Nieuwland, G.

Over the past decades numerous studies have reported declines in stony corals and, in many cases, phase shifts to fleshy macroalgae. However, long-term studies documenting changes in other benthic reef organisms are scarce. Here, we studied changes in cover of corals, algal turfs, benthic cyanobacterial mats, macroalgae, sponges and crustose coralline algae at four reef sites of the Caribbean islands of Curaçao and Bonaire over a time span of 40 yr. Permanent 9 m2quadrats at 10, 20, 30 and 40 m depth were photographed at 3- to 6-yr intervals from 1973 to 2013. The temporal and spatial dynamics in the six dominant benthic groups were assessed based on image point-analysis. Our results show consistent patterns of benthic community change with a decrease in the cover of calcifying organisms across all sites and depths from 32.6 (1973) to 9.2% (2013) for corals and from 6.4 to 1% for crustose coralline algae. Initially, coral cover was replaced by algal turfs increasing from 24.5 (1973) to 38% around the early 1990s. Fleshy macroalgae, still absent in 1973, also proliferated covering 12% of the substratum approximately 20 yr later. However, these new dominants largely declined in abundance from 2002 to 2013 (11 and 2%, respectively), marking the rise of benthic cyanobacterial mats. Cyanobacterial mats became the most dominant benthic component increasing from a mere 7.1 (2002) to 22.2% (2013). The observed increase was paralleled by a small but significant increase in sponge cover (0.5 to 2.3%). Strikingly, this pattern of degradation and phase change occurred over the reef slope down to mesophotic depths of 40 m. These findings suggest that reefs dominated by algae may be less stable than previously thought and that the next phase may be the dominance of slimy cyanobacterial mats with some sponges.

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 Curaçao 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 toward smaller coral colonies, reef complexity. Alarmingly, reefs at upper-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.

Reef slope coral communities were surveyed for long-term (20 years) changes in scleractinian coral cover, numbers of coral colonies and species richness, over the time intervals between the years 1973, 1983 and 1992. We compare such long-term structural changes in the communities at depths of 10, 20, 30 and 40 m. Our data are based on series of photographic records of permanent quadrats, a total of 36 m2 reef bottom at each depth, along four transects on the leeward coasts of the islands of Curacao and Bonaire. We summarize the changes in the permanent quadrats over time to demonstrate the main trends in the data set and, to understand the significance of the data for the reef community, test the results as effects of time and depth using mixed model ANOVA’s. Changes in numbers of coral colonies and coral cover were a function of depth. Number of coral colonies decreased significantly at depths of 10, 20 and 30 m, but not at 40 m. Coral cover decreased significantly at 10 and 20 m, but not at 30 and 40 m. Diversity (species richness) decreased through the years independent of depth. There were no consistent differences between the two 10-year time-intervals. These results confirm earlier observations of coral mortality and spatial mobility which showed the deep reef (30, 40 m) as a much more constant environment than the relatively disturbed shallower reef (10, 20 m).

Linear extension of branches in the same Acropora palmata (Lamarck, 1816) population in Curaçao was measured, employing exactly the same methods, in 1971–1973 and in 2002–2004, and the resulting coral growth rates are compared. Linear growth shows the same pattern over seasons in both periods with growth being significantly higher in summer than in winter. Growth in the 2002–2004 time interval was significantly slower than in 1971–1973. Mean monthly growth ranged from 0.69 cm (winter) to 0.81 cm (summer) in 1971–1973 and from 0.62 cm (winter) to 0.75 cm (summer) in 2002–2004. This means that linear growth rates in 2002–2004 were 7.2% lower in summer and 10.7% lower in winter compared with 1971–1973. Considering possible causative environmental factors relating to these decreases in growth rate, we cannot preclude the possibility that a change in ocean pH could be responsible for the drop in extension rate.

Coral reefs are thought to be in worldwide decline but available data are practically limited to reefs shallower than 25 m. Zooxanthellate coral communities in deep reefs (30–40 m) are relatively unstudied. Our question is: what is happening in deep reefs in terms of coral cover and coral mortality? We compare changes in species composition, coral mortality, and coral cover at Caribbean (Curacao and Bonaire) deep (30–40 m) and shallow reefs (10–20 m) using long-term (1973–2002) data from permanent photo quadrats. About 20 zoo- xanthellate coral species are common in the deep-reef communities, dominated by Agaricia sp., with coral cover up to 60%. In contrast with shallow reefs, there is no decrease in coral cover or number of coral colonies in deep reefs over the last 30 years. In deep reefs, non- agaricid species are decreasing but agaricid domination will be interrupted by natural catastrophic mortality such as deep coral bleaching and storms. Temperature is a vastly fluctuating variable in the deep-reef environ- ment with extremely low temperatures possibly related to deep-reef bleaching.