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V·M·I University Microfilms International a Beil & Howell Information Company 300 North Zeeb Road INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adverselyaffect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand corner and continuing from left to right in equal sections with small overlaps. Each original is also photographed in one exposure and is included in reduced form at the back of the book. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6" x 9" black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. V·M·I University Microfilms International A Beil & Howell Information Company 300 North Zeeb Road. Ann Arbor. M148106-1346 USA 313'761-4700 800,521-0600 Order Number 9215026 Energy allocation and reproductive effort in four cockroach species with differing modes of reproduction Koebele, Bruce Peter, Ph.D. University of Hawaii, 1991 Copyright @1991 by Koebele, Bruce Peter. All rights reserved. V·M·l 300N. Zeeb Rd. Ann Arbor,MI48106 Energy Allocation and Reproductive Effort in FourCockroach Species withDiffering Modesof Reproduction A dissertation submitted to the GraduateDivision of the Universityof Hawaii in partial fulfillment of the requirementsfor the degree of Doctor of Philosophy in Zoology December 1991 By Bruce P. Koebele DissertationCommittee: MichaelG. Hadfield, Chairperson SamuelR. Haley LeonardA. Freed Gregory A. Ahearn Franklin Chang © Copyright by Bruce P. Koebele 1991 All Rights Reserved iii Thisdissertation is dedicatedto the memory of my grandparents, Marcel and Phyllis Koebele. iv Acknowledgements There are several people I wish to thank for their help In making this dissertation a reality. First and foremost, I'd like to acknowledge the assistance and support of my advisor, Dr. Michael G. Hadfield - thanks Mike for adopting this floundering "fish person. n Thanks too to the other members of my dissertation committee, Dr. Samuel R.Haley, Dr. Leonard A. Freed, Dr. Gregory A. Ahearn, Dr. Franklin Chang, and former member Dr. James Archie for all their assistance. I am grateful to Dr. John B. Hall for the frequent use of equipment and supplies belonging to the UH Biology Program. John Strazanac, from the UH Entomology Department, was extremely helpful early on in the identification of species and providing me with information about their biology. Finally, I wish to acknowledge the continuous support and encouragement of my parents, Ralph and Marina Koebele, over the many years of my "higher education." v Abstract In an attempt to better understand how evolution from oviparity to viviparity might alter energy demand and allocation, energy budgets were determined for adultfemales offourspecies of cockroaches, the oviparous Supella /ongipa/pa, the facultative ovoviviparous B/attella germanica, the ovoviviparous Pycnosce/us indicus and the viviparous Diploptera punctata. Food consumption and feces production in all species were greatest during the early part of a reproductive cycle, immediatelyafter oviposition for S./ongipalpa and immediatelyafter offspring hatching in B. germanica, P. indicus and D. punctata. Respiration in three of the four species (all except D. punctata, where respiration was constant throughout the cycle) closely paralleled food consumption with respiration highest early in the reproductive cycle. OVOViviparity in cockroaches was characterized by the addition (to the oviparous condition) of a quiescent postoviposition gestation period, during which the female ate and respired little in comparison to preoviposition rates. Viviparity in D. punctata was characterized by a more constant rate of consumption (although still fastest in the first third of the reproductive cycle) and a constant rate of respiration. While there were significant differences in the temporal patterns and rates of energy acquisition and allocation associated with the different modes of reproduction, changes in the relative proportions of energy allocated to competing demands such as respiration, weight gain and reproductive products were slight. Multiple estimates of reproductive effort (RE), the proportion of an organism's energy budget devoted to reproduction, were calculated using seven indices based exclusively on energy budget parameters (allocation-based indices) and two mixed indices based on female body weight (soma-based indices). All but one of the allocation-based indices showed the same interspecific ranking of RE, S. longipalpa > P.indicus > B. germanica == D. punctate, while the interspecific ranking of the two soma-based indices differed from that of the allocation-based indices and from each other. SUbstituting ash-free dry vi weight or dry weight for energy equivalents reduced the absolute value of both allocation-based and soma-based RE Indices. Among allocation-based indices. those which contained more energy budget parameters yielded greater absolute RE estimates than less complicated indices. These results suggest that previous reports of agreement between allocation-based and soma-based REestimates were probably coincidental. vii Table of Contents Acknowledgements v Abstract vi List of Tables ix List of Figures , , '.' xii Chapter 1: Energy Allocation and Reproductive Effort in Four Cockroach Species with Differing Modes of Reproduction 1 Introduction 1 Materials and Methods 3 Animals 3 Measurement of EnergyBudgetParameters 3 Estimating Reproductive Effort 6 Statistical Analysis 7 Results 7 Discussion 22 Balancing EnergyBudgets 22 Food Consumption and Feces Production 26 Respiration 27 Somatic Production 30 Reproductive Effort 31 TheEvolution of Viviparity in Cockroaches 34 Summary 39 Chapter 2: An Assessment of Reproductive Effort Indices 42 Introduction 42 Materials and Methods 43 TheAnimals 43 Energy BudgetParameters and REIndices 44 Calculating REIndices 47 Statistical Analysis 48 Results and Discussion 48 Conclusions and Recommendations 54 Summary 55 Epilogue: Leftovers 56 How Real? 56 The Future of RE? 58 Appendix: Energy Budget Data for Individual Cockroach Females 61 Literature Cited 113 viii Ustof Tables Table 1-1.Feces production of Supella longipalpa, Blattella germanica, Pycnoscelus indicus, and Diploptera punctata 11 Table 1-2.Energybudget parameters for one reproductive cycle 12 Table 1-3.Assorted life history parameters 13 Table 1-4.Caloricvalue (Joules/mg) of energybudget parameters 15 Table 1-5.Percentage of ash in energybudget parameters 16 Table 1-6. Energybudget parameters for one reproductive cycle in energy equivalents 17 Table 1-7.Assimilation (food consumed - feces), respiration, and productivityefficiencies and ratescalculated using meansfrom Tables1-3 and 1-6 19 Table 1-8.Energybudget parameters (in Joules) usedto calculate reproductive effortfor Supella longipalpa, Blattella germanica, Pycnoscelus indicus and Diploptera punctata 21 Table 1-9.Fourindices of reproductive effortfor Supella longipalpa, Blattella germanica, Pycnoscelus indicus and Diploptera punctata 23 Table 1-10.Reproductive effort in cockroaches along with variouslife history parameters 32 Table 2-1. Energybudget parameters for four cockroachspecies for one completereproductive cycle 45 Table 2-2. Formulae and descriptionsof seven allocation-based and two soma-based reproductive effort (RE) indices 46 Table 2-3. Means ± standard error for seven allocation-based reproductive effort indices usinga commondata base 49 Table 2-4. Mean ± standard error for two soma-based reproductive effort indices usinga common data base 53 Table A-l. Percentage of ash in variousenergybudget parameters of Supella longipalpa, Blattella germanica, Pycnoscelus indicus and Diploptera punctata 62 Table A-2. Caloricvalues (Joules/mg) of energybudget parameters for Supella longipalpa 64 ix Table A-3. Length (days) of the reproductive cycle of Supella longipalpa 65 Table A-4. Feces production (mg dry weight) for multiple reproductive cycles of Supella longipalpa 66 Table A-5. Food consumption (mg dry weight) of Supella longipalpa for multiple reproductive cycles 67 Table A-6. Dry weights (mg) of new oothecae of Supella longipalpa for multiple reproductive cycles 70 Table A-7. Night and day rates of oxygen consumption (mL oxygen/12 hrs) for Supella longipalpa over one reproductive cycle 71 Table A-S. Weight and weight changes (mg) of Supella longipalpa over multiple reproductive cycles 72 Table A-9. Dry weight (mg) of all young and oothecal case, number of young per ootheca, and calculated dry weight of individual young of Supella longipalpa for one reproductive cycle 73 Table A-l0. Caloric values (Joules/mg) of energy budget parameters for Blattella germanica 74 Table A-11. Length (days) of the reproductive cycle, preoviposition period, and postoviposition period of Blattella germanica 75 Table A-12.
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