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Substratum Preference During Recruitment of Two Invasive Alien Corals Onto Shallow-Subtidal Tropical Rocky Shores

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Substratum Preference During Recruitment of Two Invasive Alien Corals Onto Shallow-Subtidal Tropical Rocky Shores MARINE ECOLOGY PROGRESS SERIES Vol. 330: 101–111, 2007 Published January 25 Mar Ecol Prog Ser Substratum preference during recruitment of two invasive alien corals onto shallow-subtidal tropical rocky shores Joel C. Creed1,*, Alline F. De Paula2 1Laboratório de Ecologia Marinha Bêntica, Departamento de Ecologia, Instituto de Biologia Roberto Alcântara Gomes, Universidade do Estado do Rio de Janeiro – UERJ, PHLC Sala 220, Rua São Francisco Xavier 524, CEP 20559-900, Rio de Janeiro, RJ, Brazil 2Laboratório de Celenterologia, Departamento de Invertebrados, Museu Nacional-Universidade Federal do Rio de Janeiro – Quinta da Boa Vista, São Cristóvão, 20940-040, Rio de Janeiro, RJ, Brazil ABSTRACT: Two species of azooxanthellate coral, Tubastraea coccinea Lesson, 1829 and T. tagu- sensis Wells, 1982, are alien to the rocky shores of Brazil. The influence of 5 substratum types — wood, granite, concrete, steel and ceramic tiles — on their recruitment was investigated experimen- tally. On the artificial plates, mean density of T. tagusensis varied from 202 to 512 colonies m–2 and that of T. coccinea varied from 187 to 233 colonies m–2 after 17 mo. The density of Tubastraea spp. recruits was similar to those found in coral reef environments worldwide. A strong coupling between local adult density and recruitment density was found at a scale of <1 m. Substratum type and spe- cies were important in determining density and size of the recruits of the alien corals. The density of T. tagusensis on cement was higher than on ceramic tiles or steel, but T. coccinea density did not dif- fer significantly among substratum types. The size of T. tagusensis recruits did not differ among materials, but T. coccinea recruits were smaller on steel than on granite or cement. The density of recruits also depended on the density of adults on the reef. No differences in cover or biomass of the associated biological community were observed among substrata. T. coccinea and T. tagusensis did not exhibit very strong selection for specific substrata and ably recruited to all materials. Their oppor- tunistic behavior and high fecundity indicate that these are generalist species in terms of substratum utilization, which accounts for how they can successfully disperse to and invade areas consisting of different substratum materials. KEY WORDS: Brazil · Invasive species · Scleractinia · Substratum · Tubastraea Resale or republication not permitted without written consent of the publisher INTRODUCTION 1982, were recently recorded as alien to the rocky shores of Brazil (De Paula & Creed 2004). We also ob- The orange cup coral Tubastraea coccinea Lesson, served these species on an oil rig in southwestern 1829 is the only scleractinian unintentionally intro- Brazil in the state of Rio de Janeiro and T. coccinea on duced into the western Atlantic. This azooxanthellate the hull of a support ship (authors' unpubl. data). often occurs on ship bottoms, dock pilings, buoys and Fenner & Banks (2004) and Cairns (2000) suggested attached to man-made objects (Cairns 2000, De Paula that T. coccinea has been dispersed by colonization & Creed 2004, Fenner & Banks 2004, Sammarco et al. and transport on ships’ hulls and/or oil platforms. 2004), and has invaded the Gulf of Mexico, Florida and Tubastraea is a brooding coral (Richmond 1998) with the Caribbean (Fenner & Banks 2004), expanding its a short competency period. Tubastraea spp. are cur- range from the Indo-Pacific (Cairns 2000). Two spe- rently established along 900 km of coastline off Brazil cies, T. coccinea Lesson, 1829 and T. tagusensis Wells, at 5 discrete locations within a 1000 km2 polygon. Once *Email: [email protected] © Inter-Research 2007 · www.int-res.com 102 Mar Ecol Prog Ser 330: 101–111, 2007 established, they can become the dominant benthic vant to investigations of benthic marine invasive spe- organism on shallow rocky reefs, and are engineering cies because the composition of the vector of introduc- species that change community structure. Creed tion and of local natural substratum will determine the (2006) described their deleterious effects on the most success of the introduction. Range expansion also will abundant native coral, Mussismilia hispida (Verrill), a be limited by the availability of suitable substrata and species endemic to Brazilian coastal waters, and the the composition of vectors. Furthermore, if artificial corals are considered a serious threat to regional biodi- substrata are to be used to detect invasive species or to versity and ecosystem function. monitor range expansion, appropriate substrata need The abundance of studies of coral recruitment pat- to be selected. terns that have utilized settlement plates has done In this study, we investigated the influence of 5 dif- much to highlight aspects of coral reproductive strate- ferent substratum types — 2 natural (wood, granite) gies and population dynamics, such as spatio-temporal and 3 artificial (cement, steel and ceramic tiles) — on variation in coral recruitment (Birkeland et al. 1981, the recruitment of the corals Tubastraea coccinea and Harriott & Banks 1995, Dunstan & Johnson 1998, Glas- T. tagusensis. As space is a limited resource in the som et al. 2004), influences of orientation and position marine benthos, we hypothesized that during transport of substrata (Glasby 2000, Mundy 2000), coral early and upon establishment, these corals undergo strong growth (van Moorsel 1988, Babcock & Mundy 1996) selective pressure for substratum type, and that they and the influence of community composition (Maida et will preferentially settle on materials used in vector al. 2001). However, recruitment of corals onto artificial construction or of which natural receptor reefs are settlement plates is usually used to provide a measure composed. of the relative abundance of coral recruits and recruit- ment rates (Birkeland 1977); fewer studies have com- pared recruitment rates onto different substrata, or the MATERIALS AND METHODS differential preference of the coral for settlement and subsequent recruitment (e.g. Harriott & Fisk 1987, The study was carried out from October 2003 to Fitzhardinge & Bailey-Brock 1989, Hunte & Witten- March 2005 at Ilha dos Macacos, Baía da Ilha Grande, berg 1992). Brazil (23° 04.72' S, 44° 13.50' W). The study site was a Most studies of coral recruitment have utilized exper- tropical subtidal rocky reef extending from the inter- imental surfaces not generally found in the environ- tidal to a sandy plain at 4 to 7 m depth, situated in the ment, such as glass (Lewis 1974), plexiglass (Birkeland Canal Central of the bay (Fig. 1a). Rocky reefs are 1977, Birkeland et al. 1981), ceramic/terracotta tiles typical of the region, which consists of a coastline of (Fisk & Harriott 1990, Harriott & Banks 1995, Dunstan & bays and islands with alternating rocky shores and Johnson 1998, Mundy 2000, Glassom et al. 2004), con- sandy pocket beaches. The substratum consisted of a crete or cement blocks (Birkeland 1977, Glasby 2000, reef (boulders of approximate diameter 0.5 to 5 m) Connell 2001, Lam 2003), plastic Petri dishes (Harriott & composed of granite rock. The north-facing shore is Fisk 1987) and steel (Fitzhardinge & Bailey-Brock 1989, protected from southern ocean swell and the shore in Perkol-Finkel & Benayahu 2005). Artificial substrates general is subjected to little wave action. Water tem- permit control of replication and facilitate measures of perature at the site varies from 17 to 28°C, tidal range species composition, and so have been utilized in moni- in the region is 1.4 m and rainfall is ~2500 mm yr–1, toring programs for the detection of benthic marine in- with summer wet and winter dry seasons. Secchi vasive species (e.g. Sutton & Hewitt 2004). depths are of 6 to 10 m and sea surface summer clear Schumacher (1977) suggested that a conditioning sky irradiance is 1800 to 1900 µmol m–2 s–1 (J. C. Creed period of about 1 yr is needed before corals will recruit unpubl. data). The subtidal benthic community con- to new substrata. More recently, Harriott & Banks sisted of macroalgae (multi-species turf forming, crus- (1995), Babcock & Mundy (1996) and Dunstan & John- tose coralline and foliose brown algae), heterotrophic son (1998) demonstrated that corals will settle onto nat- filter feeders (mainly sponges, bryozoans and asci- ural and artificial substrata within 4 mo of immersion. dians) and symbiotic autotrophs (corals and zoo- In previous experiments Tubastraea spp. settled onto anthid mats). These form a mosaic over the reef tiles 1 mo after deployment of plates (authors’ unpubl. (authors’ pers. obs.). data). In order to compare coral recruitment and mortality A problem inherent in the comparison of recruitment on different substrata, experimental plates were studies is that almost every author has used a different deployed on 17 October 2003 and recovered on settlement plate, and there are few studies directly 1 March 2005, 17 mo later. The substrata employed comparing the composition of these plates (Harriott & were wood, ceramic tiles, cement, steel and granite. Fisk 1987). Substratum composition is especially rele- The wood (15 mm-thick marine ply), ceramic (the Creed & De Paula: Alien coral recruitment onto artificial substrata 103 Fig. 1. Study site and material. (a) Location of the study site at Ilha Grande, Brazil; (b) experimental block showing fixed substrata unglazed side of commercially available wall tiles), demonstrated no edge effects (D. Bartholo & cement (handmade sand and cement reinforced with J. C. Creed unpubl. data). an internal steel grid) and steel (10 mm-thick ship- To quantify the biomass of the biological community building grade) plates were square (approximately on the plates, the height of the encrusting community 15 × 15 cm) with 2 holes at the corners diagonally was measured at 5 pre-defined points on each plate arranged for fixation. The granite tiles were of similar with a ruler, from which mean community height was area but rectangular, and did not have holes for fixa- calculated. To quantify the composition of the accom- tion.
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