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The Feeding Habits of Pleurobranchaea

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The Feeding Habits of Pleurobranchaea THE FEEDING HABITS OF PLEUROBRANCHAEA CALIFORNICA MACFARLAND, 1966 (OPISTHOBRANCHIA, NOTASPIDEA) IN MONTEREY BAY, CALIFORNIA A Thesis Presented to the Faculty of California State University, Stanislaus and Moss Landing Marine Laboratories In Partial Fulfillment Of the Requirements for the Degree Master of Science in Marine Science By Karen E. Battle August, 1994 ABSTRACT The natural diet of Pleurobranchaea californica was studied in Monterey Bay, California. Specimens were collected from January 29 to November 9, 1992, at depths of 30-100 meters, using an otter trawl and an interfacial trawl. Three hundred fty-six animals were collected and ranged in size from 0.01-411 grams with most of the animals weighing between 1.0 and 50 grams. The gut contents were examined and 16 different prey types were identified. Thirty-three percent of the guts were empty. Many animals had sediment in their guts, which was presumed to be a result of their ingesting or attempting to ingest prey items that live on or below the substratum. P. californica a euryphagic predator with pronounced cannibalism. Individual Index of Relative Importance values showed that opisthobranchs made up the largest portion of the diet of P. californica. The opisthobranchs in the diet included P. californica (as prey), Armina californica and Aglaja sp. The diets were similar specimens of P. californica collected at all depths, but did change depending upon the season in which the animals were collected, and their size. ii ACKNOWLEDGEMENTS I would like to thank my committee members, Dr. James Nybakken, Dr. Gregor Cailliet, and Dr. Pamela Roe for their guidance and advice throughout this study. I would also like to thank Dr. Pamela Roe for her support and help with communications between California State University, Stanislaus and Moss Landing Marine Laboratories. I also wish to express my sincere thanks to those who contributed assistance throughout this project. The collection of specimens could not have been accomplished without the knowledge and experience of Tracy Thomas. Thanks also to the students who helped on collection days, Lisa Williams for her support, Kathy Wendell for her assistance in editing, and Carolyn Comings for the use of her computer and laser printer. I would like to give special thanks to my family for their support and specifically to my parents, Bob and Valerie Battle, without whom this study could not have been done. iv TABLE OF CONTENTS Page TITLE PAGE . i ABSTRACT . ii CERTIFICATION OF APPROVAL iii ACKNOWLEDGEMENTS iv TABLE OF CONTENTS v LIST OF TABLES vi LIST OF FIGURES vii LIST OF APPENDICES xi INTRODUCTION . 1 MATERIALS AND METHODS 4 Statistical Analysis . 9 RESULTS . 12 DISCUSSION . 22 CONCLUSIONS 34 LITERATURE CITED . 35 APPENDICES 81 v LIST OF TABLES Table Page 1. Number and type of trawls used per month and the number of days trawled per month 39 2. Taxa of the prey from P. californica and the number of specimens of P. californica found with that prey type . 40 3. The Kruskal-Wallis test values for the (%Volume)(%Fullness) values of the prey 41 4. Trophic diversity and evenness summary 42 vi LIST OF FIGURES Figure Page 1. Map of Monterey Bay . 43 2. Size frequency of all individuals of P. californica collected, as the number of animals per size class 45 3. Size frequency of individuals of P. californica collected in each season and at each depth, as the number of animals per size class 47 4. The average weight (grams) of P. californica individuals, per trawl, for each month animals were collected . 49 5 . The average number of animals (P. californica), per trawl I for each month animals were collected . 51 6. Percentage of empty guts per size class (grams) . 53 7. Average percent fullness of stomach for specimens of P. californica collected at each hour . 55 vii Figure Page 8. Percentage of animals with stomach contents for specimens of P. californica collected at each hour . 57 9. Percentage of animals (P. californica) with stomach contents, in digested state 1-2 or 3-4, for specimens of P. californica collected in each hour . 59 10. Cumulative number of prey types found in the number of stomachs analyzed for the overall diet of P. californica 61 11. Cumulative number of prey types found in the number of stomachs analyzed for each depth, season and size class . 63 12. Individual Index of Relative Importance (IRI) for overall diet of P. californica 65 viii Figure Page 13. Individual Index of Relative Importance (IRI) for specimens of P. californica collected at 70-100 meters, 50-69 meters and 30-49 meters 67 14. Percent Index of Relative Importance (IRI) for specimens of P. californica collected at 70-100 meters, 50-69 meters and 30-49 meters 69 15. Individual Index of Relative Importance (IRI) for specimens of P. californica collected in Winter and Spring, and Summer and Fall . 71 16. Individual Index of Relative Importance (IRI) for specimens of P. californica which had eaten P. californica (as prey) 73 ix Figure Page 17. Percent Index of Relative Importance (IRI) for specimens of P. californica collected in Winter and Spring, and Summer and Fall. Also percent IRI for specimens of P. californica which had consumed P. californica (as prey) in Fall and Winter, and Spring and Summer 75 18. Individual Index of Relative Importance (IRI) for specimens of P. californica weighing 51-450 grams, 11-50 grams and 0.01-10 grams . 77 19. Percent Index of Relative Importance (IRI) for specimens of P. californica weighing 51-450 grams, 11-50 grams and 0.01-10 grams . 79 X LIST OF APPENDICES Appendices Page Cover page 81 A. Date and locations of trawls 82 B. Trawl dates, lengths, depths and and number of P. californica caught in each trawl . 87 C. Weight, fullness of stomach and digestion state of prey for each P. californica analyzed 92 D. Prey types eaten by P. californica, number of individual prey, percent number of prey, percent volume of prey and notes on prey . 104 xi 1 INTRODUCTION The natural history and feeding habits of most notaspideans are not well known. Cattaneo-Vietti et al (1993) suggest that the research on notaspideans limited because it is difficult to obtain a sufficient number of specimens. Willan (1984), in an overview of notaspidean diets, reported that the majority of observations on food and feeding in the literature were from incidental reportings in otherwise taxonomic studies. Willan (1984) found that members of notaspidean genera such as Tylodina, Umbraculum and Berthella were stenophagic in diet while those in other genera such as Pleurobranchaea and Pleurobranchella were euryphagic. Studies done on five species of Pleurobranchaea by Cattaneo-Vietti (1986), Cattaneo-Vietti et al (1993), Chamberlain and Behrens (1980), Ottaway (1977), Tsubokawa and Okutani (1991) and Willan (1984) demonstrated them to be carnivores but also suggested possibilities of scavenging and cannibalism. Pleurobranchaea californica (MacFarland, 1966) is the largest and most abundant notaspidean predator found off the coast of central California. This species can be found at depths ranging from 10 to 400 meters (Behrens, 1991; Chivers, 1967; Coan, 1964) and ranges in length from 8 to 210 2 millimeters (Behrens, 1991; Chivers, 1967; MacFarland, 1966). P. californica has been used extensively as an experimental animal in neurophysiological research (Davis et al, 1977; Gillette and Davis, i977; McClellan, 1982; Ram and Davis, 1977) and in avoidance conditioning experiments (Davis et al, 1980; Mpitsos and Collins, 1975; Mpitsos and Davis, 1973), but little work has actually been done on its natural history. The only research, relative to diet, that has been done on P. californica is some laboratory observations of feeding behavior (Chivers, 1967; Coan, 1964; Lee et al, 1974; Morse, 1984) and an anatomical study of its digestive system (Morse, 1984). According to Morse (1984), P. californica has sensory papillae on its oral veil. As it crawls, it holds the oral veil on the substratum until it comes in contact with a prey item. At this point, the oral veil lifted, and the oral tube and buccal mass are everted. An acid gland associated with the buccal mass immobilizes the prey to allow P. californica to ingest it (Morse, 1984). Observations of feeding behavior in captivity and morphology of the digestive system led Morse (1984) to suggest that F. californica is carnivorous and can probably ingest large prey. Morse (1984) found that one specimen of P. californica had ingested an entire opisthobranch of the genus Berthella. Chivers (1967) also found P. californica to 3 be carnivorous in captivity, eating whatever organisms were supplied, including Anthopleura egantissima (Anthozoa) and other specimens of P. californica, but the question of whether P. californica is stenophagic or euryphagic in nature, remained unanswered. In this study, I focused on establishing the natural diet of P. californica and attempted to answer whether P. californica is a generalized or specialized predator. Secondary questions included whether the diet of P. californica differed with depth, season and the size of the P. californica. 4 MATERIALS AND METHODS Specimens of Pleurobranchaea californica were collected weekly in Monterey Bay, California from January 29, 1992 to November 9, 1992. Inclement weather conditions precluded collecting specimens in December, and in January and February specimens were only collected one day out of each month. The collections were made in three separate areas in Monterey Bay (Figure 1). Area A, an extensive area located north of the Monterey Canyon and east of Soquel canyon, had muddy bottom sediments, very little debris and was the most favorable area for finding P. californica. Area B, located south of Area A and Monterey Canyon, had more coarse sandy sediment than the other areas. Drift kelp and debris were common on the bottom.
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