) and shell strength, also significantly decreased underpCO₂regimes of 500–999 µatm and 1500–1999 µatm, indicating functional exoskeletal change coincident with decreases in calcification. Overall, these results suggest that the crustacean exoskeleton can be susceptible to ocean acidification at the biomechanical level, potentially predicated by changes in ion content, when exposed to high influxes of CO₂. Future studies need to accommodate the high variability of crustacean responses to ocean acidification, and ecologically relevant ranges ofpCO₂conditions, when designing experiments with conservation‐level endpoints.