Pacific Science (1991), vol. 45, no. 1: 63-71 © 1991 by University of Hawaii Press. All rights reserved Biogeographic Relationships of a Rocky Intertidal Fish Assemblage in an Area of Cold Water Upwelling off Baja California, Mexico! CAROL A. STEPIEN, HIKARU PHILLIPS, JOSEPH A. ADLER, AND PETER J. MANGOLD 2 ABSTRACT: The rocky intertidal fish assemblage at an area of nearshore cold water upwelling at Punta Clara, northern Baja California, Mexico was sampled bimonthly for I yr. Temperatures in this upwelling region typically range from 10° to 16°C throughout the year and are significantly lower than those of surrounding areas in the warm-temperate Californian biogeographic province. The assemblage at Punta Clara is a species-rich mixture composed ofeight fishes that are primarily Californian in distribution, seven that are primarily Oregonian cold-temperate, and four that range throughout both provinces. In terms of relative numbers, 53% of the total number of fishes are Californian, 33% are Oregonian, and 14% belong to both provinces . In terms of biomass, 75% are Californian, 20% are Oregonian, and 5% belong to both provinces. Two com­ mon intertidal fishes characteristic of the Californian province (and belonging to the largely tropical and subtropical families Blenniidae and Labrisomidae) are absent, as are members of the Stichaeidae, which are characteristic of the Oregonian intertidal. Populations ofOregonian fishes in these upwelling regions off Baja California may be Pleistocene relicts maintained by cold temperatures. Alternatively, allozyme studies of two of these species suggest considerable gene flow between northern and Baja Californian populations that could be maintained by larval transport in coastal currents, such as the California Current. POINT CONCEPTION, CALIFORNIA (34S N) 14°C, and those in the Oregonian province forms a boundary or transitional zone be­ range from 9° to l2°C (see Table 1; Hubbs tween the warm-temperate Californian bio­ 1948, 1960, 1961, 1962, 1963). geographic province to the south (a region Point Conception is the southern limit of that extends to ca. 25° N, Baja California, the ranges of many species, including fishes, Mexico) and the cold-temperate Oregonian of the Oregonian province (Horn and Allen province to the north (Garth 1955, Valentine 1978). However, several nearshore fishes 1966, Brusca and Wallerstein 1979). Near­ (Hubbs 1948, 1960), invertebrates (Garth shore surface waters in the Californian prov­ 1955), and algae (Dawson 1960) that are dis­ ince typically have temperatures ranging from tributed primarily in the Oregonian province 13° to 21°C throughout the year, whereas reappear in localized nearshore areas of cold those at Point Conception range from 11.5° to water upwelling off northern Baja California, Mexico (Hubbs 1948, 1960).These Oregonian species largely skip the warmer waters of I This study was supported, in part, by N.S.F. Grant southern California and coexist with many of #BSR·8600180 (Biotic Systems and Resources) and a Sloan Postdoctoral Fellowship to C. A. Stepien. It was the Californian biota in these upwelling re­ also supported by grants from the University of Cali­ gions (Hubbs 1948, 1960, Dawson 1960, fornia U.S. Grants program for undergraduate research Briggs 1974). The upwelling areas have sur­ to J. A. Adler (no. 1988-1 and no. 1989-36) and P. J. face temperatures 3° to 9°C lower than those Mangold (no. 1989-27). Manuscript accepted 10 April of immediately adjacent areas characteristic 1990. of the Californian warm-temperate biogeo­ 2 Scripps Institution of Oceanography, A-002, Uni­ versity of California , San Diego, La Jolla, California graphic province (Hubbs 1948, 1960; see 92093. Table 1). The unique mixture ofbiota in these 63 64 PACIFIC SCIENCE, Volume 45 , J anuary 1991 TABLE 1 T EMPERATURE DATA FOR PUNTA C LARA AND COMPARISON AREAS* JAN. FEB. MAR. APR. MAY JUNE JULY AUG. SEPT. OCT. NOV. DEC. MEAN Punta Clara, Baja California, Mexi co; 310 31.9' N, 116039.7' W 1960 xxxx** 11.4 12.2 12.9 12.6 xxxx 15.6 15.0 15.2 14.6 16.8 16.6 14.3 1961 14.4 13.4 13.5 13.8*** 15.9 13.6 14.7 14.7*** 14.9 xxxx xxxx xxxx 14.3 1962 14.6 12.7 12.0 xxxx 13.4*** xxxx 17.0 14.6 15.6 16.9 12.6 14.4 14.4 1963 13.1 12.2 12.8 xxxx 12.5 14.5 14.9 16.5*** 16.7 xxxx 14.4 xxxx 14.2 Mean 14.0 12.4 12.6 13.3 13.6 14.0 15.6 15.2 15.6 15.8 14.6 15.5 14.35 La Jolla, Scripps Pier, California; 320 52.0' N, 1170 15.3' W 1961 13.8 12.6 16.2 xxxx 18.9 17.5 17.7 18.9 18.7 xxxx xxxx xxxx 16.8 1962 13.4 14.7 15.5 xxxx 17.6 xxxx 20.0 20.8 18.5 18.7 14.7 14.3 16.8 1963 13.2 12.5 15.2 xxxx 15.7 17.2 13.6 21.4 17.7 xxxx 14.8 xxxx 15.7 Mean 13.5 13.3 15.6 xxxx 17.4 17.4 17.1 20.4 18.3 18.7 14.8 14.3 16.44 Pt. Piedras Blancas, Eastern Point, California; 35 0 39.9' N, 1210 15.6' W 1961 12.1 12.2 12.2 11.4 12.5 14.2 13.2 14.7 14.8 13.0 13.9 13.0 13.1 1962 14.0 12.1 11.5 12.0 11.8 12.1 14.4 14.5 14.3 14.0 11.7 12.8 12.9 1963 11.3 12.5 11.7 12.2 11.6 12.6 13.8 13.2 14.3 14.1 12.5 13.3 12.8 Mean 12.5 12.3 11.8 11.9 12.0 13.0 13.8 14.1 14.5 13.7 12.7 13.0 12.94 Mendocino, Ca lifornia; 39 0 18.2' N, 123048.2' W 1960 10.9 10.6 9.7 10.1 10.0 9.9 10.1 10.0 10.7 11.6 11.2 11.6 10.5 1961 11.7 11.2 10.3 9.4 9.8 10.5 10.4 11.6 11.4 11.8 10.8 11.0 10.8 1962 10.4 10.8 10.1 9.1 9.0 9.0 10.2 11.1 11.0 11.8 11.2 11.1 10.4 Mean 11.0 10.9 10.0 9.6 9.6 9.8 10.2 10.9 11.0 11.7 11.1 11.2 10.58 •Based on Hubbs 1960, 1961, 1962, 1963. •• xxxx = missing data. ••• Multipl e readings for that month . upwelling areas and their disjunct distribution the water's edge, having abundant algae, and patterns have been little studied to date. ranging to 1.5 m Mean Lower Low Water In the present study, we examined the inter­ (MLLW) in maximum depth using the anes­ tidal fish assemblage at an upwelling site thetic quinaldine sulphate. Collections began off Punta Clara, Baja California, Mexico ca. 45 min before the lowest point of the tide (31° 31.9' N, 116° 39.7' W; 55 km SW of and extended to I hr after. All fishes seen were Ensenada, Mexico; Figure I) to provide base­ netted by three to five collectors (including line data for analyzing these patterns of dis­ some with snorkel gear) and immediately fro­ tribution. This site was the location ofa long­ zen on dry ice for later laboratory analysis . term study of temperature by Hubbs (1960;see Sample dates and tide levels were as follows: Table 1). In the present study, we compared 23 October 1988 (-0.5 m MLLW), 22 De­ results from I yr of bimonthly sampling with cember 1988 (-1.5 m MLLW), 17 February data from intertidal fish assemblages from the 1989 (-1.0 m MLLW), 9 April 1989 (0.0 m Oregonian and Californian biogeographic MLLW), 6 June 1989 (-1.2 m MLLW), and provinces. It is part of a larger investigation 19 August 1989 (-0.3 m MLLW). Tempera­ of gene flow between these disjunct popula­ tures at the water's edge (not in isolated tide tions of nearshore fishes. pools) ranged from IOSC (February 1989)to 14SC (December 1988). Total lengths (TL) of fishes to the nearest millimeter and weights to the nearest 0.5 g MATERIALS AND METHODS were measured as soon as possible after collec­ Fishes were collected bimonthly during tion. Determination ofprimary biogeographic minus tides for 1 yr (October 1988 through ranges of the intertidal fishes was based on September 1989) from tide pools located at data from Hubbs (1948), Williams (1957), Rocky Intertidal Fishes of Baja-STEPIEN ET AL. 65 11 5· intertidal fishes were (in decreasing order of relative abundance): spotted kelpfish, Gib­ bonsia elegans (C1inidae; 22.5% of the total number collected); woolly sculpin, Clinocottus analis (Cottidae; 19.8% of the total); crevice \. kelpfish, Gibbonsia montereyensis (Clinidae; 12.4%); rosy sculpin, Oligocottus rubellio (Cottidae; 10.1%); striped kelpfish, Gibbonsia punta Clara metzi (Clinidae; 7.9%); and opaleye, Girella nigricans (Girellidae; 7.9%) (Figure 2). Thir­ teen fishes were less common, including (in 30· decreasing order of relative abundance): the rockweed gunnel , Xererpes fucorum (Pholi­ , didae); smoothhead sculpin, Artedius lateralis (Cottidae); fluffy sculpin, Oligocottus synderi (Cottidae); reef surfperch, Micrometrus aurora (Embiotocidae) ; spotted snailfish, Liparis florae (Liparididae); California clingfish, Go­ biesox rhessodon (Gobiesocidae); coralline sculpin, Artedius corallinus (Cottidae); dwarf surfperch, Micrometrus minimus (Embioto­ cidae); cabezon, Scorpaenichthys marmoratus F IGURE 1.