Erik H. Meesters

Sub-rubble communities of Curacao and Bonaire coral reefs

The distribution and abundance of sessile organisms under coral rubble has been studied at Bonaire and Curaçao, Netherlands Antilles. Species richness under rubble is extremely high with at least 367 species of which sponges, tunicates and bryozoans are the most important. Shallow sub-rubble communities can be considered refuges as the majority of these species are crypt-obligate. Sub-rubble communities may also have a preserve function for sponges, but do not harbour enough corals to ensure a quick coral recolonization of the reef surface after a major disaster. Cryptic community composition is affected by depth and pollution, and differs substantially between the two neighbouring islands, possibly as a result of different bottom characteristics. Biomass of the sub-rubble communities may contribute considerably to total reef biomass. Diversity varies inversely with increased depth and increased rubble size, possibly indicating abiotic control (e.g. physical disturbance by wave action and reef slope substrate collapse).

Coral cavity sponges depend on reef-derived food resources: stable isotope and fatty acid constraints

The diet of cavity sponges on the narrow fringing reefs of Curaçao, Caribbean was studied. The origin and resources of the bulk food of these sponges, i.e., dissolved organic matter (DOM), were identified using stable carbon and nitrogen isotopes and fatty acid biomarkers. We found that phytoplankton and its derived DOM from the adjacent open sea and from reef overlying water is not the main source of food for most of the sponges examined nor is bacterioplankton. Interestingly, dual stable isotope signatures (δ13Corg, δ15Norg) and fatty acid biomarkers appoint coral mucus and organic matter derived from crustose coralline algae (CCA) as probable food sources for encrusting sponges. Mucus-derived DOM may contribute up to 66% to the diet of examined sponges based on results of dual isotope mixing model analysis. The contribution of CCA (as purported representative for benthic algae) was smaller with values up to 31%. Together, mucus- and CCA-derived substrates contributed for 48–73% to the diet of sponges. The presence of the exogenous fatty acid 20:4ω6 in sponges, which is abundant in coral mucus of Madracis mirabilis and in CCA, highlights these reef-derived resources as sources of nutrition for DOM feeding cavity sponges. The relatively high concentrations of exogenous 20:4ω6 in all sponges examined supports our hypothesis that the bulk of the food of the cavity sponge community is reef-derived. Our results imply that cavity sponges play an important role in conserving food and energy produced within the reef.

Long-term Shifts in Coral Communities On Shallow to Deep Reef Slopes of Curaçao and Bonaire: Are There Any Winners?

Tropical coral reefs are among the most biologically diverse and economically important ecosystems on earth. Nevertheless, we found dramatic changes in coral communities on the reef slopes of Curacao and Bonaire since 1973. Cover and abundance declined for virtually all coral species. The data show a shift from communities dominated by framework building species (e.g. Orbicella spp.) to communities consisting of small opportunistic, phenotypically plastic, species, including few remaining structural colonies. Madracis mirabilis, Porites astreoides, Diploria strigosa and Agaricia lamarcki are at present modest winners in the coral assemblage, although overall cover declined also for these species. Increased frequency and intensity of events inducing coral mortality and ongoing reduction in suitable hard substratum, provided by the remnants of large colony building species, could reduce the chance of these species to remain winners in the longer run. The observed loss in coral cover and the shift from larger structural to smaller opportunistic species reduced reef carbonate production by 67% and therewith, in combination with a trend towards smaller coral colonies, reef complexity. Alarmingly, reefs at mesophotic depths (30 - 40 m) did not escape the general degradation of the coral community. The negative effects are larger around densely populated areas where local stressors are adding to degradation caused, for instance, by region wide mass bleaching. Without proper conservation and management this already dramatic degradation will continue and turn more and more coral species into losers.

Caribbean free-living coral species co-occurring deep off the windward coast of Curaçao

We document the co-occurrence of two Caribbean species of free-living zooxanthellate corals, Meandrina danae (Milne-Edwards and Haime 1848) (Fig. 1a) and Manicina areolata (Linnaeus 1758) (Fig. 1b), in a previously not well-documented environment. The two species were encountered at three locations (ranging from 30 to 50 m depth) on the windward side of the island of Curaçao, southern Caribbean. Corals are present beyond the reef base on substrate consisting of coarse coral sand, small rubble, and fragments of shells and calcareous algae (Fig. 1c). Colony sizes (mean length) were 58 mm (SD 18, n = 133) for M. areolata and 67 mm (SD 19, n = 32) for M. danae. Maximum colony density of both species together was 30 individuals per m 2 with M. areolata colonies being 9 times as abundant. None of these species has been reported from such a deep habitat. M. danae has never previously been recorded on the island while M. areolata has been described in shallow sandy areas in Curaçao by Roos (1971). It was reported in seagrass beds and on sandy and rubble areas on the reef down to 20 m in Jamaica (Goreau and Goreau 1960) and in seagrass flats and mangrove-associated communities in the San Blas Archipelago, northern Panama (Johnson 1992). Furthermore, free-living scleractinian corals can form dense and species-rich assemblages on sandy habitats of Indo-Pacific reef systems (Hoeksema 2012). To our knowledge, this is the first documented multi-species assemblage of free-living zooxanthellate corals in the Caribbean.

Spatio–temporal variation in stable isotope signatures (δ 13 C and δ 15 N) of sponges on the Saba Bank

Sponges are ubiquitous on coral reefs, mostly long lived and therefore adaptive to changing environmental conditions. They feed on organic matter withdrawn from the passing water and they may harbor microorganisms (endosymbionts), which contribute to their nutrition. Their diets and stable isotope (SI) fractionation determine the SI signature of the sponge holobiont. Little is known of spatio–temporal variations in SI signatures of δ13C and δ15N in tropical sponges and whether they reflect variations in the environment. We investigated the SI signatures of seven common sponge species with different functional traits and their potential food sources between 15 and 32 m depth along the S-SE and E-NE side of the Saba Bank, Eastern Caribbean, in October 2011 and October 2013. SI signatures differed significantly between most sponge species, both in mean values and in variation, indicating different food preferences and/or fractionation, inferring sponge species-specific isotopic niche spaces. In 2011, all sponge species at the S-SE side were enriched in d13C compared to the E-NE side. In 2013, SI signatures of sponges did not differ between the two sides and were overall lighter in δ13C and δ15N than in 2011. Observed spatio–temporal changes in SI in sponges could not be attributed to changes in the SI signatures of their potential food sources, which remained stable with different SI signatures of pelagic (particulate organic matter (POM): δ13C −24.9‰, δ15N +4.3‰) and benthic-derived food (macroalgae: δ13C −15.4‰, δ15N +0.8‰). Enriched δ13C signatures in sponges at the S-SE side in 2011 are proposed to be attributed to predominantly feeding on benthic-derived C. This interpretation was supported by significant differences in water mass constituents between sides in October 2011. Elevated NO3 and dissolved organic matter concentrations point toward a stronger reef signal in reef overlying water at the S-SE than N-NE side of the Bank in 2011. The depletions of δ13C and δ15N in sponges in October 2013 compared to October 2011 concurred with significantly elevated POM concentrations. The contemporaneous decrease in δ15N suggests that sponges obtain their N mostly from benthic-derived food with a lower δ15N than pelagic food. Average proportional feeding on available sources varied between sponge species and ranged from 20% to 50% for benthic and 50% to 80% for pelagic-derived food, assuming trophic enrichment factors of 0.5‰ ± sd 0.5 for δ13C and 3‰ ± sd 0.5 for δ15N for sponges. We suggest that observed variation of SI in sponges between sides and years were the result of shifts in the proportion of ingested benthic- and pelagic-derived organic matter driven by environmental changes. We show that sponge SI signatures reflect environmental variability in space and time on the Saba Bank and that SI of sponges irrespective of their species-specific traits move in a similar direction in response to these environmental changes.

Quantification of chemical and mechanical bioerosion rates of six Caribbean excavating sponge species found on the coral reefs of Curaçao

Excavating sponges are among the most important macro-eroders of carbonate substrates in marine systems. Their capacity to remove substantial amounts of limestone makes these animals significant players that can unbalance the reef carbonate budget of tropical coral reefs. Nevertheless, excavating sponges are currently rarely incorporated in standardized surveys and experimental work is often restricted to a few species. Here were provide chemical and mechanical bioerosion rates for the six excavating sponge species most commonly found on the shallow reef of Curaçao (southern Caribbean): Cliona caribbaea, C. aprica, C. delitrix, C. amplicavata, Siphonodictyon brevitubulatum and Suberea flavolivescens. Chemical, mechanical and total bioerosion rates were estimated based on various experimental approaches applied to sponge infested limestone cores. Conventional standing incubation techniques were shown to strongly influence the chemical dissolution signal. Final rates, based on the change in alkalinity of the incubation water, declined significantly as a function of incubation time. This effect was mitigated by the use of a flow-through incubation system. Additionally, we found that mechanically removed carbonate fragments collected in the flow-through chamber (1 h) as well as a long-term collection method (1 wk) generally yielded comparable estimates for the capacity of these sponges to mechanically remove substratum. Observed interspecific variation could evidently be linked to the adopted boring strategy (i.e. gallery-forming, cavity-forming or network-working) and presence or absence of symbiotic zooxanthellae. Notably, a clear diurnal pattern was found only in species that harbour a dense photosymbiotic community. In these species chemical erosion was substantially higher during the day. Overall, the sum of individually acquired chemical and mechanical erosion using flow-through incubations was comparable to rates obtained gravimetrically. Such consistency is a first in this field of research. These findings support the much needed confirmation that, depending on the scientific demand, the different approaches presented here can be implemented concurrently as standardized methods.

DOC concentrations across a depth-dependent light gradient on a Caribbean coral reef

Photosynthates released by benthic primary producers (BPP), such as reef algae and scleractinian corals, fuel the dissolved organic carbon (DOC) production on tropical coral reefs. DOC concentrations near BPP have repeatedly been observed to be elevated compared to those in the surrounding water column. As the DOC release of BPP increases with increasing light availability, elevated DOC concentrations near them will, in part, also depend on light availability. Consequently, DOC concentrations are likely to be higher on the shallow, well-lit reef terrace than in deeper sections on the fore reef slope. We measured in situ DOC concentrations and light intensity in close proximity to the reef alga Dictyota sp. and the scleractinian coral Orbicella faveolata along a depth-dependent light gradient from 5 to 20 m depth and compared these to background concentrations in the water column. At 10 m (intermediate light), DOC concentrations near Dictyota sp. were elevated by 15 µmol C L−1 compared to background concentrations in the water column, but not at 5 and 20 m (high and low light, respectively), or near O. faveolata at any of the tested depths. DOC concentrations did not differ between depths and thereby light environments for any of the tested water types. However, water type and depth appear to jointly affect in situ DOC concentrations across the tested depth-dependent light gradient. Corroborative ex situ measurements of excitation pressure on photosystem II suggest that photoinhibition in Dictyota sp. is likely to occur at light intensities that are commonly present on Curaçaoan coral reefs under high light levels at 5 m depth during midday. Photoinhibition may have thereby reduced the DOC release of Dictyota sp. and DOC concentrations in its close proximity. Our results indicate that the occurrence of elevated DOC concentrations did not follow a natural light gradient across depth. Instead, a combination of multiple factors, such as water type, light availability (including the restriction by photoinhibition), and water movement are proposed to interactively determine the DOC concentrations in the close vicinity of BPP.