Rapid relative increase of crustose coralline algae following herbivore exclusion in a reef of El Salvador.

The Eastern Tropical Pacific (ETP) is one of the most isolated and least studied regions in the world. This particularly applies to the coast of El Salvador, where the only reef between Guatemala and Nicaragua, called Los Cóbanos reef, is located. There is very little published information about the reef's biodiversity, and to our knowledge, no research on its ecology and responses to anthropogenic impacts, such as overfishing, has been conducted. The present study, therefore, described the benthic community of Los Cóbanos reef, El Salvador, using the Line-Point-Intercept-Transect method and investigated changes in the benthic community following the exclusion of piscine macroherbivores over a period of seven weeks. Results showed high benthic algae cover (up to 98%), dominated by turf and green algae, and low coral cover (0-4%). Porites lobata was the only hermatypic coral species found during the surveys. Surprisingly, crustose coralline algae (CCA) showed a remarkable total cover increase by 58%, while turf algae cover decreased by 82%, in experimental plots after seven weeks of piscine macroherbivore exclusion. These findings apparently contradict the results of most previous similar studies. While it was not possible to ascertain the exact mechanisms leading to these drastic community changes, the most likely explanation is grazing on turf by small grazing macroherbivores that had access to the cages during the experiment and clearing of CCA initially covered by epiphytes and sediments. A higher CCA cover would promote the succesful settlement by corals and prevent further erosion of the reef framework. Therefore it is crucial to better understand algal dynamics, herbivory, and implications of overfishing at Los Cóbanos to avoid further reef deterioration. This could be achieved through video surveys of the fish community, night-time observations of the macroinvertebrate community, exclusion experiments that also keep out herbivorous macroinvertebrates, and/or experimental assessments of turf algae/CCA interactions.

A meta-analysis to assess long-term spatiotemporal changes of benthic coral and macroalgae cover in the Mexican Caribbean.

Coral reefs in the wider Caribbean declined in hard coral cover by ~80% since the 1970s, but spatiotemporal analyses for sub-regions are lacking. Here, we explored benthic change patterns in the Mexican Caribbean reefs through meta-analysis between 1978 and 2016 including 125 coral reef sites. Findings revealed that hard coral cover decreased from ~26% in the 1970s to 16% in 2016, whereas macroalgae cover increased to ~30% in 2016. Both groups showed high spatiotemporal variability. Hard coral cover declined in total by 12% from 1978 to 2004 but increased again by 5% between 2005 and 2016 indicating some coral recovery after the 2005 mass bleaching event and hurricane impacts. In 2016, more than 80% of studied reefs were dominated by macroalgae, while only 15% were dominated by hard corals. This stands in contrast to 1978 when all reef sites surveyed were dominated by hard corals. This study is among the first within the Caribbean region that reports local recovery in coral cover in the Caribbean, while other Caribbean reefs have failed to recover. Most Mexican Caribbean coral reefs are now no longer dominated by hard corals. In order to prevent further reef degradation, viable and reliable conservation alternatives are required.

Ongoing removals of invasive lionfish in Honduras and their effect on native Caribbean prey fishes.

The invasion of Indo-Pacific lionfish is one of the most pressing concerns in the context of coral reef conservation throughout the Caribbean. Invasive lionfish threaten Caribbean fish communities by feeding on a wide range of native prey species, some of which have high ecological and economic value. In Roatan (Honduras) a local non-governmental organisation (i.e. Roatan Marine Park) trains residents and tourists in the use of spears to remove invasive lionfish. Here, we assess the effectiveness of local removal efforts in reducing lionfish populations. We ask whether reefs subject to relatively frequent removals support more diverse and abundant native fish assemblages compared to sites were no removals take place. Lionfish biomass, as well as density and diversity of native prey species were quantified on reefs subject to regular and no removal efforts. Reefs subject to regular lionfish removals (two to three removals month-1) with a mean catch per unit effort of 2.76 ± 1.72 lionfish fisher-1 h-1 had 95% lower lionfish biomass compared to non-removal sites. Sites subject to lionfish removals supported 30% higher densities of native prey-sized fishes compared to sites subject to no removal efforts. We found no evidence that species richness and diversity of native fish communities differ between removal and non-removal sites. We conclude that opportunistic voluntary removals are an effective management intervention to reduce lionfish populations locally and might alleviate negative impacts of lionfish predation. We recommend that local management and the diving industry cooperate to cost-effectively extend the spatial scale at which removal regimes are currently sustained.

Dynamics in benthic community composition and influencing factors in an upwelling-exposed coral reef on the Pacific coast of Costa Rica.

Seasonal upwelling at the northern Pacific coast of Costa Rica offers the opportunity to investigate the effects of pronounced changes in key water parameters on fine-scale dynamics of local coral reef communities. This study monitored benthic community composition at Matapalo reef (10.539°N, 85.766°W) by weekly observations of permanent benthic quadrats from April 2013 to April 2014. Monitoring was accompanied by surveys of herbivore abundance and biomass and measurements of water temperature and inorganic nutrient concentrations. Findings revealed that the reef-building corals Pocillopora spp. exhibited an exceptional rapid increase from 22 to 51% relative benthic cover. By contrast, turf algae cover decreased from 63 to 24%, resulting in a corresponding increase in crustose coralline algae cover. The macroalga Caulerpa sertularioides covered up to 15% of the reef in April 2013, disappeared after synchronized gamete release in May, and subsequently exhibited slow regrowth. Parallel monitoring of influencing factors suggest that C. sertularioides cover was mainly regulated by their reproductive cycle, while that of turf algae was likely controlled by high abundances of herbivores. Upwelling events in February and March 2014 decreased mean daily seawater temperatures by up to 7 °C and increased nutrient concentrations up to 5- (phosphate) and 16-fold (nitrate) compared to mean values during the rest of the year. Changes in benthic community composition did not appear to correspond to the strong environmental changes, but rather shifted from turf algae to hard coral dominance over the entire year of observation. The exceptional high dynamic over the annual observation period encourages further research on the adaptation potential of coral reefs to environmental variability.

Effects of Seasonal Upwelling on Inorganic and Organic Matter Dynamics in the Water Column of Eastern Pacific Coral Reefs.

The Gulf of Papagayo at the northern Pacific coast of Costa Rica experiences pronounced seasonal changes in water parameters caused by wind-driven coastal upwelling. While remote sensing and open water sampling already described the physical nature of this upwelling, the spatial and temporal effects on key parameters and processes in the water column have not been investigated yet, although being highly relevant for coral reef functioning. The present study investigated a range of water parameters on two coral reefs with different exposure to upwelling (Matapalo and Bajo Rojo) in a weekly to monthly resolution over one year (May 2013 to April 2014). Based on air temperature, wind speed and water temperature, three time clusters were defined: a) May to November 2013 without upwelling, b) December 2013 to April 2014 with moderate upwelling, punctuated by c) extreme upwelling events in February, March and April 2014. During upwelling peaks, water temperatures decreased by 7°C (Matapalo) and 9°C (Bajo Rojo) to minima of 20.1 and 15.3°C respectively, while phosphate, ammonia and nitrate concentrations increased 3 to 15-fold to maxima of 1.3 µmol PO43- L-1, 3.0 µmol NH4+ L-1 and 9.7 µmol NO3- L-1. This increased availability of nutrients triggered several successive phytoplankton blooms as indicated by 3- (Matapalo) and 6-fold (Bajo Rojo) increases in chlorophyll a concentrations. Particulate organic carbon and nitrogen (POC and PON) increased by 40 and 70% respectively from February to April 2014. Dissolved organic carbon (DOC) increased by 70% in December and stayed elevated for at least 4 months, indicating high organic matter release by primary producers. Such strong cascading effects of upwelling on organic matter dynamics on coral reefs have not been reported previously, although likely impacting many reefs in comparable upwelling systems.

Turf algae-mediated coral damage in coastal reefs of Belize, Central America.

Many coral reefs in the Caribbean experienced substantial changes in their benthic community composition during the last decades. This often resulted in phase shifts from scleractinian coral dominance to that by other benthic invertebrate or algae. However, knowledge about how the related role of coral-algae contacts may negatively affect corals is scarce. Therefore, benthic community composition, abundance of algae grazers, and the abundance and character of coral-algae contacts were assessed in situ at 13 Belizean reef sites distributed along a distance gradient to the Belizean mainland (12-70 km): Mesoamerican Barrier Reef (inshore), Turneffe Atoll (inner and outer midshore), and Lighthouse Reef (offshore). In situ surveys revealed significantly higher benthic cover by scleractinian corals at the remote Lighthouse Reef (26-29%) when compared to the other sites (4-19%). The abundance of herbivorous fish and the sea urchin Diadema antillarum significantly increased towards the offshore reef sites, while the occurrence of direct coral-algae contacts consequently increased significantly with decreasing distance to shore. About 60% of these algae contacts were harmful (exhibiting coral tissue damage, pigmentation change, or overgrowth) for corals (mainly genera Orbicella and Agaricia), particularly when filamentous turf algae were involved. These findings provide support to the hypothesis that (turf) algae-mediated coral damage occurs in Belizean coastal, near-shore coral reefs.

Benthic primary production in an upwelling-influenced coral reef, Colombian Caribbean.

In Tayrona National Natural Park (Colombian Caribbean), abiotic factors such as light intensity, water temperature, and nutrient availability are subjected to high temporal variability due to seasonal coastal upwelling. These factors are the major drivers controlling coral reef primary production as one of the key ecosystem services. This offers the opportunity to assess the effects of abiotic factors on reef productivity. We therefore quantified primary net (Pn ) and gross production (Pg ) of the dominant local primary producers (scleractinian corals, macroalgae, algal turfs, crustose coralline algae, and microphytobenthos) at a water current/wave-exposed and-sheltered site in an exemplary bay of Tayrona National Natural Park. A series of short-term incubations was conducted to quantify O2 fluxes of the different primary producers during non-upwelling and the upwelling event 2011/2012, and generalized linear models were used to analyze group-specific O2 production, their contribution to benthic O2 fluxes, and total daily benthic O2 production. At the organism level, scleractinian corals showed highest Pn and Pg rates during non-upwelling (16 and 19 mmol O2 m(-2) specimen area h(-1)), and corals and algal turfs dominated the primary production during upwelling (12 and 19 mmol O2 m(-2) specimen area h(-1), respectively). At the ecosystem level, corals contributed most to total Pn and Pg during non-upwelling, while during upwelling, corals contributed most to Pn and Pg only at the exposed site and macroalgae at the sheltered site, respectively. Despite the significant spatial and temporal differences in individual productivity of the investigated groups and their different contribution to reef productivity, differences for daily ecosystem productivity were only present for Pg at exposed with higher O2 fluxes during non-upwelling compared to upwelling. Our findings therefore indicate that total benthic primary productivity of local autotrophic reef communities is relatively stable despite the pronounced fluctuations of environmental key parameters. This may result in higher resilience against anthropogenic disturbances and climate change and Tayrona National Natural Park should therefore be considered as a conservation priority area.

Spatial and temporal distribution of the invasive lionfish Pterois volitans in coral reefs of Tayrona National Natural Park, Colombian Caribbean.

The lionfish Pterois volitans is an invasive species throughout the Western Atlantic that disturbs functioning of local ecosystems such as coral reefs via fast and intense consumption of small fish and invertebrates. In 2009, lionfish populated the bays of Tayrona National Natural Park (TNNP), a biodiversity hotspot in the Colombian Caribbean that is strongly influenced by changing environmental conditions due to a rainy and dry season. So far, the spatial and temporal distribution of P. volitans in the bays of TNNP is unknown. Therefore, this study assessed the abundance and body lengths of P. volitans during monthly surveys throughout the year 2012 in four bays (thereof two bays where lionfish removals were undertaken) of TNNP at 10 m water depth in coral reefs using transect tools. Findings revealed lionfish abundances of 2.9 ± 0.9 individuals ha(-1) with lengths of 20-25 cm for TNNP, hinting to an established, mostly adult local population. Actual TNNP lionfish abundances are thereby very similar to those at Indo-Pacific reef locations where the invasive lionfish formerly originated from. Significant spatial differences for lionfish abundances and body lengths between different bays in TNNP suggest habitat preferences of P. volitans depending on age. Lionfish abundances were highly variable over time, but without significant differences between seasons. Removals could not reduce lionfish abundances significantly during the period of study. This study therefore recommends improved management actions in order to control the already established invasive lionfish population in TNNP.

Spatial and temporal variability of water quality in the coral reefs of Tayrona National Natural Park, Colombian Caribbean.

Tayrona National Natural Park (TNNP) is a hotspot of coral reef biodiversity in the Colombian Caribbean, located between the city of Santa Marta (>455,000 inhabitants) and several smaller river mouths (Rio Piedras, Mendihuaca, Guachaca). The region also experiences a strong seasonal variation in physical parameters (temperature, salinity, wind, and water currents) due to alternating dry seasons with coastal upwelling and rainy seasons. However, the spatial and temporal effects on water quality parameters relevant for coral reef functioning have not been investigated. Therefore, inorganic nutrient, chlorophyll a, and particulate organic carbon (POC) concentrations along with biological O2 demand (BOD), pH, and water clarity directly above local coral reefs (~10 m water depth) were monitored for 25 months in four bays along a distance gradient (12-20 km) to Santa Marta in the southwest and to the first river mouth (17-27 km) in the east. This is by far the most comprehensive coral reefs water quality dataset for the region. Findings revealed that particularly during non-upwelling, chlorophyll a and POC concentrations along with BOD significantly increased with decreasing distance to the rivers in the east, suggesting that the observed spatial water quality decline was triggered by riverine runoff and not by the countercurrent-located Santa Marta. Nitrate, nitrite, and chlorophyll a concentrations significantly increased during upwelling, while pH and water clarity decreased. Generally, water quality in TNNP was close to oligotrophic conditions adequate for coral reef growth during non-upwelling, but exceeded critical threshold values during upwelling and in relation to riverine discharge.

Benthic primary production budget of a Caribbean reef lagoon (Puerto Morelos, Mexico).

High photosynthetic benthic primary production (P) represents a key ecosystem service provided by tropical coral reef systems. However, benthic P budgets of specific ecosystem compartments such as macrophyte-dominated reef lagoons are still scarce. To address this, we quantified individual and lagoon-wide net (Pn) and gross (Pg) primary production by all dominant functional groups of benthic primary producers in a typical macrophyte-dominated Caribbean reef lagoon near Puerto Morelos (Mexico) via measurement of O2 fluxes in incubation experiments. The photosynthetically active 3D lagoon surface area was quantified using conversion factors to allow extrapolation to lagoon-wide P budgets. Findings revealed that lagoon 2D benthic cover was primarily composed of sand-associated microphytobenthos (40%), seagrasses (29%) and macroalgae (27%), while seagrasses dominated the lagoon 3D surface area (84%). Individual Pg was highest for macroalgae and scleractinian corals (87 and 86 mmol O2 m(-2) specimen area d(-1), respectively), however seagrasses contributed highest (59%) to the lagoon-wide Pg. Macroalgae exhibited highest individual Pn rates, but seagrasses generated the largest fraction (51%) of lagoon-wide Pn. Individual R was highest for scleractinian corals and macroalgae, whereas seagrasses again provided the major lagoon-wide share (68%). These findings characterise the investigated lagoon as a net autotrophic coral reef ecosystem compartment revealing similar P compared to other macrophyte-dominated coastal environments such as seagrass meadows and macroalgae beds. Further, high lagoon-wide P (Pg: 488 and Pn: 181 mmol O2 m(-2) lagoon area d(-1)) and overall Pg:R (1.6) indicate substantial benthic excess production within the Puerto Morelos reef lagoon and suggest the export of newly synthesised organic matter to surrounding ecosystems.

Bleaching susceptibility and recovery of Colombian Caribbean corals in response to water current exposure and seasonal upwelling.

Coral bleaching events are globally occurring more frequently and with higher intensity, mainly caused by increases in seawater temperature. In Tayrona National Natural Park (TNNP) in the Colombian Caribbean, local coral communities are subjected to seasonal wind-triggered upwelling events coinciding with stronger water currents depending on location. This natural phenomenon offers the unique opportunity to study potential water current-induced mitigation mechanisms of coral bleaching in an upwelling influenced region. Therefore, coral bleaching susceptibility and recovery patterns were compared during a moderate and a mild bleaching event in December 2010 and 2011, and at the end of the subsequent upwelling periods at a water current-exposed and -sheltered site of an exemplary bay using permanent transect and labeling tools. This was accompanied by parallel monitoring of key environmental variables. Findings revealed that in 2010 overall coral bleaching before upwelling was significantly higher at the sheltered (34%) compared to the exposed site (8%). Whereas 97% of all previously bleached corals at the water current-exposed site had recovered from bleaching by April 2011, only 77% recovered at the sheltered site, but 12% had died there. In December 2011, only mild bleaching (<10% at both sites) was observed, but corals recovered significantly at both sites in the course of upwelling. No differences in water temperatures between sites occurred, but water current exposure and turbidity were significantly higher at the exposed site, suggesting that these variables may be responsible for the observed site-specific mitigation of coral bleaching. This indicates the existence of local resilience patterns against coral bleaching in Caribbean reefs.

Sources and spatial distribution of heavy metals in scleractinian coral tissues and sediments from the Bocas del Toro Archipelago, Panama.

Marine ecosystems worldwide are threatened by aquatic pollution; however, there is a paucity in data from the Caribbean region. As such, five heavy metals (arsenic, cadmium, copper, zinc, mercury) were measured in tissues of the scleractinian corals Porites furcata and Agaricia tenuifolia and in adjacent sediments in the Bocas del Toro Archipelago, Panama. Samples were collected from five reef sites along a gradient of distance from an international shipping port and were analysed using inductively coupled plasma optical emission spectrometry and atomic absorption spectrophotometry for mercury. Copper and zinc were the most abundant metals and ranged from 11 to 63 mg kg(-1) and from 31 to 185 mg kg(-1) in coral tissues, respectively. The highest concentration of each metal was measured in P. furcata tissues, with copper and mercury concentrations significantly higher in P. furcata than in A. tenuifolia at every site. These results suggest that P. furcata has a higher affinity for metal accumulation and storage than A. tenuifolia. With the exception of cadmium, metal concentrations in coral tissues were generally elevated at coral reefs in closer proximity to the port; however, this pattern was not observed in sediments. Hard coral cover was lowest at reefs in closest proximity to the port, suggesting that metal pollution from port-related activities is influencing hard coral abundance at nearby coral reefs.