Effects of seawater temperature and pH on the boring rates of the sponge Cliona celata in scallop shells

Warmer, more acidic water resulting from greenhouse gas emissions could influence ecosystem processes like bioerosion of calcifying organisms. Based on summer-maxima values (temperature = 26 °C; pH = 8.1) at a collection site in New York (40°56″ N, 72°30″ W), explants of the boring sponge Cliona celata Grant, 1826 were grown for 133 days on scallop shells in seawater ranging from current values to one scenario predicted for the year 2100 (T = 31 °C; pH = 7.8). High water temperature had little effect on sponge growth, survival, or boring rates. Lower pH slightly reduced sponge survival, while greatly influencing shell boring. At pH = 7.8, sponges bored twice the number of papillar holes and removed two times more shell weight than at pH = 8.1. Greater erosion resulted in weaker scallop shells. This study suggests that lower seawater pH may increase boring rates of C. celata in shellfish, with potentially severe implications for wild and farmed shellfish populations.     

Alternate coral–bryozoan competitive superiority during coral bleaching

Bleaching of corals results from the loss of their symbiotic algae (zooxanthellae) and/or pigments. The supply of photoassimilates provided by the zooxanthellae to the coral declines during bleaching and reduces the ability to activate energy-costly processes such as maintenance, growth and reproduction. In the present study we compared the competitive outcomes, expressed as overgrowth and changes in colony sizes of Oculina patagonica (an encrusting Mediterranean stony coral) and the bryozoan Watersipora sp., growing in contact with each other, during and between bleaching events. Year-round observations of tagged colonies showed alternating competitive outcomes: O. patagonica wins over Watersipora sp. between bleaching events, but loses during bleaching events. Using the ¹⁴C-point-labeling technique on coral tissue, we examined intra-colonial translocation of photosynthetic products from the point-tissue labeling towards interaction zones. In non-bleached O. patagonica, competition resulted in preferentially oriented translocation of ¹⁴C products to the interaction zone located up to 8 cm away from the tissue-labeling site. Sites opposite the interaction zone received significantly less labeled photoassimilates compared to the interaction zone. In bleached colonies (40-85% bleached surface area), such translocation did not occur, probably explaining the failure to compete with the encrusting neighbor Watersipora sp. Our findings demonstrate the importance of colonial integration and resource orientation for the competitive superiority of O. patagonica.     

Seasonal cycle of sexual reproduction of the Mediterranean soft coral Alcyonium acaule (Anthozoa, Octocorallia)

Alcyonium acaule (Cnidaria, Octocorallia) is a common, hard-bottom soft coral in the northwestern Mediterranean Sea. This study describes sexual reproduction and the gamete development cycle of this soft coral. A population at 15–18 m depth in the Marine Protected Area of the Medes Islands (42º02′55″ N, 3º13′30″ E) was sampled from July 1994–August 1995. A. acaule is gonochoristic and a surface brooder, spawning once a year in early summer. The mean diameter of ripe spermatic sacs was 400 ± 91 (SD) μm, and the mean diameter of mature oocytes was 473 ± 37 (SD) μm. There were 30 spermatic sacs polyp^?1 in males and 14 oocytes polyp^?1 in females. Different phases of gametogenesis in female and male colonies were examined separately with respect to seasonal changes in bottom temperature and solar irradiance. The data suggest that the relatively constant temperatures in January–April are probably not related to oocyte maturation, but that rising temperatures in May could affect sperm maturation. Rapidly increasing solar irradiance in March may be the trigger for vitellogenesis and oocyte maturation, although the mechanism for this in anthozoans is not understood.     

Reproductive periodicity of the scleractinian coral Lophelia pertusa from the Trondheim Fjord, Norway

Between 2002 and 2008, samples of the cold-water scleractinian coral Lophelia pertusa were collected from the Trondheim Fjord in Norway to examine reproductive periodicity. Collections were made from three locations: Tautra, (63°35.36′N, 10°31.23′E at 40–70 m), Stokkbergneset (63°28.18′N, 09°54.73′E at 110–500 m), and Røberg (63°28.88′N, 09°59.50′E at 250 m). Populations of L. pertusa from the Trondheim Fjord initiated oogenesis in January and spawning occurred from late January to early March the following year. Gametogenic cycles of the female L. pertusa samples overlapped by approximately 2 months, with oogonia visible in January, but this was not evident in the males. This paper provides the most complete gametogenic cycle to date and spawning observations for this important structure-forming species. The results from fjord populations are compared with published and preliminary data from other regions and are discussed in the context of regional differences in physical and biological variables, particularly food supply. Differences in gametogenic cycles within a single species provide a rare opportunity (especially in deep-sea species) to examine potential drivers of reproduction.     

The effects of an abnormal decrease in temperature on the Eastern Pacific reef-building coral Pocillopora verrucosa

Coral bleaching events are associated with abnormal increases in temperature, such as those produced during El Niño. Recently, a breakdown in the coral–dinoflagellate (genus Symbiodinium) endosymbiosis has been documented in corals exposed to anomalously cold-water temperatures associated with La Niña events. Given the ecological significance of such events, as well as the threat of global climate change, surprisingly little is known about the physiological response of corals to cold stress. This study evaluated some physiological effects of continuous temperature decline in colonies of the eastern Pacific reef-building coral Pocillopora verrucosa. Twenty days of incubation at 18.5–19.0 °C resulted in a substantial decrease in holobiont lipid and Chla content, as well as an increase in Symbiodinium density. These observations suggest a combination of symbiont acclimation due to the temperature decline and reallocation of carbon toward algal growth as opposed to translocation to the host coral. With a decreased availability of symbiont-derived carbon, the coral likely catabolized storage lipids in order to survive the stress event. Despite this stress and some tissue necrosis, no mortality was noted and corals recovered quickly when returned to the ambient temperature. As these results are in marked contrast to similar studies investigating elevated temperature on this coral from this same location, Pocillopora in the Mexican Central Pacific may be more prone to long-term damage and mortality during periods of ocean warming as opposed to ocean cooling.     

The decline and recovery of four predatory fishes from the Southern California Bight

What to do about fisheries collapse and the decline of large fishes in marine ecosystems is a critical debate on a global scale. To address one aspect of this debate, a major fisheries management action, the removal of gill nets in 1994 from the nearshore arena in the Southern California Bight (34°26′30″N, 120°27′09″W to 33°32′03″N, 117°07′28″W) was analyzed. First, the impetus for the gill net ban was the crash of the commercial fishery for white seabass (Atractoscion nobilis; Sciaenidae) in the early 1980s. From 1982 to 1997 catch remained at a historically low level (47.8 ± 3.0 mt) when compared to landings from 1936–1981, but increased significantly from 1995–2004 (r = 0.89, P < 0.01) to within the 95% confidence limit of the historic California landings. After the white seabass fishery crashed in the early 1980s, landings of soupfin (Galeorhinus galeus; Triakidae) and leopard shark (Triakis semifasciata; Triakidae) also significantly declined (r = 0.95, P < 0.01 and r = 0.91, P < 0.01, respectively) until the gill net closure. After the closure both soupfin and leopard shark significantly increased in CPUE (r = 0.72, P = 0.02 and r = 0.87, P < 0.01, respectively). Finally, giant sea bass (Stereolepis gigas; Polyprionidae) the apex predatory fish in this ecosystem, which was protected from commercial and recreational fishing in 1981, were not observed in a quarterly scientific SCUBA monitoring program from 1974 to 2001 but reappeared in 2002–2004. In addition, CPUE of giant seabass increased significantly from 1995 to 2004 (r = 0.82, P < 0.01) in the gill net monitoring program. The trends in abundance of these fishes return were not correlated with sea surface temperature (SST), the Pacific Decadal Oscillation (PDO) or the El Niño/Southern Oscillation (ENSO). All four species increased significantly in either commercial catch, CPUE, or in the SCUBA monitoring program after the 1994 gill net closure, whereas they had declined significantly, crashed, or were absent prior to this action. This suggests that removing gill nets from coastal ecosystems has a positive impact on large marine fishes.     

Brooded coral larvae differ in their response to high temperature and elevated pCO2 depending on the day of release

To evaluate the effects of temperature and pCO₂ on coral larvae, brooded larvae of Pocillopora damicornis from Nanwan Bay, Taiwan (21°56.179′N, 120°44.85′E), were exposed to ambient (419–470 μatm) and high (604–742 μatm) pCO₂ at ~25 and ~29 °C in two experiments conducted in March 2010 and March 2012. Larvae were sampled from four consecutive lunar days (LD) synchronized with spawning following the new moon, incubated in treatments for 24 h, and measured for respiration, maximum photochemical efficiency of PSII (F _v/F _m), and mortality. The most striking outcome was a strong effect of time (i.e., LD) on larvae performance: respiration was affected by an LD × temperature interaction in 2010 and 2012, as well as an LD × pCO₂ × temperature interaction in 2012; F _v/F _m was affected by LD in 2010 (but not 2012); and mortality was affected by an LD × pCO₂ interaction in 2010, and an LD × temperature interaction in 2012. There were no main effects of pCO₂ in 2010, but in 2012, high pCO₂ depressed metabolic rate and reduced mortality. Therefore, differences in larval performance depended on day of release and resulted in varying susceptibility to future predicted environmental conditions. These results underscore the importance of considering larval brood variation across days when designing experiments. Subtle differences in experimental outcomes between years suggest that transgenerational plasticity in combination with unique histories of exposure to physical conditions can modulate the response of brooded coral larvae to climate change and ocean acidification.     

Latitudinal patterns of shell thickness and metabolism in the eastern oyster Crassostrea virginica along the east coast of North America

The goal of this study was to quantify growth and metabolic responses of oysters to increased temperatures like those that will occur due to global warming. Impact of temperature on eastern oyster (Crassostrea virginica) shell growth and metabolism was investigated by sampling 24 sites along the eastern North American seaboard ranging from New Brunswick, Canada, to Florida, USA, in March and August 2013. There was a positive correlation between oyster shell thickness and site temperature. At southern sites, shells were up to 65 % thicker than at the northernmost site, likely due to higher precipitation of CaCO₃ in warmer water. This was supported by laboratory experiments showing that thicker shells were produced in response to temperatures 2, 4, and 6 °C above ambient seawater temperatures (8–14 °C) in Connecticut, USA. Field experiments with oyster respiration were conducted during winter and summer at 13 sites to compare responses to thermal stress with latitude. Respiration rates were much higher during summer than winter, but the combination of summer and winter data fell along the same exponential curve with respect to temperature. At all sites, temperature-specific metabolic rates at elevated temperatures were lower than predicted, indicating significant seasonal acclimatization by C. virginica.     

Activation energy for skeletal aragonite deposited by the hermatypic coral Platygyra spp.

Average radial growth rates of the hemispherical aragonite colonies deposited by the Indo-Pacific scleractinian reef coral Platygyra spp. were determined by measuring the thickness of density variations in the skeleton that are revealed by X-radiography. Ninety-one specimens from 21 localities were examined, but only 54 of these exhibited well-defined growth-banding. The apparent temperature dependence of growth rate is linear over the range 23.9° to 29.3°C, averaging 5.4±0.94 mm/year at 24°C, 8.0±0.42 mm/year at 27°C, and 9.7 ±0.58 mm/year at 29°C (90% confidence limits). Expression of the influence of temperature on growth rate in terms of the Arrhenius equation yields an activation energy of 20,680 cal/mole, which is comparable to values for typical biological reactions, but is only half that reported for skeletogenesis in another reef coral, Pocillopora damicornis, on the basis of controlled incubation studies involving ⁴⁵Ca uptake.     

The synergistic effects of hydrogen peroxide and elevated seawater temperature on the metabolic activity of the coral <Emphasis Type="Italic">Galaxea fascicularis</Emphasis>

We examined quantitative changes in the metabolism of the coral Galaxea fascicularis caused by increases in both hydrogen peroxide (H₂O₂) concentration and seawater temperature. Seawater temperatures were maintained at 27 or 31°C in a well-controlled incubation chamber, and three levels of H₂O₂ concentration (0, 0.3, 3.0 μM) were used in experimental treatments. Gross primary production, calcification rates and respiration rates were all affected by increased H₂O₂ concentrations and high seawater temperatures. Individual treatments of high H₂O₂ or elevated seawater temperature alone caused significant declines in coral photosynthesis and calcification rates within the 3-day incubation period. The synergistic effect of high H₂O₂ combined with high seawater temperature resulted in a 134% increase in respiration rates, which surpassed the effect of either H₂O₂ or high seawater temperature alone. Our results suggest that both high H₂O₂ concentrations and elevated temperatures in seawater can strongly affect coral metabolism; however, these effects cannot be estimated by simply summing the effects of individual stress parameters.     

Effects of high temperature and CO2 on intracellular DMSP in the cold-water coral Lophelia pertusa

Significant warming and acidification of the oceans is projected to occur by the end of the century. CO₂ vents, areas of upwelling and downwelling, and potential leaks from carbon capture and storage facilities may also cause localised environmental changes, enhancing or depressing the effect of global climate change. Cold-water coral ecosystems are threatened by future changes in carbonate chemistry, yet our knowledge of the response of these corals to high temperature and high CO₂ conditions is limited. Dimethylsulphoniopropionate (DMSP), and its breakdown product dimethylsulphide (DMS), are putative antioxidants that may be accumulated by invertebrates via their food or symbionts, although recent research suggests that some invertebrates may also be able to synthesise DMSP. This study provides the first information on the impact of high temperature (12 °C) and high CO₂ (817 ppm) on intracellular DMSP in the cold-water coral Lophelia pertusa from the Mingulay Reef Complex, Scotland (56°49′N, 07°23′W), where in situ environmental conditions are meditated by tidally induced downwellings. An increase in intracellular DMSP under high CO₂ conditions was observed, whilst water column particulate DMS + DMSP was reduced. In both high temperature treatments, intracellular DMSP was similar to the control treatment, whilst dissolved DMSP + DMS was not significantly different between any of the treatments. These results suggest that L. pertusa accumulates DMSP from the surrounding water column; uptake may be up-regulated under high CO₂ conditions, but mediated by high temperature. These results provide new insight into the biotic control of deep-sea biogeochemistry and may impact our understanding of the global sulphur cycle, and the survival of cold-water corals under projected global change.     

Symbiont specificity and bleaching susceptibility among soft corals in the 1998 Great Barrier Reef mass coral bleaching event

Considerable variability in bleaching was observed within and among soft coral taxa in the order Alcyonacea (Octocorallia: Cnidaria) on the central Great Barrier Reef (GBR, latitude 18.2°–19.0°S, longitude 146.4°–147.3°E) during the 1998 mass coral bleaching event. In April 1998, during a period of high sea surface temperatures, tissue samples were taken from bleached and unbleached colonies representative of 17 soft coral genera. The genetic identities of intracellular dinoflagellates (Symbiodinium spp.) in these samples were analyzed using PCR-denaturing gradient gel electrophoresis fingerprinting analysis of the internal transcribed spacer regions 1 and 2. Alcyonaceans from the GBR exhibited a high level of symbiont specificity for Symbiodinium types mostly in clade C. A rare clade D type (D3) was associated only with Clavularia koellikeri, while Nephthea sp. hosted symbionts in clade B (B1n and B36). Homogenous Symbiodinium clade populations were detected in all but one colony. Colonies that appeared bleached possessed symbiont types that were genetically indistinguishable from those in nonbleached conspecifics. These data suggest that parameters other than the resident endosymbionts such as host identity and colony acclimatization are important in determining bleaching susceptibility among soft corals. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Symbiont shuffling during thermal bleaching and recovery in the sea anemone Entacmaea quadricolor

The sea anemone Entacmaea quadricolor simultaneously harbours multiple symbiont types from the genus Symbiodinium, while providing essential habitat for anemonefish. This anemone lives close to its upper thermal threshold and experiences bleaching under elevated temperature and light stress. Here, we determine whether E. quadricolor experienced a shuffling in the abundance of two genetically distinct symbiont types (Symbiodinium C25 and C3.25) during bleaching and recovery. Anemones were exposed to control (22.9 °C) or elevated temperature (28.5 °C) for 42 days, whereas for the following 75 days, all anemones were exposed to 22.9 °C. By day 47, a more pronounced bleaching occurred via symbiont expulsion in the elevated temperature treatment than the control, and the proportion of C25 to C25 + C3.25 increased by 6.2 and 13.2 % in the control and bleached anemones, respectively. The increased relative abundance of C25 to C3.25 after exposure to thermal stress may indicate that C3.25 performs poorly when temperature is elevated. Although no significant recovery in symbiont density was detected, a revival of the C3.25 genotype was found at day 117, which may indicate that it is either more competitive or has qualities that are beneficial to the symbiosis when thermal stress is no longer apparent. This work demonstrates the potential for this anemone species to shuffle its symbiont types in response to environmental change and could provide resilience during times of stress.     

Coefficient of variation of sea surface temperature (SST) as an indicator of coral bleaching

Coral bleaching has become a major problem on reefs around the world in recent decades. It is believed that mean temperature alone is the primary force driving this ecological phenomenon. We propose that variance in temperature in the short term is just as important as the mean. Thirty years of daily sea surface temperature (SST) data have been collected by the University of Puerto Rico at Mayaguez Marine Laboratory in La Parguera, PR. These data were collated and analyzed initially (by Amos Winter) for their relationship to coral bleaching in this area. We found that the data fell into three categories: high mean temperatures associated with severe bleaching, cooler mean temperatures associated with no bleaching, and years of high SSTs but with no coral bleaching. Here, we examined the relationship between mean temperature during those months in which bleaching occurred, temperature variance (as measured by standard deviation), and coefficient of variation (CV; i.e., SD standardized by the mean). We also derived a critical threshold temperature and level of resolution in time for calculating these statistics to clearly describe the circumstances of bleaching versus non-bleaching events, particularly at marginal bleaching temperatures. These characteristics were compared for the four warmest months of the year (July–October) for four warm bleaching years (1969, 1987, 1990, and 1995), four cool non-bleaching years (1984, 1985, 1986, and 1988), and two warm non-bleaching years (1994 and 2000). No relationship was found between the mean SST and SD in terms of predicting bleaching. The two primary statistics which, in concert, did indicate bleaching, however, were the short-term, biweekly mean temperature and its the associated CV. Bleaching occurs in association with both high temperatures and a high CV. The CV becomes a critical determinant of bleaching only when temperatures are ∼29.1–29.8°C. The warm, non-bleaching years were generally characterized by a CV of < 1.9 and a temperature range between 28.5 and 29.9°C. We conclude that increased mean SSTs alone are not sufficient to induce coral bleaching; a high variance in SST at marginal, lower bleaching temperatures can induce bleaching, and likewise, a low variance of such will not induce bleaching. This variance is most clearly described by the CV.     

Year-round reproduction in a seasonal sea: biological cycle of the introduced ascidian Styela plicata in the Western Mediterranean

The widely introduced ascidian Styela plicata is very common in the Western Mediterranean, an area that can act as a source for secondary introductions due to its high shipping activity. In order to understand the potential of this species to colonize new habitats, its reproductive features were assessed in the Western Mediterranean by means of monthly monitoring of two populations (Vilanova i la Geltrú 41°12′53″N, 1°44′11″E; Blanes 41°40′29″N, 2°47′56″E) from January 2009 to December 2010. The reproductive activity of this species was assessed through gonad histology and a gonad index. Population size-structure was measured monthly in order to study recruitment dynamics. No clear seasonal pattern was observed, and mature gametes and recruits were present all year long. Spawning was potentially continuous, although it seemed punctuated with pulses of gamete release, particularly in spring. A prolonged reproductive period is likely to confer a competitive advantage on S. plicata in temperate seas, where most species reproduce seasonally, and may promote recurrent introductions as larvae are available for settlement on transport vectors over much of the year.     

Oculina patagonica: a non-lessepsian scleractinian coral invading the Mediterranean Sea

The scleractinian coral Oculina patagonica De Angelis is a new immigrant from the Southwest Atlantic to the Mediterranean Sea, having established itself only recently along the Israeli coast. This species is the only scleractinian coral reported to have invaded a new region. In order to understand the swift establishment of this species along the Israeli coast, from 1994 to 1999 we studied its distribution, abundance, reproduction, recruitment, survival, and the effect of bleaching events on its population abundance. In addition, population studies of O. patagonica were performed at several localities along the eastern and western Mediterranean coasts. Highest abundance was recorded along the Spanish coast, reaching 30lj colonies per 10 m line transect in shallow water. Second in abundance was the Israeli coast, with a maximum of 10DŽ colonies per transect. O. patagonica was rare along the coast of Italy, and absent along the Mediterranean coast of France. During the study, both mortality and recruitment along the Israeli coast were very low. In contrast, recruitment along the Spanish coast was very high. Reproduction of the species was studied using gonadal histology. O. patagonica is gonochoric. Female gonads were first observed in May and male gonads in July, both reaching maturity in late August and early September. Naturally occurring azooxanthellate colonies of O. patagonica inhabiting small dark caves developed gonads and spawned in parallel to zooxanthellate colonies exposed to light. No gonads were found in zooxanthellate colonies that underwent bleaching during the reproduction season. The high incidence of bleaching events along the Israeli coast observed throughout the years of this study may explain the low recruitment of new colonies during the same period. In view of its current recruitment patterns, we expect further expansion of O. patagonica in range and abundance in the western Mediterranean, but very small expansion of the population in the eastern Mediterranean, due to repetitive annual bleaching events.     

Seasonal variation in the effects of ocean warming and acidification on a native bryozoan, Celleporaria nodulosa

Ocean warming and acidification are co-occurring stressors likely to affect marine biota through climate-driven change to the ocean. We investigated the effects of increased temperature and lowered pH, solely and in combination, on the growth of the endemic Australian bryozoan, Celleporaria nodulosa. Two temperatures and three pH levels were fully crossed in experimental treatments performed in winter 2008 (August) and summer 2009 (February/March). Fragments of C. nodulosa colonies (clones) were collected from Coffs Harbour, NSW, Australia, (30°18′S, 153°09′E) and elongation of colonies was assessed periodically over a 12-day incubation period. Lowered pH in winter significantly decreased growth. Elevated temperatures during the summer significantly impeded the growth of bryozoan colonies, possibly masking the effect of ocean acidification and discovering a maximal thermal tolerance at around 27 °C for C. nodulosa. The effects of decreased pH and increased temperature may be seasonally dependent and particularly acute during the summer months. Thermal stress may in fact be the initial stressor before ocean acidification, negatively affecting organisms in such a way that they are unable to survive before feeling the effects of ocean acidification.     

Temperature and salinity: two climate change stressors affecting early development of the New Zealand sea urchin Evechinus chloroticus

Temperature and salinity are important environmental factors affecting the normal functioning of marine animals, particularly animals such as sea urchins living in shallow waters and tide pools. Here, we evaluated the effect of different combinations of temperature and salinity on early embryos of the endemic New Zealand sea urchin Evechinus chloroticus. Animals were collected at Matheson’s Bay (36º18′17′′S; 174º47′51′′E) in north-eastern New Zealand in February 2013. Embryos were exposed to five salinities (29, 31, 34, 35 and 37 ppt) and two temperatures (18 and 21 °C) during the first 24 h of development. Low salinity (29 ppt) affected all parameters (fertilization, development rate, gastrulation and normal development), with ca. 50 % of embryos surviving at 29 ppt, whereas seawater temperature only affected development rate and gastrulation. An increase in temperature from 18 to 21 °C minimized the negative effect of low salinity (≤31 ppt) on development rate and gastrulation of E. chloroticus. Overall, the results of this study suggest that early embryos of E. chloroticus have developmental plasticity to withstand reductions in salinity up to 29 ppt; however, it is still unknown whether the surviving embryos will be able to complete larval development at low salinities, particularly whether the embryos and larvae are carried into extreme environments such as estuaries where salinity is even lower. Multistressor studies are very important for climate change research as multiple environmental factors will act together in the wild, having major consequences for development and recruitment of marine invertebrates.     

Rates of apical septal extension of Desmophyllum dianthus: effect of association with endolithic photo-autotrophs

It has been hypothesized that endolithic photo-autotrophs inside the skeleton of cold-water corals may have a mutualistic relationship with the coral host positively affecting coral calcification. This study investigated the effect of endolithic photo-autotrophs on the apical septal extension of the cold-water coral Desmophyllum dianthus at Fjord Comau, southern Chile (42.41°–42.15°S, 72.5°W). The fluorescent staining agent calcein was used to document the linear apical extension of septae for a period of one and a half years between 2006 and 2007. The results showed a severe reduction in extension rates associated with the presence of endolithic photo-autotrophs. Infested individuals grew about half as fast as non-infested polyps with a median value of 1.18 μm day^?1 compared to 2.76 μm day^?1. Contrary to the initial hypothesis, these results point toward a parasitic relationship between D. dianthus and its endolithic photo-autotrophs potentially impairing coral fitness. However, further data on physiological parameters and other aspects of the calcification process are necessary to confirm these findings.