Research Article |
Corresponding author: Francisco J. Fernández-Rivera Melo ( ffernandez@cobi.org.mx ) Academic editor: Maria Elina Bichuette
© 2021 Imelda G. Amador-Castro, Francisco J. Fernández-Rivera Melo, Jorge Torre.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
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. https://doi.org/10.3897/zookeys.1062.67964
|
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 in situ observations and to examine the community structure of the shallow rocky reefs of the reserve. In addition, we estimated the ecological indicators of richness, abundance, Shannon diversity, and Pielou evenness to evaluate the conservation status of the biosphere reserve. Data were collected annually from 2007 to 2017 through 2,192 underwater SCUBA transects. A total of 35 species of invertebrates and 73 species of fish were recorded. Most of the species are widely distributed along the eastern Pacific. Overall, 64% of the species found in this study are commercially important, and 11 species have been listed as protected. The abundance of commercially important invertebrate species (i.e., the sea cucumber Isostichopus fuscus and the spiny oyster Spondylus limbatus) is decreasing, while commercially important fish species have maintained their abundance with periods of increase. The ecological indicators and the abundance and size of the commercial species indicate that the reserve is in good condition while meeting its conservation objectives.
Diversity, ecological indicator, ichthyofauna, invertebrate, Midriff Islands Region, natural protected area, systematic list
The Midriff Islands Region (MIR), which is located near the central portion of the Gulf of California, is an archipelago comprising 45 islands and islets with high biodiversity. Due to this diversity, the MIR is important for conservation, and three natural protected areas have been decreed within its borders: the Archipiélago de San Lorenzo National Park; Bahía de los Ángeles, Canales de Ballenas y Salsipuedes Biosphere Reserve; and Island San Pedro Mártir Biosphere Reserve (ISPMBR; Fig.
Left panel: Map of the Gulf of California and surrounding areas. The Midriff Islands region (MIR, dotted line) includes the a Bahía de los Ángeles, Canales de Ballenas y Salsipuedes Biosphere Reserve; b Archipiélago de San Lorenzo National Park; and c island San Pedro Mártir Biosphere Reserve (ISPMBR), whereas d Bahía de Loreto National Park is located to the south of the MIR. Right panel: The limits of the ISPMBR, including Island San Pedro Mártir (shaded region) and the monitoring sites (1–6) of this study. 1 Punta Rabijunco 2 Cueva de la Reserva and 3 Los Morritos are located within the core, no-take zone (dashed region) of the ISPMBR, whereas 4 Cueva del Biólogo 5 Barra Baya 6 La Ventana and 7 Arroyo del Cartelón are located within the buffer zone.
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 MIR, there are currently multiple ecological and biogeographic information gaps and uncertainty regarding the degree to which the conservation objectives of the ISPMBR have been met. As such, the present study aimed to generate a list of conspicuous invertebrate and fish species based on in situ observations, in addition to evaluating the community structure and main species (e.g., commercial, endangered, threatened, or protected) present in the shallow rocky reefs (< 20 m depth) of the ISPMBR from 2007 to 2017.
San Pedro Mártir island is located near the southern border of the MIR, and its polygon (28°18'00"N, 112°13'30"W and 28°28'00"N, 112°23'30"W) is situated between two regions, the northern Gulf of California and central Gulf of California, which have distinct oceanographic (i.e., physical and chemical) characteristics and marine fauna (i.e., invertebrate and fish species; Walter 1960;
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 ISPMBR were monitored (Fig.
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 ISPMBR and among monitoring years. Estimations of the ecological indicators were carried out with PAST v. 4.06 (Hammer 2001), whereas statistical analyses were carried out in JAMOVI v. 1.6 (The jamovi project). When no significant differences were found between the buffer zone and core zones with regard to the three ecological indicators (Suppl. material
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 Muricidae family was the most represented in this study with three species, while 19 families were represented by one species (Table
Systematic list of invertebrates in the Island San Pedro Mártir Biosphere Reserve (ISPMBR) in the Gulf of California, Mexico. The biogeographic region of each species is shown. Abbreviations: EP = eastern Pacific; EP + WCA = eastern Pacific + western central Atlantic; EP + NEA = eastern Pacific + northeastern Atlantic; EP + SA = eastern Pacific + southern Atlantic; ECP = eastern central Pacific; SP = southeastern Pacific; CA = Central America; IP = Indo-Pacific; IP + WCA = Indo-Pacific + western central Atlantic; PO = Pacific Ocean; N/A = No information available; IUCN = International Union for Conservation of Nature; NOM-59 = Mexican law for endangered, threatened, or protected species (NOM-059-SEMARNAT-2010).
Taxa | Species | Species at risk or under protection | Biogeographic region | Commercial importance | ||
---|---|---|---|---|---|---|
IUCN | NOM-059 | |||||
MOLLUSCA | BIVALVIA | |||||
Ostreida | ||||||
Gryphaeidae | Hyotissa hyotis (Linnaeus, 1758) | Not evaluated | IP + WCA | |||
Margaritidae | Pinctada mazatlanica (Hanley, 1856) | Not evaluated | Subject to special protection | EP | • | |
Pinnidae | Pinna rugosa G. B. Sowerby I, 1835 | Not evaluated | EP + NEA | • | ||
Pteriidae | Pteria sterna (Gould, 1851) | Not evaluated | EP | • | ||
Spondylidae | Spondylus limbatus G. B. Sowerby II, 1847 | Not evaluated | Subject to special protection | IP | • | |
CEPHALOPODA | ||||||
Order Octopoda | ||||||
Octopodidae | Octopus bimaculatus Verrill, 1883 | Minor concern | PO | • | ||
GASTROPODA | ||||||
Littorinimorpha | ||||||
Strombidae | Strombus gracilior G. B. Sowerby I, 1825 | Not evaluated | EP | • | ||
Titanostrombus galeatus (Swainson, 1823) | Not evaluated | EP +SA | ||||
Neogastropoda | ||||||
Columbellidae | Strombina maculosa (G. B. Sowerby I, 1832) | Not evaluated | N/A | |||
Fasciolariidae | Triplofusus princeps (G. B. Sowerby I, 1825) | Not evaluated | EP | |||
Muricidae | Hexaplex erythrostomus (Swainson, 1831) | Not evaluated | CA | • | ||
Hexaplex nigritus (Philippi, 1845) | Not evaluated | N/A | • | |||
Hexaplex princeps (Broderip, 1833) | Not evaluated | SP | • | |||
Trochida | ||||||
Turbinidae | Turbo fluctuosus W. Wood, 1828 | Not evaluated | N/A | |||
ECHINODERMATA | ASTEROIDEA | |||||
Forcipulatida | ||||||
Heliasteridae | Heliaster kubiniji Xantus, 1860 | Not evaluated | N/A | |||
Spinulosida | ||||||
Echinasteridae | Echinaster tenuispina Verrill, 1871 | Not evaluated | N/A | |||
Valvatida | ||||||
Acanthasteridae | Acanthaster planci (Linnaeus, 1758) | Not evaluated | IP | |||
Asteropseidae | Asteropsis carinifera (Lamarck, 1816) | Not evaluated | IP | • | ||
Mithrodiidae | Mithrodia bradleyi Verrill, 1867 | Not evaluated | N/A | |||
Ophidiasteridae | Pharia pyramidata (Gray, 1840) | Not evaluated | EP | • | ||
Phataria unifascialis (Gray, 1840) | Not evaluated | EP | • | |||
Oreasteridae | Nidorellia armata (Gray, 1840) | Not evaluated | EP + WCA | • | ||
Pentaceraster cumingi (Gray, 1840) | Not evaluated | EP | • | |||
ECHINOIDEA | Arbacioida | |||||
Arbaciidae | Arbacia stellata (Blainville, 1825; Gmelin, 1791) | Not evaluated | EP | • | ||
Camarodonta | ||||||
Echinometridae | Echinometra vanbrunti A. Agassiz, 1863 | Not evaluated | EP | |||
Toxopneustidae | Toxopneustes roseus (A. Agassiz, 1863) | Not evaluated | EP | |||
Tripneustes depressus A. Agassiz, 1863 | Not evaluated | EP | ||||
Cidaroida | ||||||
Cidaridae | Eucidaris thouarsii (L. Agassiz & Desor, 1846) | Not evaluated | EP | • | ||
Diadematoida | ||||||
Diadematidae | Centrostephanus coronatus (Verrill, 1867) | Not evaluated | EP | • | ||
ECHINOIDEA | Diadematidae | Diadema mexicanum A. Agassiz, 1863 | Not evaluated | EP | ||
HOLOTHUROIDEA | ||||||
Synallactida | ||||||
Stichopodidae | Isostichopus fuscus (Ludwig, 1875) | In danger of extinction | Subject to special protection | EP | • | |
CNIDARIA | ANTHOZOA | |||||
Antipatharia | ||||||
Antipathidae | Antipathes galapagensis Deichmann, 1941 | Not evaluated | EP | |||
ARTHROPODA | MALACOSTRACA | |||||
Decapoda | ||||||
Inachoididae | Stenorhynchus debilis (Smith, 1871) | Not evaluated | SP | |||
Palinuridae | Panulirus inflatus (Bouvier, 1895) | Minor concern | ECP | • | ||
Panulirus interruptus (Randall, 1840) | Minor concern | ECP | • |
Systematic list of fish species in the Island San Pedro Mártir Biosphere Reserve (ISPMBR) in the Gulf of California, Mexico. The biogeographic region of each species is shown, as well as the average, minimum (min), and maximum (max) sizes (cm) based on the survey data. Abbreviations: EP = eastern Pacific; ECP=eastern central Pacific; C = Circumglobal; CT = Circumtropical; CT + EP = Circumtropical+eastern Pacific; IP = Indo-Pacific; IP + EP = Indo-Pacific + eastern Pacific; IUCN = International Union for Conservation of Nature; NOM-59 = Mexican law for endangered, threatened, or protected species (NOM-059-SEMARNAT-2010).
Taxa | Species | Species at risk or under protection | Biogeographic region | Commercial importance | Average size (min, max) | |||
---|---|---|---|---|---|---|---|---|
IUCN | NOM-059 | |||||||
ELASMOBRANCHII | Heterodontiformes | |||||||
Heterodontidae | Heterodontus francisci (Girard 1855) | Insufficient data | EP | • | 45.00 (30,60) | |||
Torpediniformes | ||||||||
Narcinidae | Diplobatis ommata (Jordan & Gilbert 1890) | Vulnerable | EP | • | 20.00 (20,20) | |||
Narcine entemedor Jordan & Starks 1895 | Insufficient data | EP | • | 18.33 (15,20) | ||||
Rhinopristiformes | ||||||||
Rhinobatidae | Pseudobatos productus (Ayres 1854) | Near threatened | EP | • | 66.67 (20,100) | |||
Myliobatiformes | ||||||||
Urotrygonidae | Urobatis concentricus Osburn & Nichols 1916 | Insufficient data | ECP | • | 29.63 (15,40) | |||
ACTINOPTERI | Anguilliformes | |||||||
Muraenidae | Gymnothorax castaneus (Jordan & Gilbert 1883) | Minor concern | EP | 58.64 (30,100) | ||||
Gymnothorax dovii (Günther 1870) | Minor concern | EP | No data | |||||
Muraena lentiginosa Jenyns 1842 | Minor concern | EP | 37.78 (20,60) | |||||
Blenniiformes | ||||||||
Blenniidae | Ophioblennius steindachneri Jordan & Evermann 1898 | Minor concern | EP | 9.47 (5,15) | ||||
Acanthuriformes | ||||||||
Pomacanthidae | Holacanthus clarionensis Gilbert 1890 | Vulnerable | Subject to special protection | ECP | • | 20.00 (20,20) | ||
Holacanthus passer Valenciennes 1846 | Minor concern | Subject to special protection | EP | • | 21.59 (3,40) | |||
Pomacanthus zonipectus (Gill 1862) | Minor concern | Subject to special protection | EP | • | 29.42 (15,40) | |||
Chaetodontidae | Chaetodon humeralis Günther 1860 | Minor concern | EP | • | 14.52 (5,20) | |||
Johnrandallia nigrirostris (Gill 1862) | Minor concern | EP | 13.56 (3,30) | |||||
Acanthuridae | Prionurus laticlavius Gill 1862 | Minor concern | ECP | 36.24 (5,60) | ||||
Tetraodontiformes | ||||||||
Diodontidae | Diodon holocanthus Linnaeus 1758 | Minor concern | CT | • | 24.78 (12,40) | |||
Diodon hystrix Linnaeus 1758 | Minor concern | CT+ EP | • | 24.63 (12,30) | ||||
Tetraodontidae | Canthigaster punctatissima (Günther 1870) | Minor concern | ECP | • | 6.74 (3,10) | |||
Sphoeroides lobatus (Steindachner 1870) | Minor concern | EP | 20.00 (20,20) | |||||
Balistidae | Balistes polylepis Steindachner 1876 | Minor concern | EP | • | 27.60 (3,50) | |||
Sufflamen verres (Gilbert & Starks 1904) | Minor concern | EP | 23.57 (10,30) | |||||
Centrarchiformes | ||||||||
Kyphosidae | Kyphosus vaigiensis (Quoy & Gaimard 1825) | Minor concern | C | 30.00 (25,40) | ||||
Kyphosus azureus (Jenkins & Evermann 1889) | Minor concern | ECP | 25.00 (20,30) | |||||
Kyphosidae | Kyphosus elegans (Peters 1869) | Minor concern | EP | 29.17 (25,30) | ||||
Girellidae | Girella simplicidens Osburn & Nichols 1916 | Minor concern | ECP | • | 27.32 (3,40) | |||
Cirrhitidae | Cirrhitichthys oxycephalus (Bleeker 1855) | Minor concern | IP | • | 6.04 (3,10) | |||
Cirrhitus rivulatus Valenciennes 1846 | Minor concern | ECP | • | 30.08 (10,45) | ||||
ACTINOPTERI | Perciformes | Serranidae | Alphestes immaculatus Breder 1936 | Minor concern | EP | • | 17.34 (3,30) | |
Cephalopholis panamensis (Steindachner 1876) | Minor concern | EP | • | 22.98 (5,40) | ||||
Epinephelus analogus Gill 1863 | Minor concern | EP | • | 22.50 (20,25) | ||||
Epinephelus labriformis (Jenyns 1840) | Minor concern | EP | • | 24.67 (10,40) | ||||
Mycteroperca jordani (Jenkins & Evermann 1889) | In danger of extinction | ECP | • | 81.56 (20,150) | ||||
Mycteroperca prionura Rosenblatt & Zahuranec 1967 | Insufficient data | ECP | • | 32.12 (10,50) | ||||
Mycteroperca rosacea (Streets 1877) | Minor concern | ECP | • | 30.25 (5,90) | ||||
Paralabrax auroguttatus Walford 1936 | Insufficient data | ECP | • | 30.00 (30,30) | ||||
Paralabrax maculatofasciatus (Steindachner 1868) | Minor concern | ECP | • | 22.50 (15,30) | ||||
Paranthias colonus (Valenciennes 1846) | Minor concern | EP | 20.78 (3,40) | |||||
Serranus psittacinus Valenciennes 1846 | Minor concern | EP | 7.83 (3,20) | |||||
Apogonidae | Apogon pacificus (Herre 1935) | Minor concern | EP | 5 (5,5) | ||||
A retrosella (Gill 1862) | Minor concern | ECP | 5.13 (3,10) | |||||
Carangidae | Seriola lalandi Valenciennes 1833 | Minor concern | C | • | 67.50 (60,100) | |||
Lutjanidae | Hoplopagrus guentherii Gill 1862 | Minor concern | EP | • | 33.55 (5,50) | |||
Lutjanus argentiventris (Peters 1869) | Minor concern | EP | • | 31.52 (3,50) | ||||
Lutjanus viridis (Valenciennes 1846) | Minor concern | EP | • | 25.00 (25,25) | ||||
Haemulidae | Anisotremus davidsonii (Steindachner 1876) | Minor concern | ECP | • | 40 (40,40) | |||
Anisotremus interruptus (Gill 1862) | Minor concern | EP | • | 31.32 (10,40) | ||||
Haemulon flaviguttatum Gill 1862 | Minor concern | EP | • | 26.25 (20,30) | ||||
Haemulon maculicauda (Gill 1862) | Minor concern | EP | • | 23.19 (6,30) | ||||
Haemulon sexfasciatum Gill 1862 | Minor concern | ECP | • | 29.47 (3,45) | ||||
Sparidae | Calamus brachysomus (Lockington 1880) | Minor concern | EP | • | 23.49 (15,30) | |||
Sciaenidae | Pareques fuscovittatus (Kendall & Radcliffe 1912) | Minor concern | ECP | 16.21 (10,20) | ||||
Mullidae | Mulloidichthys dentatus (Gill 1862) | Minor concern | EP | 15 (15,15) | ||||
Pomacentridae | Abudefduf troschelii (Gill 1862) | Minor concern | EP | 12.44 (5,20) | ||||
Azurina atrilobata Gill 1862 | Minor concern | EP | • | 5.76 (3,15) | ||||
Chromis limbaughi Greenfield & Woods 1980 | Minor concern | Subject to special protection | ECP | • | 4.67 (3,15) | |||
Microspathodon dorsalis (Gill 1862) | Minor concern | EP | • | 25.00 (20,30) | ||||
Stegastes acapulcoensis (Fowler 1944) | Minor concern | EP | 7.14 (5,10) | |||||
Stegastes flavilatus (Gill 1862) | Minor concern | EP | 8.50 (5,10) | |||||
Stegastes rectifraenum (Gill 1862) | Minor concern | ECP | 6.70 (3,25) | |||||
Labridae | Bodianus diplotaenia (Gill 1862) | Minor concern | EP | • | 20.10 (3,50) | |||
Halichoeres chierchiae Di Caporiacco 1948 | Minor concern | ECP | 9.63 (3,20) | |||||
Halichoeres dispilus (Günther 1864) | Minor concern | EP | 7.96 (3,25) | |||||
Halichoeres melanotis (Gilbert 1890) | Minor concern | ECP | 6.29 (3,15) | |||||
Halichoeres nicholsi (Jordan & Gilbert 1882) | Minor concern | EP | • | 13.13 (3,40) | ||||
Halichoeres notospilus (Günther 1864) | Minor concern | EP | 10.17 (3,25) | |||||
Halichoeres semicinctus (Ayres 1859) | Minor concern | EP | 16.96 (5,35) | |||||
Semicossyphus pulcher (Ayres 1854) | Vulnerable | EP | • | 38.13 (30,50) | ||||
Thalassoma grammaticum Gilbert 1890 | Minor concern | EP | 7.50 (5,10) | |||||
Thalassoma lucasanum (Gill 1862) | Minor concern | EP | 6.75 (3,20) | |||||
Scaridae | Nicholsina denticulata (Evermann & Radcliffe 1917) | Minor concern | EP | • | 19.30 (5,30) | |||
Scarus compressus (Osburn & Nichols 1916) | Minor concern | EP | 32.00 (10,60) | |||||
Scarus ghobban Forsskål 1775 | Minor concern | IP+ EP | 27.25 (8,40) | |||||
Scorpaenidae | Scorpaena mystes Jordan & Starks 1895 | Minor concern | EP | 22.28 (3,40) |
The Red List of the IUCN classified six of the species reported in this study (a sea cucumber, two rays, and three teleosts) in some risk category. The shovelnose guitarfish (Pseudobatos productus) is classified as Near Threatened, whereas the ocellated electric ray (Diplobatis ommata), clarion angelfish (Holcanthus clarionensis), and California sheephead (Semicossyphus pulcher) are classified as Vulnerable. The brown sea cucumber (Isostichopus fuscus) and gulf grouper (Mycteroperca jordani) are listed as Endangered (
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 (Pinctada mazatlanica), spiny oyster (Spondylus limbatus), clarion angelfish (H. clarionensis), king angelfish (Holacanthus passer), Cortez angelfish (Pomacanthus zonipectus), and Limbaugh’s damselfish (Chromis limbaughi), while the brown sea cucumber (I. fuscus) is listed as endangered.
The abundance of ETP species varied within the ISPMBR during the study period; for example, H. clarionensis were only recorded in 2009, and P. productus and D. ommata were recorded in two years (Table
Average abundance (standard deviation) in 60 m2 of endangered, threatened, and protected eastern tropical Pacific species in the Island San Pedro Mártir Biosphere Reserve (ISPMBR) in the Gulf of California, Mexico.
Invertebrates | 2007 | 2008 | 2009 | 2010 | 2011 | 2012 | 2013 | 2014 | 2015 | 2016 | 2017 |
n = 30 | n = 72 | n = 67 | n = 72 | n =67 | n =74 | n = 72 | n = 79 | n = 53 | n = 72 | n = 72 | |
Pinctada mazatlanica | 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) | |
Spondylus limbatus | 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) | |
Isostichopus fuscus | 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) | |
Fishes | 2007 | 2008 | 2009 | 2010 | 2011 | 2012 | 2013 | 2014 | 2015 | 2016 | 2017 |
n = 76 | n = 143 | n = 144 | n = 144 | n =141 | n =135 | n = 144 | n = 132 | n = 109 | n = 144 | n = 150 | |
Diplobatis ommata | 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) | |
Pseudobatos productus | 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) | |
Mycteroperca jordani | 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) | |
Chromis limbaughi | 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) | |
Holacanthus clarionensis | 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) | |
Holacanthus passer | 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) | |
Pomacanthus zonipectus | 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) | |
Semicossyphus pulcher | 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 ISPMBR, 45.71% are distributed in the eastern Pacific, 8.57% are distributed in the Eastern Indo-Pacific, and 5.71% are distributed in the southeastern Pacific and eastern central Pacific. Finally, no information was found on the distributions of 17.14% of the invertebrate species registered in this study.
Of the 73 registered fish species, four species are solely distributed in Mexico (Pareques fuscovittatus, Girella simplicidens, C. limbaughi, and Stegastes rectifraenum), and two of these (C. limbaughi and S. rectifraenum) are only distributed in the Gulf of California. Most of the fish species (64.38%) are widely distributed in the eastern Pacific, and 27.20% are species with a biogeographic affinity for the central eastern Pacific. A total of 2.74% of the fish species are distributed in the Indo-Pacific, and 1.37% of the fish species showed circumglobal and circumtropical distributions, respectively.
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.
Of the 108 species observed in the ISPMBR, the most abundant invertebrate species were the state pencil urchin (Eucidaris thouarsii), blue sea star (Phataria unifascialis), and yellow spotted star (Pharia pyramidata; Suppl. material
A total of 54 commercially important species (12 invertebrate and 42 fish species) for the communities of the MIR were recorded (Table
Average size in centimeters (standard deviation) of the main commercially important fish species in the Island San Pedro Mártir Biosphere Reserve (ISPMBR) in the Gulf of California, Mexico. Abbreviations: ND = Not enough data.
Species | 2007 | 2008 | 2009 | 2010 | 2011 | 2012 | 2013 | 2014 | 2015 | 2016 | 2017 |
---|---|---|---|---|---|---|---|---|---|---|---|
n = 76 | n = 143 | n = 144 | n = 144 | n = 141 | n = 135 | n = 144 | n = 132 | n = 109 | n = 144 | n = 150 | |
Balistes polylepis | 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) | |
Mycteroperca jordani | 120.00 | 84.29 | 85.07 | 94.62 | 75.00 | 82.50 | 120.00 | 22.50 | 47.50 | 0.00 | 90.00 |
(ND) | (49.95) | (24.28) | (21.06) | (37.28) | (45.00) | (ND) | (5.00) | (31.82) | (ND) | (26.46) | |
Mycteroperca prionura | 0.00 | 38.33 | 31.32 | 22.50 | 25.00 | 25.00 | 30.00 | 32.50 | 32.50 | 40.00 | 45.00 |
(ND) | (11.69) | (2.50) | (10.61) | (ND) | (13.23) | (ND) | (2.89) | (3.54) | (ND) | (ND) | |
Mycteroperca rosacea | 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) | |
Lutjanus argentiventris | 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) | (ND) |
In the ISPMBR, the average invertebrate richness was 5.50 ± 2.02 species/transect (mean ± SD). The year with the highest richness was 2008 (7.19 ± 1.93 species/transect), while the year with the lowest richness was 2012 (4.12 ± 1.50 species/transect; Fig.
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 | |
---|---|---|---|---|---|---|
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 |
Fishes | Change in S | Curve change | Change in evenness | Species loss | Species gains | |
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 |
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 ISPMBR, a total of 11 protected species (i.e., three invertebrate and eight fish species) were identified, which constitute 10% of all species surveyed in this study. The clarion angelfish (H. clarionensis) was the only species recorded in only one year (i.e., 2009). The presence of tropical species has been recorded in several studies of the Gulf of California (
Two species (S. limbatus and I. fuscus) showed decreases in abundance over the course of this study, which may have been due to commercial harvest (
The rocky reefs of the ISPMBR are essential for fishing activities. In this study, 64% of the species recorded were of commercial importance in the region. Monitoring the abundances and average sizes of these species is crucial to ensuring the sustainable use and conservation of this marine protected area (
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 MIR are scarce.
The most abundant invertebrate species in this study (E. thouarsi, P. unifascialis, and P. pyramidata) are also species that have been reported to be abundant in the Gulf of California (Luna-Salguero and Reyes-Bonilla 2010;
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 ISPMBR may be explained by the presence of low-level anthropogenic stressors and the resilience of the invertebrate and fish communities. Although this island is located in an area with abiotic stressors of medium to high intensity, the complexity of the habitats and substrates present (e.g., shallow and deep rocky reefs, mangroves, walls, and brown algae) provides protection for the different species and life stages present (
The results of this study constitute the first analysis of the community structure of the ISPMBR, with emphasis on the distribution, conservation, and use of the invertebrates and fish species present in the shallow rocky reefs of the natural protected area. A taxonomic list based on an 11-year data set is also presented.
In this study, we analyzed eleven years of survey data of the most oceanic marine protected area in the Gulf of California: the ISPMBR. We observed that invertebrate and fish fauna in the rocky coastal reefs present stability with regard to the ecological indicators considered in this study. The marine protected area is both ecologically and commercially important. A total of 108 species were recorded (35 invertebrates and 73 fishes), of which 54 are commercially important (12 invertebrate and 42 fish species). Two principal trends were observed. First, the abundance of commercially important invertebrate species is decreasing, which is probably due to their high monetary value and illegal fishing (e.g., the sea cucumber I. foscus). Second, commercially important fish species maintained their abundance overall, albeit with periods of increase. The use of long-term monitoring data can provide a more realistic picture of the dynamics inside a marine protected area, which may then be used to evaluate its performance.
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
Data type: Xlsx file.
Explanation note: Average (standard deviation) for ecological indicators of invertebrates and fishes, species richness (S), Shannon-Wiener diversity (H’), and Pielou evenness (J’).
Table S2
Data type: Xlsx file.
Explanation note: Total abundance of invertebrates. Abbreviations: n = number of transects.
Table S3
Data type: Xlsx file.
Explanation note: Total abundance of fishes. Abbreviations: n = number of transects.
Table S4
Data type: Xlsx file.
Explanation note: Results of the post-hoc analysis for ecological indicators of invertebrates and fishes, species richness (S), Shannon-Wiener diversity (H’), and Pielou evenness (J’).