Coastal Management, 32:341–351, 2004 Copyright Taylor & Francis Inc. ISSN: 0892-0753 print / 1521-0421 online DOI: 10.1080/08920750490487188
The Nearshore Fish Assemblage of the Scripps Coastal Reserve, San Diego, California MATTHEW T. CRAIG Marine Biology Research Division Scripps Institution of Oceanography La Jolla, California
F. JOEL FODRIE Integrated Oceanography Division Scripps Institution of Oceanography La Jolla, California
PHILIP A. HASTINGS Marine Biology Research Division Scripps Institution of Oceanography La Jolla, California Marine reserves are quickly becoming a primary tool in the management of coastal resources worldwide. With a growing demand for appropriate management strategies and enforcement of existing regulations, an urgent need has developed to obtain baseline data for regional faunal assemblages. In an attempt to develop a comprehensive list of fishes for one of California’s southernmost marine reserves, nearshore marine species were qualitatively sampled within the Scripps Coastal Reserve (SCR). Overall, 59 species representing 31 families were recorded during the calendar year 2002. The fish assemblage of the SCR was dominated by species typical of soft bottom communities in southern California as well as several pelagic, rocky reef, and intertidal species. The most abundant species was the speckled sanddab, Citharichthys stigmaeus. Many species were represented by juvenile or young-of-the-year age classes, while adults of C. stigmaeus were present in reproductively active stages throughout the study. The numerous habitat types within the Scripps Coastal Reserve support a diverse fish assemblage within a relatively small area and includes essential habitat
Received 15 December 2003; accepted 4 May 2004. We would like to thank B. Clement (SIO), D. Birch (SIO), J. Hyde (SIO/NMFS), H. J. Walker, Jr. (SIO), and A. Pasulka (Arizona State University) for assistance in the field, and H.J. Walker, Jr. and C. Klepadlo, SIO Marine Vertebrates Collection, for their curatorial assistance. Research was funded by a University of California Natural Reserve System Mildred E. Mathias Graduate Student Research Grant, a grant from the PADI Foundation (both to MTC), and California Seagrant (R/CZ-183). We would like to thank L. Levin (SIO), C. Byron, K. Engie, and A. Erzen for critical reviews of this manuscript. Address correspondence to Matthew T. Craig, Marine Biology Research Division, Scripps Institution of Oceanography, 9500 Gilman Dr., Mail Code 0208, La Jolla, California 92093, USA. E-mail:
[email protected]
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Citharichthys stigmaeus, Marine Reserves, MPA, near-shore fishes
Introduction Although scientists, governments, nongovernmental organizations, and in many cases the public recognize that coastal ecosystems and fisheries worldwide are experiencing a period of rapid transition, it has become apparent that much of the information necessary to set in place properly constructed management strategies does not exist. Longterm baseline datasets for most marine ecosystems are rare, and many of those that do exist, such as for invertebrates (Tegner & Dayton, 1981) and fishes (Pondella, Stephens, & Craig 2002), cover only the last quarter century at best, and rarely encompass the ecosystem as a whole. When attempting to implement coastal management strategies we are therefore faced with solving problems that are perhaps decades in the making with little or no data to identify baselines or effectively address natural variation in population and community structure. Nevertheless, resource managers face increasing pressure to act decisively in establishing measures that will maintain ecosystem function and protect essential habitat necessary to enhance the productivity of such systems. At the very least, meeting these goals requires basic community composition data for areas under consideration for protective measures. While many marine ecosystems have faced some degree of exploitation and any data compiled from them reflects a degree of alteration from human impacts, it is nonetheless important to establish current levels of diversity as a starting point for future management plans. Established in 1929 and placed into the University of California Natural Reserve System (NRS) in 1965, the Scripps Coastal Reserve (SCR) encompasses a variety of habitats that include sandy beach, sparse rocky reef, pier pilings, portions of a submarine canyon, and tide pools. Each of these habitats plays an important role in maintaining community diversity of nearshore marine organisms, including fishes, within the SCR. Although the reserve is not a designated no-take zone for fishes (McArdle, 1997), relatively little fishing is present (pers. obs.). This lack of fishing pressure within the SCR may be a result of its nested location within San Diego County’s network of Areas of Special Biological Significance, underwater parks, and ecological reserves (e.g., San Diego/La Jolla Ecological Reserve). Each of these designated areas varies in terms of protective measures for marine organisms, and the public is often not aware of which particular areas prohibit fishing. Alternatively, the limited fishing pressure may be due to the lack of visible habitat often targeted by anglers (e.g., rocky reef and kelp forest). As with most reserves in California, and despite its proximity to the Scripps Institution of Oceanography (SIO), published information pertaining to the fish community within the included area is sparse. While ichthyofaunal data are present for several widely dispersed locations in California (e.g., Quast, 1968; Feder, Limbaugh, & Turner, 1974; Ebeling et al., 1980; Stephens & Zerba, 1981; Stephens et al., 1984; DeMartini & Roberts, 1990; Holbrook et al., 1990; Hobson, 1994; Pondella & Stephens, 1994; Pondella & Allen, 1999; Hobson & Chess, 2001; Paddack & Estes, 2000), and statewide lists of fish records exist (e.g., Miller & Lea, 1972; Eschmeyer & Herald, 1983), no explicit list has been compiled for fish species that occur within the boundaries of this reserve. In order to assess the importance of the SCR as a habitat for marine fishes and to
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provide a framework for future management decisions, we qualitatively sampled the ichthyofauna of the area. Data compiled herein, coupled with existing life-history datasets and similar species lists being generated for nearby marine reserves, will help to evaluate the performance of the marine reserve system in the San Diego region, as well as to provide data that may be used in the design of marine reserves worldwide.
Materials and Methods Given the variety of habitats present within the SCR, multiple, well-established sampling techniques for surveying fish communities were employed on a quarterly or semiannual basis during the calendar year 2002 to census the fish fauna across seasons. Sampling techniques included otter trawls, gill nets, SCUBA-assisted diver surveys, and small ichthyocide collections. Overall, 24 otter trawls were conducted using a trawl with an 8 m head-rope and 1 m × 0.66 m otter boards at a range of depths (5 m, 7 m, and 10 m). Three trawls were conducted once during daylight hours and once after sunset on a quarterly basis. Following the trawl sampling, three experimental 90 m x 2.5 m gill nets with variable mesh size (2.5 cm, 3.8 cm, and 5 cm square) were deployed overnight for periods of 12–16 hours. Gill nets were set perpendicular to the shoreline and were weighted, thus extended from the bottom to the height of the net (2.4 m). Trawl and gill net sampling occurred in February, June, August, and November. A 30m beach seine was deployed twice in February adjacent to the SIO pier, however, heavy surf conditions during later study periods prohibited further use of this method. Tide pool fishes were collected from three isolated tide pools, one in the lower intertidal zone and two in the mid-intertidal zone, in February and June using the ichthyocide quinaldine (2-methylquinone). Three pools 1–3 m in diameter were inoculated with approximately 300 ml of a 1:10 solution of quinaldine to isopropanol, and all fishes were removed from each pool during the two sample periods. Visual fish surveys by SCUBA-assisted divers and above-surface surveys noting the presence of each species were conducted adjacent to the SIO pier on a monthly basis throughout the study period, and once at Dike Rock, the most conspicuous rocky reef structure within the reserve. All fishes were identified as to species and measured to the nearest 5 mm standard length (SL) in the field. The aggregate weight of each species was taken to the nearest 1 g. Fishes that were alive were returned to their habitat to minimize disturbance. Representative specimens were deposited at the SIO Marine Vertebrates Collection.
Results The marine fish assemblage of the SCR was dominated by transient species typical of soft bottom habitats in southern California, with additional representatives of rocky reef and intertidal species. A species list for all fishes recorded in the SCR is presented in Table 1. Overall, 59 species from 31 families were collected. Of these, nearly half (26) were represented by both juvenile and adult specimens (Tables 1 and 2). The otter trawl was the most effective sampling strategy, capturing a majority of the diversity within the reserve (37 species). Table 2 lists the five most abundant trawl species per sample period and the percentage of each species represented by juvenile (i.e., nonreproductive) individuals. Numerically, the fish assemblage of the SCR was dominated by the speckled sanddab, Citharichthys stigmaeus, which was the major component of both the day and night otter trawls. Individuals of C. stigmaeus were present in age classes that ranged from recruits to adults in each sample period (Figure 1), and adult females carrying visibly hydrated eggs were observed throughout the year.
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M. T. Craig et al. Table 1 List of fishes collected in the Scripps Coastal Reserve in 2002
Scientific name Heterodontidae—bullhead sharks Heterdontus francisci* (Girard, 1855) Triakidae—houndsharks Mustelus californicus (Gill, 1864) Mustelus henlei (Gill, 1863) Triakis semifasciata (Girard, 1855) Squatinidae—angel sharks Squatina californica (Ayres, 1859) Rhinobatidae—guitarfishes Platirhinoidis triseriata* (Jordan and Gilbert, 1880) Rhinobatos productus (Ayres, 1854) Zapteryx exasperata (Jordan and Gilbert, 1880) Urolophidae—round stingrays Urobatis halleri (Cooper, 1863) Gymnuridae—butterfly rays Gymnura marmorata* (Cooper, 1864) Myliobatidae—eagle rays Myliobatis californica* (Gill, 1865) Engraulidae—anchovies Anchoa delicatissima (Girard, 1854) Engraulis mordax (Girard, 1854 Clupeidae—herrings Sardinops sagax (Jenyns, 1842) Synodontidae—lizardfishes Synodus lucioceps (Ayres, 1855) Ophidiidae—cusk-eels Chilara taylori (Girard, 1858) Ophidion scrippsae (Hubbs 1916) Batrachoididae—toadfishes Porichthys myriaster* (Hubbs and Schultz, 1839) Porichtys notatus* (Girard, 1854) Atherinidae—silversides Atherinopsis californiensis (Girard, 1854) Sygnathidae—pipefishes Syngnathus spp.* (Linnaeus, 1758) Scorpaenidae—scorpionfishes Scorpaena gutatta (Girard, 1854) Sebastes sp.* (Cuvier, 1829) Cottidae—sculpins Chitonotus pugetensis (Steindachner, 1876) Clinocottus analis (Girard, 1858) Leptocottus armatus (Girard, 1854)
Common name
Habitat
Horn shark
R/S
Gray smoothound Brown smoothound Leopard shark
R/S R/S R/S
Angel shark
S
Thornback
S
Shovelnose guitarfish Banded guitarfish
S S
Round stingray
S
California butterfly ray
S
Bat ray
R/S
Slough anchovy Northern anchovy
P P
Pacific sardine
P
California lizardfish
S
Spotted cusk-eel Basketweave cusk-eel
S S
Specklefin midshipman
S
Plainfin midshipman
S
Jacksmelt
P
Pipefishes
A
California scorpionfish Rockfishes
R/S R/S
Roughback sculpin
I/R
Wooly sculpin Pacific staghorn sculpin
I/R I/R
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Table 1 List of fishes collected in the Scripps Coastal Reserve in 2002 (Continued) Scientific name Serranidae—sea basses Paralabrax clathratus (Girard, 1854) Paralabrax nebulifer* (Girard, 1854) Malacanthidae—tilefishes Caulolatilus princeps* (Jenyns, 1840) Carangidae—jacks Trachurus symmetricus (Ayes, 1855) Haemulidae—grunts Xenistius californiensis (Steindachner, 1876) Kyphosidae—sea chubs Girella nigricans* (Ayres, 1860) Sciaenidae—drums Atractoscion nobilis* (Ayres, 1860) Genyonemus lineatus* (Ayres, 1855) Menticirrhus undulatus* (Girard, 1854) Roncador stearnsi (Steindachner, 1876) Seriphus politus* (Ayres, 1860) Umbrina roncador* (Valenciennes, 1843) Embiotocidae—surfperches Cymatogaster aggregata* (Gibbons, 1854) Embiotica jacksoni (Agassiz, 1853) Hyperprosopon argenteum* (Gibbons, 1854) Micrometrus minimus (Gibbons, 1854) Phanerodon furcatus* (Girard, 1854) Pomacentridae—damselfishes Hypsypops rubicundus (Girard, 1854) Clinidae—clinids Gibbonsia elegans Hubbs 1927 Heterostichus rostratus* (Girard, 1854) Blenniidae—combtooth blennies Hypsoblennius gilberti (Jordan, 1882) Hypsoblennius jenkinsi (Jordan and Evermann, 1896) Sphyraenidae—barracudas Sphyraena argentea (Girard, 1854) Scombridae—mackerals Scomber japonicus (Houttuyn, 1782) Stromateidae—butterfishes Peprilus simillimus (Ayres, 1860) Paralichthyidae—halibut Citharichthys stigmaeus* (Jordan and Gilbert, 1882) Paralichthys californicus* (Ayres, 1859) Xystreurys liolepis (Jordan and Gilbert, 1880)
Common name
Habitat
Kelp bass Barred sand bass
R R/S
Ocean whitefish
P/S
Jack mackerel
P/S
Salema
R/P
Opaleye
R/P
White seabass White croaker California corbina Spotfin croaker Queenfish Yellowfin croaker
R/S/P S S S S S
Shiner perch Black perch Walleye surfperch Dwarf perch White seaperch
R R R R/A R
Garibaldi
R
Spotted kelpfish Giant kelpfish
I/R I/R
Rockpool blenny Mussel blenny
I/R I/R
Pacific barracuda
P
Chub mackerel
P
Pacific pompano
P
Speckled sanddab
S
California halibut Fantail sole
S S
(Table continues next page)
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M. T. Craig et al. Table 1 List of fishes collected in the Scripps Coastal Reserve in 2002 (Continued)
Scientific name
Common name
Pleuronectidae—righteye flounders Hypsopsetta guttulata* (Girard, 1856) Parophrys vetulus (Girard, 1854) Pleuronichthys decurrens* (Jordan and Gilbert, 1881) Pleuronichthys ritteri* (Starks and Morris, 1907) Pleuronichthys verticalis* (Jordan and Gilbert, 1880)
Habitat
Diamond turbot Slender sole Curlfin sole
S S S
Spotted turbot
S
Hornyhead turbot
S
Note. Species are listed in taxonomic order following Nelson (1994), and common name follows scientific name following Robins et al., 1991. Authorship of species follows Eschmeyer, 1998. Species marked with an “*” were represented by juvenile specimens. Habitat indicates typical substrate for each species as follows: R = rocky reef, S = sand, A = reef associated algae, P = pelagic, I = intertidal.
Discussion The ichthyofauna of the SCR is a typical soft bottom community for southern California, and is numerically dominated by the paralichthyid Citharicthys stigmaeus. In addition, several rocky-reef, pelagic, and intertidal species were also present (Table 1). This community differs from the ichthyofaunal community of the nearby San Diego La Jolla Ecological Reserve (SDLJER) that is dominated by rocky reef fishes, with few Table 2 Rank-order of top five species in abundance for trawl fishes in each sample period January–March Species Citharichthys stigmaeus Syngnathus spp. Scorpaena gutatta Paralichthys californicus Platyrhinoidis triseriata
April–June N
% Juvenile
524 20 11 8 7
40.4 10.0 90.9 37.5 100
Species
Citharichthys stigmaeus 723 Phanerodon furcatus 29 Hyperprosopon argenteum 26 Chilara taylori 21 Paralichthys californicus 18
July–September Species Citharichthys stigmaeus Seriphus politus Cymatogaster aggregatta Scorpaena gutatta Hyperprosopon argenteum
N
% Juvenile 18.4 100 100 0.0 50.0
October–December N
% Juvenile
491 78 68 19 18
22.5 74.3 100 47.3 100
Species Citharichthys stigmaeus Seriphus politus Scorpaena gutatta Paralichthys californicus Cymatogaster aggregatta
N 692 28 23 14 13
% Juvenile 44.2 100 52.1 14.2 46.1
N = sample size, and % juvenile indicates the proportion of juvenile specimens collected for each species.
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120 100 80
N=526
N 60 40 20
11 0
90 10 0
80
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Standard Length (mm)
Second Quarter
120 100
N=725
80 N 60 40 20
11 0
90 10 0
80
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Third Quarter
120 100
N=493
80 N 60 40 20
11 0
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Fourth Quarter
120 100
N=493
80 N 60 40 20
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Figure 1. Length frequency histograms for Citharichthyes stigmaeus collected during quarterly otter trawl sampling within the Scripps Coastal Reserve in 2002.
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representatives of soft bottom assemblages (pers. obs.). Given that the SDLJER contains relatively more rocky reef habitat in comparison with the SCR, the difference in the ichthyofaunal assemblage is not surprising. Eigenmann (1892) presented the earliest review of the fishes of San Diego, in which he documented 170 species of fishes either recorded earlier or observed by him in the late 1800s. All species collected in the current study were recorded by Eigenmann (1892) or have since been recorded in the marine waters of San Diego County in one or more subsequent studies on the fish fauna of California (e.g., Miller & Lea, 1972; Love, 1996). The Southern California Bight 1994 Pilot Project (a.k.a “bight wide surveys”), undertaken by the Southern California Coastal Water Research Project (SCCWRP), conducted otter trawl surveys of the benthic community throughout the region (Pt. Conception to the Mexican border). In total, the “bight-wide” surveys collected 87 species of fishes in 34 families (Allen et al., 2001). While many of the species collected in the present study were also taken in the bight-wide survey, more than half (31) were not. The bight-wide survey conducted trawls covering the entire shelf from 10–200 m. In the current study, the deepest trawl conducted was at the 10 m isobath, and not surprisingly many of the species that are not represented in the list of Allen et al. (2001) are shallow water and intertidal species that would not have been sampled by their protocol. Allen et al. (2001) did, however, note that C. stigmaeus numerically dominated shallow water samples, which is consistent with our findings. A striking feature of the otter trawl catch was the presence of juvenile fishes that in many cases outnumbered adult specimens throughout the year. The otter trawl is not significantly biased towards smaller fishes, and thus should not overestimate the ratio of juvenile to adult fishes for most species. Juveniles of several economically valuable species such as the white seabass, Atractoscion nobilis, the California corbina, Menticirrhus undulata, the California halibut, Paralichthys californicus, and the yellowfin croaker, Umbrina roncador, were taken within the reserve. The increased presence of juveniles was accompanied by an increase in algal wrack caught in the trawl surveys, suggesting that many of the fishes were using this habitat as a nursery ground. This phenomenon has been described in detail for A. nobilis (Allen & Franklin, 1988; 1992) and for other inshore fishes (Cross & Allen, 1993). In addition to the presence of juvenile bony fishes, several species of elasmobranch fishes, both adult and juvenile, were recorded in the reserve (Table 1). Many elasmobranchs (sharks and rays) are known to be long-lived fishes whose reproductive output is low in comparison to the more fecund bony fishes and, as such, are particularly vulnerable to overexploitation. Moser (1996) reported on the early life stages of the specklefin sanddab, C. stigmaeus. The larvae of C. stigmaeus undergo transformation at 24–38 mm SL, indicating that the smallest individuals caught in the present surveys (25–40 mm) were recently settled individuals. Moser (1996) also reported that the larvae of C. stigmaeus were collected throughout the year off California, with highest abundance in August–December. Ford (1965) reported that C. stigmaeus spawns from April to September, while Goldberg and Pham (1987) observed the presence of advanced ovarian stages in the gonads from March through October in individuals as small as 59 mm SL. As juveniles were present throughout the study period (Figure 1), our 2002 SCR data reflect a nearly year-round spawning period for C. stigmaeus, similar to that reported in Goldberg and Pham (1987). Kramer (1991) reported extreme interannual variability in the abundance of small size class C. stigmaeus in San Diego. Thus, while it is tempting to infer recruitment patterns from the size distribution data collected here, the single year data was deemed unreliable for this purpose. Miller and Lea (1972) reported a maximum length of 170 mm, indicating that the largest individuals in our survey (120 mm) fell short of the maximum length
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reported for this species. However, females were observed to be carrying visibly hydrated eggs throughout the sample period, indicating that adults in our study were mature and reproductively active. The coastal region of San Diego proximate to the Scripps Institution of Oceanography is a mosaic of marine protected areas, areas of special biological significance (ASBS), and underwater parks. These areas are designated by various state and local agencies, and all vary in the degree of governance and harvest restrictions (McArdle, 1997). Of these, the most well-known and well-protected area is the SDLJER, which prohibits the take of all marine species except the market squid, Loligo opalescens. Few quantitative data are available on the ichthyofauna of this area. Recently, however, a monitoring program was established to census seasonal and long-term trends in the abundance and diversity of conspicuous fishes that will compliment existing data and that gathered in the present study (P. Hastings and P. Dayton, current project). While the SCR is a relatively small marine reserve protecting only about one mile of coastline, a variety of nearshore aquatic habitats are contained within its boundaries. Although this area is dominated by sandy bottom habitat, there is a small rocky reef structure (Dike Rock) that is exposed during extremely low spring tides. Otherwise, the most prominent solid structure in the SCR is the SIO pier that delimits its southernmost boundary (McArdle, 1997). Additionally, a small portion of the Scripps Canyon opens into, and constitutes the deepest part of, the SCR, rapidly dropping to over 100 m. Throughout the area enclosed by the reserve, there is a noted accumulation of algal wrack (e.g., Macrocystis pyrifera, Zostera sp.), likely due to the proximity of the persistent La Jolla kelp bed and the presence of several detrital “pits” formed by predominant current patterns at the mouth of the submarine canyon. It is well established that the locations and boundaries of many marine reserves, particularly within the United States, have been delimited through political processes using such criteria as economics, logistics, or public acceptance (Halpern & Warner, 2003). There is a growing awareness among those who practice coastal management of the need to incorporate biological and ecological interactions into the design and placement of marine reserves. Additionally, management strategies must consider all stakeholder interests, and cannot ignore those groups whose economic livelihoods depend upon areas under consideration for protection. While no-take reserves appeal to conservation-minded policy makers and scientists, other stakeholders such as commercial fishermen contend that such reserves will unnecessarily limit the economic gain from harvesting marine resources. To reach a compromise, many scientists have suggested that networks of small to intermediate size reserves may function to enhance natural stocks of organisms and seed nearby areas through a “spillover” effect (Sala et al., 2002; Halpern & Warner, 2003). Results from this study support the idea that the design of even the smallest reserve should incorporate various habitat types to enhance reserve function. It is well established that increasing the total area of a reserve containing homogenous habitats protects a greater biomass or number of individuals of a few species resulting in an increase of production larval sufficient in number for sustaining fisheries stocks (Halpern & Warner, 2003). However, maximizing the variety of habitats within a proposed reserve boundary will add to the value of any management plan by increasing the overall diversity of species protected. Based upon these findings, the SCR serves as an important coastal habitat for both adult and juvenile life stages for several members of the soft bottom ichthyofaunal community, as well as for several economically and ecologically valuable species typically associated with pelagic, rocky reef, and intertidal habitats. The SCR functions as a valuable resource, enhanced by the variety of habitats and taxonomic assemblages present
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within its boundaries. The SCR also serves as one of the few buffer zones among the network of reserves in Southern California between no-use (i.e., the SDLJER) and highuse coastal regions, adding to its beneficial qualities by decreasing potential edge effects by fishermen. In concert with the several nearby reserves, most notably the SDLJER, the continued management of the SCR should greatly enhance the future existence of a healthy ichthyofaunal community in the San Diego region.
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