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University Micrcsilms International 300 N INFORMATION TO USERS This was produced from a copy of a document sent to us for microfilming. 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 material submitted. The following explanation of techniques is provided to help you understand markings or notations 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 page(s) or section, they are spliced into the film along with adjacent pages. This may have necessitated cutting through an image and duplicating adjacent pages to assure you of complete continuity. 2. When an image on the film is obliterated with a round black mark it is an indication that the film inspector noticed either blurred copy because of movement during exposure, or duplicate copy. Unless we meant to delete copyrighted materials that should not have been filmed, you will find a good image of the page in the adjacent frame. 3. When a map, drawing or chart, etc., is part of the material being photo­ graphed the photographer has followed a definite method in “sectioning” the material. It is customary to begin filming at the upper left hand corner of a large sheet and to continue from left to right in equal sections with small overlaps. If necessary, sectioning is continued again—beginning below the first row and continuing on until complete. 4. For any illustrations that cannot be reproduced satisfactorily by xerography, photographic prints can be purchased at additional cost and tipped into your xerographic copy. Requests can be made to our Dissertations Customer Services Department. 5. Some pages in any document may have indistinct print. In all cases we have filmed the best available copy. University MicrcSilms International 300 N. ZEEB ROAD. ANN ARBOR, Ml 48106 18 BEDFORD ROW. LONDON WC 1 R 4EJ. ENGLAND 79?J lf><* I KING, PUT tP T nr*ftkD NATURAL H15TTRY A.NP FITLCGY Vh 5 T I C TUf H IkTft |iMU S ANM'LICKUS (TPWNfcS) (DlPTcPAS (. H 1 F i l N Lt M i P A T ) , A U GU 5 T A CRFlK, KALAKA7UD ri(|!M », MICHIGAN. MICHIGAN GTATt U N IV .L «■' 6 ! I Y , PH. P., 1979 University’ Microfilms International 3 0 0 N ZE E B HOAD, ANN AH BO R. M) 48106 PLEASE NOTE: In all cases this material has been filmed In the best possible way from the available copy. Problems encountered with this document have been identified here with a check mark . 1. Glossy photographs________ 2. Colored illustrations________ 3. Photographs with dark background________ 4. Illustrations are poor copy__ 5. Print shows through as there 1s text on both sides _________of page 6. Indistinct, broken or small print on several pages___________ throughout 7. Tightly bound copy with print lost 1n spine________ 8. Computer printout pages with Indistinct print________ 9. Page(s) _____ lacking when material received, and not available from school or author________ 10. Page(s) /<2/ seem to be missing in numbering only as text follows 11. Poor carbon copy________ 12. Not original copy, several pages with blurred type _____ 13. Appendix pages are poor copy________ 14. Original copy with light type ______ 15. Curling and wrinkled pages ________ 16. Other University Microfilms International 300 N. 26EB RD.. ANN ARBOR. Ml 48106 '313) 761-4700 NATURAL HISTORY AND ECOLOGY OF STICTOCHIRONOMUS ANNULICRUS (TOWNES) (DIPTERA: CHIRONOMIDAE), AUGUSTA CREEK, KALAMAZOO COUNTY, MICHIGAN By Robert Howard King A DISSERTATION Submitted to Michigan State University In partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY W. K. Kellogg Biological Station Department of Entomology 1978 NATURAL HISTORY AND ECOLOGY OF STICTOCHIRONOMUS ANNULICRUS (TOWNES) (DIPTERA: CHIRONOMIDAE), AUGUSTA CREEK, KALAMAZOO COUNTY, MICHIGAN By Robert Howard King AN ABSTRACT OF A DISSERTATION Submitted to Michigan State University In partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY W. K. Kellogg Biological Station Department of Entomology 1978 ABSTRACT NATURAL HISTORY AND ECOLOGY OF STICTOCHIRONOMUS ANNULICRUS (TOWNES) (DIPTERA: CHIRONOMlDAE), AUGUSTA CREEK, KALAMAZOO COUNTY, MICHIGAN By Robert Howard King The natural history and ecology of Stictochironomus annulicrus (Townes) was conducted in Augusta Creek, Kalamazoo County, Michigan. S. annulicrus had two generations per year with a spring emergence, beginning in mid to late April, and a summer emergence, beginning in late August. Oviposition sites were localized and used by females of succeeding generations. Linear egg masses containing 100 to 800 eggs each were laid just beneath the water's surface on vegetation. The incubation period for eggs was temperature dependent, increasing with lower temperatures, but development rate and temperature were not directly proportional. Newly hatched larvae displayed a planktonic behavior for 20 to 30 minutes before constructing a tube of detritus. Larvae remain tubiculous during all instars and obtain food by collecting fine particulate detritus and associated constituents at the end of the larval tube. Instars were easily distinguished by measuring either the width of the head at the eye spots, width of labial plate, or the distance from the lower edge of the labial plate to the occipital foramen. The ratio of increase (r) did not conform to Dyar's rule. Winter form larvae were larger than summer form larvae with this size difference also apparent in pupal and adult stages. Sexual dimorphism was also apparent in pupal and adult stages with females being larger. Larvae were found in depositional areas and had a contagious distribution. Densities varied between sites and seasonally at a given site. Density estimates at 2 Kellogg Forest site on 1 August 1974 were 161/m (n = 6) and on 10 April 1974 were 338/m^ (n = 20). At Nagel site, 2 density estimates were 2115/m (n = 9 ) on 9 July 1974 and 1325/m^ (n = 26) on 26 April 1974. Larval growth rates decreased with successive instars and each instar had increased growth rates with increases in temperature. Larval growth rates, monitored in the laboratory, varied according to levels of food quality. Highest growth rates were on ground ash (Fraxinus nigra) leaves, intermediate rates were on Tipula feces, and lowest rates were on natural stream detritus. A poor relationship existed between carbon and nitrogen values of foods and larval growth rates, while a good relationship was found for ATP, respiration, and percent ash values of foods and larval growth rates. ACKNOWLEDGEMENTS I wish to thank Dr. Kenneth W. Cummins, committee chairman, academic advisor, and friend for his encouragement, guidance, support, and parience throughout this study. I also want to thank Dr. Michael Klug, Dr. Earl Werner, and Dr. Roland Fischer for serving on the committee and for their contributions to this study as well as my education. This dissertation would not have been possible without the support of my wife, Donna, who not only encouraged me to undertake the degree program but assisted me both technically and intellectually. Many others were directly or indirectly involved with this study. Among them were my colleagues Gordon Godshalk, George Spengler, David Mahan, Bob Petersen, Fred Howard, and Karen Hogg who I want to thank for their constructive inputs to this study. I wish to express my gratitude to Dr. George Lauff, Director of the W. K. Kellogg Biological Station for his support and cooperation during this study. Mary Hughes, Deloris Haire, and Art Weist are appreciated for their administrative services and friendships. Much of the financial support of this study was from the Department of Energy through Grant No. EY-76-2002. ii TABLE OF CONTENTS PAGE I. INTRODUCTION........................................ 1 II. DESCRIPTION OF STUDY AREA......................... 5 Watershed.................................. 5 Methods (Discharge and Temperature) 8 B Avenue Site............................. 9 Nagel Site................................. 11 Spring Brook #1 .................... 12 Spring Brook #2 .................... 15 Kellogg Forest............................ 15 III. LIFE HISTORY.............................. 16 Methods (Larval Morphometric Data, Degree-Day Calculations)....... 16 Eggs and Incubation....................... 17 Larvae..................................... 21 Instar Differentiation............ 23 Pupae....................................... 27 Adults..................................... 28 Emergence and AdultActivity ....... 28 Ovaposition......................... 30 Phenology.................................. 31 IV. POPULATION ECOLOGY................................. 45 iii TABLE OF CONTENTS (cont.) PAGE IV. POPULATION ECOLOGY (cont.) Methods.................................... 45 Dispersion................................. 49 Density.................................... 51 Biomass.................................... 55 V. LARVAL GROWTH CHARACTERISTICS.................... 60 Methods and Materials.................... 60 Growth Patterns........................... 65 Food Quality.............................. 78 Carbon - Nitrogen.................. 80 Percent Ash Content................ 84 ATP and Respiration................ 91 VI. SUMMARY AND CONCLUSIONS........................... 94 VII. LIST OF REFERENCES................................. 98 VIII. APPENDICES Appendix A (Discharge Data).............. 108 Appendix B (Temperature Data)............115 Appendix C (Larval Dispersion
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