Reserve effects and natural variation in coral reef communities
2008; Wiley; Volume: 45; Issue: 4 Linguagem: Inglês
10.1111/j.1365-2664.2008.01490.x
ISSN1365-2664
AutoresAlastair R. Harborne, Peter J. Mumby, Carrie V. Kappel, Craig P. Dahlgren, Fiorenza Micheli, Katherine E. Holmes, James N. Sanchirico, Kenneth Broad, Ian A. Elliott, Daniel R. Brumbaugh,
Tópico(s)Marine and coastal plant biology
ResumoJournal of Applied EcologyVolume 45, Issue 4 p. 1010-1018 Free Access Reserve effects and natural variation in coral reef communities Alastair R. Harborne, Corresponding Author Alastair R. Harborne Marine Spatial Ecology Laboratory, School of Biosciences, Hatherly Laboratory, University of Exeter, Prince of Wales Road, Exeter, EX4 4PS, UK; *Correspondence author. E-mail: a.r.harborne@ex.ac.ukSearch for more papers by this authorPeter J. Mumby, Peter J. Mumby Marine Spatial Ecology Laboratory, School of Biosciences, Hatherly Laboratory, University of Exeter, Prince of Wales Road, Exeter, EX4 4PS, UK;Search for more papers by this authorCarrie V. Kappel, Carrie V. Kappel National Center for Ecological Analysis and Synthesis, University of California–Santa Barbara, 735 State Street, Suite 300, Santa Barbara, CA 93101, USA; Hopkins Marine Station, Stanford University, Oceanview Boulevard, Pacific Grove, CA 93950-3094, USA;Search for more papers by this authorCraig P. Dahlgren, Craig P. Dahlgren Perry Institute for Marine Science, 100 N. US Highway 1, Suite 202, Jupiter, FL 33477-5112, USA;Search for more papers by this authorFiorenza Micheli, Fiorenza Micheli Hopkins Marine Station, Stanford University, Oceanview Boulevard, Pacific Grove, CA 93950-3094, USA;Search for more papers by this authorKatherine E. Holmes, Katherine E. Holmes Center for Biodiversity and Conservation, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, USA;Search for more papers by this authorJames N. Sanchirico, James N. Sanchirico Department of Environmental Science and Policy, University of California, Davis, One Shields Avenue, 2140 Wickson Hall, Davis, CA 95616, USA; andSearch for more papers by this authorKenneth Broad, Kenneth Broad Rosenstiel School of Marine and Atmospheric Science, Division of Marine Affairs and Policy, 4600 Rickenbacker, Causeway, Miami, FL 33149, USASearch for more papers by this authorIan A. Elliott, Ian A. Elliott Marine Spatial Ecology Laboratory, School of Biosciences, Hatherly Laboratory, University of Exeter, Prince of Wales Road, Exeter, EX4 4PS, UK;Search for more papers by this authorDaniel R. Brumbaugh, Daniel R. Brumbaugh Center for Biodiversity and Conservation, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, USA;Search for more papers by this author Alastair R. Harborne, Corresponding Author Alastair R. Harborne Marine Spatial Ecology Laboratory, School of Biosciences, Hatherly Laboratory, University of Exeter, Prince of Wales Road, Exeter, EX4 4PS, UK; *Correspondence author. E-mail: a.r.harborne@ex.ac.ukSearch for more papers by this authorPeter J. Mumby, Peter J. Mumby Marine Spatial Ecology Laboratory, School of Biosciences, Hatherly Laboratory, University of Exeter, Prince of Wales Road, Exeter, EX4 4PS, UK;Search for more papers by this authorCarrie V. Kappel, Carrie V. Kappel National Center for Ecological Analysis and Synthesis, University of California–Santa Barbara, 735 State Street, Suite 300, Santa Barbara, CA 93101, USA; Hopkins Marine Station, Stanford University, Oceanview Boulevard, Pacific Grove, CA 93950-3094, USA;Search for more papers by this authorCraig P. Dahlgren, Craig P. Dahlgren Perry Institute for Marine Science, 100 N. US Highway 1, Suite 202, Jupiter, FL 33477-5112, USA;Search for more papers by this authorFiorenza Micheli, Fiorenza Micheli Hopkins Marine Station, Stanford University, Oceanview Boulevard, Pacific Grove, CA 93950-3094, USA;Search for more papers by this authorKatherine E. Holmes, Katherine E. Holmes Center for Biodiversity and Conservation, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, USA;Search for more papers by this authorJames N. Sanchirico, James N. Sanchirico Department of Environmental Science and Policy, University of California, Davis, One Shields Avenue, 2140 Wickson Hall, Davis, CA 95616, USA; andSearch for more papers by this authorKenneth Broad, Kenneth Broad Rosenstiel School of Marine and Atmospheric Science, Division of Marine Affairs and Policy, 4600 Rickenbacker, Causeway, Miami, FL 33149, USASearch for more papers by this authorIan A. Elliott, Ian A. Elliott Marine Spatial Ecology Laboratory, School of Biosciences, Hatherly Laboratory, University of Exeter, Prince of Wales Road, Exeter, EX4 4PS, UK;Search for more papers by this authorDaniel R. Brumbaugh, Daniel R. Brumbaugh Center for Biodiversity and Conservation, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, USA;Search for more papers by this author First published: 09 July 2008 https://doi.org/10.1111/j.1365-2664.2008.01490.xCitations: 44AboutSectionsPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinked InRedditWechat Summary 1 No-take reserves are a common tool for fisheries management and biodiversity conservation in marine ecosystems. Despite much discussion of their benefits, data documenting many reserve effects are surprisingly scarce. Several studies have also been criticized for a lack of rigour so that changes within reserves cannot be separated from underlying natural variation and attributed unequivocally to protection. 2 We sampled both benthic (video quadrats) and associated fish communities (underwater visual censuses) in a well-enforced reserve in The Bahamas. Sampling was explicitly stratified by habitat ('Montastraea reef' and 'gorgonian plain'). To distinguish reserve effects from natural variation, we compared changes inside and outside the reserve with those seen at equivalent spatial scales in other reef systems in the Bahamian archipelago that lack reserves. Reserve-level differences in benthic or fish communities not documented in other reef systems are categorized as 'robust' effects. 3 Robust reserve effects were limited to Montastraea reefs. The reserve supported an average of ª 15% more fish species per site compared to outside the reserve. This pattern was particularly driven by more large-bodied grouper, damselfish, and butterflyfish species inside the reserve. Increases in fish biomass and differences in community structure inside the reserve were limited to large-bodied groupers. Increased grazing pressure by parrotfishes in the reserve has lowered macroalgal cover, and caused previously undocumented changes in benthic community structure compared to sites outside the reserve. 4 Some reserve-level differences in fish communities were categorized as 'misleading' because equivalent differences were seen in other reef systems, and are likely to be caused by natural intra-habitat variation. Separation of robust and misleading results was only possible because of archipelago-scale sampling. 5 Synthesis and applications. The Bahamas represents a relatively lightly fished system within the Caribbean. However, cessation of fishing has still increased the mean number of species, the abundance of the most highly prized fishes and, through trophic cascades, altered benthic community structure. In certain habitats, reserves are clearly important for conserving fisheries and biodiversity. However, reserve effects must be explicitly separated from confounding variables to ensure conservation benefits are accurately identified and reported, and not oversold to managers and local stakeholders. Introduction Protected areas are a key conservation tool in a range of ecosystems, including marine environments where they are primarily used to conserve biodiversity and manage fisheries (e.g. Sale et al. 2005). In particular, no-take marine reserves are established with the aim of conserving fish diversity, facilitating recovery of depleted stocks, and causing spill-over of individuals and larvae to fished areas (e.g. Lubchenco et al. 2003). The increasing number of marine reserves has facilitated numerous empirical studies and subsequent meta-analyses and reviews documenting the changes to fish communities, such as increases in abundance, biomass, size, and diversity, following the cessation of fishing (e.g. Côté, Mosqueira & Reynolds 2001; Halpern 2003; Micheli et al. 2004). Consequently, reserves are widely advocated to reverse trends of declining coral reef health where a range of factors, including over-exploitation of grazing fishes, has caused significant declines in coral cover and increases in macroalgal abundance (Gardner et al. 2003). Despite an apparent wealth of data, evidence for some of the expected effects of marine reserves is still equivocal, and limited by a lack of definitive experiments at appropriate spatio-temporal scales (Russ 2002). Studies clearly demonstrating reserve effects should not be confounded by factors such as insufficient spatial or temporal replication or an absence of suitable control sites outside reserves (Willis et al. 2003). For example, 'habitat effects' (habitat differences inside and outside reserves) clearly need to be separated from 'reserve effects' (García-Charton & Pérez-Ruzafa 1999; Halpern 2003). In the absence of a 'BACI' (Before/After, Control/Impact) sampling design (Underwood 1994), even within relatively well-replicated studies, it is difficult to separate reserve effects from natural variability associated with intra-habitat abiotic and biotic gradients, but this should be an important consideration during sampling design (Fraschetti et al. 2002). Finally, studies addressing the effects of reserves should be limited to locations where protection has been effectively enforced for several years. In this study, we compared fish and benthic communities inside and outside a Caribbean coral reef reserve (Exuma Cays Land and Sea Park [ECLSP], The Bahamas), stratified by habitat type, with data collected at equivalent spatial scales on reefs without reserves throughout the Bahamian archipelago. The latter data provide insights into the natural spatial variability of community structure where data before reserve establishment are not available, which is the case here and in many marine reserve studies (an 'ACI' sampling design). If differences observed between reefs inside and outside the reserve are not evident elsewhere, we can conclude that such variation can be attributed to reserve establishment. In the present study, fish data were collected to examine any direct effects of cessation of fishing, and also indirect effects caused by predator–prey relationships (Micheli et al. 2004). We used benthic community data to examine the effects of any trophic cascades caused by alterations in fish communities. For example, the reduction in fishing pressure has led to a net increase in grazing by parrotfishes, and a reduction in macroalgal cover inside the ECLSP (Mumby et al. 2006). However, Mumby et al. (2006) did not analyse benthic community structure inside and outside the reserve. Documenting changes in benthic community structure is important because of, for example, species-specific differences in the abilities of algae to compete with corals for space (McCook, Jompa & Diaz-Pulido 2001). Effects of the reserve on fish and benthic communities were documented in two hard-bottom habitat types: 'Montastraea reef' (coral-rich areas visually dominated by Montastraea spp.) and 'gorgonian plain' (low-relief areas visually dominated by fleshy brown algae and sparse gorgonians). We hypothesized that changes in fish communities would be more pronounced on Montastraea reefs because it is the favoured habitat of adults of many commercially important species (e.g. the large-bodied grouper Mycteroperca bonaci, M. tigris, M. venenosa, M. interstitialis, and Epinephelus striatus; Sluka et al. 1998) and, when fishing pressure is low, the higher topographical complexity of this habitat will support a greater biomass and number of species of fishes (Gratwicke & Speight 2005). Furthermore, because of higher densities of parrotfishes on Montastraea reefs (Harborne et al. 2006), we expected grazing-mediated changes in benthic communities to be clearer in this habitat type. Materials and methods study sites and sampling design Surveys in and around the ECLSP were conducted in October 2004. The ECLSP lies near the centre of the Bahamian archipelago (see Supplementary material Appendix S1), is large (442 km2) and was established in 1958 (Chiappone & Sullivan Sealey 2000). There is no evidence of the reserve containing especially healthy or diverse reefs before its establishment (Ray 1958). A ban on fishing has been enforced by warden patrols since 1986. Poaching inside the ECLSP has been assessed as low (supporting material associated with Mora et al. 2006). Montastraea forereef was sampled at three sites (≈ 150 × ≈ 150 m) near the centre of the ECLSP, three sites between 5·8 km and 18·1 km north of the Park and three sites around Lee Stocking Island (LSI) ≈ 70 km south of the Park (see Supplementary material Appendix S1). Gorgonian plain (the predominant forereef habitat in the area) in the ECLSP was sampled at nine sites centred on the middle of the reserve. Gorgonian plain was also sampled at six sites between 1 km and 10 km both south and north of the reserve (9 + 6 + 6 = 21 sites in total). Data from these sites were used for comparisons of communities inside and outside the ECLSP, stratified by habitat type. To assess natural variation, we sampled five additional island systems (July 2002–November 2003). These additional data facilitated intra-island comparisons at the same spatial scale (tens of kilometres) as in the Exumas, but on reefs without reserves. The additional island systems were Andros, San Salvador, and South Caicos (Turks and Caicos Islands) (Montastraea reef and gorgonian plain) and Bimini and Abaco (gorgonian plain only; see Supplementary material Appendix S1). In each island system, we sampled three groups of sites 5–10 km apart, equivalent to the group of sites inside, and the two groups of sites outside, the ECLSP. Each group of sites consisted of three (two on three occasions) replicate Montastraea reefs and/or a minimum of two gorgonian plain sites. Fishing effort at each site was estimated, and varied among islands (see Supplementary material Appendix S2). Critically, because of the presence of the ECLSP, the Exumas appeared to be the only reef system where fishing pressure varies at the scale of tens of kilometres. Any significant results within any of the other reef systems can, therefore, be attributed to natural variation rather than variable fishing pressure. community characterization At each site, 30–40 randomly placed 1 m2 quadrats were used to quantify the species composition of the benthic community. Content of quadrats was filmed in 20-cm swathes, using a high-resolution digital video camera. Following completion of all the swathes within a given quadrat, cryptic organisms and areas of high relief (e.g. under ledges) were filmed in more detail. Depth and rugosity (maximum vertical relief inside quadrat assigned to one category from 0–10 cm, 10–50 cm, 50–100 cm, 100–200 cm, and > 200 cm) were also measured for each quadrat in situ. This measure of rugosity was found to correlate well to more labour-intensive chain-transect methods (r = 0·67, P < 0·001). The digital video of each quadrat was projected onto a large monitor for identification (presence/absence) of species of scleractinian corals (minimum diameter, 1 cm), and macroalgae, macroscopic mobile invertebrates, sponges, and gorgonians to the highest taxonomic resolution possible. Data were then converted to frequency of occurrences (number of times present divided by number of quadrats) for each taxon at each site. Coral and macroalgal cover were assessed from a mean of 14·1 (Montastraea reef) or 9·4 (gorgonian plain) of the 1 m2 quadrats used for characterization of benthic communities. Coral and macroalgal cover in each 1 m2 quadrat was recorded as the mean of five randomly sub-sampled areas of 20 × 20 cm (0·04 m2). Note that we measured macroalgal cover rather than the cover of all parrotfish food items (macroalgae and large turfs) measured by Mumby et al. (2006). Presenting data on percentage cover of macroalgae (rather than macro and turf algae; cf. Mumby et al. 2006) is consistent with other patterns documented in this study. All but nocturnal (e.g. Apogonidae) and highly cryptic (Clinidae and Gobiidae) fish species were surveyed using discrete group visual fish censuses (Green & Alevizon 1989) at the same time as the benthic sampling. Species were divided into three groups and their density and size (to the nearest centimetre) estimated along belt transects at each island system. Transect size and number were optimized using data from equivalent surveys within the Caribbean (Mumby et al. 2004). Four 30 × 2 m transects were surveyed for small demersal families such as Pomacentridae; ten 30 × 4 m transects were surveyed for mid-sized demersal families such as Scaridae, and five 50 × 4 m transects were used for large demersal and pelagic fish such as Serranidae. All subsequent fish analyses (undertaken at transect-level resolution, with the exception of tests on the entire community) consider biomass; fish lengths were converted to biomass using allometric relationships (Bohnsack & Harper 1988). data analyses Percentage cover data were arcsine-transformed (Zar 1996), and other variables were square root, cube root or Box-Cox-transformed to ensure normality where possible. Differences in univariate parameters inside the ECLSP compared to sites to the north and south were tested using anova (plus Tukey's HSD pairwise comparisons), Kruskal–Wallis or Mann–Whitney tests as appropriate. Effects of the ECLSP on coral and algal cover were tested by ancova with the percentage cover of sand as a covariate to control for any variation in hard substratum area among sites. Tests for significant variation in community structure (repeated for the entire benthic and fish communities and also individual fish families) were conducted using anosim (Clarke 1993), with fish data square root transformed. The discriminating taxa were determined using Similarity Percentage (SIMPER) analysis (Clarke 1993) to examine significant anosim results further. Parrotfishes in the Montastraea reef habitat are described in detail elsewhere (Mumby et al. 2006), and are not reanalysed here. Parrotfish grazing in the gorgonian plain habitat was calculated using the model described in Mumby et al. (2006). Wherever significant reserve-level effects were detected, analogous tests were conducted on data from the other island systems to assess natural variation. Including tests involving the ECLSP, a total of 12 comparisons were possible for the Montastraea reef habitat (three groups of sites in four reef systems; Table 1), and 18 were possible for gorgonian plains (three groups of sites in six reef systems). Table 1. Definition and diagrammatic representation of terms used to categorize the effects of the Exuma Cays Land and Sea Park (ECLSP) on fish and benthic communities. Diagrammatic representation is for fish communities on Montastraea reefs. Solid circles represent replicate sites sampled. Solid arrows represent significant comparisons (P < 0·05); dotted arrows represent non-significant comparisons Reserve effect category Definition Diagrammatic representation Robust Within the Exuma Cays, only significant pairwise comparisons between the ECLSP and sites to both the north and south (fishes) or only a significant difference in the comparison between sites inside the ECLSP with those outside (benthos) and no differences at the same scale on reefs around any other island Potential* Within the Exuma Cays, only a significant pairwise comparison between the ECLSP and sites to either the north or the south and no differences at the same scale on reefs around any other island Misleading Within the Exuma Cays, only one or two significant pairwise comparisons between the ECLSP and sites to the north and south (fishes) or only a significant difference in the comparison between sites inside the ECLSP with those outside (benthos) but differences at the same scale on reefs around any other island Absent Within the Exuma Cays, no significant pairwise comparisons between the ECLSP and sites to the north and south (fishes) or no significant difference in the comparison between sites inside the ECLSP with those outside (benthos) * not possible for tests of benthic communities. categorization of results Results for differences in fish and benthic communities inside the ECLSP were defined as 'robust', 'potential', 'misleading' or 'absent' (Table 1). Benthic community structure data had to be calculated for an entire site (frequency of occurrence data are the number of times a taxon was seen divided by total number of quadrats) while individual fish transects at a site were treated as replicates. This additional statistical power allowed pairwise comparisons for fish community structure between communities inside the ECLSP and sites to the north and sites to the south. In contrast, benthic communities inside the ECLSP were compared to all sites outside the Park combined, and then, two further comparisons were made between sites to the north vs. sites in the ECLSP and to the south combined, and between sites to the south vs. sites in the ECLSP and to the north combined. For this reason, fish results could be robust, potential, misleading or absent. Effects on benthic communities could only be robust, misleading or absent. Justification of these categories can be made using the probability theory (see Supplementary material Appendix S3). Results A total of 142 fish species and 218 benthic taxa were recorded at the Montastraea reef and gorgonian plain sites. Within the Exuma Cays, there were no differences in mean depth or vertical relief in either habitat inside compared to outside the reserve (anova: P > 0·05). Montastraea reefs were found at mean depths of ≈ 9–12 m with mean vertical relief of ≈ 65–72 cm. Gorgonian plains were found at mean depths of ≈ 7–10 m with mean vertical relief of ≈ 10–27 cm. Effects of the ECLSP on the biological communities are summarized (Table 2), described in the subsequent text, and documented in full in Supplementary material Appendix S4. Table 2. Summary of the effects of the ECLSP on fish and benthic communities. Reserve effect categories defined in Table 1. Effects on fishes are presented first in each category. CS, community structure; BM, biomass Habitat Reserve effect Robust Potential Misleading None Montastraea reef • Mean number of fish species• Number of large serranid*, territorial pomacentrid† and chaetodontid species• CS and BM of large serranids• CS of scarids, and grazing by scarids (Mumby et al. 2006)• Macroalgal percentage cover• Benthic CS • CS of lutjanids • CS of chaetodontids and territorial pomacentrids†• BM of haemulids • Shannon diversity of the entire fish community• CS or BM of the entire fish community• CS or BM of acanthurids and small serranids‡• CS of haemulids • BM of chaetodontids, lutjanids, and territorial pomacentrids†• Number of benthic species or benthic Shannon diversity• Coral percentage cover Gorgonian plain – • BM of large serranids* and lutjanids • CS of chaetodontids • Mean number of fish species and Shannon diversity of the entire fish community• CS or BM of the entire fish community• CS or BM of acanthurids, haemulids, territorial pomacentrids†, scarids, and small serranids‡• CS of lutjanids and large serranids*• BM of chaetodontids• Parrotfish grazing pressure• Number of benthic species or benthic Shannon diversity• Coral and algal percentage cover• Benthic CS * large-bodied, commercially important only (Mycteroperca bonaci, M. tigris, M. venenosa, M. interstitialis and Epinephelus striatus). † Stegastes and Microspathodon only. ‡ Cephalopholis fulvus, C. cruentatus, Epinephelus guttatus, E. adscensionis. robust park effects Montastraea reef sites inside the ECLSP had significantly more fish species than those outside the Park (mean 48·67 vs. 42·33; one-tailed Mann–Whitney: P = 0·035). None of the other 11 comparisons within the four reef systems showed a significantly higher number of species at the same spatial scale (one-tailed Mann–Whitney: P > 0·05). To investigate whether species in particular fish families were driving the high number of species seen in the ECLSP, we conducted Monte Carlo simulations with the data set from all four reef systems (a conservative approach incorporating any large-scale variations in species richness). We calculated whether removing individual families reduced the difference between sites inside the ECLSP compared to those outside by a significantly (P < 0·05) greater amount than removing an equal number of species at random (10 000 permutations). Significant results were obtained for: (i) large-bodied, commercially important serranids (Mycteroperca bonaci, M. tigris, M. venenosa, M. interstitialis and E. striatus; P = 0·008); (ii) territorial pomacentrids (six Stegastes species and Microspathodon chrysurus; P = 0·042); and (iii) chaetodontids (five species; P = 0·048). Furthermore, the ECLSP has had a clear effect on mean large serranid biomass on Montastraea reefs (4750·96 g 200 m−2 inside vs. 472·47 and 1585·58 for north of the ECLSP and LSI respectively; anova: F(2,33) = 10·93, P < 0·001). Large serranid community structure was also significantly different inside the ECLSP (anosim: Global R = 0·220; P = 0·003), primarily caused by the increased abundance of E. striatus and Mycteroperca tigris (Fig. 1). Figure 1Open in figure viewerPowerPoint Mean biomass of each large-bodied grouper species (E., Epinephelus; M., Mycteroperca) inside and outside the Exuma Cays Land and Sea Park (ECLSP). Values represent percentage contribution of each species to SIMPER analysis of the dissimilarity between inside and outside the ECLSP [percentage contribution = average contribution/average dissimilarity between sites inside and outside the reserve (= 75·39%)]. n = 24 outside ECLSP and 12 inside. Only macroalgal cover on Montastraea reefs was significantly different (lower) inside the ECLSP (2·8% inside vs. 15·32% and 17·98% to the north and south, respectively; ancova: F(2,125)= 58·05, P < 0·001). Analysis of benthic community structure on Montastraea reefs inside compared to those outside the Park showed significant differences (anosim: R = 0·377, P = 0·048). The taxa driving such differences included a decrease in the frequency of occurrence of the brown macroalgae Sargassum spp. (not including S. hystrix) and Dictyota, and an increase in the abundance of S. hystrix, Lobophora variegata, and red coralline algae in the ECLSP (Table 3). Three species of corals (Montastraea franksi, Agaricia agaricites, and Millepora alcicornis) had higher frequency of occurrences within the ECLSP. Table 3. SIMPER analysis of the top 10 taxa characterising the dissimilarity between benthic communities inside and outside the Exuma Cays Land and Sea Park (ECLSP). Percentage contribution = average contribution/average dissimilarity between sites inside and outside the reserve (= 34·73%). n = 6 outside ECLSP and 3 inside the reserve Taxon Mean frequency of occurrence (%) (SE) Percentage contribution Outside ECLSP Inside ECLSP Sargassum spp. 72·97 (10·93) 12·80 (7·82) 4·14 Sargassum hystrix 36·97 (13·15) 91·60 (7·09) 3·83 Pseudopterogorgia spp. 41·24 (12·56) 69·06 (24·81) 2·96 Dictyota spp. 78·26 (12·08) 51·54 (14·04) 2·70 Montastraea franksi 13·02 (5·53) 49·37 (14·09) 2·65 Ircinia felix/strobilina 31·81 (9·93) 62·61 (13·41) 2·54 Lobophora variegata 33·14 (11·18) 48·50 (22·05) 2·41 Agaricia agaricites 39·51 (6·97) 74·23 (5·17) 2·39 Red coralline algae 43·52 (8·52) 75·23 (7·68) 2·17 Millepora alcicornis 38·29 (4·57) 46·69 (22·86) 2·09 potential park effects The community structure of lutjanids on Montastraea reefs varied between the ECLSP and LSI (anosim: R = 0·173, P = 0·021), but not between the ECLSP and sites to the north of the Park. There was significant variation in the biomass of large serranids and lutjanids on gorgonian plains between the ECLSP and sites to the north (Tukey's HSD: P < 0·05; 88·38 g 200 m−2 to the north vs. 1801·42 in the Park, and 00·00 g 200 m−2 to the north vs. 2160·22 in the Park, respectively), but not between the Park and sites to the south (Tukey's HSD: P > 0·05). misleading park effects Territorial pomacentrid community structure on Montastraea reefs inside the ECLSP differed significantly from both locations outside the reserve, but six out of nine equivalent comparisons in other reef systems lacking reserves were also significant (anosim: P < 0·05). Also on Montastraea reefs, significant differences in community structure of chaetodontids were found between ECLSP and LSI sites (anosim: R = 0·064; P = 0·010) and the biomass of haemulids between the ECLSP and north of the Park (Tukey's HSD: P < 0·05), but equivalent differences were seen in other reef systems (one and four comparisons respectively). Chaetodontid community structure d
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