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Large Recovery of Fish Biomass in a No-Take Marine Reserve

  • Octavio Aburto-Oropeza ,

    Affiliation Marine Biology Research Division, Scripps Institution of Oceanography, La Jolla, California, United States of America

  • Brad Erisman,

    Affiliation Marine Biology Research Division, Scripps Institution of Oceanography, La Jolla, California, United States of America

  • Grantly R. Galland,

    Affiliation Marine Biology Research Division, Scripps Institution of Oceanography, La Jolla, California, United States of America

  • Ismael Mascareñas-Osorio,

    Affiliation Centro para la Biodiversidad Marina y la Conservación, La Paz, BCS, Mexico

  • Enric Sala,

    Affiliations National Geographic Society, Washington, DC, United States of America, Centre d'Estudis Avançats de Blanes (CSIC), Blanes, Spain

  • Exequiel Ezcurra

    Affiliation UC-MEXUS, University of California Riverside, Riverside, California, United States of America


No-take marine reserves are effective management tools used to restore fish biomass and community structure in areas depleted by overfishing. Cabo Pulmo National Park (CPNP) was created in 1995 and is the only well enforced no-take area in the Gulf of California, Mexico, mostly because of widespread support from the local community. In 1999, four years after the establishment of the reserve, there were no significant differences in fish biomass between CPNP (0.75 t ha−1 on average) and other marine protected areas or open access areas in the Gulf of California. By 2009, total fish biomass at CPNP had increased to 4.24 t ha−1 (absolute biomass increase of 3.49 t ha−1, or 463%), and the biomass of top predators and carnivores increased by 11 and 4 times, respectively. However, fish biomass did not change significantly in other marine protected areas or open access areas over the same time period. The absolute increase in fish biomass at CPNP within a decade is the largest measured in a marine reserve worldwide, and it is likely due to a combination of social (strong community leadership, social cohesion, effective enforcement) and ecological factors. The recovery of fish biomass inside CPNP has resulted in significant economic benefits, indicating that community-managed marine reserves are a viable solution to unsustainable coastal development and fisheries collapse in the Gulf of California and elsewhere.


Overfishing has impacted marine biodiversity and ecosystems both directly (through removal of significant biomass) and indirectly (by changing ecological linkages) throughout history [1], [2]. No-take marine reserves have been proposed as one of the most successful management tools to reverse these degradation trends [e.g., 3]. Evidence supporting the positive effects of no-take reserves include a greater abundance and biomass of fish inside marine reserves than in fished areas [see meta–analysis in 4]; an exponential increase of predatory fish biomass [e.g.], [ 5,6]; and shifts in species composition and trophic cascades that result in the restoration of natural marine communities within protected areas [7][10]. While these ecological changes operate on decadal times scales through a series of transient states [5], [11], [12], initial detections of both direct effects of area closures on target species and indirect effects on other taxa through cascading trophic interactions can be observed much sooner (5 and 13 years, respectively) [13], [14].

In addition to the aforementioned conservation benefits, well-enforced marine reserves help reduce local poverty and increase the economic revenue of coastal communities [15], [16]. Protected areas with locally managed resources and stakeholder buy-in can be more successful than areas with top down, federally mandated preservation [see 17]. However, marine reserve agendas have faced considerable opposition from different sectors of the society (e.g. commercial and recreational fisheries), only 0.1 percent of the world's ocean is completely protected from extractive activities, and most reserves suffer from poor management and enforcement [18], [19]. Moreover, the long-term success of marine reserves is a social issue that requires strong local leadership, social cohesion, involvement and effective self-enforcement within the community, and inter-generational coordination [20], [21].

Most of our knowledge of the benefits produced by no-take marine reserves comes from reserves smaller than 10 km2, and from single-time comparisons between protected areas and nearby fished sites [e.g.], [ 4], [13], [22,23]. Large relative increases in fish biomass (up to 20-fold) have been observed [e.g., 5]; but since these recoveries have happened in reserves less than 1 km2, the absolute increase in biomass has been limited. Furthermore, few marine reserves have been able to restore fish biomass to values similar to unfished habitats [24], [25]. Maybe because of that, most ecological and economic benefits (via spillover of adults to nearby unprotected areas) have been found for distances of only one kilometer on average beyond the reserve's boundaries [26].

Fisheries regulations in the Gulf of California (GOC) are numerous and complex as a result of the large number of exploited marine resources, and they have not been successful in recovering fish stocks [27], [28]. Even with its relatively low population size, the GOC is no exception to worldwide coastal and marine degradation trends. Overfishing, destruction of critical habitats, and the lack of proper planning that outline conservation and fisheries management priorities threaten marine biodiversity in the GOC [e.g.], [ 28][32]. Large scale tourism developments in some areas in the GOC, though often touted as an alternative to fishing, have exacerbated problems by increasing local population sizes, fishing effort to feed tourists, and human-nature interactions [33]. These issues will become more severe as commercial and transportation development continues to grow in the region [34], but while efforts to restore degraded ecosystems (e.g. mangrove forests) are improving, no studies have demonstrated that recovery of GOC marine ecosystems is plausible.

No-take marine reserves currently represent the most widely-promoted tool for the conservation and restoration of coastal and marine ecosystems in the GOC [35], especially in the absence of strong governmental enforcement programs at both national and regional scales. However, most Marine Protected Areas (MPAs) in the GOC, are multiple use areas that include zones ranging from open access to no-take areas; although no-take areas represent less than 5% of the majority of MPAs [36], [37], and have not recorded any significant positive changes in terms of recovery of fish or economic benefits [17].

An outlier among GOC MPAs is Cabo Pulmo National Park (CPNP), an area near the southern end of the Baja California Peninsula, designated a National Park in 1995 mainly to protect its large coral communities [38]. CPNP, though one of the smallest MPAs in the region (Table 1), has the largest percentage of core (no-take) area (35%). Due to the determined action of local families, protection and enforcement as a no-take reserve has expanded to include nearly 100% of CPNP's area. In 1999, we visited 60 reefs throughout the GOC from Cabo San Lucas at the southern tip of the Baja California Peninsula to the Midriff Islands in the Upper GOC [35], including reefs inside CPNP (Fig. 1). We replicated that study ten years later. Here we report the changes in fish diversity and biomass at CPNP, relative to other MPAs and unprotected areas in the GOC, between 1999 and 2009.

Figure 1. Location of sampling sites.

A) Sites surveyed by Sala et al. in 1999 [35], and resurveyed here in 2009. Dots inside the circle represent sites surveyed in Cabo Pulmo National Park in 1999. Dots inside squares represent sites in core zones (no-take areas) of other marine protected areas; the rest represent open access sites. B) Map of 11 sites surveyed at Cabo Pulmo National Park in 2009.

Table 1. List of MPAs established by the Mexican Federal Government in the Gulf of California.


Fish species richness increased significantly at CPNP from 1999 (average = 15 species per transect) to 2009 (25 species per transect; ANOVA, p<0.01). In contrast, reefs inside no-take areas (“core zones”) in other MPAs (1999 = 22, 2009 = 18; ANOVA, p<0.0001) and in open access areas (1999 = 20, 2009 = 17, ANOVA, p<0.0001) showed a significant decrease in species richness. Additionally, the diversity of top predators (measured as the inverse of Simpson's Index) increased significantly between 1999 and 2009 at CPNP (p<0.05); while elsewhere in the GOC, it either remained the same or decreased significantly (see Table S1).

In 1999, fish biomass at CPNP was not significantly different from that in the no-take areas or core zones in other MPAs and in open access areas (Fig. 2; Student's t-test, p>0.05). Between 1999 and 2009 fish biomass increased significantly in all trophic groups at CPNP at annual rates varying between 12 and 25% (Table 2). After 10 years, total biomass at CPNP increased from 0.75 to 4.24 t ha−1, a dramatic increase of 3.49 t ha−1 that corresponds to a 463% change. Change in fish biomass for each trophic group was higher at CPNP than at other core zones or in open access areas (sign test, p = 0.03). Furthermore, while growth rates at CPNP differed significantly from the null hypothesis of zero growth for all trophic groups (p<0.03), in all other areas rates did not differ from zero in any trophic group (Table 2). Consequently, fish biomass at CPNP in 2009 was 5.4 times larger than in other core zones and open access areas (Fig. 2). In contrast, differences in fish biomass between other core zones and open access areas were not different in 1999 or 2009 (Fig. 2; Student's t-test, p = 0.15).

Figure 2. Average biomass of fish trophic groups surveyed in 1999 and 2009 in each site category in the Gulf of California.

Table 2. Changes in fish biomass between 1999 and 2009 in (a) Cabo Pulmo National Park, (b) other no-take or core zones, and (c) open-access areas.

Mean biomass within every trophic group increased significantly between 1999 and 2009 at CPNP (Table 2), and the biomass of top predators increased by 1070% (Fig. 2). The relative variance (calculated as the square of the coefficients of variation) in between-transect biomass also increased significantly for top predators (Table 2), implying that spatial aggregation of fish schools increased significantly during the intervening decade.

Differences in size class frequencies indicate that the largest fishes encountered in our surveys were within CPNP, and that there were more individuals in the largest size classes at CPNP than at other reefs in the GOC. Furthermore, for 25 of 88 species encountered in our transects (e.g., Mycteroperca spp., Lutjanus spp., and Scarus spp.), the largest individuals observed in 2009 were at CPNP.

CPNP exhibited the largest absolute recovery of biomass in a marine reserve, and the faster relative increase in biomass of top predators, with a 30% annual increase of predatory fish (Fig. 3). To the best of our knowledge, only CPNP and Cabo de Palos Marine Reserve in the Mediterranean [39] have recovered total fish biomass to values larger than 4 t ha−1, and shown ratios of biomass inside the reserve to that in the surrounding fished areas larger than 5 times more biomass inside the reserve.

Figure 3. Comparison of the magnitude and rate of change of fish biomass at Cabo Pulmo National Park relative to other marine reserves around the world.

Data from other reserves were restricted to the few studies on temporal changes of total fish biomass (including all species) inside the same reserve. In all panels numbers in parentheses above bars represent years between surveys. In panels (b) and (c), relative annual rate of change between time zero and time t is calculated as: ρ = ln(xt/x0)/t. Data sources: Mombasa [6], New Caledonia [7], Saba [43], St. Lucia [45], Sumilon and Apo [44].


Our 10-year comparison demonstrates that CPNP has been an effective marine reserve for the recovery of reef fish biomass within its boundaries. After fifteen years of protection, species richness and total biomass are greater, and top predators are more abundant. The larger densities and individual sizes of fish at CPNP (Fig. 4), combine to create an average biomass that is more than five times larger than the average biomass in open access areas in the Gulf of California (GOC).

Figure 4. Examples of the fish assemblage at Cabo Pulmo National Park (CPNP).

(a) groupers, (b) snappers, (c) jacks, and (d) parrotfishes. Photographs were taken in the summers of 2008–2010 (Photography by: Octavio Aburto).

In contrast to CPNP, core zones in other MPAs in the GOC have not yielded a significant increase in fish biomass or species richness, and are no different from open access areas. For example, two small no-take marine reserves created in 2001 at Loreto Bay National Park [40] have stabilized fish abundances (as opposed to declines observed elsewhere), but probably as a result of their small size (1.4 km2 of total no-take area), they have not resulted in the recovery of fish populations [17].

Regrettably, we only sampled two reefs inside CPNP in 1999 (compared to 11 sites in 2009), as we did not anticipate such a recovery in the reef fish assemblage. However, analyzing data from only the two sites that were visited in both 1999 and 2009 yielded qualitatively similar increases (the minimal degrees of freedom for both dates does not allow accurate tests of significances). Furthermore, we have been monitoring 45 reefs in the GOC on an annual basis for more than a decade [29], [41], [42], and have visited CPNP throughout the year since 2005 to study the behavior of large groupers (Mycteroperca jordani, M. rocacea) and characterize reef fish spawning aggregations. These observations allow us to confidently validate our results describing a fish community that rebounded remarkably from 1999 to 2009, from an area with few top predators similar to nearby open access areas to a no-take marine reserve dominated by top predators.

CPNP exhibited the largest absolute increase in biomass in a marine reserve reported in the literature [e.g.], [ 6], [ 43][45]. Previous studies have reported larger relative increases of biomass [see review in 4], but the magnitude of change was smaller. The most striking result is that full, complete recovery of a degraded fish community is possible (when placed in the right area and governed correctly), even to the level that is comparable to remote habitats that never have been impacted by fishing and other local human impacts [25], [46]. Such examples of “full” recovery are extremely rare [44], and we could not have expected that it occurred in only ten years.

The abundance of top predators and carnivores at CPNP is approaching the inverse trophic pyramid that characterizes reef fish assemblages that have faced little or no fishing pressure [25], [46]. The presence of sharks is another characteristic of healthy marine ecosystems [47], [48]. While not encountered on survey transects, large sharks (e.g. Galeocerdo cuvier, Carcharhinus leucas, Triaenodon obesus) were commonly observed at survey sites at CPNP but rarely or never observed at other reefs surveyed or at historic shark areas in the GOC [authors' pers. obs.; 33].

The ecological reasons for such a large increase in fish biomass probably include several factors: 1) the reserve was larger than the size of marine reserves studied by scientists (on average smaller than 10 km2) and thus can harbor permanent populations of large reef fishes with large home ranges, 2) the coral habitat was intact [38], 3) the reserve included spawning areas for large predators [49], and 4) it is located in an area of high productivity driven from both the spatial heterogeneity generated by long basaltic dykes that run parallel to the coast [50], and its location in the transition zone between the enclosed Gulf of California and the open waters of the Pacific Ocean.

The success of CPNP is greatly due to local leadership, effective self-enforcement by local stakeholders, and the general support of the broader community. Protected areas with locally managed resources and stakeholder buy-in can be more successful than areas with top down, federally mandated preservation [see 17]. This model is considered the most viable in rural settings where people rely on local natural resources for their livelihoods. Boat captains, dive masters, and local people in general participate in various activities to enforce the regulations of CPNP to visitors and among themselves, including surveillance, fauna protection (e.g. sea turtle nesting sites), and beach and ocean cleaning programs. These efforts have generated robust social bonds within the community [51], key elements for successfully managing aquatic resources and securing the livelihoods of the communities that depend on them [21].

The ecological successes of CPNP are steadily translating into economic benefits within the small (∼100 residents) rural village of Cabo Pulmo and the surrounding areas. A recent study found that the locally owned, small-scale tourism operators in Cabo Pulmo generated US$538,800 in 2006 and have continued to grow at a manageable rate [52]. This amount is generated by less than 30 people, working in five small businesses, and producing approximately US$18,000 per capita; an amount significantly higher than the per capita Gross National Income in Mexico. While tourism is not always the best option in ecologically sensitive areas (e.g., when excessive tourism demand for limited natural resources limits their availability for the local people and threatens ecosystem viability), these residents are showing ability for success, when local people use, manage, and benefit from their local resources.

Materials and Methods

In 2009, we completed underwater visual surveys at 73 reefs in the GOC. Of those, 37 were the same sites surveyed by Sala et al. in 1999 [35] (including 2 in CPNP, Fig. 1A), and a total of 11 were located inside CPNP (Fig. 1B). In order to ensure the compatibility of data, we utilized the same survey methods described by Sala et al. [35]. Divers swam along 50 m transects observing and documenting fish species. Divers counted and estimated the size of all fishes belonging to all species within a five meter wide belt along each transect during two passes (250 m2 total). Different behavioral groups (mobile species versus territorial species), were surveyed during each pass to ensure that individuals were only counted once. At each site, we conducted four replicate transects at 20 m depth and four transects at 5 m depth. Using this method, we completed 435 total transects in 2009.

Typically, the majority of the area within any of Mexico's MPAs' boundaries allows for extractive activities, with only a small no-take area, known as “core zone,” designated for scientific research and monitoring. Seven of the ten MPAs in the GOC protect less than 6% of their total area through core zones (Table 1). In order to test CPNP's effectiveness as the only well enforced no-take marine reserve in the GOC, we divided our sites into three categories: (1) CPNP; (2) core zones in other MPAs, and (3) open access areas. For each category and for both survey periods (1999 and 2009), we calculated species richness, size structure, and biomass of all reef fishes. We also calculated biomass for each of four broad trophic groups: top predators, carnivores, zooplanktivores, and herbivores. We limited trophic categorization to these broad and robust groups because diets of species change ontogenetically and with the environment [53]. In order to maximize comparability with existing studies, biomass is expressed as tonnes per hectare. The biomass of individual fish was calculated using the allometric length-weight conversion: W = a TLb, where parameters a and b are species-specific constants, TL is total length in mm, and W is weight in grams. Length-weight fitting parameters were obtained from FishBase [54]. Differences in total transect biomass among trophic categories and between years were tested by means of a t test for unequal variances, and by a non-parametric Mann-Whitney U-test. Because fish schooling behaviour may also differ between years, we also tested for differences in relative biomass variance (the variance of the data standardized by the mean; an indicator of Poisson aggregation) using a variance-ratio F-test.

We also surveyed peer-reviewed scientific literature to compile a database of studies that document fish biomass values inside marine reserves and the surrounding fished areas to compare the changes in the reef fish community at CPNP to other marine reserves worldwide. We included only studies of fully-protected, no-take marine reserves and the nearby fished areas, and only those studies for which effects were measured for individual reserves in order to determine: (1) total fish biomass per unit area inside reserves; (2) ratio of fish biomass on reefs inside and outside reserves; and (3) annual rate of change of both total fish biomass and biomass in top trophic levels before and after reserve implementation. As it is a standard result in calculus that relative biomass increase can also be written as the rate of change of the logarithm of the variable [(1/y).dy/dt = d ln(y)/dt)], we estimated the annual relative rates of change between time zero and time t as ρ = ln(xt/x0)/t.

Supporting Information

Table S1.

Analyses of variance of Simpsons diversity index obtained using the different species per each trophic group for every category.



We thank Shannon Yee, Catalina Lopez, Ben Ruttenberg, and one anonymous reviewer for improving earlier versions of this paper.

Author Contributions

Conceived and designed the experiments: OA-O BE ES. Performed the experiments: OA-O BE GRG IM-O ES. Analyzed the data: OA-O IM-O EE. Wrote the paper: OA-O BE GRG ES EE.


  1. 1. Pauly D, Watson R, Alder J (2005) Global trends in world fisheries impacts on marine ecosystems and food security. Philosophical Transactions of the Royal Society 360: 5–12.
  2. 2. Worm B, Barbier EB, Beaumont N, Duffy EJ, Folke C, et al. (2009) Impacts of biodiversity loss on oceans ecosystems services. Science 314: 787–790.
  3. 3. Balmford A, Gravestock P, Hockley N, McClean CJ, Roberts CM (2004) The worldwide costs of marine protected areas. Proceedings of the National Academy of Sciences 101: 9694–9697.
  4. 4. Lester SE, Halpern BS, Grorud-Colvert K, Lubchenco J, Ruttenberg BI, et al. (2009) Biological effects within no-take marine reserves: a global synthesis. Marine Ecology Progress Series 384: 33–46.
  5. 5. Russ GR, Alcala AC (2004) Marine reserves: long-term protection is required for full recovery of predatory fish populations. Oecologia 138: 622–627.
  6. 6. McClanahan TR, Graham NAJ, Calnan JM, MacNeil MA (2007) Toward pristine biomass: reef fish recovery in coral reef marine protected areas in Kenya. Ecological Applications 17: 1055–1067.
  7. 7. Wantiez L, Thollot P, Kulbicki M (1997) Effects of marine reserves on coral reef fish communities from five islands in New Caledonia. Coral Reefs 16: 215–224.
  8. 8. Shears NT, Babcock RC (2002) Marine reserves demonstrate top-down control of community structure on temperate reefs. Oecologia 132: 131–142.
  9. 9. Graham NAJ, Evans RD, Russ GR (2003) The effects of marine reserve protection on the trophic relationships of reef fishes on the Great Barrier Reef. Environmental Conservation 30: 200–208.
  10. 10. Mumby PJ, Dahlgren CP, Harborne AR, Kappel CV, Micheli F, et al. (2006) Fishing, trophic cascades, and the process of grazing on coral reefs. Science 311: 98–101.
  11. 11. Micheli F, Halpern BS, Botsford LW, Warner RR (2004) Trajectories and correlates of community change in no-take marine reserves. Ecological Applications 14: 1709–1723.
  12. 12. Barrett NS, Edgar GJ, Buxton CD, Haddon M (2007) Changes in fish assemblages following 10 years of protection in Tasmanian marine protected areas. Journal of Experimental Marine Biology and Ecology 345: 141–157.
  13. 13. Halpern BS, Warner RR (2002) Marine reserves have rapid and lasting effects. Ecology Letters 5: 361–365.
  14. 14. Babcock RC, Shears NT, Alcala NC, Barrett NS, Edgar GJ, Lafferty KD, McClanahan TR, Russ GR (2010) Decadal trends in marine reserves reveal differential rates of change in direct and indirect effects. Proceedings of the National Academy of Sciences 107: 18256–18261.
  15. 15. Leisher CP, van Beukering , Scherl LM (2007) Nature's investment bank. 52 p. Marine protected areas contribute to poverty reduction. Report to The Nature Conservancy, the Australian Government Department of the Environment and Water Resources, and the Poverty Reduction and Environment Management Program at Vrije Universiteit in Amsterdam.
  16. 16. McClanahan TR (2010) Effects of fisheries closures and gear restrictions on fishing income in a Kenyan coral reef. Conservation Biology 24: 1519–1528.
  17. 17. Cudney-Bueno R, Bourillon L, Saenz-Arroyo A, Torre-Cosıo J, Turk-Boyer P, et al. (2009) Governance and effects of marine reserves in the Gulf of California, Mexico. Ocean & Coastal Management 52: 207–218.
  18. 18. Mora C, Andréfouët S, Costello MJ, Kranenburg C, Rollo A, et al. (2006) Coral reefs and the global network of marine protected areas. Science 312: 1750–1751.
  19. 19. Wood LJ, Fish L, Laughren J, Pauly D (2008) Assessing progress towards global marine protection targets: shortfalls in information and action. Oryx 42: 340–351.
  20. 20. Russ GR, Alcala AC (1999) Management histories of Sumilon and Apo marine reserves, Philippines, and their influence on national marine resource policy. Coral Reefs 18: 307–319.
  21. 21. Gutiérrez N, Hilborn R, Defeo O (2010) Leadership, social capital and incentives promote successful fisheries. Nature 470: 386–389.
  22. 22. Roberts CM, Hawkins JP (1997) How small can a marine reserve be and still be effective? Coral Reefs 16: 150.
  23. 23. Halpern BS (2003) The impact of marine reserves: do reserves work and does reserve size matter? Ecological Applications 13: S117–S137.
  24. 24. Newman MJH, Paredes GA, Sala E, Jackson JBC (2006) Structure of Caribbean coral reef communities across a large gradient of fish biomass. Ecology Letters 9: 1216–1227.
  25. 25. Sandin SA, Smith JE, De Martini EE, Dinsdale EA, et al. (2008) Baselines and degradation in coral reef in the northern Line Islands. PLoS ONE 3: e1548.
  26. 26. Halpern BS, Lester SE, Kellner JB (2009) Spillover from marine reserves and the replenishment of fished stocks. Environmental Conservation 36: 268–276.
  27. 27. Ezcurra E, Aburto-Oropeza O, de los Angeles Carvajal M, Cudney-Bueno R, Torre J (2009) Gulf of California, Mexico. In: McLeod K, Leslie K, editors. Ecosystem-based Management for the Oceans. London: Island Press. pp. 227–252.
  28. 28. Erisman B, Paredes GA, Plomozo-Lugo T, Cota-Nieto JJ, Hastings P, Aburto-Oropeza O (2011) Spatial structure of commercial marine fisheries in Northwest Mexico. ICES Journal of Marine Science.
  29. 29. Sala E, Aburto-Oropeza O, Reza M, Paredes GA, Lopez-Lemus G (2004) Fishing down coastal food webs in the Gulf of California. Fisheries 29: 19–25.
  30. 30. Jaramillo-Legorreta A, Rojas-Bracho L, Brownell RL, Read AJ, Reeves RR, et al. (2007) Saving the vaquita immediate action, not more data. Conservation Biology 21: 1653–1655.
  31. 31. Lozano-Montes H, Pitcher TJ, Haggan N (2008) Shifting environmental and cognitive in the upper Gulf of California. Frontiers in Ecology and the Enviroment 6: 75–80.
  32. 32. Sagarin RD, Gilly WF, Baxter CH, Burnett N, Christensen J (2008) Remembering the Gulf: change to the marine communities of the Sea of Cortez since the Steinbeck and Ricketts expedition of 1940. Frontier in Ecology and the Environment 6: 372–379.
  33. 33. López-Sagástegui C, Sala E (2005) Marine Biodiversity Assessment and Human Impacts in the Ensenada de La Paz, Baja California Sur. 62 p. International Community Foundation Report.
  34. 34. Pesenti C, Dean K (2003) Sustainable coastal development. La escalera nautica: mega-tourism project on the Baja California peninsula. California: Pro peninsula. 46 p.
  35. 35. Sala E, Aburto-Oropeza O, Paredes G, Parra I, Barrera JC, et al. (2002) A general model for designing networks of marine reserves. Science 298: 1991–1993.
  36. 36. Aburto-Oropeza O, Lopez-Sagastegui C (2006) Red de reservas marinas del Golfo de California: compilación de los esfuerzos de conservación. Mexico: Greenpeace. 28 p.
  37. 37. Fraga J, Jesus A (2008) Coastal and marine protected areas in Mexico. India: Nagaraj and company. 98 p.
  38. 38. Reyes-Bonilla H (1997) A new marine reserve in the Gulf of California. Conservation Biology 11: 838.
  39. 39. García-Charton JA, Pérez-Ruzafa A, Sánchez-Jerez P, Bayle-Sempere JT, Reñones O, et al. (2004) Multi-scale spatial heterogeneity, habitat structure, and the effect of marine reserves on Western Mediterranean rocky reef fish assemblages. Marine Biology 144: 161–182.
  40. 40. Sáenz-Arroyo A, Torre J (2005) Design and evaluation of marine reserves in the islands of the Gulf of California 24): Mexico: Mexican Fund for the Conservation of Nature (A-1-00/. 38.
  41. 41. Aburto-Oropeza O, Sala E, Paredes G, Mendoza A, Ballesteros E (2007) Predictability of reef fish recruitment in a highly variable nursery habitat. Ecology 88: 2220–2228.
  42. 42. Aburto-Oropeza O, Dominguez-Guerrero I, Cota-Nieto J, Plomozo-Lugo T (2009) Recruitment and ontogenetic habitat shifts of the Yellow snapper (Lutjanus argentiventris) in the Gulf of California. Marine Biology 156: 2461–2472.
  43. 43. Roberts CM (1995) Rapid Build-up of fish biomass in a Caribbean marine reserve. Conservation Biology 9: 815–826.
  44. 44. Russ GR, Stockwell B, Alcala AC (2005) Inferring versus measuring rates of recovery in no-take marine reserves. Marine Ecology Progress Series 292: 1–12.
  45. 45. Hawkins JP, Roberts CM, Dytham C, Schelten C, Nugues MM (2006) Effects of habitat characteristics and sedimentation on performance of marine reserves in St. Lucia. Biological Conservation 127: 487–499.
  46. 46. DeMartini EE, Friedlander AM, Sandin SA, Sala E (2008) Differences in fish assemblages structure between fished and unfished atolls in northern Line Islands central Pacific. Marine ecology Progress Series 365: 199–215.
  47. 47. DeMartini EE, Friedlander AM, Holzwarth SR (2005) Size at sex change bin protogynous labroids, prey size distributions and apex predators densities at NW Hawaiian atolls. Marine Ecology Progress Series 297: 259–271.
  48. 48. Friedlander AM, Sandin SA, DeMartini EE, Sala E (2010) Spatial patterns of the structure of reef fish assemblages at a pristine atoll in the central Pacific. Marine ecology Progress Series 410: 219–231.
  49. 49. Sala E, Aburto-Oropeza O, Paredes G, et al. (2003) Spawning aggregations and reproductive behavior of reef fishes in the Gulf of California. Bulletin Marine Science 72: 103–121.
  50. 50. Squires DF (1959) Results of the puritan-amarican museum of natural history expedition to western Mexico. Bulletin of the American Museum of Natural History 118: 367–432.
  51. 51. Gámez AE (2008) Turismo y sustentabilidad en Cabo Pulmo, Baja California Sur. California: San Diego State University, Universidad Autónoma de Baja California Sur, Consejo Nacional de Ciencia y Tecnología. 314 p.
  52. 52. Martínez de la Torre JA (2008) Desarrollo local y el estado de la economía base en Cabo Pulmo. In: Gámez AE, editor. Turismo y sustentabilidad en Cabo Pulmo, Baja California Sur. California: San Diego State University, Universidad Autónoma de Baja California Sur, Consejo Nacional de Ciencia y Tecnología. pp. 133–162.
  53. 53. Harmelin-Vivien ML (2002) Energetic and fish diversity on coral reef. In: Sale PF, editor. Coral reef fishes: Dynamics and diversity in a complex ecosystem. Sydney: Academic press. pp. 265–274.
  54. 54. Froese R, Pauly D (2011) Fishbase World Wide Web electronic publication.