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Simulated diurnal pH fluctuations radically increase variance in— but not the mean of—growth in the barnacle Balanus improvisus

Journal article
Authors Louise Eriander
Anna-Lisa Wrange
Jonathan N. Havenhand
Published in ICES Journal of Marine Science
Volume 73
Issue 3
Pages 596-603
ISSN 1054-3139
Publication year 2016
Published at Department of marine sciences
Pages 596-603
Language en
Keywords crustacea, effect size, natural fluctuations, ocean acidification, penetrometry, response ratio
Subject categories Ecology


Shallow coastal waters are characterized by substantial diurnal fluctuations in pH, especially in nearshore environments. The biological effects of ocean acidification in combination with these natural fluctuations have received relatively little attention to date. We exposed multiple batches (≈ different genotypes) of newly settled barnacles, Balanus improvisus, to constant pH under “control” (pH ¼ 8.1) or “stable acidified” (pH ¼ 7.7) conditions, as well as a treatment that simulated the maximum diurnal pH fluctuations seen in the nearshore habitats where this barnacle lives (+0.2 pH units), superimposed on the stable acidified treatment (“fluctuating acidified”; 7.5 ≤ pH ≤ 7.9). We found that fluctuating acidification had no effect on mean response in growth and shell mineralogy, but caused an #20-fold increase in variance of responses, compared with stable acidification. In contrast to these results, we found no effect of fluctuating acidification on variances of response ratios for barnacle survival and shell strength. Similarly, mean survival did not vary significantly with pH. However, we observed a strong negative effect of stable and fluctuating acidification on mean shell strength. Our finding that barnacles respond differently to fluctuating pHthan to stable lowpH indicate the importance of including fluctuating acidification treatments when studying species that live in variable environments. Importantly, because phenotypic variance is the raw material for natural selection, and thus lays at the heart of evolutionary responses to environmental variability and change, our findings also highlight the need to study changes in variance of—as well as mean—responses to changing ocean climates.

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