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Dissertation the Distribution of Lotic Insect Traits in Relation to Reference Conditions and Projected Climate Change in the We DISSERTATION THE DISTRIBUTION OF LOTIC INSECT TRAITS IN RELATION TO REFERENCE CONDITIONS AND PROJECTED CLIMATE CHANGE IN THE WESTERN UNITED STATES Submitted by Matthew Ivern Pyne Graduate Degree Program in Ecology In partial fulfillment of the requirements For the Degree of Doctor of Philosophy Colorado State University Fort Collins, Colorado Summer 2014 Doctoral Committee: Advisor: N. LeRoy Poff Brian P. Bledsoe Jennifer A. Hoeting Colleen T. Webb Copyright by Matthew Ivern Pyne 2014 All Rights Reserved ABSTRACT THE DISTRIBUTION OF LOTIC INSECT TRAITS IN RELATION TO REFERENCE CONDITIONS AND PROJECTED CLIMATE CHANGE IN THE WESTERN UNITED STATES The use of species traits (e.g., life history, morphological, physiological, or ecological characteristics of an organism) to describe community responses to environmental change has become a common practice in stream ecosystems, with over 900 papers describing macroinvertebrate trait-environment relationships in streams. The use of traits provides some advantages over traditional taxonomic metrics, such as providing a mechanistic link between an organism and its environment, but also presents some challenges, such as many traits being correlated with other traits and multiple environmental variables. Various methods have been recommended to address these challenges, such as using multiple traits, posing a priori hypotheses, and evaluating streams across large-spatial scales. The vast majority of studies have not incorporated these recommendations, however, particularly in North America. My research had two general objectives: 1) describe the dominant trait-environmental relationships in natural streams in the western United States and 2) use two distinct traits-based methods to evaluate how stream aquatic insect communities are currently distributed in terms of multiple environmental variables and how species and communities may respond to climate change. Traits are often used to evaluate the ecological integrity of streams and a baseline understanding of aquatic insect trait-environment relationships is needed for the western United States. I used logistic regression, multinomial regression, and redundancy analysis to explore the ii relationships between 20 trait distributions and 83 environmental variables in 253 least-disturbed streams across 12 western states. Traits had the strongest relationships with regional climate and local stream habitat conditions (e.g., air temperature, conductivity, mean annual runoff) rather than elevation, land use, or measures of extreme hydrological events. Traits such as thermal tolerance, size, swimming strength, rheophily, voltinism, and armoring exhibited strong relationships with the environmental data and would be ideal for large-scale stream assessments. Aquatic insect communities contain many taxa that are sensitive to temperature increases and changes to runoff. Two traits, cold water preference and erosional obligate (i.e., needs to live in fast-water habitat) have been used in the past to estimate the effect of climate change on stream insect communities, but no study has accounted for both climatic and non-climatic effects on these two traits. I developed a Bayesian path analysis describing how the distributions of these two traits respond to multiple environmental gradients, not just temperature, and discovered that the distribution of cold-adapted taxa was strongly correlated with changes in air temperature in the wet, cool ecoregions, but was correlated with thermal buffers and refuges in most dry, warm ecoregions, indicating that temperature-sensitive taxa are likely on the brink of their thermal tolerance in those ecoregions. A second approach to assess community sensitivity to climate change is to determine the specific thermal tolerance of each taxon individually. I computed the thermal and stream runoff thresholds of common stream taxa and compared the World Climate Research Programme’s climate model predictions to these thresholds. I found that the stream communities most at risk to climate change were found in some dry ecoregions, concurring with the previous results, and in wet, warm ecoregions with a high proportion of spatially restricted and endemic taxa, such as northern California. These two approaches describe possible mechanisms of climate change resistance and identify sensitive ecoregions. iii ACKNOWLEDGEMENTS The research in my dissertation was made possible through the Edward and Phyllis Reed Fellowship from the Department of Biology, Colorado State University. A. T. Herlihy provided the WEMAP data, Christopher Cuhaciyan and Dan Baker developed of catchment and valley variables, and Daren Carlisle developed of the hydrology variables used in this document. I thank them for their contributions. I thank the modeling groups, Program for Climate Model Diagnosis and Intercomparison (PCMDI), and WCRP's Working Group on Coupled Modelling (WGCM) for the development, archiving, and organization of the climate models. I thank my committee chair and advisor, N. LeRoy Poff, for his guiding hand in my development as a scientist. Thank you for your advice, support, and collaboration. I would also like to thank my committee members, Brian Bledsoe, Jennifer Hoeting, and Colleen Webb, for their involvement in my degree, readily available counsel, and enthusiasm. Additionally, I thank Janice Moore and Paul Doherty for your friendship and genuine interest in me and my career. My labmates have provided many hours of consultation, commiseration, and cheer. Thank you, Julia McCarthy, Angie Moline, Julie Zimmerman, Dan Auerbach, Thomas Wilding, Ryan McShane, Kevin McCluney, David Martin, Rachel Harrington, Carolina Gutierrez, and Audrey Maheu, for all the great conversations and collaborations. I thank my family; you have been my anchor in this endeavor. My parents, in-laws, and siblings have provided so much emotional and temporal support I will never be able to repay. I am certain my children, Alexis, Nathan, Bryce, and Juliette, added at least 2 years to my PhD, but it has been worth every minute. Bekah, you make my life a joy every day and I couldn’t have done this without you. Finally, I thank my Heavenly Father for His love and guidance. iv TABLE OF CONTENTS ABSTRACT .................................................................................................................................... ii ACKNOWLEDGEMENTS ........................................................................................................... iv TABLE OF CONTENTS ................................................................................................................ v CHAPTER 1: THE USE OF MACROINVERTEBRATE TRAITS IN LOTIC SYSTEMS: A SHORT REVIEW OF THE HISTORY OF TRAIT RESEARCH IN STREAMS AND AN ASSESSMENT OF STUDY CHARACTERISTICS ..................................................................... 1 Summary ..................................................................................................................................... 1 Introduction ................................................................................................................................. 2 Advantages, challenges, and recommendations of trait research in streams .............................. 8 Review of Studies...................................................................................................................... 13 LITERATURE CITED ............................................................................................................. 24 CHAPTER 2: LOTIC INSECT TRAIT-ENVIRONMENT RELATIONSHIPS IN REFERENCE STREAMS ACROSS THE WESTERN UNITED STATES ............................... 35 Summary ................................................................................................................................... 35 Introduction ............................................................................................................................... 36 Methods ..................................................................................................................................... 40 Environmental variables ........................................................................................................ 42 Trait data ............................................................................................................................... 51 Statistical analyses ................................................................................................................ 54 A priori predictions and literature review ............................................................................. 57 Results ....................................................................................................................................... 66 1. Detecting response of single trait states ............................................................................ 66 2. Detecting shifts in multiple state distributions within a single trait .................................. 71 3. Detecting correlations between multiple traits and multiple environmental variables. ... 77 Discussion ................................................................................................................................. 79 Trait categorization ............................................................................................................... 87 Scales of traits-based bioassessment ..................................................................................... 88 LITERATURE CITED ............................................................................................................
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