Zootaxa 2922: 60–68 (2011) ISSN 1175-5326 (print edition)

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Copyright © 2011 · Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)

Checklist of conspicuous reef fishes of the Bahía de los Ángeles region,

Baja California Norte, Mexico, with comments on abundance
and ecological biogeography

ISMAEL MASCAREÑAS-OSORIO1, BRAD ERISMAN2,5, JERRY MOXLEY3,

EDUARDO BALART4 & OCTAVIO ABURTO-OROPEZA2
1
Centro para la Biodiversidad Marina y la Conservación, La Paz, Baja California Sur, Mexico. E-mail: [email protected]
2
Marine Biology Research Division, Scripps Institution of Oceanography, University of California San Diego, La Jolla CA 92093-
0202, USA. E-mail: [email protected]; [email protected]
3
Nicholas School of the Environment, Duke University, North Carolina, USA. E-mail: [email protected]
4
Centro de Investigaciones Biológicas del Noroeste, La Paz, Baja California Sur, Mexico. E-mail: [email protected]
5
Corresponding author

Abstract

A first checklist of conspicuous reef fishes observed at 15 sites in the vicinity of Bahía de los Ángeles from 2008 to 2010
is presented. A total of 70 species representing 31 families were observed. Species composition was similar to well studied
regions in the southern Gulf of California, in that most species had distributions that span the Tropical Eastern Pacific but
species endemic to Mexico or the Gulf of California ranked highest in relative abundance, frequency of occurrence, and
mean density. Several species with temperate geographic distributions were more abundant and frequent than on reefs in
the southern Gulf. Large-bodied, predatory species such as sharks and the Gulf Grouper, Mycteroperca jordani, were rare
or absent.

Key words: Reef fishes, Bahía de los Ángeles, Gulf of California, Tropical Eastern Pacific

Introduction

Bahía de los Ángeles (BLA) is a coastal bay in the Gulf of California, located on the eastern shore of the Baja Cal-
ifornia peninsula in the state of Baja California Norte, Mexico (28°55´N; 113°32´ W) (Fig. 1). The waters around
BLA include an archipelago of 17 islands that comprise a portion of the Midriff Islands and host large stretches of
rocky reef habitats (Danemann & Ezcurra 2008). These reef communities are associated with the northern Gulf of
California bioregion, where cold-water species intermix with tropical fauna typical of the warmer, southern Gulf
(Brusca et al. 2005, Hastings et al. 2010). The unique physical and biological oceanographic conditions of BLA
make it one of the most ecologically important areas in the Gulf. Complex topography and strong tidal and wind
forcing move large masses of water that create strong upwelling and high levels of primary and secondary produc-
tion (Alvarez-Borrego 2008, 2010). Consequently, the area supports some of the most productive fisheries in Mex-
ico as well as high densities of turtles, marine mammals, and seabirds (Tershy et al. 1991, Lluch-Cota et al. 2007;
Brusca 2010).
Bahía de los Ángeles is largely recognized as a priority area for the management and conservation of marine
biodiversity (Sala et al. 2002, Enríquez-Andrade et al. 2004; Brusca 2010), which was made official in 2007 by its
inclusion within a new biosphere reserve known as Reserva de la Biosfera de Bahía de los Ángeles, Canal de Bal-
lenas y Salsipuedes (DOF 2007). Despite its designation as a biosphere reserve and its economic importance with
respect to fisheries production, detailed information on the community structure and species composition of reef
fishes of BLA is scarce (but see Viesca-Lobaton et al. 2008). The present study provides the first comprehensive
checklist on conspicuous fishes that occur on shallow, rocky reefs in BLA. In addition, patterns of taxonomic rep-

60 Accepted by M. Craig: 9 May 2011; published: 17 Jun. 2011

resentation, geographic distributions, species density and abundance are commented on and compared to reefs in
the southern Gulf of California.

FIGURE 1. Map of Bahía de los Ángeles, Baja California, Mexico showing the location of the 15 surveys sites and the
region’s location within Northwest Mexico.

REEF FISHES OF THE BAHÍA DE LOS ÁNGELES REGION Zootaxa 2922 © 2011 Magnolia Press · 61
Material and methods

Underwater visual censuses using SCUBA were conducted at 15 sites each during three sampling periods (26 June
– 4 July 2008, 8–10 July 2009, 18–20 July 2010) (Figure 1). Censuses followed standard band transect methodol-
ogy described in previous studies (Stephens & Zerba 1981, Aburto-Oropeza & Balart 2001, Froeschke et al. 2005).
Each transect covered an area of 250 m2 (50 m x 5 m) conducted by a single diver swimming along an isobath of
the reef. Two transects each were performed at two isobaths (5–6 m, 15–20 m) at each site during each sampling
period (n =180 transects). For each transect, actively swimming species were recorded on the first pass, and seden-
tary or benthic, territorial species were recorded on the second pass (Aburto-Oropeza & Balart 2001, Sala et al.
2002). All fishes that passed divers from behind were omitted to avoid counting the same fish multiple times or
overestimating fish attracted to divers (Stephens & Zerba 1981, Froeschke et al. 2005). The censuses focused on
reef fishes, which are defined as fish species that use the hard substrata for protection, shelter, feeding, or reproduc-
tion (Thomson et al. 2000, Aburto-Oropeza & Balart 2001). We also considered as reef fishes the epipelagic spe-
cies that regularly visit reefs in search of food, cleaning services, and reproduction. We focused on conspicuous
species rather than cryptobenthic species, or small fishes (< ca. 5 cm in total length) that are behaviorally cryptic
and are difficult to quantify by visual surveys due to their close association with the substratum (Allen et al. 1992).
Mean density (# individuals / m2), relative abundance (# individuals of one species / total # individuals), and fre-
quency of occurrence (# transects observed / total # transects) were calculated for each species. Data were pooled
among survey periods and among depths for all such analyses.
Species nomenclature and family designations followed Eschmeyer & Fricke (2011), and sequencing of fami-
lies in Table 1 followed Nelson (2006). Recent changes in the classification of the Serranidae as proposed by Craig
& Hastings (2007) and Smith & Craig (2007) were adopted. Among these are the resurrection of the family Epi-
nephelidae as distinct from Serranidae and the placement of the genus Paranthias within Cephalopholis. Broad-
scale species distributions were based on Robertson & Allen (2008) and categorized as follows: Circumtropical
(worldwide, tropical distribution); Eastern Pacific (tropical and temperate waters of the eastern Pacific, north and
south of the equator); Eastern Pacific and Atlantic Ocean; Eastern Pacific and Indo-Pacific; Northeast Pacific (east-
ern Pacific from California to southern Mexico); Tropical Eastern Pacific (eastern Pacific from central Baja Cali-
fornia to Peru); Mexico; and the Gulf of California.

Results and discussion

A total of 70 species in 48 genera in 31 families were observed at BLA (Table 1). The most speciose families were
the Haemulidae (8 spp.), Epinephelidae (7 spp.), and the Labridae (7 spp.). Halichoeres (5 spp.), Haemulon (4
spp.), and Scorpaena (3 spp.) were the most species rich genera, and 12 genera were represented by 2 species. We
counted a total of 36,909 individuals on 180 transects, with three species (Chromis limbaughi, Stegastes rectifrae-
num, Girella simplicidens) accounting for 76.12% of all individuals observed (Table 2). S. rectifraenum (92.63%),
G. simplicidens (82.63%), and Bodianus diplotaenia (77.37%) ranked highest in frequency of occurrence, and six
additional species were observed on more than 50% of transects. Chromis limbaughi (0.348 ± 0.056 inds/m2), S.
rectifraenum (0.142 ± 0.012 inds/m2), and G. simplicidens (0.103 ± 0.017 inds/m2) ranked highest in mean density.
Our records of species richness in BLA were comparable to species richness values reported in previous stud-
ies that utilized underwater visual censuses to study reef fish assemblages in the Gulf of California (Aburto-Oro-
peza & Balart 2001, Campos-Davila et al. 2005; Alvarez-Filip et al. 2006). These results, however, are inconsistent
with previous studies by Hastings et al. (2010) and others that demonstrate that species richness in fishes varies
considerably from north to south, with twice the number of species occurring in the southern Gulf as in the north-
ern Gulf. This result is not surprising, as comprehensive estimates of reef fish diversity and community structure
require multiple sampling techniques that target both conspicuous and crypto-benthic species (Allen et al. 1992,
Ackerman & Bellwood 2000). Also, the Midriffs region of the Gulf of California is characterized by large-scale,
seasonal variations in oceanographic conditions, and many cold-water species present in shallow waters during the
winter months migrate south or move to deeper waters during the summer (Brusca et al. 2005, Alvarez-Borrego
2010, Hastings et al. 2010). Therefore, surveys conducted throughout the year are necessary to capture seasonal
dynamics in species composition and relative abundance.

62 · Zootaxa 2922 © 2011 Magnolia Press MASCARENAS-OSORIO ET AL.

TABLE 1. Taxonomic list of conspicuous reef fishes in Bahía de los Ángeles recorded by underwater visual surveys conducted
from 2008 to 2010, showing geographic distribution of each species. CT = Circumtropical; EP = Eastern Pacific; EP + IP =
Eastern Pacific and IndoPacific; EP + ATL = Eastern Pacific and Atlantic Ocean; GOC = Gulf of California; MEX = Mexico;
NEP = Northeast Pacific; TEP = Tropical Eastern Pacific.
Family Species Distribution
Heterodontidae Heterodontus mexicanus Taylor & Castro-Aguirre, 1972 TEP
Narcinidae Diplobatis ommata (Jordan & Gilbert, 1890) TEP
Rhinobatidae Zapteryx exasperata (Jordan & Gilbert, 1880) NEP
Dasyatidae Dasyatis spp. --
Urotrygonidae Urobatis concentricus Osburn & Nichols, 1916 MEX
Urolophidae Urolophus halleri Cooper, 1863 NEP
Urolophus maculatus (Garman, 1913) GOC
Myliobatidae Myliobatis californica Gill, 1865 NEP
Muraenidae Gymnothorax castaneus (Jordan & Gilbert, 1882) TEP
Muraena argus (Steindachner, 1870) TEP
Muraena lentiginosa Jenyns, 1842 TEP
Scorpaenidae Scorpaena guttata Girard, 1854 NEP
Scorpaena histrio Jenyns, 1840 EP
Scorpaena mystes Jordan & Starks, 1895 EP
Epinephelidae Alphestes immaculatus Breder, 1936 TEP
Cephalopholis colonus (Valenciennes, 1846) TEP
Cephalopholis panamensis (Steindachner, 1877) TEP
Epinephelus labriformis (Jenyns, 1840) EP
Mycteroperca jordani (Jenkins & Evermann, 1889) NEP
Mycteroperca prionura Rosenblatt & Zahuranec, 1967 GOC
Mycteroperca rosacea (Streets, 1877) GOC
Serranidae Paralabrax auroguttatus Walford, 1936 NEP
Paralabrax maculatofasciatus (Steindachner, 1868) NEP
Serranus psittacinus Valenciennes, 1846 TEP
Opistognathidae Opistognathus rosenblatti Allen & Robertson, 1991 GOC
Apogonidae Apogon spp. --
Carangidae Caranx caballus Günther, 1868 EP
Elagatis bipinnulata (Quoy & Gaimard, 1825) CT
Seriola lalandi Valenciennes, 1833 CT
Seriola rivoliana Valenciennes, 1833 CT
Lutjanidae Lutjanus argentiventris (Peters, 1869) EP
Haemulidae Anisotremus davidsonii (Steindachner, 1876) NEP
Anisotremus interruptus (Gill, 1862) TEP
Haemulon flaviguttatum Gill, 1862 EP
Haemulon maculicauda (Gill, 1862) TEP
Haemulon sexfasciatum Gill, 1862 TEP
Haemulon steindachneri (Jordan & Gilbert, 1882) EP + ATL
continued next page

REEF FISHES OF THE BAHÍA DE LOS ÁNGELES REGION Zootaxa 2922 © 2011 Magnolia Press · 63
TABLE 1. (continued)
Family Species Distribution
Microlepidotus brevipinnis (Steindachner, 1869) TEP
Microlepidotus inornatus (Gill, 1862) EP
Sparidae Calamus brachysomus (Lockington, 1880) TEP
Sciaenidae Pareques fuscovittatus (Kendall & Radcliffe, 1912) MEX
Mullidae Mulloidichthys dentatus (Gill, 1862) TEP
Kyphosidae Girella simplicidens Osburn & Nichols, 1916 GOC
Hermosilla azurea Jenkins & Evermann, 1889 NEP
Kyphosus analogus (Gill, 1862) EP
Kyphosus elegans (Peters, 1869) TEP
Chaetodontidae Johnrandallia nigrirostris (Gill, 1862) TEP
Pomacanthidae Holacanthus passer Valenciennes, 1846 TEP
Pomacanthus zonipectus (Gill, 1862) TEP
Pomacentridae Abudefduf troschelii (Gill, 1862) EP
Chromis atrilobata Gill, 1862 TEP
Chromis limbaughi Greenfield & Woods, 1980 GOC
Stegastes rectifraenum (Gill, 1862) MEX
Labridae Bodianus diplotaenia (Gill, 1862) EP
Halichoeres chierchiae di Caporiacco, 1947 TEP
Halichoeres dispilus (Günther, 1864) TEP
Halichoeres nicholsi (Jordan & Gilbert, 1882) TEP
Halichoeres notospilus (Günther, 1864) TEP
Halichoeres semicinctus (Ayres, 1859) NEP
Semicossyphus pulcher (Ayres, 1854) NEP
Scaridae Nicholsina denticulata (Evermann & Radcliffe, 1917) EP
Scarus ghobban Forsskål, 1775 EP + IP
Blennidae Ophioblennius steindachneri Jordan & Evermann, 1898 TEP
Labrisomidae Labrisomus xanti Gill, 1860 MEX
Sphyraenidae Sphyraena lucasana Gill, 1863 MEX
Balistidae Balistes polylepis Steindachner, 1876 EP + IP
Tetraodontidae Sphoeroides annulatus (Jenyns, 1842) EP
Sphoeroides lobatus (Steindachner, 1870) TEP
Diodontidae Diodon holocanthus Linnaeus, 1758 CT
Diodon hystrix Linnaeus, 1758 CT

Thirty-eight percent of species have a geographic distribution that spans the Tropical Eastern Pacific (Gulf of
California to Peru), whereas 18% occur throughout the Eastern Pacific (California, USA to Chile) and 16% occur
in the Northeast Pacific (California, USA to Gulf of California; Table 3). Five species are endemic to Mexican
waters, and six species are endemic to the Gulf of California. Only eight species are found outside the eastern
Pacific, five of which are circumtropical, and the remaining three occur in either the Indo-Pacific (Balistes polyle-
pis, Scarus ghobban) or the Atlantic Ocean (Haemulon steindachneri).

64 · Zootaxa 2922 © 2011 Magnolia Press MASCARENAS-OSORIO ET AL.

TABLE 2. List of conspicuous reef fishes observed near Bahía de los Ángeles organized by total abundance, relative abun-
dance, frequency of occurrence, and mean density (± SE).

Species Abundance Relative Abun- Frequency (%) Mean Density SE

(# inds) dance (%) (#inds/m2)
Chromis limbaughi 16536 44.80 51.58 0.348 0.056
Stegastes rectifraenum 6727 18.22 92.63 0.142 0.012
Girella simplicidens 4841 13.12 82.63 0.103 0.017
Chromis atrilobata 1639 4.44 25.26 0.035 0.009
Abudefduf troschelii 958 2.60 41.05 0.021 0.004
Halichoeres semicinctus 947 2.57 50.53 0.02 0.003
Halichoeres nicholsi 614 1.66 70 0.013 0.002
Bodianus diplotaenia 596 1.61 77.37 0.013 0.001
Mycteroperca rosacea 440 1.19 65.79 0.009 0.001
Paralabrax auroguttatus 415 1.14 30 0.009 0.003
Microlepidotus inornatus 365 0.99 14.74 0.008 0.002
Haemulon steindachneri 296 0.80 11.05 0.006 0.003
Holacanthus passer 205 0.56 51.05 0.004 <0.001
Haemulon flaviguttatum 198 0.54 13.68 0.004 0.001
Anisotremus davidsonii 179 0.48 35.26 0.004 0.001
Pomacanthus zonipectus 174 0.47 50.53 0.004 <0.001
Pareques fuscovittatus 150 0.41 20.53 0.003 0.001
Kyphosus analogus 138 0.37 5.26 0.003 0.001
Calamus brachysomus 128 0.35 33.68 0.003 <0.001
Sphoeroides annulatus 116 0.31 24.74 0.002 <0.001
Paralabrax maculatofasciatus 114 0.31 14.74 0.002 0.001
Paranthias colonus 110 0.30 11.05 0.002 0.001
Cephalopholis panamensis 94 0.25 27.37 0.002 <0.001
Serranus psittacinus 93 0.25 18.95 0.002 <0.001
Labrisomus xanti 92 0.25 8.95 0.002 0.001
Hermosilla azurea 75 0.20 6.32 0.002 0.001
Urobatis halleri 60 0.16 24.21 0.001 <0.001
Haemulon maculicauda 59 0.16 4.21 0.001 0.001
Anisotremus interruptus 57 0.15 19.47 0.001 <0.001
Balistes polylepis 57 0.15 19.47 0.001 <0.001
Halichoeres dispilus 39 0.11 4.21 0.001 <0.001
Halichoeres notospilus 34 0.09 2.63 0.001 0.001
Kyphosus elegans 34 0.09 2.11 0.001 0.001
Sphoeroides lobatus 32 0.09 5.26 0.001 <0.001
Halichoeres chierchiae 31 0.08 5.26 0.001 <0.001
Sphyraena lucasana 31 0.08 1.05 0.001 <0.001
Apogon spp. 27 0.07 0.53 0.001 0.001
Johnrandallia nigrirostris 26 0.07 10 0.001 <0.001
Semicossyphus pulcher 20 0.05 6.84 <0.001 <0.001
continued next page

REEF FISHES OF THE BAHÍA DE LOS ÁNGELES REGION Zootaxa 2922 © 2011 Magnolia Press · 65
TABLE 2. (continued)
Species Abundance Relative Abun- Frequency (%) Mean Density SE
(# inds) dance (%) (#inds/m2)
Caranx caballus 19 0.05 1.58 <0.001 <0.001
Seriola lalandi 17 0.05 3.16 <0.001 <0.001
Mulloidichthys dentatus 12 0.03 1.58 <0.001 <0.001
Nicholsina denticulata 11 0.03 4.21 <0.001 <0.001
Mycteroperca jordani 10 0.03 3.68 <0.001 <0.001
Opistognathus rosenblatti 9 0.02 1.58 <0.001 <0.001
Scorpaena plumieri mystes 9 0.02 4.74 <0.001 <0.001
Haemulon sexfasciatum 8 0.02 2.11 <0.001 <0.001
Lutjanus argentiventris 7 0.02 2.63 <0.001 <0.001
Mycteroperca prionura 6 0.02 2.63 <0.001 <0.001
Alphestes immaculatus 5 0.01 2.11 <0.001 <0.001
Epinephelus labriformis 5 0.01 2.11 <0.001 <0.001
Urobatis maculatus 5 0.01 2.11 <0.001 <0.001
Dasyatis spp. 4 0.01 2.11 <0.001 <0.001
Gymnothorax castaneus 4 0.01 2.11 <0.001 <0.001
Seriola rivoliana 4 0.01 0.53 <0.001 <0.001
Urobatis concentricus 4 0.01 2.11 <0.001 <0.001
Elagatis bipinnulata 3 0.01 0.53 <0.001 <0.001
Myliobatis californica 3 0.01 1.58 <0.001 <0.001
Ophioblennius steindachneri 3 0.01 1.05 <0.001 <0.001
Diplobatis ommata 2 0.01 0.53 <0.001 <0.001
Muraena argus 2 0.01 1.05 <0.001 <0.001
Scorpaena histrio 2 0.01 1.05 <0.001 <0.001
Diodon holocanthus 1 <0.01 0.53 <0.001 <0.001
Diodon hystrix 1 <0.01 0.53 <0.001 <0.001
Heterodontus mexicanus 1 <0.01 0.53 <0.001 <0.001
Microlepidotus brevipinnis 1 <0.01 0.53 <0.001 <0.001
Muraena lentiginosa 1 <0.01 0.53 <0.001 <0.001
Scarus ghobban 1 <0.01 0.53 <0.001 <0.001
Scorpaena guttata 1 <0.01 0.53 <0.001 <0.001
Zapterix exasperata 1 <0.01 0.53 <0.001 <0.001

With respect to biogeographic affinity, the reef fish community in BLA is typical of other regions in the Gulf of
California. As first described by Walker (1960), the Gulf is comprised mainly of species of Panamic origin that
have geographic ranges that span the entire Tropical Eastern Pacific. However, species endemic to Mexico (Ste-
gastes rectifraenum) and the Gulf of California (Chromis limbaughi, Girella simplicidens) ranked highest in rela-
tive abundance, frequency of occurrence, and mean density. Thomson & Gilligan (1983) found a similar pattern in
crypto-benthic reef fishes in the Gulf, in which eight of the 13 most abundant species were Gulf endemics. Several
species that are abundant in temperate waters of the northern Pacific coast of Baja California and Southern Califor-
nia, USA (e.g. Anisotremus davidsonii, Halichoeres semicinctus, Hermosilla azurea, Semicossyphus pulcher) are
relatively common at BLA compared to reefs in the southern Gulf (Campos-Davila et al. 2005, Alvarez-Filip et al.
2006). This phenomenon is well documented for the northern Gulf and results from past vicariance events (via
ancient seaways) during the formation of the Gulf, dispersal events during more recent glacial periods, and contin-

66 · Zootaxa 2922 © 2011 Magnolia Press MASCARENAS-OSORIO ET AL.

uous adult distributions along cold-water isobaths (Bernardi et al. 2003, Hastings et al. 2010). Conversely, circum-
tropical and Indo-pacific fauna contributed far fewer species to the reef fish community in BLA than in the
southern Gulf, where they may constitute more than 10% of the observed species community (Aburto-Oropeza &
Balart 2001).
Several large-bodied fishes of current or historical importance to commercial and sport fisheries of the region
were notably rare or absent altogether. Putative juvenile (< 50 cm in length) Mycteroperca jordani were observed at
shallow depths (5–10 m) at several sites, but large adults (> 1 m in length) were not observed. These results are
consistent with data from the commercial fishery that show that the majority of landings for M. jordani in the BLA
are comprised of landings of juveniles (Aburto-Oropeza et al. 2008), and the commercial fishery for this endemic
species has collapsed in BLA and throughout the Gulf of California (Saenz-Arroyo et al. 2005). Species of large-
bodied sharks known to occur on reefs in the northern Gulf of California (e.g. Squatina californica, Carcharhinus
leucas, Sphyrna lewini) and that historically supported commercial fisheries in the region were never observed dur-
ing any dives, which may be a result of overharvesting by commercial fisheries over the past three decades (Sala et
al. 2004).

TABLE 3. Summary of total number of conspicuous reef fishes observed near Bahía de los Ángeles by geographic distribution.

Distribution No. Species Percent

Circumtropical (CT) 5 7
Eastern Pacific & Indo-Pacific (EP + IP) 2 3
Eastern Pacific & Atlantic (EP + ATL) 1 1
Eastern Pacific (EP) 12 18
Northeast Pacific (NEP) 11 16
Tropical Eastern Pacific (TEP) 26 38
Mexico (MEX) 5 7
Gulf of California (GOC) 6 9
Total 68

Acknowledgements

We thank G. Paredes, the crew of the Don Jose, the crew of the Sandman, and Pronatura Noroeste A.C. for their
support with fieldwork. This research was funded by the Walton Family Foundation, the David and Lucile Packard
Foundation, the Marisla Foundation, the International Community Foundation, and an anonymous donor. Addi-
tional support was provided by the Center for Marine Biodiversity and Conservation and the Marine Vertebrate
Collection at Scripps Institution of Oceanography, University of California San Diego.

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