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University Microfilms International 300 North Zeeb Road Ann Arbor, Michigan 48106 USA SI INFORMATION TO USERS This material was produced from a microfilm copy of the original document. While the most advanced technological means to photograph and reproduce this document have been used, the quality is heavily dependent upon the quality of the original submitted. The following explanation of techniques is provided to help you understand markings or patterns which may appear on this reproduction. 1. The sign or "target" for pages apparently lacking from the document photographed is "Missing Page(s)". If it was possible to obtain the missing pagels) or section, they are spliced into the film along with adjacent pages. This may have necessitated cutting thru an image and duplicating adjacent pages to insure you complete continuity. 2. When an image on the film is obliterated with a large round black mark, it is an indication that the photographer suspected that the copy may have moved during exposure and thus cause a blurred image. You will find a good image of the page in the adjacent frame. 3. When a map, drawing or chart, etc., was part of the material being photographed the photographer followed a definite method in "sectioning" the material. It is customary to begin photoing at the upper left hand corner of a large sheet and to continue photoing 'from left to right in equal sections with a small overlap. If necessary, sectioning is continued again - beginning below the first row and continuing on until complete. 4. The majority of users indicate that the textual content is of greatest value, however, a somewhat higher quality reproduction could be made from "photographs" if essential to the understanding of the dissertation. Silver prints of "photographs" may be ordered at additional charge by writing the Order Department, giving the catalog number, title, author and specific pages you wish fc~jQclucecl. 5. PLEASE NOTE: Some pages may have indistinct print. Filmed as received. University Microfilms International 300 North Zeeb Road Ann Arbor, Michigan 48106 USA SI. John's Road, Tyler's Green High Wycombe, Bucks, England HP10 8HR 7820437 SARVER, DALE JERE THE ECOLOGY AND ENERGETICS OF APLYSIA JULIANA (QUav AND QAIMARD, 1832), UNIVERSITY OF HAWAII, PH.D" 1978 THE ECOLOGY AND ENERGETICS OF APLYSIA JULIANA (QUOY AND QAIMARD, 1832) A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAII IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ZOOLOGY MAY- 1978 By DALE JERE SARVER DISSERTATION COMMITTEE Michael G. Hadfield, Chairman E. A1 ison Kay Robert A. Kinzie III Richard E. Young John Stimson ACKNOWLEDGEMENTS I would like to sincerely thank Dr. Michael Hadfield for his encouragement and support throughout the course of the study and especially for his assistance in preparing various drafts of the manuscript. Dr. William Van Heukelem spent many hours helping me design experiments and interpret the often times puzzling results; for this I am very grateful. Special thanks go to Dr. Marilyn Switzer-Dunlap for raising larvae which were necessary for many of my experiments. I must also thank Brad Peebles for his help with computer programs which made data analysis much easier... l think. iii ABSTRACT The objectives of this study were: 1) to estimate recruitment rates for Aplysia juliana on a weekly basis throughout the year at two field locations on Oahu, Hawaii; 2) to examine the interrelationships between food intake, growth rate, maximum size and fecundity; 3) to establish an energy budget; and 4) to determine the interactions between recruitment, adult density, food density and season. Recruitment occurred year-round although rates were higher in winter than in summer. Recruitment occurred in pulses which lasted from one to three days and the intensity of settlement varied between locations. Aplysia juliana larvae settle on available species of the green alga Ulva and spend about three weeks feeding on the alga and growing before migrating to the adult habitat under rocks. Mortality while on the algae is very high and fewer than 2% survive to move to the adult habitat. Maximum sizes of A. juliana attained under laboratory conditions were nearly five times larger than those seen in the field. Animals caged in the field grew rapidly until they reached sexual maturity when growth stopped. Animals raised in the laboratory continued to grow for several weeks after the onset of egg laying. This difference in growth pattern is attributed to limited access to food in the field imposed by their nocturnal habits and the activity-repressive response to heavy wave action. iv Maximum sizes of animals in the field varied both between locations and within the same location at different times of the year. These differences were attributed to variation in alga density, temperature, and possibly variations in nutritional quality of the algae. Caloric content of Q. lactuca from the Sand Island study site varied from 1850 to 3150 calories per dry gram. Figured on a caloric basis, adult A. juliana, under ad libitum feeding conditions, allocated 10.69% of its nutritional energy into reproduction, 5.45% into somatic growth, 10.65% into respiration and 27.28% into feces over a 3~ month period. About 46% of the ingested calories were unaccounted for and were apparently lost to mucus produc­ tion, excretion, and activity costs. Caloric assimilation efficiency was 72.7%, caloric net growth efficiency 22.2%, caloric gross growth efficiency 16.4% wet weight gross growth efficiency 23.6% and dry weight gross growth efficiency 10.95%. The relative allocation to reproduction decreased when animals were raised on reduced food rations unless rationing was sufficiently low to prevent growth, in which case all energy above maintenance costs went to egg production. Caloric gross growth efficiency did not significantly change with lower rations. At various field sites investigated, population numbers fluctuated greatly and frequent extinctions occurred. Aplysia juliana persists under such highly variable conditions by reaching sexual maturity early, and by producing millions of eggs over an extended reproductive life. v TABLE OF CONTENTS rage ACKNOWLEDGEMENTS iii ABSTRACT iv LIST OF TABLES viii LIST OF FIGURES ix CHAPTER I. GENERAL INTRODUCTION 1 CHAPTER II. GROWTH 4 Introduction 4 He thods 5 Resu1 ts 13 Discussion 32 CHAPTER III. RECRUITMENT 40 Introduction 40 ~~ethods 42 Results 46 Discussion 59 CHAPTER IV. DEMOGRAPHY 68 Introduction 68 Methods 69 Results 70 Discussion 83 CHAPTER V. ENERGETICS 90 Introduction 90 Methods 90 Results 94 Discussion 116 vi TABLE OF CONTENTS, continued Page CHAPTER VI. SUMMARY AND CONCLUSIONS 121 APPENDIX 126 LITERATURE CITED i 33 vii LIST OF TABLES Table Page 1 Monthly fecundity estimates for animals at Sand Island 80 2 Weekly fecundity estimates for animals in a cage.at Sand Island 81 3 Energetic constants for tissue, spawn, feces and algae 95 4 Energy budget for animals fed ad libitum 105 5 Energy budget for animals fed ad libitum to sexual maturity and then 10, 20, or 30 g of algae per 2 days 107 6 Energy allocation to spawn and growth for animals fed 75% of the ad libitum rate 108 7 Comparative energy allocation at different feeding 1evel s 109 8 Table of standard deviations for mean weight values in Figure 10 127 9 Table of ranges for mean weight values in Figure 5 130 10 Individual weight values for mean growth in Figure 7 131 11 Ranges for mean caloric content of U. lactuca from Sand Island 132 viii LIST OF FIGURES Figure Page 1 Growth of juveniles on ~. fasciata and ~. reticulata from Kewalo and U. reticulata and U. lactuca from Sand Island 14 2 Estimated field growth of juveniles compared to laboratory growth 16 3 Growth of animals grown in a cage at Sand Island 17 4 Growth and survival of spotted animals released at Sand Island 19 5 Growth with ad libitum and 75% of ad libitum feeding rate using ~. lactuca as food 20 6 Growth with ad libitum feeding on U. fasciata 21 7 Growth and feeding rate with ad libitum feeding on U. reticulata 22 8 Comparison of growth and survival in animals raised in tanks with and without current 24 9 Growth and fecundity of animals in ambient light and darkness 25 10 Growth of animals fed ad libitum to sexual maturity and then fed 10, 20 or 30 g of ~. lactuca per 2 days 27 11 Relative egg production with ad libitum and 75% ad libitum feeding rate 28 12 Growth under feeding regime of .5 g of ~. lactuca per day 29 ix LIST OF FIGURES, continued Figure Page 13 Growth under 3 different feeding regimes; ad libitum, starved 8 days and then ad libitum, and ad libitum to sexual maturity, maintenance rations of 4 g per day for 6 weeks and then return to ad libitum 31 14 Size-frequency distribution of recruits found on U. reticulata from Kewalo 4i 15 Size-frequency distribution of recruits found on U. reticulata from Sand Island 48 16 Size-frequency distribution of recruits found on U. lactuca from Sand Island 49 17 Daily percent survivorship of juveniles on U. reticulata from Kewa 10 51 18 Daily percent survivorship of juveniles on U. reticulata from Sand Island 52 19 Daily percent survivorship of juveniles on U. lactuca from Sand Island 53 20 Survivorship of juveniles on U. reticulata and U. lactuca from Sand Island and U. reticulata from Kewalo 54 21 Recruitment rates per 100 g of ~. reticulata 56 22 Algal standing crop at Sand Island and recruitment rates per square meter 58 23 Monthly estimates of fecundity and density for the Sand Island pooulation 71 24 Monthly size d~stributions at Black Point 73 x LIST OF FIGURES, continued Figure Page 25 Monthly size distributions at Ala Moana 74 26 Monthly size distributions at Sand Island (7/8/75 - 2/27/76) 75 27 Monthly size distributions at Sand Island (3/29/76 - 11/10/76) 76 28 Monthly size distributions at Sand Island (12/10/76 - 4/2/77) 77 29 Monthly estimates of caloric content of U.
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