*Dirección para correspondencia

Abstract

Thermal anomalies have become more severe, frequent and well-documented across the Caribbean for the past 30 years. This increase in temperature has caused coral bleaching resulting in reef decline. At Los Roques National Park, Venezuela, temperature has been monitored at four reef sites. In mid-September 2010, seawater temperature reached 30.85°C at 5 m depth in Los Roques, an archipelago only slightly affected by previous bleaching events. For example, bleaching in Los Roques in 2005 was mild compared to the rest of the Caribbean and to the results in this study. In 2010, seawater temperatures remained above 29.0°C from mid-August until the first week of November, resulting in +16 Degree Heating Weeks by that time. Our annual survey of four reef sites indicated that 72% of 563 scleractinian colonies were partial or totally bleached (white) or pale (discolored) in October 2010. In February 2011, there were still 46% of coral colonies affected; but most of them were pale and only 2% were bleached. By February, coral cover had declined 4 to 30% per transect, with a mean of 14.3%. Thus, mean coral cover dropped significantly from 45 to 31% cover (a 34% reduction). In addition to bleaching, corals showed a high prevalence (up to 16%) of black band disease in October 2010 and of white plague (11%) in February 2011. As a consequence, coral mortality is expected to be larger than reported here. Reef surveys since 2002 and personal observations for more than 20 years indicated that this bleaching event and its consequences in Los Roques have no precedent. Our results suggest that reef sites with no previous record of significant deterioration are more likely to become affected by thermal anomalies. However, this archipelago is relatively unaffected by local anthropogenic disturbance and has a high coral recruitment, which may contribute to its recovery. Rev. Biol. Trop. 60 (Suppl. 1): 29-37. Epub 2012 March 01.

Key words: coral bleaching, mortality, loss of coral cover, Caribbean, coral reefs.

Resumen

Durante las últimas décadas las anomalías térmicas han sido más frecuentes y severas en el Caribe, quedando pocos arrecifes exentos de eventos masivos de blanqueamiento (EMB). En el Parque Nacional Los Roques, Venezuela, un archipiélago poco afectado previamente por EMB, la temperatura del agua a 5m de profundidad alcanzó 30,85°C en septiembre 2010, y fue >29,0°C entre mediados de agosto y la primera semana de noviembre en cuatro arrecifes. El 72% de 563 colonias de escleractinios estaban blanqueadas o pálidas para octubre de 2010, mientras que para febrero 2011, el 46% de las colonias aún estaban afectadas. Para febrero 2011, la cobertura béntica coralina promedio disminuyó de 45 a 31%. Además, los arrecifes mostraron una alta prevalencia (de hasta 16%) de enfermedad de banda negra en Octubre 2010, y de plaga blanca (11%) en Febrero 2011. Como consecuencia, es probable que la mortalidad coralina resulte mayor a la reportada acá. Sin embargo, Los Roques es poco afectado por perturbaciones antropogénicas y cuenta con un alto reclutamiento de corales, lo cual podría contribuir a su recuperación.

Palabras clave: blanqueamiento coralino, mortalidad, pérdida de cobertura de coral, Caribe, arrecife coralino.

Coral reefs are tropical ecosystems with high productivity and therefore they provide a myriad of ]]> et al. 2004, Rogers 2009). In the Caribbean this phenomenon has been particularly evident, as extensive reductions of coral cover and the concomitant loss of architectural complexity have occurred (Gardner et ]]> 2003, Weil et al. 2006, Alvarez-Filip et al. 2009).

Mass bleaching events (MBE, events that reduce the density of Symbiodinium in its hosts over a large geographic area) have increased in frequency and intensity (e.g. ]]> et al. 2005); and it appears that they act synergistically with other detrimental factors to produce negative consequences for coral reef organisms, populations and communities (reviewed in Brown 1997). Thus, MBE have been reported across the Caribbean as a factor that reduces coral cover (e.g. Cortés 2003, Gardner et al. 2003). However, the effects and consequences of a particular bleaching event may vary between localities depending on oceanographic processes which is partly determined by reef morphology and topography; ]]> et al. 2007, Wilkinson & Souter 2008, Brandt & McManus 2009), but the largest coral cover losses related to this MBE were detected in the eastern Caribbean, where also epizootic events came after a prolonged bleaching period (Miller et al. 2006, Muller et al. 2008). In contrast, this ]]> et al. 2010).

In 2010, another mass bleaching event was recorded worldwide (Normile 2010), although apparently not as severe as in 2005 for the Caribbean. However, coral reefs at Los Roques National Park (LRNP) were severely affected in 2010. Los Roques is a ]]> et al. 2003, Croquer et al. 2010; but see Croquer et al. 2005). The relatively good conditions of coral reefs ecosystems at LRNP (e.g. Schweizer et al. 2005) might be explained by low anthropogenic impacts, absence of runoff; and perhaps by the combination of low hurricane activity, low ]]>

Materials and Methods

]]> Study site: LRNP is located about 150 km north of the Venezuelan coast. This marine protected area encompasses 2211.4 km2 of tropical marine ecosystems including mangroves, seagrass beds, soft-bottom communities, sandy and rocky shores and coral reefs. Reef development in Los Roques is extensive showing high diversity and live cover of scleractinian corals compared with other reef areas in Venezuela (Ramírez-Villarroel 2001) and other areas of the Caribbean (Rodríguez-Ramírez et al. ]]> Acropora palmata (Zubillaga et al. 2005) and A. cervicornis.

Seawater Temperatures: seawater temperatures were monitored with four HOBO data loggers (Pro-V2), at ]]> et al. (2006). The estimation of monthly mean temperatures was done with a time series from May 2003 till July 2011 obtained from the MODIS-SCAR sensor (data obtained from http://cariaco.ws/).

]]> Surveys of coral bleaching: In order to assess the magnitude of the 2010 MBE and its effects on coral reefs in LRNP, the proportion of bleached or paled colonies as well as the live coral cover was determined along ten, 10-m long, permanent transects located at each reef site, covering a depth interval of 5 to 15 m. This was carried out in October 2010, about one month after the onset of bleaching, and then again five months after the initial survey, in February 2011. During the initial survey, four reef sites (i.e. reefs located in the slope of different cays) were sampled: two ]]> et al. 2008, Croquer et al. 2010). Coral cover was estimated from chain transects (CARICOMP 2001), and the condition of coral colonies intercepted were also recorded (coloration, diseased, surface of partial mortality). In February of 2011, two sites were resurveyed ]]>

Statistical analysis: Null hypothesis of no temporal changes in live coral cover was compared with ]]>

Results

]]> Seawater Temperature and Degree Heating Weeks: During 2010, the maximum temperature of 30.85 °C at 5 m depth was reached between the 17 and 19 of September at all four reef sites. At that depth, the seawater temperatures remained above 29.0 °C from mid-August until the first week of November 2010 (Fig. 1A).  By that time, Degree Heating Weeks (DWH) reached a maximum value of 16 °Cweeks in the superficial waters of the archipelago ]]> Fig. 1B, from NOAA) and 13 °C-weeks in waters at 5 m (data from in situ loggers).

Percentage of bleached and diseased coral colonies: In October 2010, 72% of the 563 scleractinian colonies were bleached or pale. In February 2011, there were still 46% of coral colonies affected; but the recovery of zooxanthellae density was in place, as most of them were pale, and only 2% were bleached ( Fig. 2).

The high prevalence of Black Band Disease (BBD) had no precedent since the beginning of our surveys in 2002. In fact, BBD has been only detected once before, in 2007, when it reached 1% in Dos Mosquises (DMS). In contrast, the prevalence of BBD was 6% in DMS and 16% in Cayo de Agua (AGU) in October 2010. By ]]>

Analysis of the cover change: By October 2010, mean coral cover was 42.5% ± 3.97 (± ]]> Fig. 3) and significantly from a mean of 44.9 to 30.6% (Time p< 0.05), with a concomitant loss of 17% in the number of colonies.

]]> Although there could be other causes for the rapid loss of coral cover (including the onset of diseases), a great proportion is certainly attributable to the bleaching event of 2010. This assumption is based on the relatively short time period elapsed between the high seawater temperature anomaly and both surveys. Also, the proportion of bleached or pale colonies explained about 22% of the variability obtained in the net loss of coral cover in each transect (r2= 0.2165 for the regression shown in Fig. 4).

Discussion

Considering the combined global land and ocean surface temperature, the year 2010 tied with 2005 as the warmest on record (NOAA 2010). Also, both are known as years were mass bleaching events (MBE) have occurred across the Caribbean. However, the ]]>

]]> Previous bleaching events observed in Venezuelan coral reefs have been mild compared with those that occurred during the same periods in other Caribbean localities (e.g. Lang et al. 1992, Eakin et al. 2010). In Los Roques, in particular, less than 4% of coral colonies were bleached in August and September 2005 (Rodríguez et al. 2010), and <10% across the archipelago ]]> et al. 2008). In contrast, the 2005 MBE was severe across the Caribbean, particularly in the eastern reefs, where percentage of bleached corals reached up to 80% (Ballantine et al. 2008, Oxenford et al. 2008, Brandt & McManus 2009, Croquer & Weil 2009, Miller et al. 2009, Eakin et al. 2010). ]]>

The distribution of heat in the superficial water masses along the Venezuelan coast was completely different between 2005 and 2010 (Fig. 1A in Eakin et al. 2010 versus Fig. 1B in this study) and this can be considered a factor contributing to the differences in bleaching intensity observed between 2005 and 2010. It is well known that bleaching severity is determined by a combination of intensity and duration of thermal stress and irradiance conditions (Glynn & D’Croz 1990, Lesser & Farrell 2004). In situ measurements indicated that maximum temperatures in the archipelago (excluding the internal lagoon) were about 1°C higher in 2010 than in 2005 for the same period (29.92 °C in 2005 vs. 30.85 °C in 2010). Concomitantly, Degree Heating Weeks (DHW) ]]> et al. 2008) for the reefs surveyed in this study, whereas it reached 13 °C-weeks in 2010. It has been generalized that values of over 4°C-weeks cause significant bleaching, and values over 8°C-weeks cause widespread bleaching and some mortality (Liu et al. 2006), which was also the case in this study.

]]> Another example of the variability in bleaching intensity is illustrated by the differences found between the 2005 and 2010 MBE in Venezuela. In 2005, bleaching intensity was higher in coastal rather than oceanic reefs (16% vs. <4 of bleached corals, Rodríguez et al. 2010). The opposite trend was found in 2010, when intensity seemed higher for the oceanic reefs of Los Roques compared to coastal reefs of Morrocoy (personal observations). These results contrasted with other studies that have consistently found more ]]> et al. 2004).

Coral cover was severely reduced (by 36%) at our study sites in Los Roques after the 2010 bleaching event, which was similar to the 40% mortality observed in 2005 across the Caribbean (Eakin et al. 2010). However, we suspect that the longterm consequences could be ]]> et al. 2010), as well as its detrimental consequences on coral reproduction (e.g. Szmant & Gassman 1990). Also, the onset of diseases associated to the MBE, particularly black band disease, white plague and yellow band disease, ]]> et al. 2009, Weil et al. 2009).

The loss of coral cover induced by the 2010 MBE in Los Roques has no precedent at least in 20 years. Although we expect that MBEinduced coral mortality to continue for several months after our last survey, a rapid recovery in Los Roques ]]> recovery after massive mortality events (Bastidas et al. 2006). Unfortunately, the positiveexpectations ]]>

Acknowledgments

We would like to thank Mitsui Ltd ]]>

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*Correspondencia:
Carolina Bastidas: Dept. Biología de Organismos, Universidad Simón Bolívar. Caracas 1080 VenezuelaCentro de Estudios Ecotoxicológicos en Ambientes Marinos CETOXMAR-Universidad Simón Bolívar. Caracas 1080 Venezuela. Corresponding author: Carolina Bastidas, Tel/Fax 58-212-9063415. cbastidas@usb.ve
David Bone: Dept. Biología de Organismos, Universidad Simón Bolívar. Caracas 1080 VenezuelaCentro de Estudios Ecotoxicológicos en Ambientes Marinos CETOXMAR-Universidad Simón Bolívar. Caracas 1080 Venezuela. dbone@usb.ve
Aldo Croquer: Dept. Estudios Ambientales, Universidad Simón Bolívar. Caracas 1080 VenezuelaCentro de Estudios Ecotoxicológicos en Ambientes Marinos CETOXMAR-Universidad Simón Bolívar. Caracas 1080 Venezuela. acroquer@usb.ve ]]> Denise Debrot: Centro de Estudios Ecotoxicológicos en Ambientes Marinos CETOXMAR- Universidad Simón Bolívar. Caracas 1080 Venezuela. ddebrot@gmail.com
Elia Garcia: Dept. Biología de Organismos, Universidad Simón Bolívar. Caracas 1080 Venezuela. Centro de Estudios Ecotoxicológicos en Ambientes Marinos CETOXMAR-Universidad Simón Bolívar. Caracas 1080 Venezuela. emgarcia@usb.ve
Adriana Humanes: Centro de Estudios Ecotoxicológicos en Ambientes Marinos CETOXMAR-Universidad Simón Bolívar. Caracas 1080 Venezuela. adrihumanes@gmail.com
Ruth Ramos: Centro de Estudios Ecotoxicológicos en Ambientes Marinos CETOXMAR- Universidad Simón Bolívar. Caracas 1080 Venezuela. ruthr@usb.ve
Sebastian Rodríguez: Centro de Estudios Ecotoxicológicos en Ambientes Marinos CETOXMAR-Universidad Simón Bolívar. Caracas 1080 Venezuela. sebastianr@usb.ve

1. Dept. Biología de Organismos, Universidad Simón Bolívar. Caracas 1080 Venezuela
2. Dept. Estudios Ambientales, Universidad Simón Bolívar. Caracas 1080 Venezuela
3. Centro de Estudios Ecotoxicológicos en Ambientes Marinos CETOXMAR-Universidad Simón Bolívar. Caracas 1080 Venezuela
4. Email of all authors: cbastidas@usb.ve; dbone@usb.ve; acroquer@usb.ve; ddebrot@gmail.com; emgarcia@usb.ve; adrihumanes@gmail.com, ruthr@usb.ve; sebastianr@usb.ve
5. Corresponding author: Carolina Bastidas, Tel/Fax 58-212-9063415