RESEARCH ARTICLE

CLIMATE SCIENCE 2016 © The Authors, some rights reserved;

exclusive licensee American Association for

Ocean acidification causes structural deformities the Advancement of Science. Distributed

under a Creative Commons Attribution

in juvenile coral skeletons NonCommercial License 4.0 (CC BY-NC).

10.1126/sciadv.1501130

Taryn Foster,1,2,3* James L. Falter,1,2,3 Malcolm T. McCulloch,1,2,3 Peta L. Clode2,4

Rising atmospheric CO2 is causing the oceans to both warm and acidify, which could reduce the calcification rates of
corals globally. Successful coral recruitment and high rates of juvenile calcification are critical to the replenishment
and ultimate viability of coral reef ecosystems. Although elevated PCO2 (partial pressure of CO2) has been shown to
reduce the skeletal weight of coral recruits, the structural changes caused by acidification during initial skeletal
deposition are unknown. We show, using high-resolution three-dimensional x-ray microscopy, that ocean acidifi-
cation (PCO2 ~900 matm, pH ~7.7) not only causes reduced overall mineral deposition but also a deformed and
porous skeletal structure in newly settled coral recruits. In contrast, elevated temperature (+3°C) had little effect

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on skeletal formation except to partially mitigate the effects of elevated PCO2. The striking structural deformities
we observed show that new recruits are at significant risk, being unable to effectively build their skeletons in
the PCO2 conditions predicted to occur for open ocean surface waters under a “business-as-usual” emissions
scenario [RCP (representative concentration pathway) 8.5] by the year 2100.

INTRODUCTION under high PCO2 conditions. However, to date, such research has relied
Atmospheric CO2 is set to rise to >900 parts per million (ppm) by the solely on bulk measurements of skeletal mass to assess these impacts.
end of the century under a “business-as-usual” scenario [RCP (repre- There have been no analyses of specific structural changes in the ju-
sentative concentration pathway) 8.5], with corresponding elevations in venile skeleton under acidification or warming.
both oceanic temperature (+~3°C) and partial pressure of CO2 (PCO2) Here, we used high-resolution three-dimensional (3D) x-ray mi-
(+~500 matm) (1). Both ocean temperature and PCO2 are key environ- croscopy and scanning electron microscopy (SEM) to examine the skel-
mental factors affecting coral calcification rates (2–4). There is usually etons of newly settled coral recruits of an abundant plate coral species
a parabolic relationship between temperature and calcification, with (Acropora spicifera) from the subtropical Houtman Abrolhos Islands
optimal temperatures for calcification normally defined by local con- in Western Australia. Recruits were grown for 1 month under four
ditions (5, 6), and a negative correlation between calcification and temperature-PCO2 regimes: (i) control (24°C and 250 matm, pH 8.2),
PCO2 (7, 8). However, the interactive effect of elevated temperature (ii) elevated temperature (“high T”; 27°C and 250 matm, pH 8.2),
and PCO2 on calcification appears to be variable, with both positive (iii) elevated PCO2 (“high PCO2”; 24°C and 900 matm, pH 7.7), and
(9–11) and negative (2, 11, 12) interactions reported in adult corals. (iv) elevated temperature and elevated PCO2 (“high T + PCO2”; 27°C
Adult coral calcification is already thought to have declined by 14 to and 900 matm, pH 7.7) (see also table S1 and figs. S1 and S2). X-ray
30% worldwide in recent years, with some studies suggesting that elevated microscopy was used to generate 3D images and data of the skeletons
temperature and ocean acidification are the major causes (3, 4). to discern both visual and quantitative differences in skeletal structure
Corals in their early life stages have also been shown to be sensitive between the four regimes. The 3D reconstructions enabled the extraction
to changes in temperature and PCO2 (13–18). Successful reproduction of bulk measurements (surface area and volume), cross-sectional mea-
and post-recruitment processes (particularly growth) are essential to surements (height and basal plate thickness), and internal measurements
maintaining coral reef health (19). Therefore, it is particularly impor- (corallite wall thickness and tertiary septa width)—structural infor-
tant to know how these environmental changes will affect skeletal mation that would be impossible to obtain using conventional 2D im-
growth in the early stages of development, when tiny recruits (~1 mm) aging methods (figs. S3 and S4). SEM images of the same individuals
are most vulnerable to mechanical damage, overgrowth, and preda- were also acquired to more closely examine abnormalities in the skel-
tion. Experiments examining the impact of ocean acidification on cal- etal microstructure.
cification of newly settled recruits have reported strong effects, with 20
to 60% reductions in skeletal mass under acidified conditions (20–23).
However, the combined effect of elevated temperature and PCO2 on
skeletal mass may be dependent on geographical location, with a neg- RESULTS
ative impact in the tropics (24) and a positive impact in the subtropics Coral skeletons from low PCO2 treatments (control and high T) were
(23). These studies have highlighted the vulnerability of new recruits typically radially symmetric and had six tertiary septa of similar size
to ocean acidification and, in particular, their limited ability to calcify (Fig. 1, A and E, figs. S5 and S6, and movie S1). Skeletal surfaces of the
low-PCO2 corals were also smooth and appeared solid at both the cor-
1
UWA School of Earth and Environment, University of Western Australia, Crawley, Western
allite wall and tertiary septa (Fig. 1, C, D, G, and H; see also fig. S3 for
Australia 6009, Australia. 2UWA Oceans Institute, University of Western Australia, Crawley, the exact location of these structures on the coral skeleton). In con-
Western Australia 6009, Australia. 3ARC Centre of Excellence for Coral Reef Studies, Uni- trast, high-PCO2 corals had a variety of deformities disrupting their
versity of Western Australia, Crawley, Western Australia 6009, Australia. 4Centre for Mi- symmetry. Most high-PCO2 corals had missing or unevenly sized ter-
croscopy, Characterisation and Analysis, University of Western Australia, Crawley, Western
Australia 6009, Australia. tiary septa, with recruits commonly having overgrown septa on one
*Corresponding author. E-mail: [email protected] side of the mouth and missing or stunted septa on the other side

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Fig. 1. X-ray microscopy and SEM images of 1-month-old coral skeletons under the four temperature-PCO2 treatments (A to P). Treatments
include control (A to D), high T (E to H), high PCO2 (I to L), and high T + PCO2 (M to P). 3D x-ray images: top-down view (A, E, I, and M) and side view (B, F, J, and
N). Scale bars, 500 mm. SEM images: top of the corallite wall (C, G, K, and O) and a tertiary septum (D, H, L, and P). Scale bars, 10 mm. The four images shown
in each row are of a single, representative individual from each treatment. See figs. S5 to S8 for images of the other individuals from each treatment.

(Fig. 1, I and M). Perhaps the most striking difference between high- High PCO2 also significantly contributed to reductions in overall skel-
and low-PCO2 corals was the deep pitting and porous microstructure etal deposition, as shown by high surface area–to–volume ratios (SA/vol)
of the corallite wall (Fig. 1, K and O), a feature that was consistent as well as reduced diameter and basal plate thickness in high-PCO2
across all individuals grown under high PCO2 (figs. S7 and S8). Although corals under both temperature regimes (Fig. 3 and table S2). Further-
not as severe, shallow pitting was also observed in the tertiary septa of more, the length-to-width ratios of the tertiary septa were highly re-
the high-PCO2 corals (Fig. 1, L and P, and figs. S7 and S8). Other de- duced because of malformation (that is, incomplete extension of the
formities observed only in high-PCO2 corals included small gaps in the septa) in the high PCO2 treatment, and vertical growth (height) was sim-
lattice-like structure of septa and synapticulae (horizontal connecting ilarly stunted. Temperature alone appeared to have little effect, only
structures) (Fig. 2, E and F) as well as more severe deformities, such as significantly increasing tertiary septa length-to-width ratios. Post hoc
large sections of the skeleton being completely absent (Fig. 2, G and H, comparisons showed that where there were statistically significant in-
and movie S2). Fractures were also observed in the septa and corallite teractions between temperature and PCO2 they were positive, with el-
walls of 50% of the high-PCO2 corals (Fig. 2, A to D), whereas no evated temperature significantly increasing both height and tertiary septa
fractures were recorded in the low-PCO2 corals. length-to-width ratios under high PCO2 (P = 0.033 and P = 0.001 from

Foster et al. Sci. Adv. 2016; 2 : e1501130 19 February 2016 2 of 7

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treatments, the eroded and highly porous surfaces of high-PCO2 corals

at both temperatures demonstrate that the microstructure of the cor-
allite wall was severely altered by high PCO2, even if the larger scale
structure of the wall was not.

DISCUSSION
Our results indicate that coral recruits are unable to build normal cal-
cium carbonate skeletons under high PCO2 and low pH conditions.
High PCO2 resulted in structurally compromised skeletons that were
smaller (increased SA/vol, reduced diameter and height), more fragile
(thinner basal plate, pitted or porous corallite walls), and asymmetric.
A higher frequency of fractures was also observed in the high-PCO2

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corals. Although the formation of these was likely an artifact of sample
handling and processing after the experiment, it is remarkable that
50% of the individuals grown under acidified conditions had skeletal
fractures, whereas no fractures were observed in skeletons grown un-
der low PCO2 conditions. Alternatively, the fractures may have oc-
curred during the experiment while the recruits were still alive and
growing their skeletons. Perhaps the increased porosity and reduced
structural thickness caused fractures to form in situ under the me-
chanical strains of an actively growing skeleton. Future studies will
need to examine this possibility with greater scrutiny. Consistent with
the SEM data, fractures recorded in the high-PCO2 environment high-
light a more fragile, porous skeleton. A recent study on adult corals
similarly reports that acidification causes more porous skeletal struc-
tures, but at a much higher PCO2 than that applied to the juvenile corals
studied here (PCO2 ~ 2200 and 3800 versus 900 matm) (25). Our results
highlight how acutely vulnerable the growth and development of ju-
venile corals are to the changes in ocean chemistry expected to occur
over the coming century under a business-as-usual scenario (RCP 8.5)
(1, 26). Although our low PCO2 conditions were lower than present-
day ambient conditions (~250 versus 390 matm), it is unlikely that this
lower PCO2 condition influenced the overall conclusions of this study
given that (i) the impacts of the high PCO2 treatment on the skeleton
were so severe, (ii) the difference in PCO2 between present-day and
the low PCO2 treatments (~140 matm) is much smaller than the differ-
ence between the low and high PCO2 treatments (~650 matm), and (iii)
research comparing juvenile calcification under preindustrial and current
ambient conditions has shown no significant differences (27). None-
theless, it is possible that the more preindustrial PCO2 levels used in
our controls could have led to greater differences in skeletal deposition
between the low and high PCO2 conditions.
Corals are not the only calcifiers that experience deformation of
their skeletal structure under acidified conditions. Skeletal asymmetry
and microstructural abnormalities have been observed in both sea
Fig. 2. Fractures and deformed skeletal structures in high PCO2–treated urchin juveniles and brittle star larvae exposed to acidified conditions
corals. Fractures in the septa (A and B) and corallite wall (C and D). Small
(28–31). One study suggested that acidification might affect brittle star
sections of missing septa and synapticulae (E and F). Gross deformities, with
large sections of the skeleton missing or malformed (G and H).
larval symmetry through the symmetry-controlling ion-flux mecha-
nism (29, 32). This mechanism induces left-right symmetry breakage
in sea urchin larvae through asymmetrically localized control of H+
Bonferroni pairwise comparisons between high PCO2 and high T + and K+ transport (32). It is possible that similar mechanisms are re-
PCO2 for height and tertiary septa length-to-width ratios, respectively; sponsible for the CO2-driven skeletal asymmetry observed in the coral
Fig. 3 and table S2). Elevated temperature appeared to somewhat mit- recruits studied here. In addition, features indicative of mineral disso-
igate the negative effects of high PCO2 on skeletal growth, particularly lution (that is, disorganized aragonite bundles and an overall pitted ap-
appearing to facilitate vertical growth at high PCO2. Although there pearance of the skeletal surface) have also been observed in adult corals
were no significant differences in corallite wall thickness between exposed to acidified conditions at natural CO2 vents (12). However,

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A B C
P CO2 P CO2 Temperature * P CO2
Surface area/volume

Diameter (µm)

Height (µm)
Control High T High PCO2 High T + PCO2 Control High T High PCO2 High T + PCO2 Control High T High PCO2 High T + PCO2

D E F
Temperature

Corallite wall thickness (µm)

Basal plate thickness (µm)

P CO2

Tertiary septa length/width

Temperature * P CO2

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Control High T High PCO2 High T + PCO2 Control High T High PCO2 High T + PCO2 Control High T High PCO2 High T + PCO2

Fig. 3. Quantitative output from x-ray microscopy scans of 1-month-old coral skeletons under the four temperature-PCO2 treatments (mean ± SE).
(A to F) Measurements include (A) SA/vol ratio, (B) diameter, (C) height, (D) basal plate thickness, (E) corallite wall thickness, and (F) tertiary septa length/width
ratio. Factors [temperature, PCO2, or their interaction (temperature *PCO2)] significantly contributing to differences between treatments are indicated by ★ at
the top of the graph (n = 5 individuals per treatment). See table S2 for details on statistical tests.

disordered crystals were only observed in areas of exposed skeleton which may offset the more severe effects on the skeleton in the early
(that is, parts of the skeleton not covered by tissue), whereas areas post-recruitment stages.
where a tissue layer covered the whole skeleton exhibited no such dis- In contrast to the effects of acidification, our results suggest that the
solution characteristics (12). The skeletons of juvenile corals in our initial effects of ocean warming (+3°C) on calcification in subtropical
study were completely covered in a tissue layer throughout the exper- juveniles could either be minimal or may even help to mitigate the
iment and yet still showed deep pitting on the skeletal surface, which negative impacts of elevated PCO2; however, whether the same re-
could indicate dissolution. This may be due to juveniles having a much sponse is seen in other subtropical species is yet to be tested. This result
thinner tissue layer for protection (33) compared to adult corals and is contrary to studies that have been carried out in the tropics, where
further highlights the vulnerability of juvenile skeletogenesis under high temperature has exacerbated reductions in calcification under
elevated PCO2. high PCO2, in both juvenile (24) and adult (2, 12) corals. Tropical corals
As some coral reefs currently experience elevated PCO2 (and sup- exist close to their upper thermal limit (38) and yet are presumably
pressed pH) relative to the open ocean (up to ~500 matm) (34, 35), it growing within the optimal range of temperatures for calcification. Con-
is possible that recruits are adopting mechanisms to cope with these sequently, a 3°C increase in temperature could subject them to thermal
conditions in situ. A recent study has shown that adult corals living stress and depress calcification rates below optimal levels. In contrast,
in highly dynamic thermal and chemical environments more rigor- subtropical coral larvae and new recruits appear to withstand moderate
ously control the up-regulation of pH within the calcifying fluid, thus (+3°C) increases in temperature (23, 39). This may be a result of sub-
demonstrating a greater resilience to the effects of ocean acidification tropical corals being acclimated to the wider annual range of tempera-
(36). Whether or not coral recruits that have spawned from or settled tures generally experienced at higher-latitude locations. Additionally,
in such highly variable reef environments also exhibit a greater de- the ability of new recruits to withstand moderate temperature increases
gree of physiological resilience to acidification remains to be seen. could be related to larval dispersal in the water column, where larvae
Alternatively, new recruits may already be calcifying at reduced rates must survive, often for extended periods, over long distances, and
(relative to preindustrial conditions) in these environments. The through changing thermal environments (23, 40, 41).
reduced ability to quickly produce a robust skeleton in new coral In conclusion, acidified conditions caused severely abnormal skel-
recruits can have a direct impact on survival rates. For example, re- etogenesis in new coral recruits. Although the partially mitigative effect
duced growth rates in 2-month-old coral recruits, under elevated of elevated temperature is encouraging in these subtropical recruits, taller
PCO2, led to reductions in overall survivorship due to elevated pre- but similarly fragile skeletons may not provide and sustain the structural
dation mortality, demonstrating how size-escape thresholds can support and protection required during early post-recruitment. Fur-
shift under ocean acidification (37). However, it is important to rec- thermore, this mitigative effect does not appear to be present in tropical
ognize that those individuals that do survive the smaller and more coral recruits (24). Our data show that new recruits are thus highly
susceptible size classes may be better able to cope with high PCO2 vulnerable to acidification and indicate that near-future projections for
conditions than (i) prior generations and (ii) as they get larger, ocean carbonate chemistry (1, 26) have the potential to heavily reduce

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post-recruitment success. Disruptions to normal skeletal development in driving PCO2 below atmospheric levels. A rigorous study of the varia-
new recruits could compromise the ability of coral reefs to successfully tion in carbonate chemistry at the Abrolhos is yet to be undertaken;
migrate to more suitable thermal environments, as well as inhibit the however, spot measurements taken over 2011 to 2013 show the range
replenishment of existing reefs after episodic disturbances caused by for PCO2 and pH to be ~300 to 500 matm and ~7.9 to 8.1, respectively
storms, disease outbreaks, and mass bleaching events—all of which (45). Light levels were maintained at a 12:12–hour light/dark cycle,
are expected to increase over the next century if the present CO2 trajec- with a mean (±SE) light intensity of 212 ± 8 mmol photons m−2 s−1
tory is not abated. (Biospherical Instruments, QSL-2100). The light levels applied in this
(212 mmol photons m−2 s−1) and other juvenile calcification studies
(62 to 135 mmol photons m−2 s−1) (20–22, 24, 27) are relatively low
MATERIALS AND METHODS compared to the natural light levels experienced on a shallow reef.
Higher light intensities experienced in situ may increase calcification
Collection methods relative to those recorded in laboratory experiments; however, larvae
Gravid adult colonies of the abundant plate coral species A. spicifera often show a preference during settlement for microcrevices and ledges
were collected off Basile Island in the Pelsaert group of the Houtman that offer protection from predators but are presumably exposed to lower

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Abrolhos Islands (28°52′S, 113°57′E) in Western Australia, before the light intensities. Thus, the lower light levels we used may be more repre-
2013 March mass-spawning event. Colonies were maintained in flow- sentative of their environment in the early post-recruitment phase.
through outdoor aquaria exposed to natural lighting, which received
seawater filtered through a 20-mm nylon mesh. On the night of spawn- Larval settlement
ing, fertilized eggs (cross-fertilized from six spawning colonies, repre- Larvae (n = 40) were transferred into 50-ml clear acrylic tubes, with a
senting equal contributions from all six genotypes) were transferred to 100-mm mesh at the ends to allow water exchange. Plastic transparen-
larval rearing tanks and maintained under ambient conditions until cy paper, washed and soaked in ambient seawater for 1 week, lined the
the larvae reached the planulae stage of development at 6 days post inside of the tubes. The transparency paper lining was used both for
fertilization. Larvae were then transferred to treatment tanks (four easy removal of the recruit skeletons at the end of the experiment and
replicate tanks per treatment). as a low-density substrate, easily excluded from 3D x-ray microscopy
measurements of the skeleton. Three small (0.5 cm2) crustose coralline
Experimental design algae chips (Hydrolithon), collected from the same site as the adult
All seawater entering the aquarium system was foam-fractionated and A. spicifera colonies, were added to each tube to induce settlement.
ultraviolet-sterilized to remove solids and bacteria. Seawater was treated Twelve tubes were transferred into each replicate treatment tank. Be-
in four separate sump tanks (one sump per treatment) that flowed into cause of logistical constraints, each tank housed multiple larval tubes,
the replicate tanks every 3 hours for an ~90% water change. The car- resulting in a partial pseudoreplicate design (independent tanks with
bonate chemistry was adjusted using a pH-dependent feedback sys- replicated tubes). Although the larvae had no measureable impact on
tem, which bubbled pure CO2 into the seawater when pH deviated the environmental conditions in each treatment, skeletal analyses were
0.01 pH units from set values (CO2 Set Professional, AquaMedic). The performed on individuals randomly selected across tubes and tanks
pH was checked manually in all tanks every second day. The pH was to help counter pseudoreplication within the experimental design. At
measured on the total scale (±0.03) using a Schott Handylab pH meter 7 days post settlement, recruits were inoculated with cultured clade C1
equipped with a Blueline 24 electrode and calibrated against a combi- zooxanthellae (V. Beltran, AIMS), a common clade in this region (46).
nation of NBS (pH 4 and 7) and artificial seawater (tris) buffers; the Settled juveniles were grown for 4 weeks post settlement under treat-
latter was calibrated against certified reference materials (CRMs), ment conditions. Before termination of the experiment, each individ-
provided by A. Dickson at the Scripps Institute of Oceanography ual was examined and photographed using both a stereoscope and a
(batch #7). Total Alkalinity (TA, ±5 mmol/kg) samples were taken fluorescence microscope to verify that the recruits were alive until the
once a week and measured using a single end-point titration to a spec- endpoint of the experiment. To remove organic material, the recruits
trophotmetrically determined pH of ~4.0 (42), and calibrated against were immersed in 3 to 7% sodium hypochlorite and rinsed three times
CRMs, also provided by A. Dickson (batch #105). Salinity was checked in deionized water.
twice a week using a refractometer (35.5). Aragonite saturation state
(War) and PCO2 were calculated from pH, TA, salinity, and temperature X-ray microscopy settings
using the program CO2SYS (43). Water temperature was controlled Five primary polyps were randomly selected across tubes and tanks
with heater chillers (Resun, CL 150) and monitored using Hobo Pen- from each treatment for x-ray microscopy analyses. Randomly selected
dant temperature loggers (±0.5° to ±0.9°C). The four temperature-PCO2 recruits also needed to meet four selection criteria to be usable for
conditions are outlined in table S1. The annual temperature range at the scanning: (i) were in the primary polyp stage of development, (ii) had
Abrolhos is ~19° to 24°C (44). Thus, the high-temperature treatment settled on the plastic transparency paper, (iii) had settled a suitable
(27°C) represents an elevation of 3°C above the maximum monthly distance from other recruits, and (iv) had remained attached to the
mean water temperature at the Abrolhos, which occurs around spawn- paper and undamaged throughout processing. All x-ray scans were con-
ing time (24°C). The high PCO2 treatment (PCO2 ~ 900 matm) is similar to ducted using an Xradia Versa XRM520 x-ray microscope. Scanning was
projections for the year 2100 under the RCP 8.5 scenario (~930 matm) (26). undertaken under the following conditions to produce data with
The low PCO2 treatments are closer to preindustrial (250 matm) than ~1.3-mm pixel resolution: voltage, 50 kV; power, 4 W; exposure time,
present-day atmospheric levels (~390 matm). This was likely due to 20 s; binning, 2; and angle, full 360°, with a total of 3201 projections
seawater being pumped into the experimental facility in the afternoon per sample. Reconstruction of the projections was conducted using
when photosynthesis of benthic primary producers in the region was the Xradia software XMReconstructor. Because of the long scan times

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associated with obtaining high-resolution 3D images (~19 hours per Fig. S4. Examples of manual measurements of diameter, corallite wall thickness (A and B),
tertiary septa length/width ratio (C and D), and height and basal plate thickness (E and F).
individual recruit), only five individuals per treatment were scanned.
Fig. S5. All remaining individuals from the control (that is, excluding the individual shown in
However, the small variation within each treatment in the quantitative Fig. 1).
output indicated that this replication was sufficient. Fig. S6. All remaining individuals from the high T treatment (that is, excluding the individual
shown in Fig. 1).
Quantitative data extraction Fig. S7. All remaining individuals from the high PCO2 treatment (that is, excluding the individual
shown in Fig. 1).
The software Avizo Fire was used to process the reconstructed projec- Fig. S8. All remaining individuals from the high T + PCO2 treatment (that is, excluding the
tions and extract quantitative data. The Edit New Label Field tool was individual shown in Fig. 1).
used to set threshold radiodensity values, which removed the transpar- Fig. S9. nMDS ordination in two dimensions of quantitative output from 3D x-ray microscopy
ency paper, leaving only the CaCO3 skeleton. The samples were then scans of juvenile coral skeletons, including SA/vol (A), diameter (B), height (C), basal plate
thickness (D), corallite wall thickness (E), and tertiary septa length/width ratio (F) for
volume-rendered to create a 3D image. A label analysis was conducted
individuals grown under different temperature-PCO2 regimes.
on the segmented data set to determine the surface area and volume of Fig. S10. nMDS ordination in two dimensions, pooling all measures from 3D x-ray microscopy
the sample. All other measurements were made manually, using the Mea- scans of juvenile coral skeletons.
sure tool (fig. S4). For height, corallite wall thickness, and basal plate Table S1. Physical and chemical conditions maintained in each treatment for the duration of

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thickness, the mean of three measurements per sample was determined, the experiment [mean ± SD; Foster et al. (23)].
Table S2. Two-way ANOVAs testing for significant effects of temperature, PCO2, and interactions
whereas the mean diameter was calculated from four measurements per between the two factors (temperature * PCO2) on x-ray microscopy measurements of juvenile
sample. For height and basal plate thickness, the three highest or thick- coral skeletons.
est points in the sample were selected by scrolling through the slices of Movie S1. Skeleton of a 1-month-old coral recruit grown under control (24°C and 250 matm)
the sample. For diameter and corallite wall thickness, a single slice near conditions.
Movie S2. Skeleton of a 1-month-old coral recruit grown under high PCO2 (24°C and 900 matm)
the base showing the whole diameter was selected, and measurements
conditions.
were taken at the same points and along the same axis for each sample.
The length and width were measured for all tertiary septa.

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Foster et al. Sci. Adv. 2016; 2 : e1501130 19 February 2016 7 of 7

 Ocean acidification causes structural deformities in juvenile
coral skeletons
Taryn Foster, James L. Falter, Malcolm T. McCulloch and Peta L.
Clode (February 19, 2016)
Sci Adv 2016, 2:.
doi: 10.1126/sciadv.1501130

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