Activity Patterns and Feeding Chronology of the Kelp Rockfish, Sebastes Atrovirens, in a Central California Kelp Forest
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ACTIVITY PATTERNS AND FEEDING CHRONOLOGY OF THE KELP ROCKFISH, SEBASTES ATROVIRENS, IN A CENTRAL CALIFORNIA KELP FOREST A Thesis Presented to The Faculty of the Department of Biology San Jose State University In Partial Fulfillment · of the Requirements for the Degree Master of Arts By Gilbert S. Van Dykhuizen May 1983 ABSTRACT Activity patterns, home range behavior, feeding habits, and chronology of adult kelp rockfish,·Sebastes atrovirens, were investigated in a kelp forest in Carmel Bay, California. Kelp rockfish abundance fluctuated with kelp density and canopy cover in this study area. Fish were distributed throughout the water column both day and night in the kelp season and were confined to the bottom in the non-kelp season. Tagged kelp rockfish occupied home ranges throughout the study, although several departed as a result of storms and reduced kelp density. Kelp rockfish fed opportunistically on available and abundant organisms including kelp-associated, open water, and epibenthic prey. Digestive states and rates on fish prey were determined in the laboratory, an~ kelp rockfish fully digested a medium-sized meal of juvenile Sebastes in approximately 33 hours. Initial time of feeding was estimated for field-collected kelp rockfish containing juvenile fishes. From recency of feeding indices, feeding appeared to occur at all times of day with a tendency towards dawn and night feeding. iii ACKNOWLEDGMENTS I would like to thank the members of my committee, Drs. Greg Cailliet, Ralph Larson, and Robert-Lea for their helpful comments, support, and assistance during the course of this work. Dr. Mike Foster also provided valuable suggestions along the way. Many thanks and appreciation go to the divers who assisted me, including Todd Anderson, Mickey Singer, Mark Carr, John Heine, Don Canestro, Jim Brennan, Bruce Welden, Alan Fukuyama, Mike Denega, Dave Ebert, Drs. David Schiel and Ralph Larson, Doug Maier, Michelle Whitney, Susan Dearn, Barbara Pierson, and Alistair Hamilton. The Pebble Beach Corporation kindly allowed diving access to Stillwater Cove. Peter Slattery helped identify stomach contents of kelp rockfish. Dr. David Schiel, Dr. Liz Denley, and Kevin Lohman helped with statistical procedures. Kevin Lohman and Lynn McMasters drafted the figures. Deborah Tuel typed the final manuscript. My thanks and appreciation are extended to all these fine people. Thanks also go to Larry Jones, Ken DeLopst, and Preston Watwood for maintaining diving equipment and general assistance. The staff and faculty at Moss Landing Marine Laboratories were also helpful and supportive during my stay at the lab. The use of Moss Landing Marine Laboratories' HP 9825 computer system was an important aid in the completion of this project. Finally, I would like to thank the David and Lucille Packard Foundation for financial assistance in purchasing equipment and supplies. iv TABLE OF CONTENTS ABSTRACT iii ACKNOWLEDGMENTS iv LIST OF FIGURES vi LIST OF TABLES vii INTRODUCTION 1 MATERIALS AND METHODS 4 Study Site 4 Seasonal and Diel Activity 5 Home Range Behavior 8 Feeding Habits and Chronology 9 Digestive States and Rates 10 RESULTS 14 Seasonal ~nd Diel Activity 14 Home Range Behavior 15 Feeding Habits and Chronology 16 Digestive States and Rates 18 DISCUSSION 20 LITERATURE CITED 26 v LIST OF FIGURES Figure 1 Map showing study area. 32 2 Map of Arrowhead Point. 34 3 Map illustrating transect sampling procedure. 36 4 Flow diagram illustrating the three-factor ANOVA model. 38 5 Cumulative abundance versus number of transects. 40 6a Canopy cover estimates at Arrowhead Point. 42 6b Mean Macrocystis density and percentage of plants with greater or less than five stipes at Arrowhead Point. 42 7 Number of fish per 40 m2 transect in kelp versus non-kelp season. 44 8 Diel activity in kelp versus non-kelp season. 46 9 Mi c ~habitat use in ke 1 p versus non-ke 1 p season. 48 10 Cumulative number of prey types versus number of stomachs. 50 11 Percent number and frequency of occurrence for dawn, noon, dusk, and night collected stomachs. 52 12 Fullness-recency data versus time of collection. 54 13 Estimated initial feeding time of field-collected specimens. 56 LIST OF TABLES Table 1 Three-factor ANOVA comparing seasons, time of day, and behaviors. 57 2 Three-factor ANOVA comparing seasons, time of day, and microhabitat. 58 3 Home range observatons for 14 tagged kelp rockfish. 59 4 State of digestion indices. 61 INTRODUCTION The subtidal kelp environment of the central California coast supports a variety of fishes, the ~ost numerous in abundance and biomass being the rockfishes (Scorpaenidae: genus Sebastes). Rockfishes are also a major constituent of sport and commercial fisheries along California and the eastern north Pacific coast. Rockfishes have been included in many investigations of kelp forest fishes in southern California (Larson, 1972; Burge and Schultz, 1973; Feder et al., 1974; Ebeling and Bray, 1976; Hobson and Chess, 1976; Coyer, 1979; Ebeling et al., 1980; Ebeling et al., 1980; Larson, 1980; Love, 1980; Hobson et al., 1981). Studies in central California have dealt with the spatial partitioning and food habits of rockfishes (Miller and Geibel, 1973; Hallacher, 1977; Roberts, 1979; Love and Westfall, 1981). Of these investigations few have focused on the behavior and ecology of a single species (Love, 1980; Larson, 1980). Concentrating on a single species often leads to a more detailed understanding of an organism's role in its environment. In addition, the effects of seasonal and diel variation in diet and habitat can ultimately provide insight into the growth and reproduction of a species. This information will also help in assessing the status of local populations of a species with respect to fishing pressure by divers and fishermen. One of the more common kelp forest rockfishes is the kelp rockfish, Sebastes atrovirens, which has· been studied in both central and southern 2 California (Larson, 1972; Ebeling and Bray, 1976; Hobson and Chess, 1976; Hallacher, 1977; Roberts, 1979). The kelp rockfish ranges from Punta San Pablo, Baja California, to Timber Cove, California and is found primarily in kelp forests to· depths of 45 meters (Miller and Lea, 1972). The adults reach a reported maximum length of 425 mm (Miller and Lea, 1972). Juveniles of this species recruit to the kelp canopy in late summer in central California (Singer, 1982; Carr, 1983). Adults are found throughout the water column and feed on kelp-associated invertebrates and juvenile fishes (Larson, 1972; Roberts, 1979). Hobson and Chess (1976) observed kelp rockfish to feed nocturnally and usually idle near the bottom during daylight hours. Rockfishes are known to maintain home ranges and defend territories. Carlson and Haight (1972) showed yellowtail rockfish, 2· flavidus, to h~ve a home range and also return to this home range from as far away as 22.5 km. Larson (1980) found the gopher rockfish, 2· carnatus, and the black and yellow rockfish, 2· chrysomelas, two species segregated by depth, to defend territories. Information concerning the home range or territorial behavior of the kelp rockfish is lacking. Since the activity and feeding habits of a species may change over its geographic range (Fox and Morrow, 1981), the purpose of this investigation was to observe the activity patterns and feeding chronology of the kelp rockfish and compare these findings with those found for southern California kelp rockfish. 3 The major questions addressed were: 1) Are there seasonal abundance and activity patterns, and what factors contribute to changes in these patterns? 2) Do kelp rockfish maintain" a home range or a territory similar to other rockfish species? 3) What are feeding habits and feeding chronology of the kelp rockfish in central California and how do they differ from individuals of this species in southern California? 4 MATERIALS AND METHODS Study Site This study was conducted in ~acrocystis pyrifera kelp habitat off Arrowhead Point in Stillwater Cove, Carmel Bay, California (36°33'30"N, 121°55'30''W) (Figs. 1, 2). Stillwater Cove is protected from northwesterly swell conditions but is exposed to severe southerly storms which commonly occur from October through February. During the summer months, June through September, M. pyrifera forms a thick canopy which, along with many of the kelp plants, is removed during the winter storms. In addition to~· pyrifera, there are uncommon occurrences of other brown algae such as Pterygophera californica, Cystoseira osmundacea, and Desmarestia spp. The botto~ topography consists of large granitic boulders and vertical rock outcroppings extending approximately 4 m above the sea floor. Sand channels are interspersed among the boulders and outcroppings. The depth at the study site varies from 9 m at the large outcrops to 12.5 m, MLLW, on the sea floor. The reef substrate is encrusted with various erect and crustose sponges, tunicates, and bryozoans as well as a thick mat of red articulating coralline algae (Foster, 1982). ~· pyrifera canopy cover and density were measured over the year to determine the fluctuation of this habitat. ~· pyrifera density was estimated by running three 20m-long by 1m-wide transects during six 5 different sampling dates over the year. Both the number of plants per m2 and number of stipes per plant were counted. Percent canopy cover in the study area was estimated on a scale of 1 to 4 with: 1 = 0-25%; 2 = 25-50%; 3 = 50-75%; 4 = 75-100%. Seasonal and Diel Activity Seasonal and diel activities were observed in depth-stratified transects made day and night over the course of 17 months. Three buoys were anchored by pitons in rocks along the kelp forest-sand channel border. Concrete nails with numbered strips of bicycle handlebar tape were placed at one meter intervals along the bottom of the 40 m by 20 m study area (Fig. 3).