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Diptera:Tipulidae) Ai' ABSTRACT OF THE THESIS OF Thomas Leavitt Dudley for the degree ofMaster of Science 28 September 1982 in Entomology presented on Population and Production Ecology of Lipsothrix spp. Title: (Diptera: Tipulidae) Abstract approved: Redacted for Privacy /Dr. N.H. Ariderson Craneflies of the genus Lipsothrix inhabit decomposing wood in streams. The life cycles and ecological relationships of L. nigrilinea (Doane) and L. fenderi Alexander were investigated to determine how they exploit and respond to the wood habitat and their role in degradationof woody debris in western Oregon. Surveys of the western states and provinces provided data on geographical distribution of the four western species, and a key is given to the five species known from Americanorth of Mexico. The non-adult stages are spent within single logs, primarilyred alder (Alnus rubra), in headwater streams where disturbanceby abrasion is minimized. The two species are sympatric in soft logs in constant contact with water. L. fenderi larvae are found in a wider variety of wood types, including harder wood, other species of wood, andin marginal habitats in which they are more susceptible to desiccation and interactions with the semi-terrestrial community. Habitat selection may relate to the specificity of ovipositing females. Biological and behavioral characteristics of the life stages are described. Both species have a basic biennial life cycle with an ephemeral (1-8 days) adult stage. This long cycle may be possible because because the habitat is relatively predictable and constant. L. fenderi is smaller than L. nigrilinea, and some individuals emerge as annuals, probably in response to an indirect thermal cue in thefall. L. nigrilinea uses combined cues of temperature and descendingwater level, which more directly preclict suitable conditions for emergencein spring and summer. Some L. nigrilinea take three years or longerif they do not receive the water level cue in a portionof the log. A large degree of variability exists in size of bothspecies, and especially L. nigrilinea, within a single log. Though growth rate differences are correlated with temperature and foodquality, variability in timing of oviposition probably accountsfor the majority of the difference. The life history traits of L. fenderi are more conducive to increasing reproductive quantity whencompared to L. nigrilinea, which may be related to the associationof L. fenderi with relatively less predictable habitats. The mean biomass of L. nigrilinea was near 12.0mg/100 cm2, with low variation, while L. fenderi had low biomass in thefall (1.40 mg/100 cm2) and peaked prior to emergence (12.65 mg/100cm2). Wood hardness was the major determinant oflarval densities and feeding rates of both species. Egestion rates were estimated at 88% of dry weight perday in firm wood and 223% per day in highly decayed material. Egestion is increased by a factor of two between low (3°C) andhigh (15°C) temperatures encountered in the field. The total ecological impact will be the greatest where the highest abundance ofthe most suitable habitat is found, as in small streams during themiddle stages of forest succession. Population and Production Ecology of Lipsothrix spp. (Diptera:Tipulidae) by Thomas Leavitt Dudley A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Master of Science September, 1982 Commencement June, 1983 APPROVED: Redacted for Privacy Professor of Entomology in charge of major Redacted for Privacy Head of Department of Entomology Redacted for Privacy Dean of Gradua School 28 September 1982 Date thesis is presented Thomas L. Dudley Typed by Suzi Sargent for ACKNOWLEDGEMENT Many people have given me support and advice during the occasionally difficult course of this project. I particularly appreciate the patience and assistance of my advisor, Norm Anderson, and the encouragement of Ken Cummins. Gordon Pritchard and George Byers gave me sage advice from afar. The members of my committee, Peter McEvoy, Bruce Menge and George Beaudreau, have been helpful, as have been countless other mentors and providers of factsand semi-facts. I am grateful to several people for their assistance, including Ed Grafius, Tom Seibert, Mary Jo Wevers, Chuck Hawkins, Peggy Wilzbach and Diane Belnavis. Carla D'Antonio has helped in innumerable ways. I suppose that if I had it to do over again,I would do a few things differently, yet as with most things in life, one learns more from experience and from friends than canbe measured by the end product. This work was supported in part by National Science Foundation Grant No. DEB 78-10-594. TABLE OF CONTENTS Page INTRODUCTION 1 BACKGROUND 3 Wood in Streams: Role and Utilization 3 Biology of Tipulidae 6 MATERIALS AND METHODS 9 Field Analysis of Habitats, Habits and Life Cycles 9 Experimental Procedures for Laboratory Rearing 11 Natural Substrates 11 Artificial Media 12 Obtaining and Handling Experimental Animals 12 Laboratory Studies of Growth and Feeding 13 Growth and Development Patterns 13 Feeding and Processing Rates 14 Field Studies 15 Colonization and Growth in Non-Colonized Wood 15 Processing Rates 15 Studies of Adults 16 RESULTS AND INTERPRETATION 18 Characterization of Organisms and Habitats 18 Keys to the Immature Stages of the North American Species of the genus Lipsothrix 18 Habitat Characteristics of Lipsothrix nigrilinea 19 Geographic Distribution 19 Local Distribution 20 Relationship with Forest Successional Stages 21 Habitat Characteristics of Lipsothrix fenderi. 22 Geographic Distribution 22 Local Distribution 23 Relationship with Forest Successional Stages 23 Biology and Behavior of Lipsothrix spp 24 Eggs 24 Larval Morphology 24 Location Within the Wood Habitat 25 Larval Gallery - Locomotion and Orientation 27 Feeding 27 Larval Defensive or Escape Behavior 28 Pupal Preparation 29 Pupae 30 Adult Emergence 30 Adult Biology 31 Mating Behavior 32 Oviposition Behavior and Habitat Suitability 33 TABLE OF CONTENTS (continued) Page Reproductive Allocation and Parental Care 34 Environmental Modification of Growth Rates 36 Thermal Effects on Growth 36 Field Evidence 36 Temperature Studies in the Laboratory 38 Food Quality and Growth. 39 The Role of Food Quality 39 Laboratory Studies of Food Quality 40 Food Choice by Larvae 41 Life Cycle Analysis 42 L. nigrilinea 43 Variability - A Statement of the Problem 43 Model for the Life Cycle of L. nigrilinea 43 Evidence for a Biennial Life Cycle 44 Emergence Pattern and Pupation Cues 45 Environmentally Induced Extension of Life Cycles 48 Regional and Sex-Related Variability in Growth and Emergence 50 L. fenderi 51 Model for the Life Cycle of L. fenderi 51 Length of Life Cycle 52 Emergence Pattern 52 Factors Resulting in Variability 53 Size and Reproduction 54 Population Size and Dynamics 56 Larval Density 56 Mortality Factors 57 Abiotic Mortality 58 Biological Mortality 60 Adult Mortality 61 Biomass and Production 62 Impact of Lipsothrix on Wood Degradation 63 Laboratory Studies 64 Feeding Rates in Artificial Media 64 Processing Rates from Natural Substrates in the Laboratory 64 Field Studies of Processing 66 Ecosystem Role in Relation to Physical Processing 67 DISCUSSION 70 Overview of Life Cycles: Cohort Asynchrony within and between Years 70 The Ecosystem Role of Lipsothrix spp 72 Particle Reduction of Woody Debris 72 Secondary Production 74 Evolutionary Consequences of Xylophagy 74 TABLE OF CONTENTS (continued) Page Woody Debris as a Habitat 74 Evolutionary Advantage of Life Cycle Extension 75 Comparative Life History Adaptations in Relation to Habitat 76 Lipsothrix Biology and the Implications for Stream Community Ecology 80 LITERATURE CITED 82 APPENDICES 149 LIST OF FIGURES Figure Page 1. Map of transect sites 93 2. Distributions of western species of Lipsothrix 94 3. County distributions of L. nigrilinea and L. fenderi 95 4. General morphology of L. nigrilinea larva 96 5. Male and female pupae of L. nigrilinea 96 6. General morphology of L. fenderi larva 97 7. Male and female pupae of L. fenderi 97 8. Key characters of larvae and pupae of L. hynesiana, L. shasta and L. sylvia 98 9. Typical stream inhabited by Lipsothrix spp 99 10. Pathways of wood degradation with respect to microhabitat 100 11. Colonization of uninhabited experimental logs by L. nigrilinea and L. fenderi in various stream types 101 12. Diagrammatic representation of zonation of L. nigrilinea, L. fenderi and Austrolimnophila badia 102 13. Larva of L. nigrilinea in gallery 103 14. Pupation turret of L. nigrilinea 103 15. Adults of L. fenderi and L. nigrilinea 104 16. Effect of canopy cover, as a temperature index, on growth of L. nigrilinea larvae in the field for six months 105 17. Growth of L. nigrilinea larvae at various temperatures in the laboratory 106 18. In vitro weight change of Lipsothrix spp. with respect to temperature 107 19. In vitro weight change of L. nigrilinea with respect to food quality 108 20. Distribution of each size class of L. nigrilinea over one year 109 Figure Page 21. Date of collection and weight of L. nigrilinea pupae 110 22. Date of collection and weight of L. nijrilinea 111 23. Emergence pattern of L. nigrilinea 112 24. Induction of pupation of L. nigrilinea in response to water level 113 25. Size class distribution of L. fenderi larvae over time 114 26. Date of collection and weight of L. fenderi pupae 115 27. Date of collection and weight of L. fenderi larvae from Dinner Creek 116 28. Emergence pattern of L. fenderi 117 29. L.nigrilinea female adult dry weight vs. percent allocation to reproductive tissues 118 30. Densities of L. fenderi and L. nigrilinea 119 31. Larval densities of L. nigrilinea and L. fenderi from transect collections over time 120 32. Biomass of L. fenderi and L. nigrilinea from transect collections over time 121 33. Processing in vitro by L. nigrilinea in response to temperature and food quality 122 34. Processing of logs in the laboratory with respect to temperature and wood hardness 123 35.
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