Assessment of the initial state of two reserves, Micro áreas Ecoturísticas Litorales (MAEL), in Gran Canaria, Canary Islands

Assessment of the initial state of two reserves, Micro áreas Ecoturísticas Litorales (MAEL), in Gran Canaria, Canary Islands Tesis de Máster presentada por Javier R. Vidal López Dirigida por Fernando Tuya y Arturo Boyra Tutorizada por Antonio Juan González Ramos Tutor Directores Tesinando Las Palmas de Gran Canaria, julio de 2014 3 CONTENTS 1. ABSTRACT……………………………………………………………………………..4-6 2. INTRODUCTION……………………………………………………………………6-8 3. MATERIAL AND METHODS…………………………………………………8-12 3.1 Study area and sampling design……………………………………………..8 3.2 Selection of fish species, trophic and functional groups………………………………………………………………………………………….10 3.3 Data analysis…………………………………………………………………………..12 4. RESULTS……………………………………………………………………………13-23 4.1 Commercially-targeted species……………………………………………...13 4.2 Taxonomic, trophic and functional diversity………………………….20 4.3 Correlation between taxonomic, trophic and functional diversity.................................................................................................................22 5. DISCUSSION……………………………………………………………………..24-26 5.1 Overall results…………………………………..................................................24 5.2 Cabrón-Risco Verde……………………………………………………………….24 5.3 Canteras-Confital…………………………………............................................25 5.4 Link between taxonomic, trophic and functional diversity……………………………………………………………………………………….26 6. ACKNOWLEDGES……………………………………………………………………26 7. REFERENCES……………………………………………………………………..27-32 8. APPENDIX………………………………………………………………………………33 4 1. Abstract The establishment of ' Micro áreas ecoturísticas litorales ' (MAELs) is a new strategy of marine conservation and management, based on a bottom-up governance strategy. In this study, we assessed the initial state ('state 0') of two MAELs at Gran Canaria Island (Canary Islands): 'El Cabrón' and 'Las Canteras', located in the east and in north-east coast of Gran Canaria Island, respectively. Specifically, we evaluated differences in the abundance and biomass of target fish species between inside (MAELI) and outside (MAELO) these two proposed reserves; five commercially-targeted fish species were selected: the parrotfish, Sparisoma cretense , the white sea-bream, Diplodus sargus cadenati , the common two-banded sea-bream, Diplodus vulgaris , the island grouper, Mycteroperca fusca , and salema, Sarpa salpa . Fish assemblages were sampled at 7 times within each management category in during summer and autumn of 2013. Univariate tests provided an overall moderate ‘reserve effect’ for the initial state of both MAELs. Mycteroperca fusca and Diplodus vulgaris showed greater abundances and biomasses, respectively, within the ‘El Cabrón’ MAELI compared to the adjacent MAELO. Sparisoma cretense , Diplodus sargus cadenati , Diplodus vulgaris and Sarpa salpa showed greater abundances and biomasses within the 'Las Canteras' MAELI compared to the adjacent MAELO. Furthermore, we tested whether taxonomic diversity (through calculation of the Shannon diversity index for the entire fish assemblage) may be a surrogate for the trophic and functional diversity of the fish assemblage to adequately support the capacity of MAMPs to preserve marine biodiversity. Our data suggested a neat link between them. Keywords: Micro áreas marinas protegidas, MAMPs, functional diversity, taxonomic diversity, trophic diversity, target species and reserve effect. Resumen ' Micro áreas ecoturísticas litorales ' (MAELs) es una nueva estrategia de gestión y conservación marina, basada en la estructura de decisión bottom-up. En este estudio, evaluamos el estado inicial ('estado 0') de dos MAELs en Gran Canaria (Islas Canarias). 'El Cabrón' y 'Las Canteras', localizadas en este y noreste de la isla de Gran Canaria, respectivamente. Específicamente, evaluamos diferencias en abundancia y biomasa de especies de peces objetivo-comerciales, dentro (MAELI) y fuera (MAELO) de las dos reservas propuestas; Las cinco especies comerciales estudiadas fueron: Sparisoma cretense, Diplodus sarguscadenati, Diplodus vulgaris, Mycteroperca fusca y Sarpa salpa. Las poblaciones de peces 5 fueron muestreadas durante 7 tiempos comprendidos entre verano y otoño de 2013. Tests univariantes mostraron un moderado 'efecto reserva' durante el estado inicial de las dos MAELS. Mycteroperca fusca y Diplodus vulgaris mostraron mayores abundancias y biomasas, respectivamente, dentro ‘El Cabrón’ MAELI comparadas con la zona adyacente MAELO. Sparisoma cretense , Diplodus sargus cadenati , Diplodus vulgaris y Sarpa salpa mostraron mayores abundancias y biomasas dentro de 'Las Canteras' MAELI comparado con la zona adyacente MAELO . Además, testamos si la diversidad taxonómica (mediante el cálculo del índice de diversidad de Shannon para la población de peces) puede estar relacionada con la diversidad trófica y funcional de la población de peces para evaluar la capacidad de preservar la biodiversidad marina de las MAMPs. Encontramos en nuestros datos una unión entre ellas. Palabras clave: Micro áreas marinas protegidas, MAMPs, diversidad funcional, diversidad taxonómica, diversidad trófica, especies objetivo y efecto reserva. 6 2. Introduction The United Nations (UN) for Food and Agriculture Organization's (FAO) estimate that 75% of the world's fisheries are actually overexploited (Ray Hilborn et al., 2005). In particular, shallow-water fishery resources are being subjected to strong anthropogenic pressures, including overexploitation in the last decades (Lauck et al., 1998; Castilla., 2000). In fact, the main source of erosion of marine ecosystem biodiversity is overfishing (Jackson et al., 2001), but there are others sources of biodiversity loss such as pollution (European agency, 2006), invasion by alien species (Gollasch., 2006) and catastrophes induced by global warming (Harley et al., 2006). A solution to preserve coastal resources consists in the establishment of Marine protected areas (MPAs). Two main objectives have been identified when addressing the purposes of MPAs: ensuring sustainable use of economic resources, and protecting biodiversity – valuable species, habitats and landscapes (Salm et al. 2000): In turn, the number of MPA has been increasing in the last decades, to preserve and manage coastal resources and their habitats, and so coastal ecosystems and their biodiversity (Fraschetti et al., 2011). The efficacy of MPAs has been widely discussed (Harmelin-Vivien et al., 1995; Guidetti et al., 2005; Micheli et al ., 2005), what depends on a range of different factors (Barret et al., 2007), as the size of no-take and adjacent buffering areas (Claudet 2008; 2010), the time since protection (Micheli et al., 2004), connectivity with adjacent zones (Vega Fernandez et al., 2008) and, of course, the effective level of enforcement and compliance by local administrations and users (Claudet, 2010). A new strategy of marine conservation and management promote the establishment of Micro áreas ecoturísticas litorales (MAELs), which are based on a bottom-up governance philosophy. In this case, local communities boost the establishment and declaration of protected areas via local administrations, rather than relying on legal authorities with competence in fisheries and conservation management The overall goal of MAELs is to contribute to the conservation of the biological diversity and productivity of the oceans, including ecosystem processes, but promoting sustainable uses such as eco-tourism, traditional fishing, scientific research and so to improve local economy. Traditionally, management of coastal ecosystems have followed a (top-down approach), where governmental bodies decide on specific regulations. Frequently, MPA governing bodies have not taken full responsibilities in their attempts at management; in turn, managers fail to recognize and encompass 7 stakeholder opinions in their attempts at realizing a successful MPA (Himes, 2007). A different strategy, promotes stakeholder participation, via a bottom-up strategy, where local communities, users and local-administrations collaborate since the earliest stage of creation of these reserves. It has been widely recognized that public participation and local community involvement is an essential factor contributing to the success of MPAs (Fiske, 1992; Wolfenden, 1994). For example, changes in policies concerning how exploitation of marine resources in the Philippines should be implemented have shifted from a centralized bureaucracy to co-management among local communities, local administration, and the national government (Alcala et al., 2006). Galicia (NW Spain) has pioneered co-management initiatives proposing the creation of a marine reserve, designed and developed by the fishers in partnership with biologists and social scientists, environmentalists and members of the autonomous Government of Galicia (Perez, 2013). Typically, reserves established via bottom-up governance approaches are of reduced size. The positive effects of small-sized reserves, i.e. with a similar size to the MAELs, have been demonstrated in many cases (Lester et al., 2009; Afonso et al., 2011; Hort et al., 2013), we show the positive the effects of smallsized reserves around the world (Appendix 1). Moreover, a small size do facilitate co-management between stakeholders (e.g. fishers, users, divers, administration), favoring a sustainable management. A goal of any marine reserve is to evaluate the expected benefits, either from an ecological point of view, or also through social and economic metrics. In the literature, there are many different population parameters as bioindicators to assess such expected ‘reserve effect’. Many biological studies have focused on one or several target species and, in many circumstances, reported increased abundances and larger sizes inside MPAs (e.g. Barret et al., 2006; Tuya et al., 2006; Brito et al., 1997, 1998, 2001). However, conservation of particular species is questioned, because it depends on species-specific life traits (Villamor et al., 2012). Moreover, understanding species’ role in nature is limited; less 1% have been studied (Wilson, 2000), a species can be considered to be functionally redundant when the community contains functionally-analogous species, so that its disappearance from the community entails no measurable loss of functionality (Duarte, 2000). Relationships between species, their biodiversity and ecosystem function are important for predicting the ecological and economic impact of human 8 interventions (Armsworth et al., 2007). Indices based only on the taxonomic identity provide an incomplete view of biodiversity (Villeger et al., 2010). A recent consensus point out the importance of particular taxa rather than species richness to explain ecosystem processes in aquatic communities (O’Connor et al., 2008). A step further in biodiversity assessment needs to consider the role of each species in ecosystems or species responses to environmental conditions. This can be somehow approached through the estimation of trophic and functional diversity of biotic communities in conjunction with taxonomic diversity studies (Mc Gill et al., 2006). Thereby, taxonomic studies that traditionally have focused on the identity of species may be complemented with trophic and functional diversity approaches to adequately support the capacity of MAELs to preserve marine biodiversity. In this study, our goal was to assess the initial state ('state 0') of two MAELs at Gran Canaria Island, by comparing several descriptors inside and outside these two proposed reserves. This included the abundances and total biomasses of several target species (univariate responses), as well as estimators of the taxonomic diversity (through Shannon diversity index) and trophic and functional diversity (multivariate responses). We ultimately aimed to unravel whether taxonomic, trophic and functional diversity were correlated within the study systems. 3. Material and methods 3.1 Study area and sampling design This study focused on two recently proposed Micro áreas ecoturísticas litorales (MAELs) at Gran Canaria Island. The first, 'El Cabrón', is located in the east side of Gran Canaria Island. The second, 'Las Canteras', is located in north-east side of the island (Fig 1). Both zones are biogeographically and climatically similar. At both locations, two adjacent areas were studied, one within the proposed MAEL: no take zone (MAELI), where the exploitation of benthic and demersal resources will be prohibited, hereafter so-called 'Cabrón' and 'Canteras', respectively, and two adjacent areas outside these protected areas (MAELO), hereafter so-called 'Risco Verde' and 'Confital', respectively, where fishing activities are allowed. Data collection were undertaken during Summer-Autumn of 2013 (Table 1), at 7 random times. From an environmental point of view, both areas at each location 9 were similar in terms of depth, type of bottom, wave climate and oceanography. At Cabrón and Risco Verde, sampling was performed between 10 and 18 m depth, on rocky bottoms of similar structural complexity, to minimize the possible effect of the habitat (so-called 'habitat effect', sensu García-Charton and Pérez-Ruzafa., 1999). At ‘Canteras’ and ‘Confital’, sampling took place between 3-5 m depth, on rocky bottoms of similar structural complexity. Table 1. Sampled localities and times to compare fish assemblages between MAELI and MAELO at Gran Canaria Island. Locality (UTM) Cabrón Risco Verde Canteras Confital Date 27º52’N 15º23’W 27º51’N 15º23’W 28º08’N 15º26’W 28º09’N 15º26W T1 26/07/2013 8/06/2013 T2 10/08/2013 22/07/2013 T3 12/09/2013 30/07/2013 T4 22/09/2013 7/08/2013 T5 30/09/2013 12/08/2013 T6 7/10/2013 2/09/2013 T7 15/10/2013 12/10/2013 16 Las Canteras Vs Confital Abundance Biomass Diplodus sargus cadenati df MS F P df MS F P Locality 1 77.709 22.650 0.001 1 3068.685 33.378 0.001 Time 6 4.115 3.312 0.009 6 112.901 1.896 0.104 LoxTi 6 3.430 2.761 0.022 6 91.936 1.544 0.184 Res 42 1.242 42 59.520 Total 55 55 Diplodus vulgaris df MS F P df MS F P Locality 1 2.448 9.483 0.025 1 208.878 6.504 0.038 Time 6 0.176 0.549 0.780 6 35.177 0.837 0.564 LoxTi 6 0.258 0.805 0.564 6 32.117 0.764 0.642 Res 42 0.321 42 42.021 Total 55 55 Sparisoma cretense df MS F P df MS F P Locality 1 14.488 14.479 0.005 1 5.084 5.085 0.046 Time 6 0.890 0.713 0.638 6 2852.681 1.011 0.433 LoxTi 6 1.000 0.801 0.573 6 4444.058 1.575 0.171 Res 42 1.249 42 2820.191 Total 55 55 Sarpa salpa df MS F P df MS F P Locality 1 72.470 24.679 0.004 1 24461.08 8.993 0.025 Time 6 3.552 0.766 0.606 6 3523.73 1.171 0.344 LoxTi 6 2.936 0.633 0.707 6 2720.00 0.904 0.510 Res 42 4.636 42 3008.84 Total 55 55 17 Figure 3. (A, B) Abundance and (C, D) total biomass (± SE) of the white seabream, Diplodus sargus cadenati , at areas within MAELI (black bars) or MAELO (gray bars) at each sampling time. 18 Figure 4. (A, C) Abundance and (B, D) total biomass (± SE) of the common twobanded sea-bream , Diplodus vulgaris , at areas within MAELI (black bars) or MAELO (gray bars) at each sampling time. 19 Figure 5. (A) Abundance and (B) total biomass (±SE) of the island grouper, Mycteroperca fusca, at areas within MAELI (black bars) or MAELO (gray bars) at each sampling time. Figure 6. (A) Abundance and (B) total biomass (±SE) of the Salema, Salpa salpa at areas within MAELI (black bars) or MAELO (gray bars) at each sampling time. 20 Figure 7. (A, B) Abundance and (C, D) total biomass (± SE) of the parrotfish, Sparisoma cretense , at areas within MAELI (black bars) or MAELO (gray bars) at each sampling time. 4.2 Taxonomic, trophic and functional diversity With regard to taxonomic diversity, no significant differences were detected between protected (MAELI) and non-protected (MAELO) areas at ‘Cabrón-Risco Verde’ (Table 5, Fig. 8-A). However, taxonomic diversity varied significantly between protected and non-protected area (p= 0.036, Table 4, Fig. 8-A) at ‘Canteras’. Differences in trophic diversity between protected (MAELI) and non-protected (MAELO) areas were not found (Table 4, Fig. 8-B). In turn, analysis of trophic 21 composition did not show significant differences (Appendix 2). In ‘CanterasConfital’, we found that functional diversity was significantly greater (Table 5, Fig 8-C) in the MAELI than the MAELO, but the functional composition did not vary (Appendix 2). Figure 8. Taxonomic diversity (A), trophic diversity (B), and functional diversity (C) on the two MAMPs studied. Cabrón Las Canteras Taxonomic Diversity 0.0 0.2 0.4 0.6 0.8 1.0 Cabrón Canteras Trophic diversity 0.0 0.2 0.4 0.6 0.8 Cabrón Las Canteras Functional diversity 0.0 0.2 0.4 0.6 0.8 A B C 22 Table 5. Results of 2-way ANOVAs testing for differences between areas and times, for the taxonomic, trophic and functional diversity at each location. Pvalues < 0.05 are considered significant. Cabrón Vs Risco Verde Canteras Vs Confital Taxonomic Diversity df MS F P df MS F P Locality 1 0.007 1.138 0.326 1 0.166 9.289 0.026 Time 6 0.009 3.570 0.007 6 0.025 1.761 0.141 LoxTi 6 0.006 2.501 0.041 6 0.018 1.242 0.303 Res. 42 0.003 42 0.014 Total 55 55 Trophic Diversity df MS F P df MS F P Locality 1 0.0000 0.002 0.969 1 0.016 1.251 0.305 Time 6 0.0011 0.510 0.797 6 0.011 1.533 0.187 LoxTi 6 0.0023 1.097 0.384 6 0.013 1.863 0.119 Res. 42 0.0021 42 0.007 Total 55 55 Functional Diversity df MS F P df MS F P Locality 1 0.015 1.132 0.330 1 0.160 31.57 0.002 Time 6 0.010 1.637 0.165 6 0.014 2.13 0.077 LoxTi 6 0.013 2.116 0.081 6 0.005 0.78 0.594 Res. 42 0.006 42 0.007 Total 55 55 4.3 Correlation between taxonomic, trophic and functional diversity Taxonomic, trophic and functional diversity were positively correlated (Fig. 9). 23 Functional Diversity 0.30 0.35 0.40 0.45 0.50 0.55 0.60 Taxonomic Diversity 0.65 0.70 0.75 0.80 0.85 0.90 0.95 1.00 1.05 y=0.442+0.295x;Rsqr = 0.489,p=0.005 0.25 0.30 0.35 0.40 0.45 0.50 0.55 0.60 0.65 0.3 0.4 0.5 0.6 0.7 0.8 0.9 y= -0.403 +2.207x;Rsqr = 0.666,p=0.001 Trophic Diversity 0.48 0.50 0.52 0.54 0.56 0.58 0.60 0.62 Taxonomic Diversity 0.65 0.70 0.75 0.80 0.85 0.90 0.95 1.00 1.05 y=-0.120+1.381x;Rsqr =0.585 ,p= 0.001 0.30 0.35 0.40 0.45 0.50 0.55 0.60 0.65 0.3 0.4 0.5 0.6 0.7 0.8 0.9 y=-0.120+1.381x;Rsqr =0.585 ,p= 0.001 Functional Diversity 0.30 0.35 0.40 0.45 0.50 0.55 0.60 Trophic Diversity 0.48 0.50 0.52 0.54 0.56 0.58 0.60 0.62 y=0.442+0.295x;Rsqr = 0.489,p=0.005 0.25 0.30 0.35 0.40 0.45 0.50 0.55 0.60 0.65 0.30 0.35 0.40 0.45 0.50 0.55 0.60 0.65 y=0.337+0.382x;Rsqr =0.243 ,p= 0.073 Figure 9. Relationship between taxonomic, trophic and functional diversity. Linear regression models tested the significance of this relation separately for ‘El Cabrón’ (A, C, D) and ‘Las Canteras’ (B, D and F). 24 5. Discussion 5.1 Overall results Our study aimed to test for differences in the abundances and biomasses of target species between areas that will be soon be implemented as marine protected areas and adjacent, un-protected, areas that may act as controls; this is an attempt to quantify the initial state ('state 0') to get baseline data to assess the so-called 'reserve effect' in the future. Such 'reserve effect' using a range of species have been demonstrated by a range of studies that showed that the abundances and total biomasses of certain species differed between protected and un-protected areas (Barret et al., 2006; Tuya et al., 2006; Brito et al. 1999, 1998, 2001, 2006, among others). To adequately assess the effectiveness of MPAs, before-after control–impact (BACIP) approaches are highly necessary (Edgar and Barret 1997; Edgar et al., 2004). In this sense, our study evaluated the 'reserve effect' immediately before the implementation of the start of the enforcement. Without a doubt, our data will help out to implement a proper BACIP protocols in the future. Furthermore, we analyzed community-level differences between 'protected' (at 'state 0') and adjacent areas. We used our data to calculate species (taxonomic), trophic and functional diversity, reducing our multivariate data into single diversity values. Some authors have demonstrated that diversity are often higher inside than outside protected areas (Barret et al., 2007; Claudet et al., 2006) and even trophic and functional diversity can response to protection more rapidly than species (taxonomic) diversity (Villamor et al., 2012). Recent syntheses and empirical studies have highlighted that functional traits predict the effects of global changes on ecosystem services better than species diversity per se (Cadotte et al., 2011) and many ecosystem processes and services depend more on functional diversity than species diversity (Nystrom., 2006). Importantly, our study has demonstrated that, at the study locations in Gran Canaria Island, there is a clear connection between the 3 ways biodiversity of nearshore fishes was quantified, i.e. at the taxonomic, trophic and functional levels. 5.2 Cabrón-Risco Verde Mycteroperca fusca , a top predator inhabiting shallow rocky reefs of the Macaronesia, was unique among the four studied species in the sense that we found larger abundances and total biomasses inside relative than outside the protected areas (at time 0, of course). In the Canary Islands, Tuya et al. (2006) found the greatest mean abundances and total biomasses of this species at El 25 Hierro Island ( ca . mean abundances of 1.5-2 ind 100 m2), particularly inside the 'Mar de Las Calmas' MPA. We found similar abundance values for this species, even at the state 0 of implementation. This result is indicative of the good status of this fish at this area, as this species is slow-growing, large-sized, with low population turnover rates (Zabala et al., 1997; La Mesa et al., 2002; Bodilies et al., 2003) and is heavily targeted by both professional and recreational fishermen in the Canarias (Bas et al., 1995; Falcon et al., 1996; Tuya et al., 2006). A similar outcome has been described in the Mediterranean Sea, where the effects of protection from fishing near the coast have lead to increments in the abundance and biomass of another Serranid, the dusky grouper, Epinephelus marginatus (Zabala et al., 1997; La Mesa et al., 2002). The sea-bream, Diplodus vulgaris , is a species targeted by both recreational and commercial fisheries in the Mediterranean (Coll et al., 2004; Lloret et al., 2004) and the Atlantic (Velga et al., 2010). This fish showed a higher biomass inside than outside the protected area. Small protected areas, such as MAEL can therefore offer an alternative for the sustainable development of this and similar species (Alós et al., 2011). 5.3 Canteras-Confital The target species: Sparisoma cretense , Diplodus sargus , Diplodus vulgaris and Sarpa salpa , showed larger abundances and total biomasses between the future protected area and the neighboring un-protected area. This may be attributed to eased control of fishing activities; by law, fishing is prohibited inside beaches, what is also facilitated by the large number of users that somehow make difficult extraction of resources within the beach. As a result, this area can be a great site to conserve and regenerate fish population. Sparisoma cretense , Diplodus vulgaris and Diplodus sargus are highly prized in both local recreational and commercial fisheries across the Macaronesian region, and especially throughout the Canarian Archipelago (Bortone et al., 1991; Bas et al., 1995). The larger abundances and total biomasses recorded for Salpa salpa might be the result of a larger fishing pressure outside the protected area. 5.4 Link between taxonomic, trophic and functional diversity Estimation of functional diversity is relevant to assess the health state of coastal resources, where there is increasing interest in clarifying the role of natural and human impacts. In this study, we only found higher functional diversity in ‘Las Canteras’ than in the adjacent un-protected area. 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Polunin & Roberts (1993) Agardy (1993) Roberts & Hawkins (1997) Reserve Saba Marine Park Ambergris Caye Reserva Biosfera Sian Ka'an (Quintana Roo) Parque Marino Isla Saba Gran Barrera de Arrecifes Anse Chastanet Country Netherlands Antilles Belize Méjico Antillas Neerlandesas Australia St. Lucía Size (ha) 20 20 320000 2.6 Age 4 4 7 2 Indicators: ecologic/fishing Total density of fishes + + n.a. n.a. n.a. + Total biomass of fishes + + n.a. n.a. n.a. + Density of predators fishes + + n.a. n.a. n.a. + Biomass of predators fishes + + n.a. n.a. n.a. + Species diversity n.a. n.a. n.a. n.a. n.a. n.a. Species size average + + n.a. n.a. n.a. + Predators size average + + n.a. n.a. n.a. + Density of commercial target fishes n.a. n.a. n.a. Indicators:socio-economic Tourist uses n.a. n.a. + + + n.a. Divers n.a. n.a. n.a. + + + Fishing local benefits + + n.a. n.a. n.a. n.a. Economic local benefits n.a. n.a. n.a. n.a. n.a. n.a. Enviroment local benefits n.a. n.a. n.a. n.a. n.a. n.a. 2 Russ et al. (2004) Parnell et al. (2005) Melita A. Samoilys et al. 2007 Harmelin -Viven et al. (2008) Apo Island La Jolla Handumon Pandano n Asina n Bilangbilangan Batasa n Banyuls Cabo de Palos Cabrer a Carry-leRouet Medes Tabarc a Filipinas Californiam, US Bohol, Filipinas Francia España Francia Españ a 22.5 216 50 20 66 10.5 21 650 1898 8680 85 418 1400 >18 12 3 7 8 8 >10 n.a. - + + + + + n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. + + + + + + n.a. 0 + + + + + n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. + n.a. n.a. n.a. n.a. n.a. n.a. + + + + n.a. 0 n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. 0 n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. 0 n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. 3 Claudet et al. (2008) La Restinga La Graciosa Cabo de Palos Tabarca San Antonio Columbretes Medes Islands CerbereBanyuls Cap Couronne CarryleRouet Bouches de Bonifacio Siis Mal di Ventre España Francia Italia 180 1225 270 120 110 1883 93 65 210 85 1200 529 14 13 13 22 15 18 25 34 14 26 17 15 n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. + + + + + + + + + + + + n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. n.a. 4 Appendix 2. Results of two way ANOVAs testing for differences between localities, times, for the species, trophic and functional composition each marine reserve. *Significant difference at P <0.05. Cabrón Vs Risco Verde Canteras Vs Confital Taxonomic composition df MS F P df MS F P Locality 1 0.976 4.309 0.087 1 1.116 3.981 0.098 Time 6 0.287 2.543 0.032 6 0.275 1.488 0.199 LoxTi 6 0.227 2.006 0.091 6 0.280 1.518 0.195 Res. 42 0.113 42 0.185 Total 55 55 Trophic composition df MS F P df MS F P Locality 1 0.004 0.226 0.624 1 0.187 1.295 0.305 Time 6 0.032 1.317 0.281 6 0.045 0.514 0.799 LoxTi 6 0.016 0.658 0.677 6 0.145 1.648 0.168 Res. 42 0.024 42 0.089 Total 55 55 Functional composition df MS F P df MS F P Locality 1 0.009 0.048 0.835 1 0.378 1.671 0.246 Time 6 0.147 1.731 0.139 6 0.057 0.573 0.750 LoxTi 6 0.188 2.219 0.062 6 0.226 2.284 0.051 Res. 42 0.084 42 0.099 Total 55 55 5