Checklist of Conspicuous Reef Fishes ... by Gulf of California Marine Program

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‐OSORIO, BRAD ERISMAN, JERRY MOXLEY, EDUARDO BALART AND OCTAVIO ABURTO‐OROPEZA

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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: otolino@gmail.com Marine Biology Research Division, Scripps Institution of Oceanography, University of California San Diego, La Jolla CA 920930202, USA. E-mail: berisman@ucsd.edu; maburto@ucsd.edu 3 Nicholas School of the Environment, Duke University, North Carolina, USA. E-mail: jerry.moxley@duke.edu 4 Centro de Investigaciones Biológicas del Noroeste, La Paz, Baja California Sur, Mexico. E-mail: ebalart04@cibnor.mx 5 Corresponding author 2

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 California 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 production (Alvarez-Borrego 2008, 2010). Consequently, the area supports some of the most productive fisheries in Mexico 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 Ballenas 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-

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

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 methodology 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 sedentary 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 reproduction (Thomson et al. 2000, Aburto-Oropeza & Balart 2001). We also considered as reef fishes the epipelagic species 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 frequency 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 families 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 Epinephelidae as distinct from Serranidae and the placement of the genus Paranthias within Cephalopholis. Broadscale 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 (eastern Pacific from California to southern Mexico); Tropical Eastern Pacific (eastern Pacific from central Baja California 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 rectifraenum, 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 studies that utilized underwater visual censuses to study reef fish assemblages in the Gulf of California (Aburto-Oropeza & 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 northern 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.

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

Scorpaena guttata Girard, 1854

NEP

Scorpaena histrio Jenyns, 1840

Scorpaena mystes Jordan & Starks, 1895

Alphestes immaculatus Breder, 1936

TEP

Cephalopholis colonus (Valenciennes, 1846)

TEP

Cephalopholis panamensis (Steindachner, 1877)

TEP

Epinephelus labriformis (Jenyns, 1840)

Mycteroperca jordani (Jenkins & Evermann, 1889)

NEP

Mycteroperca prionura Rosenblatt & Zahuranec, 1967

GOC

Mycteroperca rosacea (Streets, 1877)

GOC

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

Elagatis bipinnulata (Quoy & Gaimard, 1825)

Seriola lalandi Valenciennes, 1833

Seriola rivoliana Valenciennes, 1833

Lutjanidae

Lutjanus argentiventris (Peters, 1869)

Haemulidae

Anisotremus davidsonii (Steindachner, 1876)

NEP

Anisotremus interruptus (Gill, 1862)

TEP

Haemulon flaviguttatum Gill, 1862

Haemulon maculicauda (Gill, 1862)

TEP

Haemulon sexfasciatum Gill, 1862

TEP

Haemulon steindachneri (Jordan & Gilbert, 1882)

EP + ATL

Scorpaenidae

Epinephelidae

Serranidae

continued next page

REEF FISHES OF THE BAHÍA DE LOS ÁNGELES REGION

TABLE 1. (continued) Family

Species

Distribution

Microlepidotus brevipinnis (Steindachner, 1869)

TEP

Microlepidotus inornatus (Gill, 1862)

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)

Kyphosus elegans (Peters, 1869)

TEP

Chaetodontidae

Johnrandallia nigrirostris (Gill, 1862)

TEP

Pomacanthidae

Holacanthus passer Valenciennes, 1846

TEP

Pomacanthus zonipectus (Gill, 1862)

TEP

Abudefduf troschelii (Gill, 1862)

Chromis atrilobata Gill, 1862

TEP

Chromis limbaughi Greenfield & Woods, 1980

GOC

Stegastes rectifraenum (Gill, 1862)

MEX

Bodianus diplotaenia (Gill, 1862)

Halichoeres chierchiae di Caporiacco, 1947

TEP

Halichoeres dispilus

TEP

Halichoeres nicholsi (Jordan & Gilbert, 1882)

TEP

Halichoeres notospilus

TEP

Halichoeres semicinctus (Ayres, 1859)

NEP

Semicossyphus pulcher (Ayres, 1854)

NEP

Nicholsina denticulata (Evermann & Radcliffe, 1917)

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)

Sphoeroides lobatus (Steindachner, 1870)

TEP

Diodon holocanthus Linnaeus, 1758

Diodon hystrix Linnaeus, 1758

Pomacentridae

Labridae

Scaridae

Diodontidae

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 polylepis, Scarus ghobban) or the Atlantic Ocean (Haemulon steindachneri).

MASCARENAS-OSORIO ET AL.

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

Abundance (# inds)

Relative Abun- Frequency (%) dance (%)

Mean Density (#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

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

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

0.25

27.37

0.002

<0.001

Serranus psittacinus

0.25

18.95

0.002

<0.001

Labrisomus xanti

0.25

8.95

0.002

0.001

Hermosilla azurea

0.20

6.32

0.002

0.001

Urobatis halleri

0.16

24.21

0.001

<0.001

Haemulon maculicauda

0.16

4.21

0.001

Anisotremus interruptus

0.15

19.47

0.001

<0.001

Balistes polylepis

0.15

19.47

0.001

<0.001

Halichoeres dispilus

0.11

4.21

0.001

<0.001

Halichoeres notospilus

0.09

2.63

0.001

Kyphosus elegans

0.09

2.11

0.001

Sphoeroides lobatus

0.09

5.26

0.001

<0.001

Halichoeres chierchiae

0.08

5.26

0.001

<0.001

Sphyraena lucasana

0.08

1.05

0.001

<0.001

Apogon spp.

0.07

0.53

0.001

Johnrandallia nigrirostris

0.07

0.001

<0.001

Semicossyphus pulcher

0.05

6.84

<0.001

<0.001 continued next page

REEF FISHES OF THE BAHÍA DE LOS ÁNGELES REGION

TABLE 2. (continued) Species

Abundance (# inds)

Relative Abun- Frequency (%) dance (%)

Mean Density (#inds/m2)

Caranx caballus

0.05

1.58

<0.001

Seriola lalandi

0.05

3.16

<0.001

Mulloidichthys dentatus

0.03

1.58

<0.001

Nicholsina denticulata

0.03

4.21

<0.001

Mycteroperca jordani

0.03

3.68

<0.001

Opistognathus rosenblatti

0.02

1.58

<0.001

Scorpaena plumieri mystes

0.02

4.74

<0.001

Haemulon sexfasciatum

0.02

2.11

<0.001

Lutjanus argentiventris

0.02

2.63

<0.001

Mycteroperca prionura

0.02

2.63

<0.001

Alphestes immaculatus

0.01

2.11

<0.001

Epinephelus labriformis

0.01

2.11

<0.001

Urobatis maculatus

0.01

2.11

<0.001

Dasyatis spp.

0.01

2.11

<0.001

Gymnothorax castaneus

0.01

2.11

<0.001

Seriola rivoliana

0.01

0.53

<0.001

Urobatis concentricus

0.01

2.11

<0.001

Elagatis bipinnulata

0.01

0.53

<0.001

Myliobatis californica

0.01

1.58

<0.001

Ophioblennius steindachneri

0.01

1.05

<0.001

Diplobatis ommata

0.01

0.53

<0.001

Muraena argus

0.01

1.05

<0.001

Scorpaena histrio

0.01

1.05

<0.001

Diodon holocanthus

<0.01

0.53

<0.001

Diodon hystrix

<0.01

0.53

<0.001

Heterodontus mexicanus

<0.01

0.53

<0.001

Microlepidotus brevipinnis

<0.01

0.53

<0.001

Muraena lentiginosa

<0.01

0.53

<0.001

Scarus ghobban

<0.01

0.53

<0.001

Scorpaena guttata

<0.01

0.53

<0.001

Zapterix exasperata

<0.01

0.53

<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 (Stegastes rectifraenum) and the Gulf of California (Chromis limbaughi, Girella simplicidens) ranked highest in relative 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 California, 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 路 Zootaxa 2922 漏 2011 Magnolia Press

MASCARENAS-OSORIO ET AL.

uous adult distributions along cold-water isobaths (Bernardi et al. 2003, Hastings et al. 2010). Conversely, circumtropical 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 largebodied 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 during 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)

Eastern Pacific & Indo-Pacific (EP + IP)

Eastern Pacific & Atlantic (EP + ATL)

Eastern Pacific (EP)

Northeast Pacific (NEP)

Tropical Eastern Pacific (TEP)

Mexico (MEX)

Gulf of California (GOC)

Total

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. Additional 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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