Possible recovery of Acropora palmata (Scleractinia:Acroporidae) within the Veracruz Reef System, Gulf of Mexico: a survey of 24 reefs to assess the benthic communities
Posible recuperación de Acropora palmata (Scleractinia:Acroporidae) en el sistema arrecifal de Vera Cruz, Golfo de México: evaluación de comunidades bentónicas en 24 arrecifes
Elizabeth A. Larson1*, David S. Gilliam1, Mauricio Lόpez Padierna1 & Brian K. Walker1
Abstract
Recent evidence shows that Acropora palmata within the Veracruz Reef System, located in the southwestern Gulf of Mexico, may be recovering after the die off from the flooding ]]>
A. palmata. The 28 named reefs in the system are divided into 13 northern and 15 southern groups by the River. Between 2007 and 2013, we surveyed 24 reefs to assess the benthic communities. Seven of the 11 reefs surveyed in the northern group and all in the southern group had A. palmata. ]]>
Key words:Acropora palmata, population recovery, Veracruz, Mexico, colony size class frequency.
Resumen
Evidencia reciente indica que las poblaciones de Acropora palmata
del sistema arrecifal de Vera Cruz, ubicado al suroeste del Golfo de México, podría estarse recuperando después de las inundaciones del río Jamapa e intrusiones de agua fría que afectaron la región en los años setenta. Desde la disminución, pocos estudios han documentado el estado A. palmata. Consiste de 28 arrecifes divididos 13 al norte y 15 al sur del río. Entre el 2007 y 2013 se ]]>
A. palmata. Las colonias se encontraron por lo general en el barlovento de los corales en agua sómeras a lo largo del borde del arrecife. También registramos los diámetros de las colonias y condición a lo largo de transectos tipo conturón en dos arrecifes al norte (Anegada de ]]>
Palabras clave:Acropora palmata, recuperación poblacional, Veracruz, Mexico, distribución de frecuencia de tamaño.
Acropora palmata and A. cervicornis (Lamarck, 1816) historically have been major reef framework builders in the greater Caribbean, Gulf of Mexico and Florida (Adey, 1978; Neigell & Avise, 1983). Their complex three-dimensional structure provides habitat to numerous invertebrate and vertebrate species and coastal protection during storms and hurricanes. Between the 1970’s and 1980’s populations across the species range were plagued by disease and in ]]>
]]>
Since the listing, many populations have not returned to self-sustainable levels despite increased restoration activities and protection (Knowlton et al., 1990; Aronson & Precht, 2001; Bruckner, 2002; Miller, Bourque & Bohnsack, 2002; Acropora Biological Review Team, 2005); however, there are remnant populations throughout the Acropora spp. range. Large Acropora spp. populations have been reported in Florida (Vargas-Angel, Thomas & Hoke, 2003; Williams, Miller & Kramer, 2008; Walker, Larson, Moulding & Gilliam, 2012); Punta Rusia, Dominican Republic (Lirman et al., 2010); Roatan, Honduras (Keck, Houston, Purkis & Riegl, 2005); Venezuela (Zubillaga, Márquez, Cráquer & ]]>
A. palmata in Veracruz, Mexico.
The Veracruz Reef System (VRS) is located within the Parque Nacional Sistema Arrecifal Veracruzano ]]>
Since the late 1800’s, A. palmata and A. cervicornis have dominated the shallow-water coral reefs of the VRS (Heilprin, 1890). However, similar to other Caribbean populations, researchers observed local declines through the 1970’s and 1980’s. Mortality in this region was attributed to disease and the flooding of the nearby Jamapa and Papaloapan rivers (Rannefeld, 1972; Tunnell, 1988; 1992). In 1971, Rannefeld (1972) documented up to a 100% loss at Enmedio for both species and by 1973, abundance had decreased dramatically on many reef slopes (Tunnell, 1988; 1992). However, Jordan-Dahlgren (1992) documented re-colonization of A. palmata to old standing dead colonies in the late 1980’s. Another study by Lara, Padilla, García and Espejel (1992) observed Acropora spp. at 82% of the reefs surveyed (n=17) and recorded them as dominant (>20% live cover) on the fore reef edge and inner fore reef zones. Despite ]]>
A. palmata were reported as low (0–0.3% cover) at six reefs in the VRS (Jones et al., 2008), some of which had previously been documented as having colonies (Lara et al., 1992).
This study was part of a larger, ]]>
Acropora. Using the knowledge gained of these populations during the benthic surveys, additional
A. palmata assessments were completed to document the current status, health and possible recovery of A. palmata within the VRS. Population recovery herein is defined by an increase in colony abundance, size and distribution and the potential to sexually reproduce with low prevalence of recent mortality in comparison to previous studies in the ]]>
Materials and Methods
Benthic characterization assessment: Reefs within the VRS were visited between 2007 and ]]>
A. palmata. These data were utilized to document the distribution of A. palmata within the VRS and target sites for the A. palmata assessment.
Acropora palmata assessment: Five ]]>
A. palmata was found in greatest abundance were used to further investigate the status of A. palmata within the VRS. These sites were on two reefs in the northern group (Verde and Anegada de ]]>
A live area index (LAI) was used to compare the amount of live tissue between sites and across years to better understand the population structure within the VRS. LAI indices typically use colony length, width and percent live tissue ]]>
]]>
The resulting size frequency distribution was used to describe the population structure. Soong and Lang (1992) reported that A. palmata was not fertile until colonies were >60cm2
, and as size of the colony increased as did the frequency of fertility, 31% of the colonies between 250 and 1 000cm2, 43% of the colonies between 1 000 and 4 000cm2 and 88% of the colonies >4 000cm2 were fertile. In addition to ]]>
m2 were reproductively mature. Therefore, our chosen size classes, similar to those of Schärer et al. (2008), are as follows: small colonies with no reproductive potential (<60cm2), ]]>
m2) and large mature colonies with moderate to high potential to spawn (>1 600cm2).
Results
Benthic characterization assessment: Based on the point intercept surveys and site descriptions, A. palmata was present at 20 of the 24 reefs ]]>
A. palmata was present at seven of the 11 reefs and 13 of the 28 sites (Fig. 1). In the southern group, A. palmata
was present at all reefs (n=13) and 21 of the 27 sites (Fig. 2). Where A. palmata was present, colony abundance varied by site ranging from one colony to multiple large overlapping colonies. ]]>
Five reefs were of particular interest because A. palmata was present in greater abundance. In the northern group, large overlapping A. ]]>
colonies were found at Verde and Anegada de Adentro reefs. Colonies were observed on the reef flat at Anegada de Adentro and on the leeward slope of Verde. Three reefs in the southern group, Rizo, Sargazo and Periférico also had greater abundance. Colonies at Rizo were observed on the reef flat and on the upper fore reef of Sargazo and Periférico.
Acropora palmata assessment: Belt transects targeting A. palmata
were completed between 2011 and 2013 at Verde, Anegada de Adentro, Rizo, Periférico and Sargazo reefs, surveying a total area of 21 420m² (Table 1). A total of 1 804 colonies were measured across all sites and all years. The number of colonies ranged from 40 in 2011 at Rizo to 764 at Verde in 2012 (Table 1).
]]>
Colony density (±SD) ranged from 0.02±0.03colonies/m² to 0.28±0.08colonies/m² (Table 1). The greatest abundance of A. palmata was found at Anegada de Adentro where the density was ]]>
Table 1). The increased number of colonies at Rizo was likely due to the VRS being directly impacted by ]]>
Mean colony length (±SD) for all reefs was 58±73cm. Anegada de Adentro had significantly larger colonies than the rest of the reefs (122±163cm —Kruskal-Wallis, p<0.05) and the largest colony recorded (length= 1 320cm). At Verde colony length increased from 2011 to 2012. At Rizo ]]>
Table 1).
Mean percent live tissue (±SD) per colony for all reefs across all years was 89±18%. Mean percent live tissue per colony varied by reef and was greatest at Rizo in 2011, 96% (Table 1). However, from ]]>
Predation by the threespot damselfish, Stegastes planifrons (Cuvier 1830), snail, Coralliophilia ]]>
(Lamarck 1816) and bearded fireworm, Hermodice carunculata (Pallas 1766) were present in both groups. Damselfish predation, through the creation of algal gardens was the most common type, observed on 95 colonies. Its impact was difficult to quantify and was only recorded as present or absent on a colony since it was not considered a cause of recent mortality. The most common cause of recent mortality was predation by snails, affecting 35 colonies throughout the three years of data collection. Half of the colonies with snail predation were observed at Sargazo. Fireworm predation was recorded on ]]>
Disease prevalence was low. Although it was recorded at four of the five reefs, it was only observed on 2.7% of the colonies across the entire study. Disease was not observed at Rizo. Rapid tissue loss (RTL) was the most commonly recorded condition (Williams & Miller, 2005), observed affecting ]]>
Mean LAI ranged from 828cm² to 23 448cm² per reef (Fig. 3). The reefs in the ]]>
Fig. 3).
In 2011, for all reefs combined, 6% of the measured colonies were small (<60cm²), 66% were medium (60-1 600cm²) and 28% were large (>1 600cm²). From 2011 to 2012, there was a small shift from small colonies to medium and large at Verde. At Rizo, there was a decrease in the number of large colonies ]]>
Fig. 3, 4).
Discussion
This study is the most comprehensive study of A. palmata within the Veracruz Reef System to date. Its distribution through the VRS indicates that
A. palmata may be recovering from the 1970’s and 1980’s die off. Acropora palmata was present at a majority of the sites surveyed in both reef groups. For ]]>
A. palmata assessment.
In order for a population to be recovering it needs to be dominated by ‘healthy’ colonies, which are characterized by a broad distribution of colony size frequencies ]]>
The A. palmata populations in the VRS have a high spawning potential, 38% of the colonies are considered mature (>1 600cm²), of those colonies 53% were >4 000cm2. According to Soong and Lang (1992) there is a high potential (88%) that colonies >4 000cm2 are fertile. All sites were dominated by medium sized colonies; only 4% of the colonies were classified as small across all years. This size frequency distribution indicates that these populations have a high potential for colonies to spawn, whereas two ]]>
Although the A. palmata populations ]]>
A. palmata ]]>
These data suggest that A. palmata in the VRS is healthy, abundant, broadly distributed, and has a high spawning potential, whereas most of the Caribbean populations are not. Hence, the next question may be, why has this species been able to ]]>
Acknowledgments
We thank the staff of the Parque Nacional Sistema Arrecifal Veracruzano, M. Rangel Avalos and J. Santander Monsalvo for critical logistical support. Nova Southeastern University Oceanographic Center graduate students, V. Brinkhuis, S. Bush, A. Costaregni, N. D’Antonio, P. Espitia, D. Fahy, A. Halperin, M. Sathe, J. Walczak, and C. Walton, provided essential help with data collection. Partial funding for this research was provided by multiple ]]>
References ]]>
Acropora Biological Review Team. (2005). Atlantic Acropora status review document. Report to National Marine Fisheries Service, Southeast Regional Office. [ Links ]
Adey, W. H. (1978). Coral reef morphogenesis: a multidimensional model. Science, 202, 831-837. [ Links ]
Aronson, R. B., Bruckner, A. W., Moore, J.,. Precht W. F., & Weil, E. (2008a). IUCN Red List of Threatened Species: Acropora cervicornis. Retrieved from www.iucnredlist.org. [ Links ] Aronson, R. B., Bruckner, A. W., Moore, J., Precht W. F., & Weil, E. (2008b). IUCN Red List of Threatened Species: Acropora palmata. Version 2012.2. Retrieved from www.iucnredlist.org. [ Links ]
Aronson, R. B., & Precht, W. F. (2001). White-band disease and the changing face of Caribbean coral reefs. Hydrobiologia, 460, 25-38. [ Links ]
Blake, E. S. (2011). Tropical Cyclone Report. Tropical Cyclone Harvey. National Oceanic and Atmospheric Administration, USA. [ Links ]
Bruckner, A. W. (2002). Proceedings of the Caribbean Acropora Workshop: Potential Application of the U.S. Endangered Species Act as a Conservation Strategy. NOAA Technical Memorandum NMFS-OPR-24. [ Links ]
Diario Oficial, D. (1992). Decreto por el que se declara área natural protegida con el carácter de Parque Marino Nacional, la zona conocida como Sistema Arrecifal Veracruzano, ubicada frente a las Costas de los municipios de Veracruz, Boca del Río y Alvarado del estado de Veracruz Llave, con superficie de 52,238-91-50 hectáreas. Diario Oficial de la Federacion, Mexico. [ Links ]
Dustan, P. & Halas, J. C. (1987). Changes in the reef-coral community of Carysfort reef, Key Largo, Florida: 1974 to 1982. Coral Reefs, 6, 91-106. [ Links ]
Emery, K. O. (1963). Coral reefs off Veracruz, Mexico. Geofisica internacional, 3, 11-17. [ Links ]
Grober-Dunsmore, R., Bonito, V., & Frazer, T. K. (2006). Potential inhibitors to recovery of Acropora palmata populations in St. John, US Virgin Islands. Marine Ecology Progress Series, 321, 123-132. [ Links ]
Heilprin, A. (1890). The Corals and Coral Reefs of the Western Waters of the Gulf of Mexico. Proceedings of the Academy of Natural Sciences of Philadelphia, 42, 303-316. [ Links ]
Horta-Puga, G. (2007). Environmental Impacts, p. 126-141. In Tunnell J. W. Jr., Chavez E. A., & Withers K. (eds.). Coral Reefs of the Southern Gulf of Mexico. Texas: A&M University Press, College Station. [ Links ]
Jones, J., Withers, K., & Tunnell, J. W. Jr. (2008). Comparison of Benthic Communities on Six Coral Reefs inthe Veracruz Reef System (Mexico). Proceedings of the 11th International Coral Reef Symposium, Fort Lauderdale, FL USA, 757-760. [ Links ]
Jordan-Dahlgren, E. (1992). Recolonization Patterns of Acropora palmata in a Marginal Environment. Bulletin of Marine Science, 51, 104-117. [ Links ]
Keck, J., Houston, R. S., Purkis, S., & Riegl, B. M. (2005). Unexpectedly high cover of Acropora cervicornis on offshore reef in Roatan (Honduras). Coral Reefs, 24, 509. [ Links ]
Kimberlain, T. B. (2011). Tropical Cyclone Report. Hurricane Nate. National Oceanic and Atmospheric Administration, USA. [ Links ]
Knowlton, N. (2001). The future of coral reefs. Proceedings of the National Academy of Science of the United States of America, 5419-5425. [ Links ]
Knowlton, N., Lang, J. C. & Keller, B. D. (1990). Case study of natural population collapse: post-hurricane predation on Jamaican Staghorn coral. Smithsonian Contributions to the Marine Sciences, 36. [ Links ]
Lara, M., Padilla, C., Garcia, C., & Espejel, J. J. (1992). Coral Reef of Veracruz Mexico I. Zonation and community. Proceedings of the 7th International Coral Reef Symposium, Guam., 1, 535-544. [ Links ]
Lirman, D., Bowden-Kerby, A., Schopmeyer, S., Huntington, B., Thyberg, T., Gough, M., Gough, T., Gough, R., & Gough, Y. (2010). A window to the past: documenting the status of one of the last remaining ‘megapopulations’ of the threatened staghorn coral Acroporacervicornis in the Dominican Republic. Aquatic Conservation: Marine and Freshwater Ecosystems, 20, 773-781. [ Links ]
Mayor, P.A., Rogers, C.S. & Hillis-Starr, Z.M. (2006). Distribution and abundance of elkhorn coral, Acropora palmata, and prevalence of white-band disease at Buck island reef national monument, St. Croix, US Virgin Islands. Coral Reefs, 25, 239-242. [ Links ]
Miller, M. W., Bourque, J., & Bohnsack, J. (2002). An analysis of the loss of Acroporid corals at Looe Key, Florida, USA 1983-2000. Coral Reefs, 21, 179-182. [ Links ]
Neigell, J. E., & Avise, J. C. (1983). Clonal diversity and population structure in a reef-building coral, Acropora cervicornis: self recognition analysis and demographic interpretation. Evolution, 37, 437-453. [ Links ]
NOAA. (2006). Endangered and threatened species: final listing determinations for Elkhorn coral and Staghorn coral Federal Register 71: 26852-26861. [ Links ]
Rangel Avalos, M. A., Jordan, L. K. B., Walker, B. K., Gilliam, D. S., Carvajal Hinojosa, E., & Spieler, R. E. (2007). Fish and Coral Reef Communities of the Parque Nacional Sistema Arrecifal Veracruzano (Veracruz Coral Reef System National Park) Veracruz, Mexico: Preliminary Results. Proceedings of the 60th Annual Gulf and Caribbean Fisheries Institute Meeting, Punta Cana, Dominican Republic, 60, 427-435. [ Links ]
Rannefeld, J. W. (1972). The stony corals of Enmedio Reef off Veracruz, Mexico. M.S Thesis. Texas A & M University, College Station. [ Links ]
Richmond, R. H. (1997). Reproduction and recruitment in corals: critical links in the persistence of reefs, p. 175-197. In Brikeland, C. (eds.). Life and Death of Coral Reefs. New York: Chapman & Hall. [ Links ]
Schärer, M., Nemeth, M., Valdivia, A., Miller, M., Williams, D., & Diez, C. (2008). Elkhorn coral distribution and condition throughout the Puerto Rican Archipelago. Proc. of the 11th Int. Coral Reef Symp., Fort Lauderdale, FL USA., 815-819. [ Links ]
Soong, K., & Lang, J. C. (1992). Reproductive integration in reef corals. Biological Bulletin, 183, 418-431. [ Links ]
Tunnell, J. W. J. (1988). Regional comparison of southwestern Gulf of Mexico to Caribbean sea coral reefs. Proceedings of the 6th International Coral Reef Symposium, Townsville, Australia, 3, 303-308. [ Links ]
Tunnell, J. W. J. (1992). Natural Versus Human Impacts to Southern Gulf of Mexico Coral Reef Resources. Proceedings of the 7th International Coral Reef Symposium, Guam., 1, 300-306. [ Links ]
Vargas-Angel, B., Thomas, J. D., & Hoke, S. M. (2003). High-latitude Acropora cervicornis thickets off Fort Lauderdale, Florida, USA. Coral Reefs, 22, 465-473. [ Links ]
Walker, B. K., Larson, E. A., Moulding, A. L., & Gilliam, D. S. (2012). Small-scale mapping of indeterminate arborescent acroporid coral (Acropora cervicornis) patches. Coral Reefs, 31, 885-894. [ Links ]
Williams, D. E., & Miller, M. W. (2005). Coral disease outbreak: pattern. prevalence and transmission in Acropora cervicornis. Marine Ecology Progress Series, 301, 119-128. [ Links ]
Williams, D. E., Miller, M. W., & Kramer, K. L. (2008). Recruitment failure in Florida Keys Acropora palmata, a threatened Caribbean coral. Coral Reefs, 27, 697-705. [ Links ]
Williams, D. E., & Miller, M. W. (2011). Attributing mortality among drivers of population decline in Acropora palmata in the Flordia Keys (USA). Coral Reefs, 31, 369-382. [ Links ]
Zubillaga, A. L., Márquez, L. M., Cráquer, A., & Bastidas, C. (2008). Ecological and genetic data indicate recovery of the endangered coral Acropora palmata in Los Roques, Southern Caribbean. Coral Reefs, 27, 63-72. [ Links ] ]]>
1. Nova Southeastern University Oceanographic Center, 8000 N. Ocean Dr. Dania Beach, FL, 33004 USA; goergen@nova.edu
Received 03-XI-2013 Corrected 12-III-2014 Accepted 24-III-2014
