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Latin american journal of aquatic research

versión On-line ISSN 0718-560X

Lat. Am. J. Aquat. Res. vol.41 no.3 Valparaíso jul. 2013 

Research Article


Zoning of the Mejillones Peninsula marine protected coastal area of multiple uses, northern Chile

Zonificación del área marina costera protegida de múltiples usos de la península de Mejillones, norte de Chile


Raúl Ulloa1, Adolfo Vargas2,3 Cristian Hudson2 & Marcelo M. Rivadeneira4

1 Centro Regional de Investigación Pesquera, Instituto Nacional de Pesca Dirección General de Investigación Pesquera Pacifico Norte Miguel Alemán sur N°605, C.P. 85400, Guaymas, Sonora, México
2 Programa Magíster en Ciencias Aplicadas mención Biología Pesquera Departamento Ciencias del Mar, Universidad Arturo Prat, P.O. Box 121, Iquique, Chile
Centro de Investigación y Desarrollo de Profesionales Marinos Pacífico Ltda. Vivar 1218, Iquique, Chile
4 Centro de Estudios Avanzados en Zonas Áridas (CEAZA) & Facultad de Ciencias del Mar Universidad Católica del Norte, Av. Bernardo Ossandón 877, CP. 1781681, Coquimbo, Chile
Corresponding author: Marcelo M. Rivadeneira (

ABSTRACT. Marine protected areas of multiple uses (MPA-MU), are an important management tool to protect biodiversity and regulate the use of coastal marine resources. However, robust conservation plans require an explicit consideration of not only biological but also social components, balancing the protection of biodiversity with a sustainable exploitation of marine resources. Here we applied the decision-making algorithm MARXAN to provide a zoning analysis at the Mejillones Peninsula MPA-MU in northern Chile, one of largest MPA's of the Humboldt Current Marine Ecosystem. We set conservation goals for coarse and fine-filter conservation targets that were crossed out against different threats and pressure factors from human activities across the area. We identified a portfolio of sites for conservation, within the Mejillones Peninsula MPA-MU, representing different ecological systems with different levels of human impacts and vulnerability. These results may serve as a foundational guideline for the future administration of the MPA-MU.

Keywords: MPA-MU, MARXAN, conservation plans, marine coastal ecosystems, northern Chile.

RESUMEN. Las áreas marinas protegidas de múltiples usos (AMCP-MU), son una importante herramienta de manejo para la protección de la biodiversidad y la regulación del uso de recursos marinos costeros. Sin embargo, planes robustos para la conservación, deben considerar explícitamente no sólo componentes biológicos sino además sociales, equilibrando la protección de la biodiversidad con la explotación sustentable de los recursos marinos. En este trabajo se aplica el algoritmo de toma de decisiones MARXAN, para entregar un análisis de zonificación del AMCP-MU, de la península de Mejillones, en el norte de Chile, una de las AMP más grandes del Ecosistema Marino del sistema de la Corriente de Humboldt. Se fijaron metas de conservación para objetivos de conservación de filtro grueso y fino, que fueron cruzados contra diferentes amenazas y factores de presión, generados por las actividades humanas registradas en la zona. Se identificó un portafolio de sitios prioritarios de conservación, dentro de la AMCP-MU, representativos de diferentes sistemas ecológicos, sometidos a diferentes niveles de impacto humano y vulnerabilidad. Estos resultados pueden proveer las guías fundacionales para el futuro plan de administración de la AMCP-MU de la península de Mejillones.

Palabras clave: AMCP-MU, MARXAN, planes de conservación, ecosistemas marinos costeros, norte de Chile.



Marine ecosystems are fundamental to the sustainable development of coastal countries, providing a large variety of environmental resources. Currently, these ecosystems face a growing menace of overexploitation of fisheries, pollution, habitat destruction, among other human-driven impacts (Jackson, 2001; Jackson et al., 2001; Jackson & Sala, 2001; Rivadeneira et al., 2010). In this scenario, Marine Protected Areas (MPA) arises as one of most practical strategies for the conservation of the marine biodiversity (Allison et al., 1998). MPA not only seek to protect biodiversity but also allows sustainable exploitation of the ecosystem, providing social, economic, and research benefits (Salm et al., 2000).

In contrast to terrestrial ecosystems, MPA networks are worldwide poorly developed and less than 1% of the global ocean is currently protected (Chape et al., 2005). The situation is no better in Latin America and Caribbean regions, where despite there are over 700 MPA established, only 1.5% of coastal and shelf waters areas are protected (Guarderas et al., 2008) . Moreover, vast areas of the Humboldt Current Large Marine Ecosystem, included Chile, are largely unprotected (Guarderas et al., 2008).

The current environmental policy of Chile recognizes eight categories of areas for conservation of biodiversity (Sierralta et al., 2011). Among them, marine protected costal area of multiple uses (MPA-MU) are mixed-used MPA, aimed to conserve biodiversity, protect marine species, reduce conflicts of use, allowing commercial and recreational activities, including fishing and tourism (Sierralta et al., 2011). Currently, there are four MPA-MU established for the Chilean coast (Tognelli et al., 2009) . Despite the existing MPA network that seems to protect a large fraction of the marine biodiversity, many species with small geographic ranges are not covered. In addition, the success of a MPA depends on the existence of zoning plans, able to resolve conflicts between local users and the environment, balancing the protection of marine biodiversity with a sustainable exploitation of marine resources and uses of the coastal seascape (Villa et al., 2002; Douvere, 2008). However, to date, such zoning analysis has been implemented only at Isla Grande de Atacama MPA-MU (Gaymer et al., 2008; Rojas-Nazar et al., 2012).

The Mejillones Peninsula (Fig. 1), has been previously identified as a priority site for marine conservation (Tognelli et al., 2005, 2008, 2009; Miethke et al., 2007), and is one of the recently proposed MPA-MU for the Chilean coast (Hudson et al., 2008). The Mejillones Peninsula MPA-MU, is placed in the coastal area of the hyper-arid Atacama Desert, and it is the third largest MPA of the southeastern Pacific, covering more than 400 km2 along a stretch of ca. 100 km of coast (Wood, 2007). The marine ecosystem is characterized by nearly continuous upwelling events, which introduce cold and nutrient-rich waters to coastal areas (Sobarzo & Figueroa, 2001; Thiel et al., 2007). The high marine productivity and relative isolation from human activities, begets a great diversity and abundance of marine species. The area is a vertebrate hotspot of endemism (Tognelli et al., 2005, 2008), and it is well recognized for the presence of large colonies of sea lions, transit of dolphins, great abundance and diversity of seabirds, and the seasonal arrival of sea turtles and whales (Aguayo & Maturana, 1973; Guerra et al., 1987; Rendell et al., 2004; Vilina et al., 2006; Guerra-Correa et al., 2008).


Figure 1. Spatial distribution of the coarse filter for conservation in the study area.


In spite of the relative isolation of the Mejillones Peninsula, different human activities can be identified, all of which can potentially interfere on marine biodiversity and conservation plans, including both legal and illegal artisanal fisheries, aquaculture centers, and non-regulated tourism from the nearby cities (Hudson et al., 2008). Therefore, conservation plans for the MPA-MU must take into account the integration of human stakeholders in order to find optimal solutions that balance biodiversity protection with socio-cultural and economic realities. In this work, we carried out a spatial zoning analysis to identify specific areas for conservation across the Mejillones Peninsula MPA-MU, using optimization algorithms to reduce human threats and maximize the conservation goals.


Biological and anthropogenic database

The study area included 435 km2, 98.5 km of coastline, encompassing all marine coastal shelf habitats (<200 m depth), from the intertidal to 2.7 km offshore within the coastal shelf. We compiled an exhaustive georeferenced database, documenting different facets of the marine biodiversity and human activities (see below) in the study area. The information data was obtained from four main sources: a) Published scientific literature, including papers, thesis, technical reports from Chile governmental services (e.g., Ministerio del Medio Ambiente de Chile, Instituto de Fomento Pesquero, Fondo de Investigación Pesquera, Comisión Nacional Forestal, and Centro Nacional de Datos Hidrográficos y Oceano-gráficos), b) Unpublished information, available from Chilean public services (e.g., Servicio Nacional de Marine protected area zoning Pesca, Subsecretaría de Pesca, Servicio Hidrográfico y Oceanográfico de la Armada), c) the traditional knowledge of local fishermen regarding fisheries, reproduction and recruitment zone, and sea bottom types, and d) new information from field samplings in intertidal and subtidal benthic habitats, carried out during November 2008. Field surveys allowed a community-level characterization of the biodiversity, and also cross-validate the information provided by the traditional knowledge of fishermen.

The georeferenced information was assigned to 10,873 hexagonal planning units (PU) of 4 ha each (0.04 % of total area of MPA) (Fig. 1). For a geographic homologation we used the zone 19 UTM andDatum WGS-84.

Conservation targets and goals

Conservation targets were defined using two complementary approaches, fine and coarse filters, following previous studies (Day & Roff, 2000; Beck & Odaya, 2001; Roberts et al, 2003; Vierros, 2004). Fine filters were selected according two criteria: i) species included in any threatened category according to UICN, CITES, CONAF or MMA, ii) particular traits associated to species; justifying further conservation efforts (e.g., endemic, migratory, keystone, umbrella, and flagship species, Barua, 2011) (Table 1). Because fine filters are typically available only for a very restricted subset of marine biodiversity (i.e., higher vertebrates), we also used coarse filters that may provide a broader picture of ecological processes occurring in the area, including ecosystem engineers, areas of reproduction/resting, areas with existing conservation/management plans, and areas of high primary productivity (Table 2). Ecosystem engineers (Jones et al., 1994), such as kelps, mussels, scallops, and tunicates were included, because they provide a habitat to a vast diversity of other species. Reproductive/resting areas of migratory or flag species imply the existence of a high population abundance were bottom-up/top-down processes may be enhanced. Similarly, the restricted or reduced human perturbations expected in the existing management and exploitation area for benthic resources (MEARB's) and the La Rinconada marine reserve may imply an indirect protection of marine biodiversity (Fernández & Castilla, 2005; Navarrete et al., 2010).


Table 1. Criteria used to define fine-filters and conservation goals. Conservation categories according to IUCN and MMA (DD: data deficient, LC: least concern, NT: near threat, VU: vulnerable, EN: endangered).


Table 2. Coarse-filters for conservation, indicating the geographic extend (PU’s) of each target and the conservation goals set.


Table 3. Different anthropogenic threats identified, and number of planning units (PU’s) and costs assigned.


In order to ensure the viability and persistence of each conservation target we assigned a conservation goal to each one (Groves et al., 2000; CONABIO, 2009; Ulloa et al., 2006). It should be remarked that these goals were consensual with social actors of the Mejillones Peninsula, including artisanal fishermen, academic researchers and public authorities. We set conservation goals for each target, with values varying between 20% (less relevant) and 100% (irreplaceable).

Pressure factors and threats

Threats to biodiversity were selected based on known impacts of anthropogenic activities on ecological systems. The aim was to use high quality data to characterize pressure factors in order to select sites that could still be valuable to invest in conservation or restoration. The data used by the prioritization algorithm for representing threats is often referred as costs, following the logic that areas suffering from negative impacts are more difficult to protect and require higher conservation investment. Cost information is used to distribute conservation priorities to sites amenable to long-term persistence of conservation features (Chan et al., 2006).

A threat was defined as any human activity affecting negatively marine biodiversity. Threats were identified based on compiled information for five categories: fishing, tourism, human settlements, shipping zones, and aquaculture (Table 4). The spatial extension of each threat was digitalized into a geographic information system. Coasts (arbitrary units) were assigned to each threat, ranging from 500 (lowest) to 10.000 (highest).

Those values were estimated considering the relationship between the spatial extension of the threat and the number of users or elements associated (e.g., number of fishermen, local population, frequency of use of space, volume of production). Thus, despite fishing activities are carried out along all the area, we assigned low to mid costs because of the relatively low levels of fishermen involved in the activity. In contrast, we assigned a large cost to aquaculture, due to the large volume of production involved.

Identifying priority areas for conservation

The identification of priority sites was carried out based on biological variables, and on pressure factors, using the simulated annealing algorithm in the MARXAN software (Ball & Possingham, 2000), a robust and widely used analytic approach (McDonnell et al., 2002; Stewart & Possingham 2002). The program was run with 1.000.000 iterations and 10.000 runs, using the adaptive annealing schedule with normal iterative improvement at the end of each run (Cook & Auster, 2005; Chan et al., 2006). The type of data and criteria used in the process of assigning conservation goals to biodiversity elements and cost values to represent pressure factors followed the methodology used by different authors (Ball & Possingham, 2000; Groves et al., 2000; Ulloa et al., 2006). The selection frequency of a planning unit provides a fundamental measure of conservation value for the unit (Stewart et al., 2007) indicating its relative importance to meet given targets. Planning units included in all the MARXAN solutions are considered irreplaceable, and thus were designated as high priority.


A total of 76 conservation targets, 58 fine filter and 18 coarse filters were identified (Tables 2 and 3). For coarse filters, larger goals (100%) were assigned for reproductive/resting areas of flag or migratory turtles, Marine protected area zoning Marine protected area zoning seabirds and sea mammals (Table 2). However, we also assigned high goals (>60%) to seaweed and invertebrate beds, MEARB's and the sole genetic reserve in the region. For fine filters, larger goals were assigned to endangered, endemic and flag species. These include the turtles Chelonia mydas and Lepidochelys olivacea; the dolphins Delphinus delphis and Tursiops truncates, and also the tunicate Pyura praeputialis. In general, the conservation goals were distributed across the entire study region, but particularly in coastal shallower areas (Fig. 1).

We assigned a cost value to each one of the nine types of human threats identified along the Mejillones Peninsula region (Table 3), with larger values indicating a higher potential damage to biodiversity. The largest costs were located in nearshore areas, where intertidal and subtidal benthic areas are subject to human activities, mainly aquaculture, artisanal fisheries, and non-regulated tourism (Table 3, Fig. 2). The spatial integration of the costs of the threats showed that 9% of PU yielded very high costs (e.g., 5000).


Figure 2. Spatial distribution of the threats for conservation along the study area.


The zoning analysis yielded a portfolio of 211 PU more frequently selected (5.001-10.000), encompassing 2.4% (844 ha) of the total area evaluated. These areas are spatially clustered in several shallow-water hotspots along the study region (Fig. 3). The two largest hotspots are located at the northern (Caleta Lobería-Punta Angamos) and southern ends (Isla Santa María-La Rinconada) of the Mejillones Peninsula. The analysis also identified several smaller and discontinuous spots highly selected across the MPA-MU.


Figure 3. Portfolio of sites identified as conservation priority according MARXAN.



The Mejillones Peninsula has been recognized as a priority the area for the conservation of marine biodiversity in the Chile (Tognelli et al., 2005, 2008, 2009; Miethke et al., 2007). Therefore, our zonation analysis represents an important step for the establishment of administrative policies of the Mejillones Peninsula MPA-MU.

The definition of conservation goals, in coastal ecosystems, is one the most debated questions in coastal planning, due its complexity and the limited understanding of the ecological processes and its scale-dependency (Roberts et al., 2003). Quantitative goals has been introduced only recently, but mainly based on the expert criteria, and the abundance of the conservation targets (Beck & Odaya, 2001; DeBlieu et al., 2005; Ulloa et al., 2006). In this vein, our conservation proposed goals, accordingly to four criteria, allowed us to reduce the uncertainty introduced by the approaches based on the expert criteria (Ballantine, 1997), or the abundance of the conservation target. This is particularly relevant because the conservation status is available for a small fraction of species (i.e., mainly vertebrates), leaving out the vast majority of non-vertebrate biodiversity. Indeed, the total number of species used to set fine filters was strongly biased towards seabirds and mammals, representing ca. 11% of total marine richness reported for the study area (Hudson et al., 2008). We mitigated this problem by using coarse-filters, which reflect population, community or ecosystem-level properties. Given the high diversity of invertebrates and vertebrates associated to these species, we ensured the secondary protection of many species for which we lack assessments of conservation status

The two areas identified as target for conservation, at the northern and southern ends of the MPA, have been previously recognized as important areas for different ecological processes relevant for conservation plans. For instance, these areas are traditional spots for cetacean watching, such as the common dolphin Delphinus delphis (Guerra et al., 1987), and resting/reproductive areas of seabirds (Vilina et al., 2006). On the other hand, these areas show high abundances of ecosystem engineers (Jones et al., 1994), including kelps, tunicates, scallop and mussel beds, which serve as habitat for a large number of invertebrate and vertebrates (Cerda & Castilla, 2001; Thiel & Ullrich, 2002; Vega et al., 2005; Vasquez et al., 2006). For instance, Cerda & Castilla (2001) reported more than 96 putative species living within the beds of the tunicate P. praeputialis, and invasive species but with a very restricted geographic range (i.e., endemic) along the southeastern Pacific coast (Castilla et al., 2004). Hence, setting high conservation goals for these species may ensure the indirect protection of a vast diversity of marine species.

Because of the spatial clustering of the portfolio of priority PU's, connections between areas may be not warranted. Roberts & Hawkins (2000) and Sala et al. (2002) have suggested that isolated MPA are of limited use for conservation since they protect a limited fraction of the marine biodiversity, and therefore the distance among MPA's should enable the dispersal among populations. The existence of a permanent off-shore upwelling plume at the region would potentially reduce the larval dispersal alongshore, although larval retention may be enhanced by the existence of eddies (Thiel et al., 2007). However, recent phylogeographic studies have revealed no genetic break within or near the MPA-MU zone in species with very contrasting dispersal capabilities (Cardenas et al., 2009a, 2009b; Tellier et al., 2009), which suggest that populations should be genetically connected across the region. In addition, different oceanographic processes (i.e., Ekman trans port, wind velocity, slick occurrence), and the recruitment of intertidal barnacles are structured at spatial scales of ca. 50 km (Lagos et al., 2008). Altogether, these pieces of evidence suggest a strong connectedness among high priority PU's, ensuring the conservation potential of the network.

How viable would be our zoning scheme on a long-term basis? This is a question often overlooked by conservation plans (Pressey et al., 2007; Game et al., 2008) partly because of the state-invariant nature of the zoning algorithms such as MARXAN (i.e., conservation targets and threats are temporally invariant). For instance, coarse-scale and fine-scale filters may be deeply altered in occurrence and location during the El Niño events (e.g., Tomicic, 1985) via changes in the structure of local communities (Vásquez et al., 2006; Sielfeld et al., 2010), or inducing shifts in the geographic ranges of the species (Riascos et al., 2008; Carstensen et al., 2010). Moreover, these changes in distribution may not be necessarily transient, but represent the long-term shifts in the geographic distribution of species (Rivadeneira & Fernández, 2005), and hence potentially altering the diversity and composition of species at the MP. While the MP area is far away from the biogeographic breaking zones, described for different marine taxa across the Pacific coast of South America (Lancellotti & Vásquez, 1999, 2000; Camus, 2001), which may reduce the potential impacts of the shifts of the geographic ranges of species, a real evaluation of this threat remains untested. Future studies may improve our model by considering uncertainties introduced by increasing threats (e.g., increasing fishing effort, and aquaculture center and touristic activities).

The Mejillones Peninsula, encompassing an extension of 435 km2, is the third largest marine protected area of the Humboldt Current Large Marine Ecosystem (Wood, 2007), harboring more than 500 species, from diatoms to whales (Hudson et al., 2008). Our analysis could set the foundational basis for robust conservation plans of the Mejillones Peninsula Marine Protected Area. A similar zoning approach may be undertaken, at a much larger spatial scales, in order to design a regional/nationwide network of MPA's able to balance the growing needs for robust plans of conservation of marine biodiversity, and the sustainable use of coastal spaces and the management of marine resources.


We deeply appreciate the logistic help, continuous encouragement and valuable comments provided by our old friends Pedro Pizarro, John W. Bobbitt and Dirk Shafer. Two anonymous reviewers provided extremely helpful and constructive suggestions and comments that improved the final version of this manuscript. This project was funded by the Ministerio del Medio Ambiente de Chile (i.e., formerly CONAMA). The research of M.M.R. was partially funded by projects FONDECYT 11070147 and 1110582.



Aguayo, A. & R. Maturana. 1973. Presencia del lobo marino común Otaria flavescens en el litoral chileno. Biol. Pesq., 6: 49-75.

Allison, G.W., J. Lubchenco & M.H. Carr. 1998. Marine reserves are necessary but not sufficient for marine conservation. Ecol. Appl., 8: 79-92.

Ball, I.R. & H.P. Possingham. 2000. MARXAN (V1.8.2). Marine reserve design using spatially explicit annealing. The Great Barrier Reef Marine Park Authority, Townsville, 69 pp.

Ballantine, W.J. 1997. Design principles for systems of 'no-take' marine reserves. In: T. Pitcher (ed.). The design and monitoring of marine reserves. Fisheries Centre, University of British Columbia, British Columbia, Vancouver, pp. 1-9.

Barua, M. 2011. Mobilizing metaphors: the popular use of keystone, flagship and umbrella species concepts. Biodivers. Conserv., 20: 1427-1440.

Beck, M.W. & M. Odaya. 2001. Ecoregional planning in marine environments: identifying priority sites for conservation in the northern gulf of Mexico. Aquat. Conserv. Mar. Freshw. Ecosyst., 11: 235-242.

Camus, P.A. 2001. Marine biogeography of continental Chile. Rev. Chil. Hist. Nat., 74: 587-617.

Cardenas, L., J.C. Castilla & F. Viard. 2009a. A phylogeographical analysis across three biogeogra-phical provinces of the south-eastern Pacific: the case of the marine gastropod Concholepas concholepas. J. Biogeogr., 36: 969-981.

Cardenas, L., A.X. Silva, A. Magoulas, J. Cabezas, E. Poulin & F.P. Ojeda. 2009b. Genetic population structure in the Chilean jack mackerel, Trachurus murphyi (Nichols) across the south-eastern Pacific Ocean. Fish. Res., 100: 109-115.

Carstensen, D., J.M. Riascos, O. Heilmayer, W.E. Arntz & J. Laudien. 2010. Recurrent, thermally-induced shifts in species distribution range in the Humboldt Current upwelling system. Mar. Environ. Res., 70:293-299.

Castilla, J.C., R. Guiñez, A.U. Caro & V. Ortiz. 2004. Invasion of a rocky intertidal shore by the tunicate Pyura praeputialis in the bay of Antofagasta, Chile. Proc. Natl. Acad. Sci. USA, 101: 8517-8524.

Cerda, M. & J.C. Castilla. 2001. Diversity and biomass of macroinvertebrates in intertidal matrices of the tunicate Pyura praeputialis (Heller, 1878) in the bay of Antofagasta, Chile. Rev. Chil. Hist. Nat., 74: 841-853.

Comisión Nacional para el Conocimiento y Uso de la Biodiversidad (CONABIO). 2009. Capital natural de México: estado de conservación y tendencias de cambio. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, México D.F., 2: 821 pp.

Cook, R.R. & P.J. Auster. 2005. Use of simulated annealing for identifying essential fish habitat in a multispecies context. Conserv. Biol., 19: 876-886.

Chan, K.M.A., M.R. Shaw, D.R. Cameron, E.C. Underwood & G.C. Daily. 2006. Conservation planning for ecosystem services. PLoS Biol., 4: 2138-2152.

Chape, S., J. Harrison, M. Spalding & I. Lysenko. 2005. Measuring the extent and effectiveness of protected areas as an indicator for meeting global biodiversity targets. Philos. Trans. Roy. Soc., Ser. B, 360: 443-455.

Day, J.C. & J.C. Roff. 2000. Planning for representative marine protected areas: a framework for Canada s Oceans. Report prepared for World Wildlife Fund Canada, Toronto, 77 pp.

DeBlieu, J., M. Beck, D. Dorfman & P. Ertel. 2005. Conservation in the Carolinian ecoregion: an ecoregional assessment. The Nature Conservancy, Arlington, 60 pp.

Douvere, F. 2008. The importance of marine spatial planning in advancing ecosystem-based sea use management. Mar. Policy, 32: 762-771.

Fernández, M. & J.C. Castilla. 2005. Marine conservation in Chile: historical perspective, lessons, and challenges. Conserv. Biol., 19: 1752-1762.

Game, E.T., M.E. Watts, S. Wooldridge & H.P. Possingham. 2008. Planning for persistence in marine reserves: A question of catastrophic importance. Ecol. Appl., 18: 670-680.

Gaymer, C.F., U. Rojas, F.A. Squeo, G. Luna-Jorquera, A. Cortés, G. Arancio, C. Dumont, M. Cortés, D. Hiriart & D. López. 2008. AMCP-MU Isla Grande de Atacama: flora y fauna marina y terrestre. In: F.A. Squeo, G. Arancio & J.R. Gutiérrez (eds.). Libro rojo de la flora nativa y los sitios prioritarios para su conservación: región de Atacama. Ediciones Universidad de La Serena, La Serena, pp. 223-249.

Groves, C., L. Valutis, D. Vosick, B. Neely, K. Wheathon, J. Touval & B. Runnels. 2000. Diseño de una geografía de la esperanza: manual para la planificación de la conservación ecorregional. The Nature Conservancy, Arlington, 186 pp.

Guarderas, A.P., S.D. Hacker & J. Lubchenco. 2008. Current status of marine protected areas in Latin America and the Caribbean. Conserv. Biol., 22: 1630-1640.

Guerra-Correa, C.G., C.M. Guerra-Castro, P.D. Bolados, A. Silva & P. Garfias. 2008. Sea turtle congregations in discrete temperate shoreline areas in cold northern Chilean coastal waters. NOAA Technical Memorandum NMFS SEFSC Sea turtle congregations in discrete temperate shoreline areas in cold northern Chilean coastal waters, pp. 211-212.

Guerra, C., C.K. Van Waerebeek, G. Portflitt & G. Luna. 1987. Presencia de cetáceos frente a la segunda región de Chile. Estud. Oceanol., 6: 87-96.

Hudson, C., R. Ulloa, A. Vargas, M.M. Rivadeneira, G. Guzmán & D. Schiappacasse. 2008. Estudio de línea base de la biodiversidad marina del sitio prioritario península de Mejillones. Comisión Nacional del Medio Ambiente (CONAMA), Antofagasta, 115 pp.

Jackson, J.B.C. 2001. What was natural in the coastal oceans? Proc. Nat. Acad. Sci. USA., 98: 5411-5418.

Jackson, J.B.C., M.X. Kirby, W.H. Berger, K.A. Bjorndal, L.W. Botsford, B.J. Bourque, R.H. Bradbury, R. Cooke, J. Erlandson, J.A. Estes, T.P. Hughes, S. Kidwell, C.B. Lange, H.S. Lenihan, J.M. Pandolfi, C.H. Peterson, R.S. Steneck, M.J. Tegner & R.R. Warner. 2001. Historical overfishing and the recent collapse of coastal ecosystems. Science, 293: 629-638.

Jackson, J.B.C. & E. Sala. 2001. Unnatural oceans. Sci. Mar., 65: 273-281.

Jones, C.G., J.H. Lawton & M. Shachak. 1994. Organisms as ecosystem engineers. Oikos, 69: 373-386.

Lagos, N.A., J.C. Castilla & B.R. Broitman. 2008. Spatial environmental correlates of intertidal recruitment: a test using barnacles in northern Chile. Ecol. Monogr., 78: 245-261.

Lancellotti, D.A. & J.A. Vasquez. 1999. Biogeographical patterns of benthic macroinvertebrates in the southeastern Pacific littoral. J. Biogeogr., 26: 1001-1006.

Lancellotti, D.A. & J.A. Vasquez. 2000. Zoogeography of benthic macroinvertebrates of the Chilean coast: contribution for marine conservation. Rev. Chil. Hist. Nat., 73: 99-129.

Pressey, R.L., M. Cabeza, M.E. Watts, R.M. Cowling & K.A. Wilson. 2007. Conservation planning in a changing world. Trends Ecol. Evol., 22: 583-592.

McDonnell, M.D., H.P. Possingham, I.R. Ball & E.A. Cousins. 2002. Mathematical methods for spatially cohesive reserve design. Environ. Monit. Assess, 7: 107-114.

Miethke, S., S. Reichle, E. Armijo, Z. Ferdaña, L. Sotomayor, A. Chatwin, B. Ramírez & R. DeAndrade. 2007. Coastal and marine conservation priorities in Chile. In: A. Chadwin (ed.). Priorities for coastal and marine conservation in South America. The Nature Conservancy, Arlington, Virginia, pp. 25-29.

Navarrete, S., S. Gelcich & J.C. Castilla. 2010. Monitoreo de largo plazo en el ecosistema marino costero de Las Cruces, Chile: definiendo líneas base para construir alfabetización ecológica en un mundo que cambia. Rev. Chil. Hist. Nat., 83: 143-157.

Rendell, L., H. Whitehead & R. Escribano. 2004. Sperm whale habitat use and foraging success off northern Chile: evidence of ecological links between coastal and pelagic systems. Mar. Ecol. Prog. Ser., 275: 289-295.

Riascos, J.M., T. Ashton & A. Pacheco. 2008. First record of Cymatium keenae Beu, 1970 (Mollusca: Ranellidae) from Antofagasta bay, northern Chile, in connection with El Niño events. Helgol. Mar. Res., 62: 107-110.

Rivadeneira, M.M. & M. Fernández. 2005. Shifts in southern endpoints of distribution in rocky intertidal species along the southeastern Pacific coast. J. Biogeogr., 32: 203-209.

Rivadeneira, M.M., C.M. Santoro & P.A. Marquet. 2010. Reconstructing the history of human impacts on coastal biodiversity in Chile: constraints and opportunities. Aquat. Conserv. Mar. Freshw. Ecosyst., 20: 74-82.

Roberts, C.M., G. Branch, R.H. Bustamante, J.C. Castilla, J. Dugan, B.S. Halpern, K.D. Lafferty, H. Leslie, J. Lubchenco, D. McArdle, M. Ruckelshaus & R.R. Warner. 2003. Application of ecological criteria in selecting marine reserves and developing reserve networks. Ecol. Appl., 13: 215-228.

Roberts, C.M. & J.P. Hawkins. 2000. Fully-protected marine reserves: a guide. World Wildlife Fund, Washington. D.C. and University of York, New York, 137 pp.

Rojas-Nazar, U., C.F. Gaymer, F.A. Squeo, R. Garay-Flühmann & D. López. 2012. Combining information from benthic community analysis and social studies to establish no-take zones within a multiple uses marine protected area. Aquat. Conserv. Mar. Freshw. Ecosys., 22: 74-86.

Sala, E., O. Aburto-Oropeza, G. Paredes, I. Parra, J.C. Barrera & P.K. Dayton. 2002. A general model for designing networks of marine reserves. Science, 298: 1991-1993.

Salm, R.V., J.R. Clark & E. Siirila. 2000. Marine and coastal protected areas: a guide for planners and managers. International Union for Conservation of Nature and Natural Resources, Washington D.C., 21: 371 pp.

Sielfeld, W., J. Laudien, M. Vargas & M. Villegas. 2010. El Niño induced changes of the coastal fish fauna off northern Chile and implications for ichthyogeography. Rev. Biol. Mar. Oceanogr., 45: 705-722.

Sierralta, L., R. Serrano. J. Rovira & C. Cortés. 2011. Las áreas protegidas de Chile, Ministerio del Medio Ambiente, Santiago, 35 pp.

Sobarzo, M. & D. Figueroa. 2001. The physical structure of a cold filament in a Chilean upwelling zone (Peninsula de Mejillones, Chile, 23°S). Deep-Sea Res. Part I, 48: 2699-2726.

Stewart, R.R. & H.P. Possinghan. 2002. A framework for systematic marine reserve design south Australia: a case study. In: J.P. Beumer, A. Grant & D.C. Smith (eds.). Proceedings of the world congress on aquatic protected areas- what works best and how do we know? Australian Society for Fish Biology, Brisbane, pp. 132-143.

Stewart, R.R., I.R. Ball & H.P. Possingham. 2007. The effect of incremental reserve design and changing reservation goals on the long-term efficiency of reserve systems. Conserv. Biol., 21: 346-354.

Tellier, F., A.P. Meynard, J.A. Correa, S. Faugeron & M. Valero. 2009. Phylogeographic analyses of the 30°S south-east Pacific biogeographic transition zone establish the occurrence of a sharp genetic discontinuity in the kelp Lessonia nigrescens: Vicariance or parapatry? Mol. Phylogenet. Evol., 53: 679-693.

Thiel, M., E.C. Macaya, E. Acuña, W.E. Arntz, H. Bastias, K. Brokordt, P.A. Camus, J.C. Castilla, L.R. Castro, M. Cortes, C.P. Dumont, R. Escribano, M. Fernandez, J.A. Gajardo, C.F. Gaymer, I. Gomez, A.E. Gonzalez, H.E. Gonzalez, P.A. Haye, J.E. Illanes, J.L. Iriarte, D.A. Lancellotti, G. Luna-Jorquera, C. Luxoroi, P.H. Manriquez, V. Marin, P. Muñoz, S.A. Navarrete, E. Perez, E. Poulin, J. Sellanes, H.H. Sepulveda, W. Stotz, F. Tala, A. Thomas, C.A. Vargas, J.A. Vasquez & J.M.A. Vega. 2007. The Humboldt Current System of northern and central Chile: oceanographic processes, ecological interactions and socioeconomic feedback. Oceanogr. Mar. Biol. Annu. Rev., 45: 195-344.

Thiel, M. & N. Ullrich. 2002. Hard rock versus soft bottom: the fauna associated with intertidal mussel beds on hard bottoms along the coast of Chile, and considerations on the functional role of mussel beds. Helgol. Mar. Res., 56: 21-30.

Tognelli, M.F., M. Fernandez & P.A. Marquet. 2009. Assessing the performance of the existing and proposed network of marine protected areas to conserve marine biodiversity in Chile. Biol. Conserv., 142: 3147-3153.

Tognelli, M.F., P.I.R. de Arellano & P.A. Marquet. 2008. How well do the existing and proposed reserve networks represent vertebrate species in Chile? Divers. Distrib., 14: 148-158.

Tognelli, M.F., C. Silva-Garcia, F.A. Labra & P.A. Marquet. 2005. Priority areas for the conservation of coastal marine vertebrates in Chile. Biol. Conserv., 126: 420-428.

Tomicic, J.J. 1985. Efectos del fenómeno El Niño 1982-83 en las comunidades litorales de la península de Mejillones. Invest. Pesq., Chile, 32: 209-213.

Ulloa, R., J. Torre, L. Bourillón & N. Alcantar. 2006. Planeación ecorregional para la conservación marina: golfo de California y costa occidental de Baja California Sur. Informe Final. The Nature Conservancy Guaymas, 153 pp.

Vásquez, J.A., J.M.A. Vega & A.H. Buschmann. 2006. Long term variability in the structure of kelp communities in northern Chile and the 1997-98 ENSO. J. Appl. Phycol., 18: 505-519.

Vega, J.M.A., J.A. Vásquez & A.H. Buschmann. 2005. Population biology of the subtidal kelps Macrocystis integrifolia and Lessonia trabeculata (Laminariales, Phaeophyceae) in an upwelling ecosystem of Northern Chile: interannual variability and El Niño 1997-1998. Rev. Chil. Hist. Nat., 78: 33-50.

Vierros, M. 2004. Biodiversity issues for consideration in the planning, establishment and management of protected area sites and networks: some considerations on marine and coastal protected areas network design. Secretariat of the Convention on Biological Diversity, Montreal, Technical Series, 15: 164.

Vilina, Y., H. Cofré & C. Pizarro. 2006. Reporte final aves acuáticas de Chile. Water bird conservation of the Americas. Bird Life International, 40 pp.

Villa, F., L. Tunesi & T. Agardy. 2002. Zoning marine protected areas through spatial multiple-criteria analysis: the case of the Asinara Island National Marine Reserve of Italy. Conserv. Biol., 16: 515-526.

Wood, L.J. 2007. MPA Global: A database of the world's marine protected areas. Sea Around Us Project, UNEP-WCMC & WWF. Reviewed: 20 October 2011.


Received: 29 November 2011; Accepted: 10 June 2013


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