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The Pennsylvania State University The Graduate School School of Ecosystem and Science Management ANALYSIS OF HISTORICAL AND CONTEMPORARY CONSUMPTION OF AQUATIC MACROINVERTEBRATES IN DARTER SPECIES OF A HIGHLY DIVERSE STREAM A Dissertation in Wildlife and Fisheries Science by Casey Renee Bradshaw 2015 Casey R. Bradshaw Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy December 2015 The dissertation of Casey Bradshaw was reviewed and approved* by the following: Jay R. Stauffer, Jr. Distinguished Professor of Ichthyology Dissertation Advisor Chair of Committee Elizabeth W. Boyer Associate Professor of Water Resources Michael A. Rutter Associate Professor of Statistics Jeanette L. Schnars Director, Regional Science Consortium and Research Director of PA Sea Grant Gregory Hoover Special Signatory, Department of Entomology Michael G, Messina Head and Professor of the Department of Ecosystem Science and Management *Signatures are on file in the Graduate School iii ABSTRACT Darters are small, benthic, insectivorous fish that comprise majority portion of vertebrate communities within riffles of North American streams. The objectives of this study were first, to determine diet overlap among sympatric darter species at a site with high darter diversity (French Creek at Venango) and a site with low diversity of darters (Woodcock Creek). The hypothesis for this objective is that darters living sympatrically in French Creek at Venango will partition their diet due to increased competition and therefore become more specialized in their food choices. The second objective was to compare diet of both common species of darters and currently listed species from historic collections in French Creek to contemporary collections and determine if diet has changed over time. The hypothesis is that diet will be similar from both collections given the French Creek watershed has remained relatively unaltered throughout history. I approached this objective by including currently listed species which were collected in the 1980's and early 1990's; before they were deemed either threatened or endangered. Those specimens were able to be dissected and food items analyzed. By comparing diets among common species from historic and present collections inferences were made about the diet of threatened and endangered species of darter in French Creek currently. Summary of major findings include dietary trends for historic to current darter collections have remained similar, indicating species under current conservation protection have a diet that has also remained relatively unchanged. There were obvious seasonal trends in dietary components for darters and differences in diet between Woodcock Creek and French Creek. Lastly, darters found in French Creek, especially in late summer, had a broader diet than those from earlier seasons and from Woodcock Creek, which indicates food is probably not a limited resource and therefore both intraspecific and interspecific competition is low. French Creek remains one of the few streams in the United States that holds such a high abundance and diversity of fish species, including iv many under conservation protection. It remains an important refuge for species in the upper Allegheny River, both terrestrial and aquatic. With undeniable threats to the biodiversity in French Creek, including invasive species (i.e. round gobies), global climate change, riparian zone alterations and non-point source pollution, it is important to investigate aquatic community structures and ensure preservation of one of North America's last aquatic treasures. v TABLE OF CONTENTS List of Figures .......................................................................................................................... vi List of Tables ........................................................................................................................... viii Acknowledgements .................................................................................................................. x Chapter 1 Introduction To Darters ........................................................................................... 1 Darter Communities in French Creek .............................................................................. 5 Chapter 2 Methods ................................................................................................................... 8 Site Selection: French Creek Watershed .......................................................................... 10 Field Collections .............................................................................................................. 16 Fish Processing ................................................................................................................ 17 Benthic Macroinvertebrate Processing ............................................................................ 18 Data Analysis ................................................................................................................... 18 Methods Used in Historic Fish Collections ..................................................................... 20 Chapter 3 Results ..................................................................................................................... 21 Abiotic Measurements from the Stream .......................................................................... 21 Benthic Macroinvertebrate Collections from the Stream ................................................. 21 Electivity by Species ........................................................................................................ 26 Etheostoma blenniodes ............................................................................................. 26 Etheostoma zonale.................................................................................................... 27 Etheostoma caeruleum ............................................................................................. 28 Etheostoma flabellare .............................................................................................. 29 Etheostoma nigrum .................................................................................................. 30 Etheostoma variatum................................................................................................ 30 Percina macrocephala ............................................................................................. 31 Percina caprodes...................................................................................................... 31 Percina maculata, Etheostoma camurum and Etheostoma tippecanoe ................... 32 Diet Comparison Using Principle Component Analysis and ANOVA ............................ 38 Chapter 4 Discussion ............................................................................................................... 51 Summary of Major Findings ............................................................................................ 67 Literature Cited ........................................................................................................................ 68 vi LIST OF FIGURES Figure 1-1. Photograph of French Creek in Venango, PA (N 41○46.366'/W 80○ 06. 397') looking downstream. Historically this location has housed up to 14 species of darters within a 200meter stretch of stream……………………………………….. 9 Figure 2-2. Photograph of Woodcock Creek in Saegertown, PA (N 41○ 41.859'/W 80○06.401') looking upstream. This location was used for the low species richness site, normally housing four to six species of darters within a 200 meter stretch of stream ............................................................................................................................ .. 9 Figure 2-3: Map of French Creek Watershed showing sub-watersheds and major tributaries of French Creek. © Chris Shaffer, Allegheny College GIS Manager ............. 10 Figure 2-4: Aerial view of Union City Dam on French Creek ................................................ 12 Figure 2-5: Aerial view of Woodcock Creek Lake Dam on Woodcock Creek, a tributary to French Creek ................................................................................................................ 12 Figure 2-6: Delineated map of the French Creek Watershed created by Allegheny College using ArcGIS Desktop version 10.3 from 2013 land use data. Land use percentages are as follows: Water = 1.33%, Developed Area = 7.46%; Forest = 51.50%; Grasslands = 1.33%, Agriculture = 32.95%; Wetlands 5.42% .......................... 15 Figure 3-1: PCA scores plot showing the relationship between darter species in terms of PC 1 (x axis) versus PC 2 (y axis) from Woodcock Creek in early summer (A) and French Creek in early summer (B) (May to June). Darter species were plotted and color coded, diet was broken into taxonomic level of Order ........................................... 41 Figure 3-2: PCA scores plot showing the relationship between darter species in terms of PC 1 (x axis) versus PC 2 (y axis) from Woodcock Creek in early summer (A) and French Creek in early summer (B) (July to September). Darter species were plotted and color coded, diet was broken into taxonomic level of Order..................................... 42 Figure 3-3: PCA scores plot showing the relationship between diet contents in Etheostoma blenniodes in terms of PC 1 (x axis) versus PC 2 (y axis) from both Woodcock and French Creek and early to late summer. Seasons and