The Distribution of Pelagic Juvenile Rockfish of the Genus Sebastes in the Upwelling Region Off Central California Ralph J
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LARSON ET AL.: DISTRIBUTION OF PELAGIC JUVENILE ROCKFISH CalCOFl Rep., Vol. 35, 1994 THE DISTRIBUTION OF PELAGIC JUVENILE ROCKFISH OF THE GENUS SEBASTES IN THE UPWELLING REGION OFF CENTRAL CALIFORNIA RALPH J. LARSON WILLIAM H. LENARZ AND STEPHEN RALSTON Department of Biology National Manne Fishenes Semce San Francisco State University 31 50 Paradlse Dnve San Francisco. California 94132 Tiburon, California 94920 ABSTRACT en la composici6n por especies y tallas. Los juveniles Many species of central California rockfishes conclude peligicos mas pequeiios se encontraron a menudo ale- their pelagic stage during the spring-summer upwelling jados de la costa, en la regibn asociada a1 frente de la sur- period, when advection of surface waters could carry gencia, lo que sugiere que fueron transportados por la them away from nearshore postpelagic habitats. We ex- corriente de surgencia. Sin embargo, pocos juveniles amined the distributions of late-stage pelagic juvenile peligicos fueron encontrados del lado hacia mar aden- rockfish in April and May/June of 1987 and 1988, based tro del frente de la surgencia. Los juveniles peligicos de on midwater-trawl surveys between Point Reyes and mayor tamaiio se encontraron a menudo pr6ximos a la Monterey Bay, California. Distributional patterns were costa, aun en 10s periodos de actividad de la surgencia. complex, and changed rapidly with changing oceano- Lo anterior sugiere que 10s juveniles peligicos cambian graphic conditions and with changes in species and size su comportamiento durante sus fases tardias, lo que les composition. The smallest pelagic juveniles often ap- permite desplazarse hacia la costa, o bien, permanecer peared offshore, in the region of the upwelling front, alli a pesar de la surgencia. Asimismo encontramos evi- which suggested that they had been advected offshore dencia de un proceso mas pasivo de advecci6n y reten- at some time during upwelling. However, few pelagic cibn de juveniles peligicos de todas las tallas, incluyendo juveniles were found offshore of the upwelling front. movimiento hacia la costa durante periodos de aflo- Larger pelagic juveniles were often found close to shore, jamiento de la surgencia. Este aflojamiento podria ser un even when upwelling was active. This suggests that later- aspect0 importante per0 no determinante para el es- stage pelagic juveniles undergo behavioral changes that tablecimiento de 10s juveniles del rocot del hibitat enable them to move toward shore or remain there in peligico a1 post-peligico. En tal caso, la advecci6n du- spite of upwelling. We also found evidence of more pas- rante la surgencia podria no tener un efecto negativo sive advection and retention of pelagic juveniles of all en el asentamiento y reclutamiento final del rocot. sizes, including onshore movement during relaxation of upwelling. It appears that relaxation of upwelling may INTRODUCTION be a sufficient but not necessary aspect of the settlement The problems of drift and retention in marine ani- ofjuvenile rockfish from pelagic to postpelagic habitats. mals with pelagic larval stages are widely recognized (e.g., If so, advection during upwelling may not have a neg- Norcross and Shaw 1984; Bakun 1985, 1986; Sinclair ative effect on the settlement and ultimate recruitment 1988). To complete their life cycles, pelagic individuals of rockfish. must either remain in areas suitable for the next stage in the life cycle, or travel from the pelagic zone to these RESUMEN areas. Coastal upwelling systems pose a particular prob- Muchas especies de rocot del centro de California lem in this respect, since surface waters are transported concluyen su estadio peligico durante el period0 de sur- offshore, possibly carrying propagules away f?om the areas gencias de primavera-verano, cuando la advecci6n de las inhabited by postpelagic juveniles. Parrish et al. (1981), aguas superficiales podria transportarlos lejos de 10s hibitats Bakun and Parrish (1982), and Bakun (1985) suggested post-peligicos cercanos a la costa. Examinamos 10s pa- that the potential for offshore advection of larvae has se- trones de distribucibn de 10s estadios tardios de 10s ju- lected for reproductive patterns that minimize the ex- veniles peligicos de 10s rocot durante abril y mayo/junio posure of larvae to upwehg condtions, and Roughgarden de 1987 y 1988. Los ejemplares fueron obtenidos por et al. (1988) described the negative effect of offshore ad- medio de arrastres a media agua en un irea compren- vection in barnacles. dida entre Punta Reyes y la Bahia de Monterey, Dozens of species of rockfish in the genus Sebastes in- California. Los patrones de distribucibn encontrados habit the shelf and coastal zones in the region of maxi- fueron complejos y cambiaron ripidamente de acuerdo mum upwelling between Cape Blanco, Oregon, and a las condiciones oceanogrificas y de acuerdo a cambios Point Conception, California (Chen 1971). Many of these species bear young in winter and early spring, [Manuscript received January 22, 1994.1 before upwehng begins (Kendall and Lenarz 1987; Wyllie 175 LARSON ET AL.: DISTRIBUTION OF PELAGIC JUVENILE ROCKFISH CalCOFl Rep., Vol. 35, 1994 Echeverria 1987). However, the larvae and juveniles of months, and appear to grow at a rate of about 0.5 mm many species remain pelagic well into the springsummer per day (Woodbury and Ralston 1991). They trans- upwelling period (Kendall and Lenarz 1987; Moser and form into pelagic juveniles at least 20 mm long before Boehlert 1991), so that juveniles of these species must leaving the pelagic zone for juvenile habitats. The ju- settle from the pelagic zone when upwelling is active venile habitats vary among species at several scales (Love (Anderson 1983; Carr 1983, 1991). Thus, while vivi- et al. 1991). Species of Sebastes commonly leave the parity and winter spawning in rockfishes could be pelagic zone for habitats that are shallower than those seen as adaptations to the negative effects of offshore occupied by adults, often recruiting to areas quite near advection during upwelling (Parrish et al. 1981; Ainley shore (Love et al. 1991). Observations by divers demon- et al. 1993), the late pelagic stages of many rockfish strate that nearshore species show considerable special- are exposed to upwelling at the time they settle. If this ization of microhabitats (Carr 1983, 1991; Love et al. offshore advection does affect rockfish, spatial and 1991). Some deepwater species make extensive sur- temporal variation in upwelling could have important face-to-bottom migrations (Boehlert 1977); these species effects on both year-class strength and the geographical were not commonly encountered as pelagic juveniles in structure of rockfish populations. this study. Although many species clearly leave the pelagic Some evidence suggests that upwelling does have an zone for association with the benthos, the distinction advective effect on pelagic-stage rockfish. Moser and between pelagic and benthic habitat may be less clear in Boehlert (1991) noted the offshore distribution of some species such as shortbelly and chilipepper rockfish, whose Sebastes larvae, and their figures indicate relatively greater postpelagic juveniles and adults form schools that asso- offshore distributions off northern California, where sea- ciate more loosely with the bottom. Nevertheless, these ward jets are common. Simpson (1987) also illustrated species become invulnerable to midwater trawls, pre- the offshore distribution of Sebastes larvae in a filament sumably by shifting habitat. Essentially all of the species of advected water associated with an offshore eddy. covered in this study settle from the pelagic zone to more Brodeur et al. (1985) found Sebastes larvae unusually close benthic habitats that are shoreward of their pelagic dis- to shore during the El Nifio spring of 1983, but much tributions. farther offshore soon after an upwelling event later that Several aspects of the ecology of pelagic juveniles vary summer. Hobson and Howard (1989) found that mass among species. The timing and duration of parturition, strandings of juvenile shortbelly rockfish near shore in along with the duration of the pelagic stage, influence northern California were correlated with onshore trans- the timing and duration of their appearance as pelagic port, suggesting that the usual offshore distributions were juveniles in midwater trawls and their recruitment to maintained by normal offshore transport. Finally, Norton postpelagic habitats. A preliminary overview of the sea- (1987) has suggested that year-class strength of widow sonal occurrence of pelagic and newly settled juveniles rockfish is correlated with onshore transport, and Ainley of species encountered frequently in our study is pre- et al. (1993) found that year-class strength in the suite sented in table 1. Some species are taken in midwater of species found off central California (dominated by trawls for limited periods; others are taken over longer shortbelly rockfish) was affected negatively by strong up- periods of time. The peak seasonal occurrence of pelagic welling in January-February. juveniles varies by a month or more from year to year. In this paper we examine the distribution of late-stage The size of pelagic juveniles also varies among species pelagic juvenile rockfishes off central California. Our (appendix), and the relative sizes of pelagic and settled purpose is to infer the effects of upwelling on these fish individuals are concordant (comparing data in the ap- by comparing their distributions with oceanographic