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INFORMATION TO USERS This reproduction was made from a copy of a manuscript sent to us for publication and microfilming. While the most advanced technology has been used to pho tograph and reproduce this manuscript, the quality of the reproduction is heavily dependent upon the quality of the material submitted. Pages in any manuscript may have indistinct print. In all cases the best available copy has been filmed. The following explanation of techniques is provided to help clarify notations which may appear on this reproduction. 1. Manuscripts may not always be complete. When it is not possible to obtain missing pages, a note appears to indicate this. 2. When copyrighted materials are removed from the manuscript, a note ap pears to indicate this. 3. Oversize materials (maps, drawings, and charts) are photographed by sec tioning the original, beginning at the upper left hand corner and continu ing from left to right in equal sections with small overlaps. Each oversize page is also filmed as one exposure and is available, for an additional charge, as a standard 35mm slide or in black and white paper format. * 4. Most photographs reproduce acceptably on positive microfilm or micro fiche but lack clarity on xerographic copies made from the microfilm. For an additional charge, all photographs are available in black and white standard 35mm slide format. * ♦For more information about black and white slides or enlarged paper reproductions, please contact the Dissertations Customer Services Department. TJhiversity Mcrafflms International 8603064 Thanthianga, Clement BIOLOGY OF CALLOSOBRUCHUS MACULATUS The Ohio State University Ph.D. 1985 University Microfilms International300 N. Zeeb Road, Ann Arbor, Ml 48106 BIOLOGY OF CALLOSOBRUCHUS MACULATUS DISSERTATION Presented in Partial Fulfillment of the Requirement for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Clement Thanthianga The Ohio State University 1985 Dissertation Committee Approved by: Dr. Rodger Mitchell Dr. Roy A. Stein Dr. Valayamghat Rhagavan Dr. Thomas E. Hetherington AAdvis’or Department of Zoology ACKNOWLEDGEMENTS I would like to thank my adviser, Dr. R. Mitchell, for his guidance and understanding during my study inspite of my many shortcomings. He gives me every opportunity to make me feel at home not only in this University but in the United States of America. He is not my adviser alone but also my benefactor during my years at the Ohio State University. I extend my deep appreciation to my Dissertation Committee members for their understanding throughout this work. I wish to thank Dr. Roy A. Stein, who first welcomed me as a student in Zoology Department, for his advice and valuable suggestions in this reseach. I also extend my acknowledgement to Drs. V. Raghavan and T. E. Hetherington, who read through the draft copies of the dissertation, giving me constructive solutions to many problems. Lastly, I extend my acknowledgement to Dr. Roy A. Tassava whose set an example as a teacher and to many good friends whose names I will not forget. ii VITA January 1, 1954 Born - Sialsuk Village, Mizoram, India. 1970-1974 BSc(Hons), North-Eastern Hills University, Shillong, India. 1974-1976 MSc,, North-Eastern Hills University, Shillong, India 1 1976-1982 Lecturer, Pachhunga Univ. College, Alzawl, Mizoram, India. iii TABLE OF CONTENTS PAGE ACKNOWLEDEMENTS ..................................... ii VITA ................................................ iii LIST OF TABLES ........... v LIST OF FIGURES ..................................... viii INTRODUCTION ........................................ 1 LITERATURE REVIEW ................................... 3 Biology of C_. maculatus ............................ 25 Materials and methods ........................ 25 Adult stage ................................... 27 1. Emergence and mating ................ 27 2. Intrinsic determinants of fecundity .. 33 3. Oviposition behavior ................ 55 4. A model for host selection .......... 71 Develpment of passive stages ................. 76 1. Development of single larvae in Berkin 76 2. Effects of food on development ...... 80 3. Experimental analysis of competition . 96 The consequences of oviposition decisions .... Ill SUMMARY ............................................. 115 LITERATURE CITED .................................... 118 iv LIST OF TABLES TABLE PAGE 1. Influence of mating regimen on fecundity females (n=10) given 50 beans for oviposition.................................. 35 2. The effect of length of development from egg to adult and competition within the bean on fecundity. The means for each set were tested (t-test) against the early emerging females.................................... 37 3. Effect of feeding on the fecundity of pairs placed on 50 berkin beans.................. 39 4. Pattern of oviposition based on counts from five females taken at each time interval from a cohort of isolated pairs on 50 beans. Negative values can be obtained because the eggs deposited during an interval were estimated by subtracting the average at the start of the period from average at the end of the period.......................... 43 5. The average (s.d.) fecundity for females (n=10) given different number of beans for oviposition. ............................. 51 6. The eggs deposited by females given 0, 1, and 10 beans and then tansferred to a dish with 50 beans for the rest of her life. The sum is the total eggs laid before and after transfer. Averages (n=10) were compared (t-test) to controls on 50 beans (mean 77.4, s.d. 11.1, n=5)............... 52 v 7. The density of eggs per bean at which the females first begin to add a second egg to beans. The transferred females were placed on 50 beans after being inhibited by having too few oviposition sites (see Fig 8). There were no differences between the means (t-tests).................................. 59 8. Outcomes of experiments to determine the discrimination of two sizes of Berkin beans. There are 10 replicates for each experiment. 68 9. Selection of oviposition sites by beetles given choices between 15 rough (chickpea) and 15 smooth (soybean) beans for 24 h. There were no differences (t-test) between the means for 10 replicates................ 69 10. Selection of oviposition sites by beetles (n=10) given choices between Berkin beans and the alternate hosts for 30 h ...... 70 11. Development in days and weight (mg) of C_. maculatus feeding alone in beans. Larvae were weighed (n=10) at the end of the molt and pupae were weighed when first formed... 79 12. Composition of the major hosts of C_. maculatus from Duke (1981). Components that show major differences are given... 83 13. Life tables for C_. maculatus on its principle hosts. With Berkin as controls, confidence limit for percentages were used to test for differences in survival and t-tests for differences in fecundities. High egg mortality was due to high temperatures, up to 30 C, shortly after oviposition... 84 14. Frequencies of various interactions between competing larvae when larvae first come into contact............................... 93 15. Average (s.d.) weight (mg) of 20 larvae alone and 10 pairs of competing larvae of C_. maculatus segregated by the position of their burrow............................... 94 vi 16. Average (s.d.) weight (mg) of larvae growing singly and in pairs in chickpea and pigeon pea (n=20)................................. 95 17. Weights of larvae in beans glued together to simulate conditions of competition within. a single bean. Controls were beans with 1 larva ground and the surface glued...... -98 18. Experiments to alter the transmission of cues between two larvae of C_. maculatus from bean to bean. The experiments were set up while the eggs were fresh and the larvae weighed when they were estimated to be 14 days old............................. 108 19. Weights (s.d.) of larvae in beans glued together after larvae had started to feed alone. Weights were taken 12 days after hatching for 20 larvae in beans glued in the first instar and at 13 days for larvae joined in the second instar.............. 109 20. Tests to determine the reversibility of the competitive behavior...................... 110 vii \ LIST OF FIGURES FIGURE PAGE 1. Basic life cycle of C_. maculatus with the active non-feeding stage (adult) separated from the passive and feeding stages (egg, larva, and pupa) by the double line. As shown in Fig. 3, passive stages must live in the single bean selected by their mother...................................... 5 2. Female C_. maculatus ovipositing on a bean (X 20)...................................... 8 3. Development of C_. maculatus in a bean of the Berkin variety. Larvae are shown at the end of the first (A), second (B) and third (C) larval instars and the pupal stage (D) to show the burrowing pattern of a larva alone in a bean (X 20)..................... 19 4. The basic life cycle (Fig. 1) expanded to indicate the interactions proposed or inferred from published reports........... 23 5. The first three days of emergences from an even aged cohort of 500 C_. maculatus. Black horizontal bars indicate dark periods..... 29 6. The daily pattern of emergences (points) and the frequency of mating pairs (bars) found at each 2-h sampling period. The records for the first three days are pooled. Black horizontal dars indicate dark periods..... 32 7. Diel pattern of oviposition by C_. maculatus based on 10 replicates taken every 12 h from a cohort of isolated pairs................. 42 viii 8. Eggs