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Succession of Crustose Coralline Red Algae (Rhodophyta) on Coralgal Reefs Exposed to Physical Disturbance in the Southwest Atlantic

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Succession of Crustose Coralline Red Algae (Rhodophyta) on Coralgal Reefs Exposed to Physical Disturbance in the Southwest Atlantic Helgol Mar Res (2013) 67:687–696 DOI 10.1007/s10152-013-0354-3 ORIGINAL ARTICLE Succession of crustose coralline red algae (Rhodophyta) on coralgal reefs exposed to physical disturbance in the southwest Atlantic Rodrigo Mariath • Rafael Riosmena Rodriguez • Marcia A. O. Figueiredo Received: 4 November 2011 / Revised: 25 February 2013 / Accepted: 3 April 2013 / Published online: 14 May 2013 Ó Springer-Verlag Berlin Heidelberg and AWI 2013 Abstract Biological and physical disturbances create the abundant. Physical disturbance increased crust recruitment conditions for species succession in any biological eco- and the low-light environment created by sediments. The system. In particular, coral reefs are susceptible to this data demonstrated coexistence among crustose coralline process because of the complexity of their ecological species and a tolerance to physical disturbance, which relationships. In the southwest Atlantic, nearshore reefs are seemed to favor the thinner crusts of P. conicum over mostly coated by a thin layer of coralline crusts rather than thick-crust species during succession. The succession pat- stony corals. However, little is known about the succession tern observed in this subtropical Brazilian coral reef differs of crustose coralline algae. We studied this process by from that described for shallow tropical reef communities. means of a series of experimental and control discs exposed to physical disturbance. Our results showed that Keywords CCA Á Coral reef Á Benthic communities Á the dominant species in natural conditions, Pneophyllum Brazil conicum, had early recruits and later became dominant on the discs, replicating the community structure of the actual reef. This species had mature reproductive structures and Introduction available spores from the beginning of the colonization experiments. Thicker crusts of Porolithon pachydermum Brazilian coral reefs differ from the classical example of and Peyssonnelia sp. were found on the discs after the Great Barrier Reef, for example, mainly in the impor- 112 days, and significantly increased their cover over the tant role of crustose coralline algae (hereafter CCA) as a succeeding months; and after 1 year, P. conicum was less living cover (Table 1), in addition to their role in the construction of reef structure. The nearshore bank reefs are surrounded and even filled with muddy siliciclastic sedi- Communicated by I. Bartsch. ments. The coral fauna is not diverse but is rich in endemic species, and their offshore reefs may be shaped into unique R. Mariath mushroom columns (Lea˜o and Dominguez 2000). On the Programa de Po´s-Graduac¸a˜o em Botaˆnica, Escola Nacional de windward side of the bank reefs is an abundant algal rim Botaˆnica Tropical, Instituto de Pesquisas Jardim Botaˆnico de Rio de Janeiro, Rua Pacheco Lea˜o 2040, composed by CCA together with vermetid gastropods Rio de Janeiro, RJ 22460-030, Brazil (Lea˜o 1996). Recently, human impacts have affected the cover of reef-building corals in this region (Dutra et al. & R. R. Rodriguez ( ) 2004), mostly through large increases in sediment deposi- Programa de Investigacio´n en Bota´nica Marina, Departamento de Biologı´a Marina, Universidad Auto´noma tion (Lea˜o and Kikuchi 2005). The reef-building corals and de Baja California Sur, 23080 La Paz, B.C.S, Mexico CCA are relatively little studied (Lea˜o 1996). e-mail: [email protected] Species composition and cover of CCA and corals on the emergent banks of Brazilian reefs have been observed M. A. O. Figueiredo Instituto de Pesquisas Jardim Botaˆnico de Rio de Janeiro, to change markedly at different intervals (Lea˜o and Kiku- Rua Pacheco Lea˜o 915, Rio de Janeiro, RJ 22460-030, Brazil chi 2005; Figueiredo et al. 2008). Turf algae, fleshy algae 123 688 Helgol Mar Res (2013) 67:687–696 Table 1 Abundance of crustose coralline algae on diverse reef flats in the southwestern Atlantic Place Relative abundance Abrolhos reefs 32–79 % in Figueiredo (1997); 4–36 % in Villac¸a and Pitombo (1997); 2–15 % in Figueiredo and Steneck (2002); 3–40 % in Figueiredo (2006); 10–20 % in Taˆmega and Figueiredo (2007) Rocas Atoll 36–60 % in Gherardi and Bosence (2001); 30–50 % in Villas Boas et al. (2005) Coroa Vermelha and Ponta Grande 30–40 % in Costa et al. (2002) Recife de Fora (studied site) 30–49 % in present work and CCA are the most abundant algal elements on the experimental evaluation of succession has been done to coastal reefs. CCA may reach their maximum development determine whether their succession sequence eventually on the seaward offshore reef (Costa et al. 2002), where they produces a community that resembles the natural coralline are probably exposed to higher grazing pressure (Steneck abundance on reef in a sediment-disturbed environment. 1988; Steneck and Dethier 1994). The main aim of the present study was to evaluate the Ecological experiments have shown that CCA may be species abundance of CCA in several succession stages, in one of the first colonizers of hard substrates (Quinn 1982; relation to a continuous physical disturbance. We hypoth- Steneck 1986). The success of this colonization depends on esized that CCA species will interact at the beginning of the growth strategy of each species in the competition for succession, and later only one species with a thick thallus space (Paine 1984; Steneck et al. 1991; Keats and Mane- will become dominant, provided that macroalgae and veldt 1994; Keats et al. 1994) in relation to physical con- trapped sediment are removed. ditions (Littler and Littler 1984; Kendrick 1991; Steneck and Dethier 1994) or biological disturbances (Paine 1984; Steneck et al. 1991). Figueiredo (1997) showed that CCA Materials and methods in the southwest Atlantic have a similar degree of cover on wave-exposed offshore reefs after 3–5 months of coloni- Study site zation, in shallow zones dominated by Porolithon pachy- dermum (Foslie) Foslie and Lithophyllum congestum The study was carried out within the Municipal Marine (Foslie) Foslie (Figueiredo and Steneck 2002; Figueiredo Park of Recife de Fora in Porto Seguro, southeastern 2006; Figueiredo et al. 2008). Brazil. The park is centered on an island 6 km from the Coralline crusts with a thin thallus such as Mesophyllum coast of the first conservation unit in this area, established spp. are primarily found under macroalgal canopies and in 1997, located at 16°230S and 16°260S and 39°000W and low-light environments, contrasting with the massive or 38°580W (Fig. 1), and covers a total area of approximately complexly branched species such as Porolithon pachy- 17.5 km2. The depth of the surrounding reef ranges from 6 dermum and Lithophyllum congestum in sunlit shallow to 8 m nearshore and may reach up to 15 m in the northeast zones with little fleshy algae (Figueiredo and Steneck offshore area (Costa et al. 2002; Prates 2006). 2002). Thick crusts are generally considered to be com- The study site selected for establishing the succes- petitively superior to thinner crusts (Steneck 1986; Dethier sion experiments, named Ponto Oeste (16°2403600S; et al. 1991), but the reverse might be expected in highly 38°5900800W), is a sheltered nearshore reef (Fig. 1b). grazed environments (Steneck et al. 1991). As the taxon- Visually, the reef flat was dominated by massive brain omy of this algal group is complex, there are limited corals of Siderastrea stellate (Verril 1868) and Mussismilia studies on their ecological relationships (Ringeltaube and braziliensis (Verril 1867) and branching corals of Mussis- Harvey 2005), especially on sediment disturbance, which milia hartii (Verril 1868) and Millepora alcicornis may influence their succession sequence on reefs. (Linnaeus 1758), with CCAs growing among and over the Porolithon pachydermum and Lithophyllum congestum corals. The reef flat is cut with grooves and is constantly dominate the higher levels of Caribbean algal ridges, with a submerged, with depths ranging from 0.5 to 3 m on the reef smaller standing crop of fleshy algae (Adey and Vassar flat and 8 m near the soft bottom. 1975; Adey 1998). The only study of succession among CCA demonstrated their adaptation to lower-light levels Transects (Adey and Vassar 1975). This study showed that Neo- goniolithon megacarpum gradually replaces other crustose The linear point intercept method (LPI) was deployed species when a reduction in wave action or grazing allows along three 10 m transect lines, which were laid 1 m from a massive development of fleshy algae. However, no each other and oriented parallel to the reef border. 123 Helgol Mar Res (2013) 67:687–696 689 Fig. 1 Location of the Marine Park of Recife de Fora, state of Bahia, Brazil. a Coastline of southern Bahia; arrow indicates the position of the Marine Park; b Marine Park; arrow indicates the survey location The number of times an organism found under 10 random diameter and 6 mm thick, handmade plastic), which were sampling points on each transect line was counted and its placed in the same area as the transect lines (Fig. 2). They overall proportion was calculated (method used to quantify were randomly distributed on horizontal surfaces of dead CCA by Figueiredo and Steneck 2002; Villas Boas et al. coralline sunlit areas of the reef flat close to the edge, and 2005;Taˆmega and Figueiredo 2007). Nandon and Stirling fixed in place by means of underwater epoxy putty (Tubolit (2006) indicated that LPI was the most cost-effective MEN) (Figueiredo 1997). We used artificial discs because means of measuring coral cover and at least as precise and they offer the advantages of being replicable, independent, accurate as the belt transect and the line transect methods. controllable and biota free (West and Crump 1974; Because CCA were partially hidden by canopy algae and Fletcher et al. 1985). The experiment started in February branching corals, they were removed or displaced prior to 2007, and ten discs were randomly removed at each time record.
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