Gertraud M. Schmidt

Large-amplitude internal waves benefit corals during thermal stress.

Tropical scleractinian corals are particularly vulnerable to global warming as elevated sea surface temperatures (SSTs) disrupt the delicate balance between the coral host and their algal endosymbionts, leading to symbiont expulsion, mass bleaching and mortality. While satellite sensing of SST has proved a reliable predictor of coral bleaching at the regional scale, there are large deviations in bleaching severity and mortality on the local scale that are poorly understood. Here, we show that internal waves play a major role in explaining local coral bleaching and mortality patterns in the Andaman Sea. Despite a severe region-wide SST anomaly in May 2010, frequent upslope intrusions of cold sub-pycnocline waters due to breaking large-amplitude internal waves (LAIW) mitigated coral bleaching and mortality in shallow waters. In LAIW-sheltered waters, by contrast, bleaching-susceptible species suffered severe bleaching and total mortality. These findings suggest that LAIW benefit coral reefs during thermal stress and provide local refugia for bleaching-susceptible corals. LAIW are ubiquitous in tropical stratified waters and their swash zones may thus be important conservation areas for the maintenance of coral diversity in a warming climate. Taking LAIW into account can significantly improve coral bleaching predictions and provide a valuable tool for coral reef conservation and management.

Metabolic plasticity of the corals Porites lutea and Diploastrea heliopora exposed to large amplitude internal waves

The metabolic plasticity of the two mounding coral species Porites lutea (Milne-Edwards and Haime, 1860) and Diploastrea heliopora (Lamarck, 1816) was investigated in the Similan Islands (Thailand), an offshore Andaman Sea island group subjected to large amplitude internal waves (LAIW). Nutrient concentrations were highly correlated with LAIW intensity and contributed to 3- and 10-fold higher symbiont densities in P. lutea and D. heliopora, respectively, along with elevated pigment concentrations, protein content, host tissue, and symbiont biomass. The comparison of LAIW-exposed and LAIW-sheltered island faces, and LAIW-intense and LAIW-weak years suggests a species-specific metabolic plasticity to LAIW, where D. heliopora benefits more from increased nutrient and organic matter availability than P. lutea. The ubiquitous LAIW in Southeast Asia and beyond may provide so far unexplored clues to coral acclimatization to disturbances on various scales, and hence, a potential key to coral resilience to climate change.

Differential Impact of Monsoon and Large Amplitude Internal Waves on Coral Reef Development in the Andaman Sea

The Andaman Sea and other macrotidal semi-enclosed tropical seas feature large amplitude internal waves (LAIW). Although LAIW induce strong fluctuations i.e. of temperature, pH, and nutrients, their influence on reef development is so far unknown. A better-known source of disturbance is the monsoon affecting corals due to turbulent mixing and sedimentation. Because in the Andaman Sea both, LAIW and monsoon, act from the same westerly direction their relative contribution to reef development is difficult to discern. Here, we explore the framework development in a number of offshore island locations subjected to differential LAIW- and SW-monsoon impact to address this open question. Cumulative negative temperature anomalies – a proxy for LAIW impact – explained a higher percentage of the variability in coral reef framework height, than sedimentation rates which resulted mainly from the monsoon. Temperature anomalies and sediment grain size provided the best correlation with framework height suggesting that so far neglected subsurface processes (LAIW) play a significant role in shaping coral reefs.

Internal pH regulation facilitates in situ long-term acclimation of massive corals to end-of-century carbon dioxide conditions

The resilience of tropical corals to ocean acidification depends on their ability to regulate the pH within their calcifying fluid (pHcf). Recent work suggests pHcf homeostasis under short-term exposure to pCO2 conditions predicted for 2100, but it is still unclear if pHcf homeostasis can be maintained throughout a corals lifetime. At CO2 seeps in Papua New Guinea, massive Porites corals have grown along a natural seawater pH gradient for decades. This natural gradient, ranging from pH 8.1–7.4, provides an ideal platform to determine corals’ pHcf (using boron isotopes). Porites maintained a similar pHcf (~8.24) at both a control (pH 8.1) and seep-influenced site (pH 7.9). Internal pHcf was slightly reduced (8.12) at seawater pH 7.6, and decreased to 7.94 at a site with a seawater pH of 7.4. A growth response model based on pHcf mirrors the observed distribution patterns of this species in the field. We suggest Porites has the capacity to acclimate after long-time exposure to end-of-century reduced seawater pH conditions and that strong control over pHcf represents a key mechanism to persist in future oceans. Only beyond end-of-century pCO2 conditions do they face their current physiological limit of pH homeostasis and pHcf begins to decrease.

Benthic reef primary production in response to large amplitude internal waves at the Similan Islands (Andaman Sea, Thailand).

Coral reefs are facing rapidly changing environments, but implications for reef ecosystem functioning and important services, such as productivity, are difficult to predict. Comparative investigations on coral reefs that are naturally exposed to differing environmental settings can provide essential information in this context. One prevalent phenomenon regularly introducing alterations in water chemistry into coral reefs are internal waves. This study therefore investigates the effect of large amplitude internal waves (LAIW) on primary productivity in coral reefs at the Similan Islands (Andaman Sea, Thailand). The LAIW-exposed west sides of the islands are subjected to sudden drops in water temperature accompanied by enhanced inorganic nutrient concentrations compared to the sheltered east. At the central island, Ko Miang, east and west reefs are only few hundred meters apart, but feature pronounced differences. On the west lower live coral cover (-38 %) coincides with higher turf algae cover (+64 %) and growth (+54 %) compared to the east side. Turf algae and the reef sand-associated microphytobenthos displayed similar chlorophyll a contents on both island sides, but under LAIW exposure, turf algae exhibited higher net photosynthesis (+23 %), whereas the microphytobenthos displayed reduced net and gross photosynthesis (-19 % and -26 %, respectively) accompanied by lower respiration (-42 %). In contrast, the predominant coral Porites lutea showed higher chlorophyll a tissues contents (+42 %) on the LAIW-exposed west in response to lower light availability and higher inorganic nutrient concentrations, but net photosynthesis was comparable for both sides. Turf algae were the major primary producers on the west side, whereas microphytobenthos dominated on the east. The overall primary production rate (comprising all main benthic primary producers) was similar on both island sides, which indicates high primary production variability under different environmental conditions.

Autotrophic and heterotrophic responses of the coral Porites lutea to large amplitude internal waves.

SUMMARY Large amplitude internal waves (LAIW) cause frequent and severe changes in the physico-chemical environment of Andaman Sea coral reefs and are a potentially important source of disturbance for corals. To explore the coral response to LAIW, prey capture disposition and photosynthesis were investigated in relation to changes in seawater temperature, pH, flow speed and food availability in LAIW simulation studies under controlled laboratory conditions, using Porites lutea as a model organism. Although food presence stimulated polyp expansion, we found an overriding effect of low temperature (19°C) causing retraction of the coral polyps into their calices, particularly when pH was altered concomitantly. Decreases in pH alone, however, caused the expansion of the polyps. The exposure history of the colonies played a crucial role in coral responses: prior field exposure to LAIW yielded lower retraction levels than in LAIW-inexperienced corals, suggesting acclimatization. Low temperature (19°C) exposure did not seem to influence the photosynthetic performance, but LAIW-experienced corals showed higher values of maximum dark-adapted quantum yield (Fv/Fm) of photosystem II than LAIW-inexperienced controls. Collectively, these data suggest that P. lutea, the dominant hermatypic coral in the Andaman Sea, can acclimatize to extreme changes in its abiotic environment by modulating its mixotrophic nutrition, through polyp expansion and potential feeding, as well as its photosynthetic efficiency.

Coral Growth and Bioerosion of Porites lutea in Response to Large Amplitude Internal Waves

The Similan Islands (Thailand) in the Andaman Sea are exposed to large amplitude internal waves (LAIW), as evidenced by i.a. abrupt fluctuations in temperature of up to 10°C at supertidal frequencies. Although LAIW have been shown to affect coral composition and framework development in shallow waters, the role of LAIW on coral growth is so far unknown. We carried out a long-term transplant experiment with live nubbins and skeleton slabs of the dominating coral Porites lutea to assess the net growth and bioerosion in LAIW-exposed and LAIW-protected waters. Depth-related, seasonal and interannual differences in LAIW-intensities on the exposed western sides of the islands allowed us to separate the effect of LAIW from other possible factors (e.g. monsoon) affecting the corals. Coral growth and bioerosion were inversely related to LAIW intensity, and positively related to coral framework development. Accretion rates of calcareous fouling organisms on the slabs were negligible compared to bioerosion, reflecting the lack of a true carbonate framework on the exposed W faces of the Similan Islands. Our findings show that LAIW may play an important, yet so far overlooked, role in controlling coral growth in tropical waters.

Reduced heterotrophy in the stony coral Galaxea fascicularis after life-long exposure to elevated carbon dioxide

Ocean acidification imposes many physiological, energetic, structural and ecological challenges to stony corals. While some corals may increase autotrophy under ocean acidification, another potential mechanism to alleviate some of the adverse effects on their physiology is to increase heterotrophy. We compared the feeding rates of Galaxea fascicularis colonies that have lived their entire lives under ocean acidification conditions at natural carbon dioxide (CO2) seeps with colonies living under present-day CO2 conditions. When provided with the same quantity and composition of zooplankton as food, corals acclimatized to high CO2 showed 2.8 to 4.8 times depressed rates of zooplankton feeding. Results were consistent over four experiments, from two expeditions and both in field and chamber measurements. Unless replenished by other sources, reduced zooplankton uptake in G. fascicularis acclimatized to ocean acidification is likely to entail a shortage of vital nutrients, potentially jeopardizing their health and survival in future oceans.

Large-amplitude internal waves sustain coral health during thermal stress

Ocean warming is a major threat for coral reefs causing widespread coral bleaching and mortality. Potential refugia are thus crucial for coral survival. Exposure to large-amplitude internal waves (LAIW) mitigated heat stress and ensured coral survival and recovery during and after an extreme heat anomaly. The physiological status of two common corals, Porites lutea and Pocillopora meandrina, was monitored in host and symbiont traits, in response to LAIW-exposure throughout the unprecedented 2010 heat anomaly in the Andaman Sea. LAIW-exposed corals of both species survived and recovered, while LAIW-sheltered corals suffered partial and total mortality in P. lutea and P. meandrina, respectively. LAIW are ubiquitous in the tropics and potentially generate coral refuge areas. As thermal stress to corals is expected to increase in a warming ocean, the mechanisms linking coral bleaching to ocean dynamics will be crucial to predict coral survival on a warming planet.

Gaining Ground—Cleaning with Mesenterial Filaments by the Cold-Water Coral Desmophyllum Dianthus?

The solitary cosmopolitan cold-water coral Desmophyllum dianthus (Esper, 1794) (A, scale bar approximately 1 cm) occurs as a deep-water emergent species in Patagonian fjords, Chile, and can form extensive banks (Forsterra et al. 2005). This coral functions as a bioengineer by providing a 3D habitat for a diverse benthic community (Forsterra et al. 2005). With skeletal growth, the living tissue is retracted by the organism to the upper part of the calyx, leaving the lower skeleton uncovered. Endolithic algae are then able to enter the tissue covered skeleton from below and start to encrust the tissue covered part of the skeleton (Forsterra et al. 2005, 2008). They are visible as a greenish coat on the skeleton (A). The algae are able to bore into the skeleton (Forsterra et al. 2005) and thus could interfere with the coral’s calcifying process. This process could result in a parasitic behavior (Forsterra et al. 2012) rather than the previously hypothesized (Forsterra and Haussermann 2008) symbiotic or even mutualistic relationship. This infection by algae may be the primary reason for the peculiar behavior we observed on several specimens both in situ (A) and in our aquaria system (B–D, scale bar approximately 1 cm). After an acclimatization time in our aquaria system, wild collected corals were well adapted and well fed, and secondarily expanded their tissue toward their lower skeleton (about 1 cm in 4 mo, CJ and GMS pers obs), recovering an area that had been without any tissue. Concomitantly, the corals everted their mesenterial filaments between this re-entered outer-skeleton and the covering tissue layer (A, B; C, D: both enlargements of B). During the eversion of the mesenterial filaments the new tissue generally appeared unusually pleated (C, D) in contrast to a less common smooth tissue surface (A). This was not a short-term or rare mechanism: corals maintained this behavior in the aquaria over multiple months. Several tropical coral species are known to eject their mesenterial filaments to digest prey too large to be swallowed (Yonge 1930). The tropical, branching coral Acropora pulchra (Brook, 1891) has even been observed to clean the substrate before it expands its tissue and skeleton (Roff et al. 2008). Our preliminary observations suggest a combination of both mechanisms in this cold-water coral. Mesenterial filaments could be digesting the growing algae below their tissue, but also may be cleaning the substrate (their own skeleton) prior to further tissue expansion.