Corresponding author: Francisco J. Fernández-Rivera Melo (
Academic editor: Maria Elina Bichuette
San Pedro Mártir island is of high biological, ecological, and fishery importance and was declared a biosphere reserve in 2002. This island is the most oceanic in the Gulf of California, and information on its rocky reefs is scarce. The present study aimed to generate the first list of conspicuous invertebrate and fish species based on
Amador-Castro IG, Fernández-Rivera Melo FJ, Torre J (2021) Marine diversity in the biosphere reserve of the most oceanic island in the Gulf of California: San Pedro Mártir. ZooKeys 1062: 177–201.
The Midriff Islands Region (
Left panel: Map of the Gulf of California and surrounding areas. The Midriff Islands region (
A large number of studies of ecological indicators and inventories of invertebrate and fish species of the rocky reefs of the Gulf of California have been published, most of which have been conducted in natural protected areas (e.g.,
Despite the various studies that have been conducted in the
San Pedro Mártir island is located near the southern border of the
From 2007 to 2017, the rocky reefs of three sites within the core zone (Punta Rabijunco, Cueva de la Reserva, and Los Morritos) and four sites within the buffer zone (Arroyo del Cartelón, Cueva del Biólogo, Barra Baya, and La Ventana) of the
Visual surveys were conducted following the underwater monitoring protocols for kelp forests of the Partnership for Interdisciplinary Studies of Coastal Oceans (
The invertebrate species were classified based on the information available in the World Register of Marine Species (WoRMS 2019), and data on their distributions were obtained from
The data collected during sampling were used to estimate the ecological indicators of abundance, richness (S), diversity (i.e., Shannon-Wiener diversity index; ln; H’), and evenness (Pielou evenness index; J’), which are the most commonly used indices that have been used to evaluate community structure in the Gulf of California. These indices were used to compare community structure between the buffer and core zones of the
During the eleven years of this study, a total of 2,192 transects were surveyed (730 and 1,462 invertebrate and fish transects, respectively). Overall, 31,766 invertebrate individuals belonging to 35 species, 20 genera, and 27 families were recorded. The
Systematic list of invertebrates in the Island San Pedro Mártir Biosphere Reserve (
Taxa | Species | Species at risk or under protection | Biogeographic region | Commercial importance | ||
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NOM-059 | |||||
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Not evaluated | |||||
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Not evaluated | Subject to special protection |
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• | ||
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Not evaluated | • | ||||
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Not evaluated |
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• | |||
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Not evaluated | Subject to special protection |
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• | ||
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Minor concern |
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• | |||
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Not evaluated |
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• | |||
Not evaluated | ||||||
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Not evaluated | N/A | ||||
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Not evaluated |
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Not evaluated |
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Not evaluated | N/A | • | ||||
Not evaluated |
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• | ||||
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Not evaluated | N/A | ||||
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Not evaluated | N/A | ||||
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Not evaluated | N/A | ||||
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Not evaluated |
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Not evaluated |
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• | |||
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Not evaluated | N/A | ||||
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Not evaluated |
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Not evaluated |
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Not evaluated | • | ||||
Not evaluated |
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Not evaluated |
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Not evaluated |
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Not evaluated |
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Not evaluated |
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Not evaluated |
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In danger of extinction | Subject to special protection |
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• | ||
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Not evaluated |
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Not evaluated |
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Minor concern |
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• | |||
Minor concern |
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• |
Systematic list of fish species in the Island San Pedro Mártir Biosphere Reserve (
Taxa | Species | Species at risk or under protection | Biogeographic region | Commercial importance | Average size (min, max) | |||
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NOM-059 | |||||||
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Insufficient data |
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• | 45.00 (30,60) | ||||
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Vulnerable |
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• | 20.00 (20,20) | ||||
Insufficient data |
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• | 18.33 (15,20) | |||||
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Near threatened |
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• | 66.67 (20,100) | ||||
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Insufficient data |
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• | 29.63 (15,40) | ||||
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Minor concern |
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58.64 (30,100) | |||||
Minor concern |
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No data | ||||||
Minor concern |
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37.78 (20,60) | ||||||
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Minor concern |
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9.47 (5,15) | |||||
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Vulnerable | Subject to special protection |
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• | 20.00 (20,20) | |||
Minor concern | Subject to special protection |
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• | 21.59 (3,40) | ||||
Minor concern | Subject to special protection |
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• | 29.42 (15,40) | ||||
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Minor concern |
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• | 14.52 (5,20) | ||||
Minor concern |
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13.56 (3,30) | ||||||
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Minor concern |
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36.24 (5,60) | |||||
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Minor concern |
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• | 24.78 (12,40) | ||||
Minor concern | • | 24.63 (12,30) | ||||||
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Minor concern |
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• | 6.74 (3,10) | ||||
Minor concern |
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20.00 (20,20) | ||||||
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Minor concern |
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• | 27.60 (3,50) | ||||
Minor concern |
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23.57 (10,30) | ||||||
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Minor concern | C | 30.00 (25,40) | |||||
Minor concern |
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25.00 (20,30) | ||||||
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Minor concern |
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29.17 (25,30) | |||||
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Minor concern |
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• | 27.32 (3,40) | ||||
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Minor concern |
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• | 6.04 (3,10) | ||||
Minor concern |
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• | 30.08 (10,45) | |||||
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Minor concern |
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• | 17.34 (3,30) | ||
Minor concern |
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• | 22.98 (5,40) | |||||
Minor concern |
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• | 22.50 (20,25) | |||||
Minor concern |
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• | 24.67 (10,40) | |||||
In danger of extinction |
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• | 81.56 (20,150) | |||||
Insufficient data |
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• | 32.12 (10,50) | |||||
Minor concern |
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• | 30.25 (5,90) | |||||
Insufficient data |
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• | 30.00 (30,30) | |||||
Minor concern |
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• | 22.50 (15,30) | |||||
Minor concern |
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20.78 (3,40) | ||||||
Minor concern |
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7.83 (3,20) | ||||||
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Minor concern |
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5 (5,5) | |||||
Minor concern |
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5.13 (3,10) | ||||||
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Minor concern | C | • | 67.50 (60,100) | ||||
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Minor concern |
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• | 33.55 (5,50) | ||||
Minor concern |
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• | 31.52 (3,50) | |||||
Minor concern |
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• | 25.00 (25,25) | |||||
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Minor concern |
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• | 40 (40,40) | ||||
Minor concern |
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• | 31.32 (10,40) | |||||
Minor concern |
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• | 26.25 (20,30) | |||||
Minor concern |
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• | 23.19 (6,30) | |||||
Minor concern |
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• | 29.47 (3,45) | |||||
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Minor concern |
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• | 23.49 (15,30) | ||||
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Minor concern |
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16.21 (10,20) | |||||
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Minor concern |
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15 (15,15) | |||||
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Minor concern |
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12.44 (5,20) | |||||
Minor concern |
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• | 5.76 (3,15) | |||||
Minor concern | Subject to special protection |
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• | 4.67 (3,15) | ||||
Minor concern |
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• | 25.00 (20,30) | |||||
Minor concern |
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7.14 (5,10) | ||||||
Minor concern |
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8.50 (5,10) | ||||||
Minor concern |
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6.70 (3,25) | ||||||
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Minor concern |
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• | 20.10 (3,50) | ||||
Minor concern |
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9.63 (3,20) | ||||||
Minor concern |
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7.96 (3,25) | ||||||
Minor concern |
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6.29 (3,15) | ||||||
Minor concern |
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• | 13.13 (3,40) | |||||
Minor concern |
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10.17 (3,25) | ||||||
Minor concern |
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16.96 (5,35) | ||||||
Vulnerable |
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• | 38.13 (30,50) | |||||
Minor concern |
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7.50 (5,10) | ||||||
Minor concern |
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6.75 (3,20) | ||||||
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Minor concern |
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• | 19.30 (5,30) | ||||
Minor concern |
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32.00 (10,60) | ||||||
Minor concern | 27.25 (8,40) | |||||||
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Minor concern |
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22.28 (3,40) |
The Red List of the
In the Official Mexican Standard NOM-059-SEMARNAT-2010, six species (three invertebrate and four fish species) are listed as in need of special protection: the pearl oyster (
The abundance of
Average abundance (standard deviation) in 60 m2 of endangered, threatened, and protected eastern tropical Pacific species in the Island San Pedro Mártir Biosphere Reserve (
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n = 30 | n = 72 | n = 67 | n = 72 | n =67 | n =74 | n = 72 | n = 79 | n = 53 | n = 72 | n = 72 | |
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0.10 | 0.14 | 0.06 | 0.08 | 0.16 | 0.05 | 0.10 | 0.04 | 0.04 | 0.03 | 0.15 |
(0.40) | (0.51) | (0.24) | (0.28) | (0.54) | (0.28) | (0.42) | (0.25) | (0.19) | (0.17) | (0.55) | |
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0.27 | 0.22 | 0.04 | 0.01 | 0.07 | 0.07 | 0.03 | 0.09 | 0.02 | 0.00 | 0.00 |
(0.69) | (0.86) | (0.27) | (0.12) | (0.31) | (0.34) | (0.17) | (0.40) | (0.14) | (0.00) | (0.00) | |
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0.77 | 0.64 | 0.21 | 0.22 | 0.27 | 0.05 | 0.38 | 0.05 | 0.11 | 0.15 | 0.01(0.12) |
(1.79) | (1.08) | (0.45) | (0.48) | (0.62) | (0.28) | (0.88) | (0.22) | (0.42) | (0.39) | (0.12) | |
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n = 76 | n = 143 | n = 144 | n = 144 | n =141 | n =135 | n = 144 | n = 132 | n = 109 | n = 144 | n = 150 | |
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0.01 | 0.00 | 0.01 | 0.00 | 0.00 | 0.00 | 0.00 | 0.00 | 0.00 | 0.00 | 0.00 |
(0.11) | (0.00) | (0.12) | (0.00) | (0.00) | (0.00) | (0.00) | (0.00) | (0.00) | (0.00) | (0.00) | |
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0.03 | 0.00 | 0.00 | 0.00 | 0.00 | 0.01 | 0.00 | 0.00 | 0.00 | 0.00 | 0.00 |
(0.16) | (0.00) | (0.00) | (0.00) | (0.00) | (0.09) | (0.00) | (0.00) | (0.00) | (0.00) | (0.00) | |
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0.01 | 0.13 | 0.12 | 0.19 | 0.05 | 0.18 | 0.01 | 0.05 | 0.02 | 0.00 | 0.02 |
(0.11) | (0.61) | (0.38) | (0.71) | (0.25) | (1.58) | (0.08) | (0.27) | (0.13) | (0.00) | (0.14) | |
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0.08 | 0.16 | 0.59 | 0.24 | 0.74 | 0.10 | 0.04 | 0.00 | 0.00 | 0.06 | 0.03 |
(0.58) | (1.00) | (3.31) | (1.12) | (4.13) | (0.75) | (0.35) | (0.00) | (0.00) | (0.67) | (0.41) | |
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0.00 | 0.00 | 0.02 | 0.00 | 0.00 | 0.00 | 0.00 | 0.00 | 0.00 | 0.00 | 0.00 |
(0.00) | (0.00) | (0.25) | (0.00) | (0.00) | (0.00) | (0.00) | (0.00) | (0.00) | (0.00) | (0.00) | |
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4.17 | 2.40 | 2.34 | 2.20 | 2.63 | 2.03 | 2.17 | 2.49 | 2.06 | 2.24 | 1.75 |
(6.56) | (2.79) | (2.34) | (2.39) | (3.06) | (2.65) | (2.26) | (3.15) | (2.06) | (3.93) | (1.70) | |
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0.18 | 0.03 | 0.13 | 0.07 | 0.04 | 0.04 | 0.03 | 0.07 | 0.07 | 0.06 | 0.07 |
(0.86) | (0.22) | (0.68) | (0.37) | (0.20) | (0.23) | (0.16) | (0.31) | (0.35) | (0.34) | (0.34) | |
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0.00 | 0.00 | 0.00 | 0.00 | 0.00 | 0.00 | 0.01 | 0.00 | 0.00 | 0.03 | 0.01 |
(0.00) | (0.00) | (0.00) | (0.00) | (0.00) | (0.00) | (0.08) | (0.00) | (0.00) | (0.18) | (0.12) |
Of the 35 invertebrate species registered in the
Of the 73 registered fish species, four species are solely distributed in Mexico (
The year with the highest total richness of invertebrates was 2008 (33 species), while the lowest richness value was present in 2017 (21 species). The highest total recorded abundance of invertebrates was 6,359 individuals in 2013. For reef fishes, the years with the highest recorded richness values were 2009 and 2016 with 50 species recorded in each year, while the lowest number of species (37) was recorded in 2014. The highest fish abundance during the study period was 26,332 individuals, which was recorded in 2008 (Fig.
Total abundance and number of
Of the 108 species observed in the
A total of 54 commercially important species (12 invertebrate and 42 fish species) for the communities of the
Average size in centimeters (standard deviation) of the main commercially important fish species in the Island San Pedro Mártir Biosphere Reserve (
Species | 2007 | 2008 | 2009 | 2010 | 2011 | 2012 | 2013 | 2014 | 2015 | 2016 | 2017 |
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n = 76 | n = 143 | n = 144 | n = 144 | n = 141 | n = 135 | n = 144 | n = 132 | n = 109 | n = 144 | n = 150 | |
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23.75 | 23.33 | 26.67 | 26.92 | 28.07 | 27.33 | 30.12 | 27.55 | 27.46 | 26.57 | 28.39 |
(5.82) | (6.57) | (5.40) | (5.84) | (5.23) | (6.94) | (4.15) | (5.58) | (7.22) | (5.66) | (6.06) | |
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120.00 | 84.29 | 85.07 | 94.62 | 75.00 | 82.50 | 120.00 | 22.50 | 47.50 | 0.00 | 90.00 |
( |
(49.95) | (24.28) | (21.06) | (37.28) | (45.00) | ( |
(5.00) | (31.82) | ( |
(26.46) | |
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0.00 | 38.33 | 31.32 | 22.50 | 25.00 | 25.00 | 30.00 | 32.50 | 32.50 | 40.00 | 45.00 |
( |
(11.69) | (2.50) | (10.61) | ( |
(13.23) | ( |
(2.89) | (3.54) | ( |
( |
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21.86 | 25.90 | 25.77 | 30.56 | 31.78 | 32.84 | 30.69 | 33.51 | 35.49 | 28.61 | 34.08 |
(11.31) | (11.11) | (9.82) | (12.98) | (9.72) | (10.74) | (11.99) | (9.78) | (11.84) | (13.40) | (12.45) | |
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20.00 | 30.00 | 29.31 | 31.47 | 32.50 | 31.43 | 31.64 | 30.60 | 35.00 | 30.87 | 34.00 |
(10.00) | (5.53) | (8.21) | (4.0) | (5.34) | (5.73) | (7.69) | (7.94) | (3.97) | (8.48) | ( |
In the
Ecological indicators of invertebrate species richness (S), Shannon-Wiener diversity (H’), and Pielou evenness (J’) by transect. The inferior and superior sides of each blue rectangle represent the first and third quartiles (P25 and P75), respectively, and the median is represented by the horizontal black line. The points indicate the values of each data point, while the line surrounding each box plot shows the probability density.
In the case of fish, the average S value (mean ± SD) was 8.02 ± 2.43 species/transect. The highest S value was observed in 2010 (9.05 ± 2.09 species/transect), while the lowest S value was observed in 2017 (6.43 ± 2.13 species/transect; Fig.
Ecological indicators of fish species richness (S), Shannon-Wiener diversity (H‘), and Pielou evenness (J’) by transect. The inferior and superior sides of each blue rectangle represent the first and third quartiles (P25 and P75), respectively, and the median is represented by the horizontal black line. The points indicate the values of each data point, while the line surrounding each box plot shows the probability density.
When yearly changes in the rank abundance curves (Table
Differences in the alpha diversity metrics of invertebrates and fishes between 2007 and 2017.
Invertebrates | Change in S | Curve change | Change in evenness | Species loss | Species gains | |
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2007 | 2008 | 0.214 | 0.107 | -0.006 | 0.000 | 0.214 |
2008 | 2009 | -0.032 | 0.134 | -0.022 | 0.129 | 0.097 |
2009 | 2010 | -0.100 | 0.104 | -0.002 | 0.200 | 0.100 |
2010 | 2011 | 0.036 | 0.115 | -0.020 | 0.107 | 0.143 |
2011 | 2012 | -0.034 | 0.138 | 0.068 | 0.172 | 0.138 |
2012 | 2013 | -0.120 | 0.157 | -0.045 | 0.160 | 0.040 |
2013 | 2014 | 0.120 | 0.094 | 0.008 | 0.040 | 0.160 |
2014 | 2015 | -0.036 | 0.102 | 0.032 | 0.179 | 0.143 |
2015 | 2016 | 0.036 | 0.138 | -0.032 | 0.143 | 0.179 |
2016 | 2017 | -0.250 | 0.175 | 0.027 | 0.250 | 0.000 |
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2007 | 2008 | 0.000 | 0.143 | -0.008 | 0.154 | 0.154 |
2008 | 2009 | 0.109 | 0.108 | 0.012 | 0.091 | 0.200 |
2009 | 2010 | -0.073 | 0.093 | -0.006 | 0.164 | 0.091 |
2010 | 2011 | 0.019 | 0.146 | 0.035 | 0.113 | 0.132 |
2011 | 2012 | -0.019 | 0.129 | -0.026 | 0.132 | 0.113 |
2012 | 2013 | -0.040 | 0.124 | -0.016 | 0.120 | 0.080 |
2013 | 2014 | -0.149 | 0.123 | 0.035 | 0.213 | 0.064 |
2014 | 2015 | 0.048 | 0.126 | -0.027 | 0.071 | 0.119 |
2015 | 2016 | 0.196 | 0.116 | 0.015 | 0.039 | 0.235 |
2016 | 2017 | -0.017 | 0.137 | -0.016 | 0.172 | 0.155 |
Yearly ranks of the ten most abundant species of invertebrates
The distribution of a species is determined by both abiotic and biotic conditions, in addition to the accessibility of areas based on the dispersal limits of the species and the region in which it originally evolved. Abiotic and biotic conditions vary greatly among the different regions of the Gulf of California, and a south-north diversity gradient has been identified. This gradient has been reported by Thompson (1979),
In the
Two species (
The rocky reefs of the
Studies of invertebrates in the Gulf of California have focused on generating lists of both conspicuous and cryptic species of various invertebrate taxa (e.g., cnidarians, echinoderms, mollusks, and crustaceans) in the northern, central, and southern regions (
The studies that have been published on ichthyofauna in the
The most abundant invertebrate species in this study (
As
The H’ and J’ indices are useful for monitoring the conservation status of ecosystems because they consider the total number of species and the homogeneity with which their abundances are distributed. Both components of community structure have been interpreted against a background of important ecological processes (Magurran 2003). It may be deduced that a complex community with a greater number of interactions and stability is present when species diversity is high compared to that when species diversity is low (
The diversity and evenness results allowed us to identify trends for both the invertebrate and fish communities in this study. Between 2007 and 2011, a decrease in invertebrate diversity was observed, while a gradual increase in fish diversity was recorded (Fig.
Disturbance plays a central role in structuring communities, and the prevalence of human-induced disturbance has resulted in wide-ranging effects on biodiversity and ecosystem functioning and species abundance in particular (Matthews and Whittaker 2014). The stability of the ecological indicators evaluated in the
The results of this study constitute the first analysis of the community structure of the
In this study, we analyzed eleven years of survey data of the most oceanic marine protected area in the Gulf of California: the
This project was carried out with the support of the Comisión Nacional de Áreas Naturales Protegidas (CONANP), the Fondo Mexicano para la Conservación de la Naturaleza A.C. (FMCN), the World Wildlife Fund-Fundación Carlos Slim (WWF-FCS), The David and Lucile Packard Foundation, Marisla Foundation, Sandler Family Foundation, and International Community Foundation. We thank D. Torres, J. Torres, F. Pelayo, L. Encinas, N. Bustamante, R. Solano, S. Delgado, and U. Becerra of the Grupo de Monitoreo Submarino y de Cambio Climático of Bahía de Kino. We also thank the Sociedad Cooperativa Buzos Líderes. We thank A.L. MacTavish for the English edition.
Table S1
Xlsx file.
Average (standard deviation) for ecological indicators of invertebrates and fishes, species richness (S), Shannon-Wiener diversity (H’), and Pielou evenness (J’).
Table S2
Xlsx file.
Total abundance of invertebrates. Abbreviations: n = number of transects.
Table S3
Xlsx file.
Total abundance of fishes. Abbreviations: n = number of transects.
Table S4
Xlsx file.
Results of the post-hoc analysis for ecological indicators of invertebrates and fishes, species richness (S), Shannon-Wiener diversity (H’), and Pielou evenness (J’).