SCU-Yl-79-003 c~ 2 ECOLOGYAND GROWTHOF JUVENILE CALIFORN1A SPINYLOBSTZR, PAHULIRUS ZÃTERRUPTUS RANDALL! by JohnMarlin Engle gP>g. ~',',~g E3P t $ga g, .,!':. Pg,"."g!ggpf USCSC-TD-03-79 November 1979 DISSERTATION SERIES ECOLOGY AND GROWTH OF JUVENILE CALIFORNIA SPINY LOBSTER, PANULZRUS ZNTZRRUPTUS RANDALL! by JohnMarlin Engle gP~>g~>~,'.8 gg~ 6;,»'-.5:.B~N9ff USCSC-~3-79 November 1979 This work is the result of research sponsored by the NOAAOffice of Sea Grant, Department of Commerce, under grant no. USC NOAA 04-8-M01-186. The U. S. government is authorized to produce and distribute reprints for governmental purposes, notwithstanding any copyright notation that may appear hereon. Institute for Marine and Coastal Studies University of Southern California University Park Los Angeles, California 90007 November 1979 ECOLOGY AM3 GROWTH OF JUVENILZ CALIFORNIA SPINY LOBSTER, PANULIRUS INTERRUPTUS RAZDALL! by John Marlin Engle A Dissertation Presented to the FACULTY OF THE GRADUATE SCHOOL UNIVERSITY OF SOUTHERN CALZFORXLC In Partial Fulfillment of the Requirements for the Degree DOCTOR OF PHILOSOPHY Biology! April 1979 Copyright John 2 larlin Engle 1979 UNlVKRSlTY OF SOUTHERN CALIFORNIA THK ORADUATKSCHOOl UHIVKhStTV PAhK LOS AHQKLSS, CAI I I'OhNI A 00007 This dissertation,written by JOHN MARLIN ZNGLZ under the direcfion of hler DissertationCom- mittee, and approeedby all its members,has beenpresented fo and accepted by TheGraduate School,in partialfulfi'llmenf of requirementsof fhe degreeof DOCTOR OF PHILOSOPHY DSNS Date 2i.l«d V DISSERTATION COMMITTEE DEDIC ATION This dissertation is dedicated to a good person--a person who lived unselfishly for his family and friends, and who died be- cause his work was his life--my father. Morris M. Engle, Jr. AC KNOW LED GE IVMNTS l wish to express my gratitude to Dr. Kristian Fauchald, my diss .tation committee chairman, for guidance, encouragement, and constructive criticism given throughout the study to his only "non-worm" student. Thanks are due to my other committee mem- bers--Drs. Richard Brusca, Robert Given, Andrew Pilmanis, and Russell Zimmer--for their suggestions and review of the manu- script,. Many fellow graduate students and friends assisted me in this research. I especially wish to thank the following persons who provided valuable field assistance and frequently endured consider- ably less than desirable water conditions in. order to help as dive partners: Terry Audesirk, Marty Meisler, Peggy O'Donnell, Rick Selby, Cindy Shoemaker, Ray Wells, and Joe Wible. Others con- tributing diving assistance, technical advice, helpful discussions, or highly valued support and friendship include: Jerry Audesirk, Rick Beclavitt, Tina Behrents, Sandy Benson, Garry Brown, Margaret Callahan, Jim. Coyer, John Dmohowski, Jane Dykzeul, Bill Gorham, Greg Hageman, Anne Harrington, Beth Hart, Mike Haufler, Bill Herrnkind, Dave Jahn, Bob Lavenberg, Andy Lissner, Muffy McDonaid, Robin IViacPadden, Zizn Morin, Marianne Ninos, John Pilger, Buss Schmidt, Miguel Angel Tenorio, Tim Tricas, and Rick Vance. I thank Dr. Russell Zimmer and Dr. Robert Given for the use of the exceUent facilities of the Catalina Marine Science Center CMSC! provided by the Institute for Marine and Coastal Studies IMCS!. Mr. Larry Loper was invaluable for his efforts in the design, construction, and repair of various pieces of equipznent. Ship time was donatedby the California Departznent of Pish and Game, Scripps Institution of Oceanography, and the Tatman Founda- tion. Pier facilities for night-lighting were provided by Marineland of the Pacific. Dr. Patrick Parker and his staff analyzed tissue samples for carbon isotope ratios at the University of Texas Marine Institute. I would like to acknowledge the financial support received from IMCS through CMSC employment and Sea Grant traineeships, and frozn the USC Department of Biological Sciences in the form of r teaching assistantships. Mr. Ronald Linsky was instruznental in initiating federal funding of this research. The study was sponsored in part by NOAA Office of Sea Grant, Department of Co~merce, under Grant No. USDC 04-6- l.58-443.18 to the University of Southern Calif ornia. zv Acknowledgements are due to Ms. Karin Fouts for drafting the figures and to iUrrs. O'Detta Hawkins for typing the final copy. IViuffy--special thanks for your invaluable assistance in preparation of the manuscript and for your encouragement and patient under- standing. Finally, I would like to thank Pan, for making it all possible. TABLE OF CONTENTS Page DEDICATION . ACKNOWLEDGIVlENTS . LIST OF FIGURES LI.ST OF TABLES LIST OF APPENDICES INTRODUC TION Juvenile Habitat 5 Lobster Growth 89 The Molt Cycle Growth and Ecology . 10 Fishery Management and Mariculture Perspectives 11 Juvenile Growth 12 Juvenile Abundance 14 Juvenile Activity Patterns and Gregariousness . 15 Juvenile Food Habits 17 Juvenile Mortality 18 ~ tgy fP h ~ B dll. 19 Distribution 19 Behavioral Ec ology 20 Life History 22 Growth 29 The California Lobster Fishery . 33 MATERIALS AND METHODS 36 Juvenile Habitat 36 Puerulus Recruitment . 36 Puerulus Metamorphosis 41 Puerulus Habitat Preference 41 Juvenile Habitat Surveys 42 Page Juvenile Habitat Preference . 46 Surf Grass Distribution Surveys 46 The Primary Study Site.. 47 The Bird Rock Surf Grass Ecosy stem 49 Stable Carbo~ Isotope Analysis 50 Juvenile Growth 52 Field Capture Techniques 52 Lobster Measurements 53 Length-Weight Relationships 54 Sex Determination 55 Size- Frequency Distributions 56 Field Molt Increments . 57 Mark-Recapture Growth . 58 Laboratory Cultur e 59 Growth Analysis 62 Juvenile Abundanc e . 62 Juvenile Activity Patterns 64 Juvenile Food Habits . 66 Juvenile Mortality 68 RESUI TS . 69 Juvenile Habitat 69 Puerulus Recruitment . 69 Night- Lighting . 69 73 Puerulus Metamorphosis 74 Bird Rock Recruitment 76 Age at Settlement 78 Sea Water Temperatures 79 Puerulus Habitat Preference 84 Laboratory Substrate- Choice Tests 84 Field Observations 84 Juvenile Habitat Surveys 86 Relationship of Body Color to Habitat 92 Juvenile Habitat Preference . 94 Laboratory Mbstrate-Choice Tests 94 Micr ohabitat Observations 96 Surf Grass Distribution Surveys 96 Bird Rock and Isthmus Reef Habitat Surveys . 101 The Bird Rock Surf Grass Ecosystem 105 Stable Carbon Isotope Analysis 109 Juvenile Growth . 114 Sex Ratios . 114 Length-Weight Relationships . 114 Measurement Precision 114 Regression Equations 116 Size- Fr equency Distributions 121 Monthly Mean Size Analysis 126 Field Molt Incr ements 136 Mark-Recapture Growth 138 The Marking System . 138 Molt Stage Determination 145 Recapture of Marked Individuals 145 Laboratory Culture .. 151 Growth Analysis 159 Field Single Molt Determination . 159 Effect of Appendage Regeneratio~ on Molt Incre ent . 163 Effect of Premolt Size on Molt Increment . 163 Effect of Temperature on Molt Increment . 171 Growth Equations . 171 Molt Frequency Predictions 175 The Generalized Pattern of Growth 177 Juvenile Abundance 181 Catch per Unit Effort . 181 Mark-Recapture Population Size Estimates 186 Population Density 188 Aggregations 191 Juvenile Activity Patterns 191 Den Residency 191 Diel Activity Rhythms 192 Home Range 195 Seasonal Movement Patterns . 197 Juvenile Food Habits 199 Juvenile Mortality . 206 Mortality of Laboratory-Raised Juveniles . 206 CPUZ and Mark-Recapture Mortality Trends 207 Predation . 207 Page DISC USSION . 212 Puerulus Recruitment 212 Early Juvenile Habitats 216 220 Juvenile Growth. 222 Length- Weight Relationships 222 Year Class Growth . 224 Field versus Laboratory Growth 229 Factors Affecting Growth 230 Adult Age- Size Relationships 236 Juvenile Food Habits 239 Juvenile Population Dynamics 244 Juvenile Survival and Growth Tactics 249 SUMMARY AND CONC LUSIONS 254 LITERATURE CITED . 261 APPENDICES 274 276 282 IV 297 LIST OF FIGURES Figure 1. Temporal sequences of life history phen'oznena in ~ ~ ~ 23 2. Location of sampUng sites for puerulus larvae and y" 3. Geographic locations of subtidal and intertidal I I 'I I' ~ ~ ~ ~ 4 d Ph ll dix spp.. 43 4. Mean semimonthly surface sea water temperatures at Bird Rock, Santa Catalina Island, during 1975 and 1976 82 I ' fj 'I P. sites at western Santa Catalina Island . 89 d' ~f Rl Cavern Point to Arrow Point, Santa Catalina Island . ~P' y habitats at Bird Rock, Santa Catalina Island.... 102 8. Relationship between carapace length and wet weight I I df I'j 'I ~ ~ ~ 119 9. Monthly size frequencies for 1 mrs size classes of ~ ~ ~ IR I 122 10. Size frequencies for 0. 6 mm size classes of a11 laboratory-raised and fie1d-captured juvenile P. 127 Figure Page kl kl»' collected at Bird Rock during 1975 and 1976 129 12. Monthly mean sizes of laboratory-raised and fieM P 134 13. Appearance of the holes punched in the uropods and telson of typical laboratory-reared juvenile P. ~ 141 Growth of 15 representative marked juvenile P. ~ 149 15. Mean increases in size per year for marked I P.~ 152 16. Mean intermolt times for 2 mm size groups of ~ ~ ~ I P aquaria 157 Relationship between initial size and total increase in size after two years of growth in laboratory aquaria . 160 18. Mean single molt percentage increases in carapace length for 2 mm size groups of laboratory-raised d ' dd I 164 19. Mean single molt increments for 2 mm size groups of laboratory-raised and field-captured juvenile P.'t t 167 20. Instar sizes of laboratory- raised and field-captured P.'t tu 21, 11" I" P 182 22. Monthly means and ranges of catch per diver-hour I I 'I P. ~ d' d II 1 184 23. Catch per diver-hour and mark-recapture popula- tion density estimates for two settlement year 189 Xl LIST OF TABLES Table Page 1. Comparison af annual natural growth rates and critical age predictions for the California spiny 33 2 f~hll d 3 f pueruH. ~ ~ ~ 2 40 3. Night-light sampling for the puerulus larvae of P. 3 C ll d dM 2 of the Pacific, Los Angeles 2' V f 2 P.~ collected at Marineland on September 4-6, 1975, to reach three postsettlement pigmentation stages and the metamorphic molt 75 5. Monthly captures of 7-12.