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Xerox University Microfilms 300 North Zeeb Road Ann Arbor 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 Pagg(s)". If it was possible to obtain the missing page(s) 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. 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Silver prints of "pho,tographs" may be ordered at additional charge by writing the Order Department, giving the catalog number, title, author and specific pages you wish reproduced. 5. PLEASE NOTE: Some pages may have indistinct print. Filmed as received. Xerox University Microfilms 300 North Zeeb Road Ann Arbor. Michigan 48106 76-16,433 FUJII, Jack Koji, 1940- EFFECTS OF AN ENTOMOGENOUS NEMATODE, NEOAPLECTANA CARPOCAPSAE WEISER, ON THE FORMOSAN SUBTERRANEAN TERMITE, COPTOTERMES FOm10SANUS SHlRAKI, WITH ECOLOGICAL AND BIOLOGICAL STUDIES ON C. FORMOSANUS. University of Hawaii, Ph.D., 1975 Entomology Xerox University Microfilms, Ann Arbor, Michigan 48106 EFFECTS OF AN ENTOMOGENOUS NEMATODE, l.J"EOAPLECTANA CARPOCAPSAE WEISER, ON THE FORMOSAN SUBTERRANEAN TERMITE, COPTOTERMES FORMOSANUS SHlRAKI, WITH ECOLOGICAL AND BIOLOGICAL STUDIES ON C. FORMOSANUS 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 ENTOMOLOGY DECEMBER 1975 By Jack Koji Fujii Dissertation Committee: Minoru Tamashiro, Chairman Henry Y. Nakasone John W. Beardsley Martin Sherman Wallace C. Mitchell iii ACKNOWLEDGEMENTS I gratefully acknowledge Mr. Jerry Wakatsuki. Agriculturist of the Oahu Sugar Co., who allowed me to set-up termite traps in their cane fields; Dr. Asher Ota and Mr. Yuki Inouye of the Hawaii S\lgar Planters' Association who were instrumental in setting-up termite traps on Kauai- -Grove Farm Plantation; Mr. Po-yung Lai for his valuable assistance in portions of this research. I am also indebted to the Office of Naval Research Grant NOOOl4-67 -A-0387 -006 which supported the present study. Special thanks are due to Iny wife, Gail and my sons, Scott and Todd for their patience and understanding. iv ABSTRACT The effects of an entomogenous nematode, the DD-136 strain of Neoaplectana carpocapsae Weiser, in the Formosan subterranean termite, Coptotermes formosanus Shiraki, were studied. Gross symptomatology, histopathology and the course of infection of N. carpocapsae in g. formosanns were determined. The primary mode of nematode entry into termites anesthetized with carbon dioxide was via the anus although they were also able to enter via the mouth. Nernatodes were able to penetrate the termite alimentary tract into the hemocoel in the regions of the fore-, mid- and hindgut. The bacteria, Achromobacter neinato~ehilus Poinar and Thomas, associated with the nematode invaded the termite Inuscle, hemolymph, fat body and nervous tissues. The nematode in turn invaded the termite nervous tissue, fat body, salivary gland. muscle tissue and sternal gland. There was a significant difference in the body weight of termites from different colonies. Concentration-mortality studies of ~. carpocapsae in g. formosanus from two colonies were studied. The results indicated that weight differences between termites from different colonies had no significance in the susceptibility studies. The LC values were 2,666 and 3,472 dauerlarvae, respectively. SO LTSO values were also deterr.ained for termites exposed to various concentrations of nematodes. v Seasonal abundance of ~. formosanus workers, soldiers and alate nymphs from two field colonies were studied over a 18 month period. There was no definite seasonal pattern of the termite populations in the colonies studied. Growth stages based on pronotum width and the number of antennal segments were determined for ~. formosanus workers, soldiers and alate nymphs from two colonies. The soldier caste did not have any growth stages and represented a terminal molt. Seasonal fluctuation of the relative abundance of the worker and alate nymph growth stages revealed a definite pattern of development for each successive growth stage. Approximately 60 and 40 weeks were required for the smallest growth stage to become the largest for the workers and alate nymphs respectively. The rate of movement of the worker caste within their galleries in the field was determined by the use of stained termites. The attractiveness of Douglas fir wood of various densities to the termites in the field was studied. Termites were attracted to the lighter, less dense wood. In addition, as expected, there was a direct correlation with termite numbers and wood consumption. vi TABLE OF CONTENTS Page ACKNOWLEDGEMENTS. .•..•.....•.....•••....•..•.•....••• iii ABSTRACT. •.•••.••••••••.•••...••.•••.••••.•..•••••....•• iv LIST OF TABLES. • •• .•. •• •. ••• •. • •. • •• ••••. •• •• •. •. •. .• •• viii LIST OF ILLUSTRATIONS. ..••••..•••••••••.••••..•••••••.•• xi INTROnUCTIOI\l .•..•...........•.....•.................... 1 MATERIALS AND METHODS Collecting TerITlite s.•.•••••••.••••••••..•..•••.•.••.. 11 Preparation of the InoculuITl. ••••••••••.•••.•.•.••••••.. 16 Histopathology and Course of Infection. ••.••.••..•••..... 18 Susceptibility Studi ."l •••••••••••••••••••••••••••••••••• 20 Ecological and Field Biology Studies Seasonal Abundance. .••••••••.•••.••.•••••.•..•.• 22 TerITlite Growth Stages Workers. e ~ •••••• 'j. ••••• •• •• ••• ••• ••• ••• •• • 26 Soldiers ~ ................... 27 Alate NYIllphs .............................. 27 Seasonal Fluctuation of the Growth Stages Worlters. .................................. ~8 Alate Nymphs .. II ••••••••••••••••••••••••••• 28 TerITlite Movement Studies. •. •••• ••• •• •• •••. • •. ••. 28 RESULTS AND DISCUSSIONS G ros s Syrnptolnatology . 35 Course of Infection and Histopathology.••••.•••.•••••.••. 36 Susceptibility Studies Foraging Worker Body Weights.••••••.•••••••••.•. 60 Carbon Dioxide Exposure on Recovery TiITle .•.••••. 62 Susceptibility of Workers to NeITlatodes LC50 Studies.. tJ •••••••••••••••• 0 ••••••••••• 65 LTSO Studies II •••••••••••• 69 Ecological and Field Biological Studies Seasonal Abundance fool •••••••• 74 Foraging Workers . 76 Soldiers . 92 Alate NYlnphs 0 •••••••••••••• 100 vii TABLE OF CONTENTS (Continued) Page General Biology. .. .. .. .. .. 101 Foraging Workers. •• •• •.• •• •• ••• •• ••• .• ••• • 102 Soldiers................................... 112 Alate Nynlphs .............................. 114 Seasonal Fluctuation of the Relative Abundance of Foraging Worker Growth Stages .••••••••.•.• 120 Seasonal Fluctuation of the Relative Abundance of Alate Nymph Growth Stages. •• •• •.•••. •••••. 125 Termite Movenlent Studies. • • ••••••••••••.••...••. 134 Relationship of Wood Weight and the Number of Foraging Workers. .•••••••••••••••.•••••••• 1. 38 Relationship of Foraging Worker Numbers and Wood Consumed •••••••••••••.•••••.•.••.• 144 SUMMARY AND CONCLUSIONS ..•••. 0. •••••••••••••••••••••• 149 LITERATURE CITED. ••.•.••••••••••••••••••••.•••••.••.•.• 155 viii LIST OF TABLES Table Page I THE NUMBER OF COPTOTERMES FORMOSANUS SHIRAKI WORKERS OBSERVED IN SAGGITAL SECTIONS HARBORlNG NEMATODES AFTER VARIOUS EXPOSURE PERIODS TO THE DD-136 STRAIN OF NEOAPLECTANA CARPOCAPSAE WEISER DAUERLARVAE••••..•.••...•.. 38 II LOCATION OF NEMATODES IN TERMITES AT HOURLY INTERVALS AFTER EXPOSURE .•.•.••••.••••.•.•.•••.. 39 III ANALYSIS OF VARIANCE OF A SPLIT-PLOT EXPERIMENT ON FORAGING WORKER BODY WEIGHTS OF COPTOTERMES FORMOSANUS SHlRAKI FROM THREE CO.·;~ONIES.•••.................. 61 IV ANALYSIS OF VARIANCE OF A SPLIT-PLOT EXPERIMENT ON THE RECOVERY TIME OF COPTOTERMES FORMOSANUS SHIRAKI FORAGING WORKERS FROM THREE COLONIES EXPOSED TO CARBON DIOXIDE FOR VARIOUS PERIODS ..•••.•••..•.. 63 V MEAN RECOVERY TIME FOR COPTOTERMES FORMOSANUS SHIRAKI FORAGING WORKERS FROM THREE COLONIES EXPOSED TO FOUR CARBON DIOXIDE EXPOSURE PERIODS.•............... 63 VI PERCENTAGE MORTALITY OF UH-l AND EWA COPTOTERMES FORMOSANUS SHIRAKI WORKERS AFTER SEVEN DAYS EXPOSURE TO THE DD-136 STRAIN OF NEOAPLECTANA CARPOCAPSAE WEISER DAUERLARVAE (COMBINED VALUES FOR 3 TESTS) ..... 66 VII CUMULATIVE CORRECTED MEAN PERCENT MORTALITIES OF UH-l AND EWA COPTOTERMES FORMOSANUS SHlRAKI WORKERS EXPOSED TO
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